taxonID	type	description	language	source
03C0F319043EFFC8FEF6D524FDE4FAD6.taxon	description	urn: lsid: zoobank. org: act: 99 B 77227 - 3 F 9 B- 4 F 26 - 87 DF- 0191 AF 3 EF 857	en	Lazaridis, Georgios, Tsoukala, Evangelia, Kostopoulos, Dimitris S. (2022): Validation of a prematurely abolished new Propotamochoerus Pilgrim, 1925 species (Mammalia, Suidae) from SE Mediterranean. Comptes Rendus Palevol 21 (26): 531-549, DOI: 10.5852/cr-palevol2022v21a26, URL: https://doi.org/10.5852/cr-palevol2022v21a26
03C0F319043EFFC8FEF6D524FDE4FAD6.taxon	materials_examined	HOLOTYPE. — Cranium LGPUT KRY 3820 preserving the right P 1 - M 3 and the left P 2 - M 3. PARATYPES. — Mandible LGPUT KRY 1094 (fragment LGPUT KRY 3490 is part of this mandible) with right p 2 - m 3, c and right and left i 1 - i 2; upper canines, C: LGPUT KRY 1092, LGPUT KRY 1193; p 2: LGPUT KRY 1096. REPOSITORY. — Museum of Geology-Palaeontology-Palaeoanthropology, Aristotle University of Thessaloniki (LGPUT).	en	Lazaridis, Georgios, Tsoukala, Evangelia, Kostopoulos, Dimitris S. (2022): Validation of a prematurely abolished new Propotamochoerus Pilgrim, 1925 species (Mammalia, Suidae) from SE Mediterranean. Comptes Rendus Palevol 21 (26): 531-549, DOI: 10.5852/cr-palevol2022v21a26, URL: https://doi.org/10.5852/cr-palevol2022v21a26
03C0F319043EFFC8FEF6D524FDE4FAD6.taxon	etymology	ETYMOLOGY. — The species is named after its area of distribution around the Aegean Sea.	en	Lazaridis, Georgios, Tsoukala, Evangelia, Kostopoulos, Dimitris S. (2022): Validation of a prematurely abolished new Propotamochoerus Pilgrim, 1925 species (Mammalia, Suidae) from SE Mediterranean. Comptes Rendus Palevol 21 (26): 531-549, DOI: 10.5852/cr-palevol2022v21a26, URL: https://doi.org/10.5852/cr-palevol2022v21a26
03C0F319043EFFC8FEF6D524FDE4FAD6.taxon	materials_examined	TYPE LOCALITY. — Kryopigi, Chalkidiki Peninsula, Greece.	en	Lazaridis, Georgios, Tsoukala, Evangelia, Kostopoulos, Dimitris S. (2022): Validation of a prematurely abolished new Propotamochoerus Pilgrim, 1925 species (Mammalia, Suidae) from SE Mediterranean. Comptes Rendus Palevol 21 (26): 531-549, DOI: 10.5852/cr-palevol2022v21a26, URL: https://doi.org/10.5852/cr-palevol2022v21a26
03C0F319043EFFC8FEF6D524FDE4FAD6.taxon	distribution	GEOGRAPHIC DISTRIBUTION. — Greece: Kryopigi and Polychrono (Kassandra, Chalkidiki Peninsula), Ravin des Zouaves- 5 (Axios Valley), Samos, Thermopigi (Serres); North Macedonia: Vozarci, Kalnitsa; Bulgaria: Kalimantsi; Possible occurrences: Achladi, Maramena (Greece); Gravitelli, Casino, Baccinello V 3 (Italy); Salihpasalar, (Turkey).	en	Lazaridis, Georgios, Tsoukala, Evangelia, Kostopoulos, Dimitris S. (2022): Validation of a prematurely abolished new Propotamochoerus Pilgrim, 1925 species (Mammalia, Suidae) from SE Mediterranean. Comptes Rendus Palevol 21 (26): 531-549, DOI: 10.5852/cr-palevol2022v21a26, URL: https://doi.org/10.5852/cr-palevol2022v21a26
03C0F319043EFFC8FEF6D524FDE4FAD6.taxon	description	AGE. — Late Miocene (Turolian, MN 11 - MN 13).	en	Lazaridis, Georgios, Tsoukala, Evangelia, Kostopoulos, Dimitris S. (2022): Validation of a prematurely abolished new Propotamochoerus Pilgrim, 1925 species (Mammalia, Suidae) from SE Mediterranean. Comptes Rendus Palevol 21 (26): 531-549, DOI: 10.5852/cr-palevol2022v21a26, URL: https://doi.org/10.5852/cr-palevol2022v21a26
03C0F319043EFFC8FEF6D524FDE4FAD6.taxon	diagnosis	DIAGNOSIS (after Fortelius et al. 1996; van der Made et al. 1999; Montoya et al. 2006; Geraads et al. 2008; Chit Sein et al. 2009; van der Made 2010; and present study). — A species of Propotamochoerus with widely separated temporal lines running parallel to each other in their caudal part; zygomatic arches originated above the M 1; small I 1 lacking a distal cusp; elongated and narrow I 2 and I 3; presence of a long diastema between C-P 1 and a shorter between P 1 - P 2; moderately large M 3; relatively small male lower canines with the posterior face narrower than both the labial and the even wider and more convex lingual one; broad p 3 / P 3; short p 4 with small to absent metaconid and reduced precristid; presence of a small hexaconid on the third lobe of m 3. DIFFERENTIAL DIAGNOSIS. — Propotamochoerus aegaeus n. sp. differs from P. palaeochoerus in having wider parietals; temporal lines that do not merge; longer anteroposteriorly zygomatic; slender muzzle above the M 1; shorter diastema between P 2 and P 1; longer I 2 and I 3; longer P 2 compared to P 3 but equally broad in its anterior and posterior part; shorter p 4; broader p 3; and a distinct hexaconid in the talonid of the m 3. It differs from P. hysudricus in the slightly larger size (6 % on average in linear cranial and dental measurements); the more separated caudal parts of the temporal lines; the longer and slender muzzle above the M 1; the presence of diastemata between C-P 1 and P 1 - P 2; the longer P 2 than P 3; the larger M 3 / m 3; the smaller lower male canine; the presence of diastema between p 1 - p 2; the broader p 3 / P 3 and the presence of a hexaconid in the talonid of m 3. It differs from P. hyotherioides in the longer I 2, shorter and broader P 3 and P 4, and the broader p 3. It differs from P. provincialis in the smaller size; more elongated I 2, I 3; the presence of diastemata between C-P 1 and P 1 - P 2; the bi-rooted P 1; the longer P 2 than P 3; and the shorter p 4, m 2 and both M 3 / m 3. Propotamochoerus aegaeus n. sp. is nearly 30 % larger in linear dental measurements than P. wui, which also lacks a hexaconid on the third lobe of m 3.	en	Lazaridis, Georgios, Tsoukala, Evangelia, Kostopoulos, Dimitris S. (2022): Validation of a prematurely abolished new Propotamochoerus Pilgrim, 1925 species (Mammalia, Suidae) from SE Mediterranean. Comptes Rendus Palevol 21 (26): 531-549, DOI: 10.5852/cr-palevol2022v21a26, URL: https://doi.org/10.5852/cr-palevol2022v21a26
03C0F319043EFFC8FEF6D524FDE4FAD6.taxon	description	DESCRIPTION The cranium KRY 3820 is almost complete and belongs to an old individual based on its heavily worn molars (Figs 1; 2; Tables 1; 2). In most respects (measurements 1 - 3, 7, 12, 18, 19 in Table 1), its overall size appears intermediate between those of male crania of P. hysudricus and P. palaeochoerus. The snout is broken anterior to the canines, which are also missing. Nevertheless, the supra canine flange (also known as canine alveolar crest or canine boss) is preserved in the right side and bears a relatively large alveolus of triangular crosssection. Its posterior edge is placed above the posterior end of the first premolar. Although slightly deformed, the cross section of the snout is almost square. However, the cheek region is slightly concave. The supra canine flange is large, rather short with rough surface and curved anteroposteriorly. The zygomatic arches are wide, and deep at their central part. They flare outwards from the facial surface at a right angle. Anteriorly, they extend to the level above the anterior part of the M 1. On both the dorsal and ventral surfaces of the prezygomatic shelf there are wide but shallow fossae. The orbits are small compared to the size of the cranium and associated with large and deep lachrymal notches. The dorsal surface of the skull is flat; slightly concave anteroposteriorly in the fronto-nasal region and weakly convex in the fronto-parietal one. The parietal lines run in parallel caudally, reaching a minimum distance of about 40 mm just in front of the nuchal crest. The supraorbital grooves are deep and extend anteriorly to a level above the P 2. The infraorbital foramen is located above the contact of P 3 / P 4. The posterior part of the skull is strongly elevated and the occiput is fan-shaped with strongly concave central part. The condyles are small compared to the size of the cranium. The U-shaped choanae open posterior to the M 3 s. The paroccipital processes are placed behind the auditory bullae and they do not fuse to them. A diastema of 10 mm exists between C and P 1, and a smaller diastema (c. 4 mm) between P 1 and P 2. The incisors are not preserved. The upper canine alveolus has a triangular shape (DAPmax = 24 mm; DTmax = 19 mm) with the posterior face longer than the lateral one; the canines themselves are not preserved but the isolated and extremely worn upper canines KRY 1092, and KRY 1193 likely belong to the cranium KRY 3820. In their preserved proximal part, they show an oval cross section and remnants of a thick cement cover. P 1 is relatively narrow, bi-rooted and slightly worn. P 3 is wider at its posterior half. Although heavily worn, the M 2 and M 3 show a small, low cusp (ectostyle) on the buccal side and between the two lobes. The talon of the M 3 is offline compared to the anteroposterior axis of the tooth and of the entire dentition, fully shifted towards the palatine and occupied by a large pentacone. The mandible is robust and rather high but partially preserved (Fig. 2; Table 2), retaining the left and right i 1 and i 2, the right canine and the right p 2 - m 3. The left mandibular corpus posterior to the symphysis is missing. The symphysis ends posteriorly at the level of p 2. The ventral part of the right mandibular corpus posterior to the m 1 is missing, as well as most of the vertical ramus and part of the mandibular condyle. The corpus is much wider (thicker) below the premolars and gradually thins towards the molars. The teeth are heavily worn, similarly to those of the cranium KRY 3820 and the perfect teeth occlusion suggests they belong to the same individual. The alveolus of p 1 is poorly preserved but it seems that p 1 is closer to c than to p 2, although accurate measurements cannot be taken. The preserved incisors are set horizontally. The lower canine strongly curves outwards in its preserved part, however keeping its dorsal surface roughly at the same level as that of the incisors. It has a sub-triangular crosssection with the posterior face slightly narrower than the labial or lingual ones (9.4 mm, 11.2 mm, and 12.7 mm respectively; measurement taken just above the alveolus). The premolars are less worn than the molars and they appear broader at their posterior half. It is highly possible that all the specimens from Kryopigi belong to the same senile individual. The large supra canine flanges, the fairly large upper canines with traces of a thick cement cover, the strongly inflated and flaring zygomatic arches and the relatively large lower canines suggest a male individual. COMPARISONS The medium size, the morphological features of the facial cranium, the morphology and placement of the zygomatic arches, the development and convergence pattern of the parietal lines, the shape of the snout, the long symphysis and the canine placement are suitable for the genus Propotamochoerus, according to the diagnosis provided by Pickford (1988). Three Asian (P. hysudricus, P. hyotherioides, and P. wui) and two European (P. palaeochoerus, and P. provincialis) species of late Miocene Propotamochoerus are currently recognized by most scholars. Unfortunately, cranial anatomy is known only for P. palaeochoerus, and P. hysudricus (Pilgrim 1926; Mottl 1966; Schmidt-Kittler 1971; Pickford 1988). The cranium of P. palaeochoerus from Johnsdorf, Austria, differs from that of the Kryropigi taxon in the larger (especially longer) supra canine flanges; the more posterior origin of the zygomatic arch (above M 2 instead of M 1 in Kryopigi); the converging temporal lines behind the postorbital processes of the frontals, tightly spaced along the parietal; the more constricted laterally parietals; the less compressed zygomatic-temporal join on the temporal fossa (dorsal view); the well concave nuchal crest (in dorsal view); the less widen rostral part of the nasals; and the more posteriorly extended and well curved supraorbital grooves (Fig. 3). P. hysudricus cranium from Siwaliks differs from the Kryopigi one in the more converging temporal lines towards the occipital; the more posterior origin of the zygomatic arch on the face (above M 2); the less compressed zygomatic-temporal joint on the temporal fossa (dorsal view); the concave profile of the nuchal crest (in dorsal view); the narrower rostral part of the nasals; the shallower depressions on prezygomatic shelf; and, possibly, in the longer rostral part of the snout (in front of the supra canine flange) (Fig. 3). Metrically, the cranium from Kryopigi appears longer than that of P. hysudricus. (measurement 1 in Fig. 4) and possibly closer to the cranial length of P. palaeochoerus. The length of the supra canine flange is larger than that of P. hysudricus, but significantly smaller in comparison with that of P. palaeochoerus (measurement 2 in Fig. 4). The partially preserved palatal length (measurement 3 in Fig. 4) is intermediately placed to the other two species, but it was probably closer to P. palaeochoerus. The length of the zygomatic arch (measurement 4 in Fig. 4), and the length of the cheek teeth (P 2 - M 3; measurement 5 in Fig. 4) surpass both P. hysudricus and P. palaeochoerus. The height of the occipital (measurement 6 in Fig. 4) is similar to that of P. hysudricus and smaller than that of P. palaeochoerus. The height (measurement 7 in Fig. 4) and width (measurement 8 in Fig. 4) of the zygomatic arches, and the orbital width (measurement 9 in Fig. 4) are close to those of P. palaeochoerus. Furthermore, the Kryopigi taxon displays a narrower snout above the M 2 and likely larger at P 3 (measurement 10 and 11 in Fig. 4, respectively); although this may be exaggerated due to distortion. The breadth between the supra canine flanges is intermediate to the compared species. The palatal breadth (measurement 13 in Fig. 4) is about 27 % larger than in P. hysudricus and similar to that of P. palaeochoerus. The length from the orbit to the hindmost part of the skull (measurement 14 in Fig. 4) and to the foremost part of the skull (measurement 15 in Fig. 4; partially preserved) exceeds the dimensions of P. hysudricus. Although the foremost part of the skull in the studied cranium is missing, it can be estimated to be 3 % to 15 % longer than that of the standard. Premolar and molar series length (measurement 16 and 17 in Fig. 4) are 17 % and 4 % larger than in P. hysudricus, respectively. Taking into account the restricted evidence available (e. g. Pearson 1928; Mottl 1966; Schmidt-Kittler 1971; Pickford 1988, 2013; van der Made & Han 1994; van der Made et al. 1999), it appears that the Kryopigi taxon also differs from P. hysudricus in the presence of diastemata between C-P 1 and between P 1 - P 2. In the P. palaeochoerus upper dentition, the C-P 1 diastema is slightly shorter or equal to that between P 1 - P 2, which is different from the Kryopigi, Samos (Thenius 1950), Maramena (Hellmund 1995) and Vozarci (Geraads et al. 2008) Propotamochoerus, all exhibiting a very short to absent P 1 - P 2 diastema but a significant C-P 1 one. As in P. provincialis from its type locality (Pickford 2013: fig. 8), Propotamochoerus lower dentition from Samos (Thenius 1950), Ravin de Zouaves 5 (de Bonis & Bouvrain 1996), Vozarci (Geraads et al. 2008), and likely Kryopigi show a long p 1 - p 2 diastema, compared to the c-p 1 one. No data are known to us for P. hyotherioides or P. palaeochoerus, whereas limited data for P. hysudricus and P. wui (Pickford 1988; van der Made & Han 1994) rather indicate a shorter c-p 1 diastema. In any case, diastema length appears to be highly variable in recent Suidae, depending on both ontogenetic age and sex (e. g. van der Made 1991) and hence, the discriminant value of the observed features may be limited. Van der Made et al. (1999: fig. 4; see also Montoya et al. 2006: fig. 6; van der Made 2010: figs 16; 18) have already shown that the Samos, and Maramena Propotamochoerus are characterized by a small I 1 without distal cusp and elongated and narrow I 2 and I 3, features that altogether differentiate the Balkan taxon form other species, apart the significantly smaller P. wui. According to data provided by van der Made et al. (1999: fig. 1) the lower male canine of the Kryopigi taxon, as well as that from Ravin de Zouaves 5, are significantly smaller than the canine of P. hyotherioides, smaller on the average than the single known specimen of P. hysudricus and close to the minimum values obtained for P. palaeochoerus. In both the Kyropigi and Ravin des Zouaves 5 Propotamochoerus (Fig. 2 C-D), the lingual side of the lower male canine is slightly wider and more convex than the labial one and both are slightly wider than the posterior side. A metrical comparison of the upper third molar (Fig. 5) shows that the Kryopigi taxon, along with those from Vozarci (Northern Macedonia) and Kalimantsi (Bulgaria) form a rather distinct cluster between the relatively smaller P. hysudricus and P. palaeochoerus and the larger P. hyotherioides and P. provincialis from Montpellier. A M 3 from Casino Basin, and two M 3 from Baccinello V 3 (Italy) are plotted with the larger specimens of Propotamochoerus from the Balkans and well below the range of P. provincialis. Two M 3 from Maramena locality are plotted close to the largest specimens of P. aegaeus n. sp. but they appear relatively narrower. The hypothesis of same multivariate mean (based on the length and maximum width) of the M 3 of Propotamochoerus from the Balkans and the rest of the other species in comparison (with more than two observations) is rejected in all cases apart from the Maramena suid and marginally from P. hyotherioides (Table 3). No significant morphological differences in M 3 have been observed among different species and samples. A PCA byIannucci et al. (2020: figs 2; 3) based on the length and width of P 3, P 4 and M 3 is interpreted as indicating close proximity between the Balkan group and P. provincialis. Nevertheless, data on P. provincialis from its type locality are missing from their analysis (partly because P 3 and P 4 are not recorded there; see Pickford 2013). A repetition of the PCA (Fig. 6) using the same matrix but with the inclusion of the two M 3 attributed to this species by Pickford (2013) from the “ sables marins de Montpellier ” provides exactly the same results as in Iannucci et al. (2020: fig. 2). The first component explains more than 90 % of the variance, affected positively mostly by the M 3 dimensions (Appendices 1; 2). P. provincialis from Montpellier is well distinct from the SE Mediterranean cluster, just because of its significantly larger M 3 (compare also with Fig. 5). The Casino specimen groups with the Balkan taxon whereas the Venta del Moro, Spain specimen keeps an intermediate position between the Balkan taxon and P. provincialis from Montpellier (Fig. 6). By removing the absolute size effect (Appendix 16), it appears that only P. palaeochoerus is clearly separated from other Propotamochoerus species due to its proportionally longer upper premolars, whereas both the Balkan taxon and P. provincialis overlap significantly in the morphospace with the smaller Sus arvernensis Croizet & Jobert, 1828. A PCA with the length and width of p 3, p 4, m 2, and m 3 in a number of Propotamochoerus species (Fig. 7) shows that the new material from Kryopigi is plotted together with that from other Greek, Bulgarian and North Macedonian localities in a distinct space between P. hysudricus and P. provincialis. On this diagram, the first principal component displays a great variance (Appendix 3) and is positively affected by all the variables, especially the length and width of m 2, and m 3 (Appendix 4). Thus, species with larger molars are plotted to the right side of the diagram. The PC 2 axis is positively affected mostly by the length of p 4 and negatively by the width of p 3, m 2, and m 3 (Appendix 4). The Balkan taxon is overall larger than P. wui (see also Fig. 8) and well distinguished from P. palaeochoerus by its shorter p 4 and slightly wider p 3, m 2, and m 3 (see also Fig. 8). The mean of P. hyotherioides indicates a rather similar size to the Balkan taxon, both distinguished from P. provincialis by their shorter molars (see also Fig. 8). As for the upper toothrow, transformed data show (Appendix 17) that only P. palaeochoerus can be clearly distinguished in the dental morphospace mostly by its p 4 proportions; the Balkan taxon greatly overlaps with both P. provincialis, and P. hysudricus. Scatter plots (Fig. 8) of individual teeth proportions (length against width) provide some additional information. The Balkan group of samples forms a cluster that overlaps partially with P. hysudricus and less with P. palaeochoerus (especially in p 4 and m 1). Propotamochoerus hyotherioides is not so well represented in these diagrams. However, it displays some differences such as narrower premolars plotted outside of the 95 % confidence interval of the Balkan taxon; in molars the difference is not so sharp. Propotamochoerus provincialis from Montpellier forms a well-defined cluster that occupies mainly the area of the largest dimensions in the diagrams (especially for m 2), overlapping significantly with P. palaeochoerus in the p 4 and m 1 size. Apart from the p 3, the Balkan taxon discriminates in all other examined teeth from P. provincialis. In the p 4 and especially the m 2, the separation of the two groups is clear as most P. provincialis teeth are plotted outside the 95 % ellipses of the Balkan taxon. Specimens from the Casino Basin are plotted close to both the Balkan taxon and P. provincialis, except for the p 4 which is grouped with the former species. Specimens from Maramena are plotted inside the confidence interval of the rest Balkan samples for m 1 and m 3 but outside for the p 3 and m 2. The probability of same multivariate variance in the lower teeth (p 4, m 1, m 2, m 3) between the Balkan taxon and all the compared Propotamochoerus species is summarized in Table 4 (pairwise comparisons per tooth among all species are provided as supplementary information, Appendices 9 - 12. A statistically significant difference for all compared teeth is found between the Balkan taxon and Propotamochoerus provincialis, as well as between the Balkan taxon and P. wui. With the rest of the species, the null hypothesis for same multivariate variance is rejected in at least one tooth (Table 4). Even by using a single Balkan sample (in this case Vozarci, which has the most specimens), the results (Appendices 13 - 15) hold well, with one exception in the p 3 of P. provincialis (the null hypothesis cannot be rejected; see supplementary information, Appendix 13). According to the PCA loadings and original metrical data, two of the main metrical differences of the Balkan species from the rest of the compared Propotamochoerus taxa concern the width of the p 3 and the length of the p 4; this is summarized in the scatter diagram of Figure 9. In this plot, the Balkan taxon, P. wui, P. provincialis, P. hysudricus and P. palaeochoerus are well separated, whereas the mean of P. hyotherioides is plotted between the last two taxa indicating some overlap among those species. P. provincialis from the type locality shows a longer p 4 compared to the Balkan taxon. Iannucci et al. (2020) also recognize a size difference in the p 4 length between the Balkan taxon and the west European P. provincialis but they interpret it as trivial and possibly related to ecological adaptations. In fact, the average length difference of 1.5 mm represents about 11 % of size difference between the two samples. Although intraspecific dental morphological variability cannot be controlled by the available data (Pickford 1988; van der Made & Han 1994; van der Made et al. 1999), the Balkan taxon seems to differ from P. palaeochoerus, P. hysudricus and P. wui in the presence of a distinct hexaconid in the third lobe of m 3. Hexaconids occur in the m 3 s of P. hyotherioides and P. provincialis (van der Made & Han 1994; Pickford 2013); compared to the latter taxon, the hexaconid of m 3 in the Ravin des Zouaves 5, Polychrono (Fig. 2 D-E), Maramena, and Vozarci lower dentitions is less developed and lingually restricted. The p 4 of P. hysudricus, P. palaeochoerus, P. hyotherioides, P. wui, and to a lesser degree that of P. provincialis, show a distinct metaconid (synonym of Innenhügel; Pickford 1988, 2013; see van der Made & Han 1994; van der Made et al. 1999), which is smaller to absent in the studied taxon (see also Thenius 1950; de Bonis & Bouvrain 1996; Geraads et al. 2008; and pers. obs. on the Maramena suid). Additionally, the precristid of the p 4 is reduced in the Balkan taxon as well as in P. provincialis (Pickford 2013).	en	Lazaridis, Georgios, Tsoukala, Evangelia, Kostopoulos, Dimitris S. (2022): Validation of a prematurely abolished new Propotamochoerus Pilgrim, 1925 species (Mammalia, Suidae) from SE Mediterranean. Comptes Rendus Palevol 21 (26): 531-549, DOI: 10.5852/cr-palevol2022v21a26, URL: https://doi.org/10.5852/cr-palevol2022v21a26
