FIGURE 1. Geographic and geological location of the outcrops in which remains of the Spanish podocnemidid turtle Neochelys zamorensis have been identified, all of them located in the Entrala Formation (Lutetian, middle Eocene) of the Zamora Province (Castile and Leon Autonomous Community, northwestern Spain, Duero Basin). A, Geographic location of the Duero Basin in the Iberian context. B, Geological locations of the outcrops. C, Stratigraphic position of the outcrops. The numbers in B and C represent the position of the outcrops: 1, type locality of the species, i.e., the El Escobar fossil site (Sanzoles Municipality); 2, Infiernos hill (Monfarracinos Municipality), where the Cubillos 1 and Tardemayas 1 fossil sites are located; 3, Valdegallina locality (Zamora Municipality), where the Valdegallina 1, Valdegallina 2, and Valdegallina 3 fossil sites are located; 4, Teso de la Mora fossil site (Torres del Carrizal Municipality). A and B are modified from the fig. 2 and C of fig. 3 in Ortega et al. (2022).
FIGURE 2. Shell remains of the Spanish podocnemidid turtle Neochelys zamorensis from its type locality, the Lutetian (middle Eocene, Entrala Formation) fossil site of El Escobar (Sanzoles Municipality, Zamora Province). The carapacial elements are shown in dorsal view and the plastral remains in ventral one. A-B, STUS 12121 (holotype of the species). C-D, STUS 12120. E, STUS 8797. F-G, STUS 8698. H-I, STUS 10063. J, STUS 10061. The dotted lines indicate broken edges, the continuous black lines correspond to the margins of the plates, the borders of the scutes are represented by thicker gray lines, and the gray surfaces correspond to sediment. Lowercase abbreviations correspond to the plates. The abbreviations of the scutes are in capital letters.
FIGURE 3. Relatively complete to partial shells the Spanish podocnemidid turtle Neochelys zamorensis, including both carapacial and plastral elements, from the Lutetian (middle Eocene, Entrala Formation) of the Valdegallina locality (Zamora Municipality, Zamora Province). A-B, STUS 17944. C-D, STUS 8640. E-F, STUS 14510. G-H, STUS 8795. The carapacial elements are shown in dorsal view and the plastral remains in ventral one. STUS 17944 comes from the Valdegallina 3 fossil site, and the others from the Valdegallina 1 fossil site. The dotted lines indicate broken edges, the continuous black lines correspond to the margins of the plates, the borders of the scutes are represented by thicker gray lines, and the gray surfaces correspond to sediment and non-shell osseous elements. Lowercase abbreviations correspond to the plates. The abbreviations of the scutes are in capital letters.
FIGURE 4. Partial carapaces the Spanish podocnemidid turtle Neochelys zamorensis from the Lutetian (middle Eocene, Entrala Formation) of the Valdegallina 1 fossil site (Valdegallina locality, Zamora Municipality, Zamora Province). A, STUS 9095. B, STUS 6285. C, STUS 12061. D, STUS 6287. E, STUS 12076a. F, STUS9161. G, STUS 10611. They are shown in dorsal view. The dotted lines indicate broken edges, the continuous black lines correspond to the margins of the plates, the borders of the scutes are represented by thicker gray lines, and the gray surfaces correspond to sediment. Lowercase abbreviations correspond to the plates. The abbreviations of the scutes are in capital letters.
FIGURE 5. Partial carapaces the Spanish podocnemidid turtle Neochelys zamorensis from the Lutetian (middle Eocene, Entrala Formation) of several fossil sites in the Zamora Province. A, STUS 6290. B, STUS 6289. C, STUS 6391. D, STUS 8845. E, STUS 12076b. F, STUS 10612. G, STUS 8638. H, STUS 16329. They are shown in dorsal view. STUS 6391 comes from the Tardemayas 1 fossil site (south margin of the Infiernos Hill, Monfarracinos Municipality), and the others from the Valdegallina 1 fossil site (Valdegallina locality, Zamora Municipality). The dotted lines indicate broken edges, the continuous black lines correspond to the margins of the plates, the borders of the scutes are represented by thicker gray lines, and the gray surfaces correspond to sediment. Lowercase abbreviations correspond to the plates. The abbreviations of the scutes are in capital letters.
FIGURE 6. Relatively complete to partial plastra the Spanish podocnemidid turtle Neochelys zamorensis from the Lutetian (middle Eocene, Entrala Formation) of the Valdegallina locality (Zamora Municipality, Zamora Province). A, MGM-5862N. B, STUS 6288. C, STUS 10609. D, MGM-5863N. E, STUS 6334. F, MGM-5864N. They are shown in ventral view. MGM-5862N, MGM-5863N, and MGM-5864N come from the Valdegallina 3 fossil site, and the others from the Valdegallina 1 fossil site. The dotted lines indicate broken edges, the continuous black lines correspond to the margins of the plates, the borders of the scutes are represented by thicker gray lines, and the gray surfaces correspond to sediment. Lowercase abbreviations correspond to the plates. The abbreviations of the scutes are in capital letters.
FIGURE 7. Partial plastra the Spanish podocnemidid turtle Neochelys zamorensis from the Lutetian (middle Eocene, Entrala Formation) of the Valdegallina locality (Zamora Municipality, Zamora Province). A, MGM-5865N. B, STUS 14255. C, STUS 10623. D, STUS 8777. E, STUS 10523. F, STUS 8639. G, STUS 10606. H, STUS 12133. I, STUS 10607. J, STUS 6286. K, STUS 6321. They are shown in ventral view. MGM-5865N comes from the Valdegallina 3 fossil site, and the others from the Valdegallina 1 fossil site. The dotted lines indicate broken edges, the continuous black lines correspond to the margins of the plates, the borders of the scutes are represented by thicker gray lines, and the gray surfaces correspond to sediment. Lowercase abbreviations correspond to the plates. The abbreviations of the scutes are in capital letters.
FIGURE 8. Partial plastra the Spanish podocnemidid turtle Neochelys zamorensis from the Lutetian (middle Eocene, Entrala Formation) of the Valdegallina 1 fossil site (Valdegallina locality, Zamora Municipality, Zamora Province). A, STUS 10625. B, STUS 10624. C, STUS 9097. D, STUS 9096. E, STUS 9117. F-G, STUS 6156. H-I, STUS 9101. J-K, STUS 9102. L-M, STUS 10507. N-O, STUS 9113. P-Q, STUS 17840. A-E are shown in ventral view, but F-Q in both dorsal and ventral ones. The dotted lines indicate broken edges, the continuous black lines correspond to the margins of the plates, the borders of the scutes are represented by thicker gray lines, and the gray surfaces correspond to sediment. Lowercase abbreviations correspond to the plates. The abbreviations of the scutes are in capital letters.
FIGURE 9. Nuchal plates of the Spanish podocnemidid turtle Neochelys zamorensis from the Lutetian (middle Eocene, Entrala Formation) of several fossil sites in the Zamora Province. A, STUS 6004. B, STUS 6002. C, STUS 8754. D, STUS 17532. E, STUS 6226. F, STUS 6315. G, STUS 8755. H, STUS 1009. I, STUS 9123. J, STUS 8947. K, STUS 6003. L, STUS 6224. M, STUS 6225. N, STUS 8946. O, STUS 8749. P, STUS 8753. Q, STUS 9107. R, STUS 9124. S, STUS 14207. T, STUS 8945. U, STUS 14206. V, STUS 9108. W, STUS 8751. X, STUS 9122. Y, STUS 6295. They are shown in dorsal view. STUS 6004, STUS 6002, STUS 6001, and STUS 6003 come from the Tardemayas 1 fossil site (south margin of the Infiernos Hill, Monfarracinos Municipality), and the others from the Valdegallina 1 fossil site (Valdegallina locality, Zamora Municipality). The dotted lines indicate broken edges, the continuous black lines correspond to the margins of the plates, and the borders of the scutes are represented by thicker gray lines. Lowercase abbreviations correspond to the plates. The abbreviations of the scutes are in capital letters.
FIGURE 10. Epiplastra of the Spanish podocnemidid turtle Neochelys zamorensis from the Lutetian (middle Eocene, Entrala Formation) of several fossil sites in the Zamora Province. A, STUS 9120. B, STUS 5826. C, STUS 5828. D, STUS 6088. E, STUS 6089. F, STUS 8757. G, STUS 8756. H, STUS 6283. I, STUS 8954. J, STUS 9117. K, STUS 6087. L, STUS 6280. M, STUS 8759. N, STUS 10420. O, STUS 6086. P, STUS 8953. Q, STUS 9103. R, STUS 9115. S, STUS 6300. T, STUS 6337. U, STUS 10608. V, STUS 5754. W, STUS 8787. X, STUS 5755. Y, STUS 13633. Z, STUS 5878. They are shown in ventral view. STUS 5826, STUS 5828, STUS 6088, STUS 6089, STUS 6087, STUS 6086, and STUS 6337 come from the Tardemayas 1 fossil site (south margin of the Infiernos Hill, Monfarracinos Municipality); STUS 6300 from the Valdegallina 2 fossil site (Valdegallina locality, Zamora Municipality); STUS 5754 and STUS 5755 from the Cubillos 1 fossil site (west margin of the Infiernos Hill); and the others from the Valdegallina 1 fossil site (Valdegallina locality). The dotted lines indicate broken edges, the continuous black lines correspond to the margins of the plates, and the borders of the scutes are represented by thicker gray lines. Lowercase abbreviations correspond to the plates. The abbreviations of the scutes are in capital letters.
FIGURE 11. Entoplastra of the Spanish podocnemidid turtle Neochelys zamorensis from the Lutetian (middle Eocene, Entrala Formation) of several fossil sites in the Zamora Province. A, STUS 17723. B, STUS 6091. C, STUS 5831. D, STUS 6098. E, STUS 6093. F, STUS 6392. G, STUS 6092. H, STUS 6096. I, STUS 6372. J, STUS 6282. K, STUS 6094. L, STUS 6095. M, STUS 6170. N, STUS 8949. O, STUS 8948. P, STUS 9118. Q, STUS 6169. R, STUS 9114. S, STUS 8950. T, STUS 6097. U, STUS 9109. V, STUS 8764. W, STUS 6281. X, STUS 10501. Y, STUS 8765. Z, STUS 5756. AA, STUS 6100. AB, STUS 5880. AC, STUS 16409. AD, STUS 10500. AE, STUS 8763. AF, STUS 9110. AG, STUS 13627. They are shown in ventral view. STUS 6091, STUS 5831, STUS 6098, STUS 6093, STUS 6392, STUS 6092, STUS 6096, STUS 6372, STUS 6282, STUS 6094, STUS 6095, STUS 6097, STUS 6100, and STUS 5880 come from the Tardemayas 1 fossil site (south margin of the Infiernos Hill, Monfarracinos Municipality); STUS 5756 from the Cubillos 1 fossil site (west margin of the Infiernos Hill); STUS 16409 from the Teso de la Mora fossil site (Torres del Carrizal Municipality); and the others from the Valdegallina 1 fossil site (Valdegallina locality, Zamora Municipality). The dotted lines indicate broken edges, the continuous black lines correspond to the margins of the plates, and the borders of the scutes are represented by thicker gray lines. Lowercase abbreviations correspond to the plates. The abbreviations of the scutes are in capital letters.
FIGURE 12. Xiphiplastra of the Spanish podocnemidid turtle Neochelys zamorensis from the Lutetian (middle Eocene, Entrala Formation) of several fossil sites in the Zamora Province. A-B, STUS 5841. C-D, STUS 6107. E-F, STUS 5836. G-H, STUS 6120. I-J, STUS 6128. K-L, STUS 6130. M-N, STUS 6117. O-P, STUS 6126. Q-R, STUS 6127. S-T, STUS 6123. U-V, STUS 6262. W-X, STUS 5843. Y-Z, STUS 6125. AA-AB, STUS 6268. AC-AD, STUS 6121. AE-AF, STUS 8748. AG-AH, STUS 6122. AI-AJ, STUS 5840. AK-AL, STUS 8745. AM-AN, STUS 6102. AO-AP, STUS 6104. AQ-AR, STUS 6119. AS-AT, STUS 8741. AU-AV, STUS 6343. AW-AX, STUS 6118. AY-AZ, STUS 9099. BA-BB, STUS 6317. BC-BD, STUS 6103. BE-BF, STUS 9112. BG-BH, STUS 9100. BI-BJ, STUS 8746. BK-BL, STUS 8743. BM-BN, STUS 8956. BO-BP, STUS 6342. BQ-BR, STUS 8750. They are shown in both dorsal and ventral views. STUS 5841, STUS 6107, STUS 5836, STUS 6120, STUS 6128, STUS 6130, STUS 6117, STUS 6126, STUS 6127, STUS 6123, STUS 5843, STUS 6125, STUS 6121, STUS 6122, STUS 5840, STUS 6102, STUS 6104, STUS 6119, STUS 6118, and STUS 6103 come from the Tardemayas 1 fossil site (south margin of the Infiernos Hill, Monfarracinos Municipality); and the others from the Valdegallina 1 fossil site (Valdegallina locality, Zamora Municipality). The dotted lines indicate broken edges, the continuous black lines correspond to the margins of the plates, and the borders of the scutes are represented by thicker gray lines. Lowercase abbreviations correspond to the plates. The abbreviations of the scutes are in capital letters.
FIGURE 13. Xiphiplastra of the Spanish podocnemidid turtle Neochelys zamorensis from the Lutetian (middle Eocene, Entrala Formation) of several fossil sites in the Zamora Province. A-B, STUS 6269. C-D, STUS 8955. E-F, STUS 17840. G-H, STUS 10509. I-J, STUS 6363. K-L, STUS 8747. M-N, STUS 5757. O-P, STUS 5753. Q-R, STUS 10508. S-T, STUS 6341. U-V, STUS 9098. W-X, STUS 17526. They are shown in both dorsal and ventral views. STUS 5757 and STUS 5753 come from the Cubillos 1 fossil site (west margin of the Infiernos Hill, Monfarracinos Municipality); and the others from the Valdegallina 1 fossil site (Valdegallina locality, Zamora Municipality). The dotted lines indicate broken edges, the continuous black lines correspond to the margins of the plates, and the borders of the scutes are represented by thicker gray lines. Lowercase abbreviations correspond to the plates. The abbreviations of the scutes are in capital letters.
FIGURE 14. Evaluation of the sexual dimorphism in the anal notch of the Spanish podocnemidid turtle Neochelys zamorensis from the Lutetian (middle Eocene, Entrala Formation) of the Zamora Province. A, shape differences. B, box-plot of centroid size of the anal notch region. Dark blue concerns the first sexual morphotype and light blue represents the second sexual morphotype. Abbreviations: M1, morphotype I; M2, morphotype II.
FIGURE 15. Evaluation of sexual dimorphism of the xiphiplastral process of the Spanish podocnemidid turtle Neochelys zamorensis from the Lutetian (middle Eocene) of the Zamora Province. A, shape differences. B, box-plot of centroid size of the xiphiplastral process. Dark blue concerns the first sexual morphotype and light blue represents the second sexual morphotype. Abbreviations: M1, morphotype I; M2, morphotype II.
FIGURE 16. Linear regression between Procrustes coordinates (dependent variable) and logged-centroid size values (independent variable) of the anal notch (A) and the xiphiplastral process (B) of the Spanish podocnemidid turtle Neochelys zamorensis from the Lutetian (middle Eocene) of the Zamora Province. Dark blue concerns the first sexual morphotype and light blue represents the second sexual morphotype.
