taxonID	type	format	identifier	references	title	description	created	creator	contributor	publisher	audience	source	license	rightsHolder	datasetID
40ECC7AE014C52939CDE8DB216EE5223.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1669854	https://doi.org/10.3897/fr.29.190187.figure3	Figure 3. Upper dentition of the gerbils Debruijnimys primaevus sp. nov. from La Piquera (A – O; Early Pliocene, Duero Basin, Spain), Sifón de Librilla 413 (P; Early Pliocene, Fortuna Basin, Spain) and Puerto de la Cadena (Q; Early Pliocene, Fortuna Basin, Spain) and Debruijnimys julii from Baza- 1 (R; Early Pliocene, Guadix-Baza Basin, Spain), in occlusal view. A. UCM-LPQ 1 - DER-DEB- 1, right M 1 of D. primaevus sp. nov. from La Piquera (debris); B. UCM-LPQ 1 - DER-DEB- 2, left M 1 of D. primaevus sp. nov. from La Piquera (debris); C. UCM-LPQ 1 - DER-DEB- 3, right M 1 of D. primaevus sp. nov. from La Piquera (debris); D. UCM-LPQ 1 - DER-DEB- 4, right M 1 of D. primaevus sp. nov. from La Piquera (debris); E. UCM-LPQ 1 - DER-DEB- 5, left M 1 of D. primaevus sp. nov. from La Piquera (debris); F. UCM-LPQ 1 - N 1 - DEB- 1, left M 1 of D. primaevus sp. nov. from La Piquera (level 1); G. UCM-LPQ 1 - N 2 - DEB- 2, left M 1 of D. primaevus sp. nov. from La Piquera (level 2); H. UCM-LPQ 1 - N 2 - DEB- 3, left M 1 of D. primaevus sp. nov. from La Piquera (level 2); I. UCM-LPQ 1 - N 3 - DEB- 1, right M 1 of D. primaevus sp. nov. from La Piquera (level 3); J. UCM-LPQ 1 - N 4 - DEB- 1, left M 1 of D. primaevus sp. nov. from La Piquera (level 4); K. UCM-LPQ 1 - N 4 - DEB- 2, left M 1 of D. primaevus sp. nov. from La Piquera (level 4); L. UCM-LPQ 1 - N 4 - DEB- 3, right M 1 of D. primaevus sp. nov. from La Piquera (level 4); M. UCM-LPQ 1 - N 4 - DEB- 3, right M 1 of D. primaevus sp. nov. from La Piquera (level 4); N. UCM-LPQ 1 - N 4 - DEB- 5, left M 1 of D. primaevus sp. nov. from La Piquera (level 4); O. UCM-LPQ 1 - N 4 - DEB- 4, right M 1 of D. primaevus sp. nov. from La Piquera (level 4); P. IPS- 101286, left M 1 of D. primaevus sp. nov. from Sifón de Librilla 413; Q. MAM / DA / 2012-0019 / A 5 ' 06 - 1, right M 1 of D. primaevus sp. nov from Puerto de la Cadena; R. BA 1-2001 - R 11 / 1, right M 1 of D. julii from Baza- 1.	Figure 3. Upper dentition of the gerbils Debruijnimys primaevus sp. nov. from La Piquera (A – O; Early Pliocene, Duero Basin, Spain), Sifón de Librilla 413 (P; Early Pliocene, Fortuna Basin, Spain) and Puerto de la Cadena (Q; Early Pliocene, Fortuna Basin, Spain) and Debruijnimys julii from Baza- 1 (R; Early Pliocene, Guadix-Baza Basin, Spain), in occlusal view. A. UCM-LPQ 1 - DER-DEB- 1, right M 1 of D. primaevus sp. nov. from La Piquera (debris); B. UCM-LPQ 1 - DER-DEB- 2, left M 1 of D. primaevus sp. nov. from La Piquera (debris); C. UCM-LPQ 1 - DER-DEB- 3, right M 1 of D. primaevus sp. nov. from La Piquera (debris); D. UCM-LPQ 1 - DER-DEB- 4, right M 1 of D. primaevus sp. nov. from La Piquera (debris); E. UCM-LPQ 1 - DER-DEB- 5, left M 1 of D. primaevus sp. nov. from La Piquera (debris); F. UCM-LPQ 1 - N 1 - DEB- 1, left M 1 of D. primaevus sp. nov. from La Piquera (level 1); G. UCM-LPQ 1 - N 2 - DEB- 2, left M 1 of D. primaevus sp. nov. from La Piquera (level 2); H. UCM-LPQ 1 - N 2 - DEB- 3, left M 1 of D. primaevus sp. nov. from La Piquera (level 2); I. UCM-LPQ 1 - N 3 - DEB- 1, right M 1 of D. primaevus sp. nov. from La Piquera (level 3); J. UCM-LPQ 1 - N 4 - DEB- 1, left M 1 of D. primaevus sp. nov. from La Piquera (level 4); K. UCM-LPQ 1 - N 4 - DEB- 2, left M 1 of D. primaevus sp. nov. from La Piquera (level 4); L. UCM-LPQ 1 - N 4 - DEB- 3, right M 1 of D. primaevus sp. nov. from La Piquera (level 4); M. UCM-LPQ 1 - N 4 - DEB- 3, right M 1 of D. primaevus sp. nov. from La Piquera (level 4); N. UCM-LPQ 1 - N 4 - DEB- 5, left M 1 of D. primaevus sp. nov. from La Piquera (level 4); O. UCM-LPQ 1 - N 4 - DEB- 4, right M 1 of D. primaevus sp. nov. from La Piquera (level 4); P. IPS- 101286, left M 1 of D. primaevus sp. nov. from Sifón de Librilla 413; Q. MAM / DA / 2012-0019 / A 5 ' 06 - 1, right M 1 of D. primaevus sp. nov from Puerto de la Cadena; R. BA 1-2001 - R 11 / 1, right M 1 of D. julii from Baza- 1.	2026-06-04	Agustí, Jordi;Piñero, Pedro;Blain, Hugues-Alexandre;Martín-Perea, David M.;Sevilla, Paloma;Laplana, Cesar		Zenodo	biologists	Agustí, Jordi;Piñero, Pedro;Blain, Hugues-Alexandre;Martín-Perea, David M.;Sevilla, Paloma;Laplana, Cesar			
40ECC7AE014C52939CDE8DB216EE5223.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1669855	https://doi.org/10.3897/fr.29.190187.figure4	Figure 4. Upper dentition of the gerbils Debruijnimys primaevus sp. nov. from La Piquera (A – E, I; Early Pliocene, Duero Basin, Spain), Botardo D (F; Early Pliocene, Guadix-Baza Basin, Spain) and Sifón de Librilla 413 (G, J; Early Pliocene, Fortuna Basin, Spain) and Debruijnimys julii from Baza- 1 (H; Early Pliocene, Guadix-Baza Basin, Spain), in occlusal view. A. UCM-LPQ 1 - DER-DEB- 6, left M 2 of D. primaevus sp. nov. from La Piquera (debris); B. UCM-LPQ 1 - N 1 - DEB- 3, right M 2 of D. primaevus sp. nov. from La Piquera (level 1); C. UCM-LPQ 1 - N 1 - DEB- 4, right M 2 of D. primaevus sp. nov. from La Piquera (level 1); D. UCM-LPQ 1 - N 1 - DEB- 5, right M 2 of D. primaevus sp. nov. from La Piquera (level 1); E. UCM-LPQ 1 - N 2 - DEB- 6, right M 2 of D. primaevus sp. nov. from La Piquera (level 2); F. IPS- 116139, right M 2 of D. primaevus sp. nov. from Botardo D; G. IPS- 101291, left M 2 of D. primaevus sp. nov. from Sifón de Librilla 413; H. BA 1-2001 - R 11 / 3, right M 2 of D. julii from Baza- 1; I. UCM-LPQ 1 - N 2 - DEB- 7, right M 3 of D. primaevus sp. nov. from La Piquera (level 2); J. IPS- 101289, right M 3 of D. primaevus sp. nov. from Sifón de Librilla 413.	Figure 4. Upper dentition of the gerbils Debruijnimys primaevus sp. nov. from La Piquera (A – E, I; Early Pliocene, Duero Basin, Spain), Botardo D (F; Early Pliocene, Guadix-Baza Basin, Spain) and Sifón de Librilla 413 (G, J; Early Pliocene, Fortuna Basin, Spain) and Debruijnimys julii from Baza- 1 (H; Early Pliocene, Guadix-Baza Basin, Spain), in occlusal view. A. UCM-LPQ 1 - DER-DEB- 6, left M 2 of D. primaevus sp. nov. from La Piquera (debris); B. UCM-LPQ 1 - N 1 - DEB- 3, right M 2 of D. primaevus sp. nov. from La Piquera (level 1); C. UCM-LPQ 1 - N 1 - DEB- 4, right M 2 of D. primaevus sp. nov. from La Piquera (level 1); D. UCM-LPQ 1 - N 1 - DEB- 5, right M 2 of D. primaevus sp. nov. from La Piquera (level 1); E. UCM-LPQ 1 - N 2 - DEB- 6, right M 2 of D. primaevus sp. nov. from La Piquera (level 2); F. IPS- 116139, right M 2 of D. primaevus sp. nov. from Botardo D; G. IPS- 101291, left M 2 of D. primaevus sp. nov. from Sifón de Librilla 413; H. BA 1-2001 - R 11 / 3, right M 2 of D. julii from Baza- 1; I. UCM-LPQ 1 - N 2 - DEB- 7, right M 3 of D. primaevus sp. nov. from La Piquera (level 2); J. IPS- 101289, right M 3 of D. primaevus sp. nov. from Sifón de Librilla 413.	2026-06-04	Agustí, Jordi;Piñero, Pedro;Blain, Hugues-Alexandre;Martín-Perea, David M.;Sevilla, Paloma;Laplana, Cesar		Zenodo	biologists	Agustí, Jordi;Piñero, Pedro;Blain, Hugues-Alexandre;Martín-Perea, David M.;Sevilla, Paloma;Laplana, Cesar			
40ECC7AE014C52939CDE8DB216EE5223.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1669856	https://doi.org/10.3897/fr.29.190187.figure5	Figure 5. Lower dentition of the gerbils Debruijnimys primaevus sp. nov. from La Piquera (A – I, M – R, T, U; Early Pliocene, Duero Basin, Spain), Botardo D (J, S; Early Pliocene, Guadix-Baza Basin, Spain) and Sifón de Librilla 413 (K; Early Pliocene, Fortuna Basin, Spain), and Debruijnimys julii from Baza- 1 (L; Early Pliocene, Guadix-Baza Basin, Spain), in occlusal view. A. UCM-LPQ 1 - N 5 B-DEB- 1, right m 1, holotype of D. primaevus sp. nov. from La Piquera (level 5); B. UCM-LPQ 1 - N 4 - DEB- 6, left m 1 of D. primaevus sp. nov. from La Piquera (level 4); C. UCM-LPQ 1 - N 2 - DEB- 5, left m 1 of D. primaevus sp. nov. from La Piquera (level 2); D. UCM-LPQ 1 - DER-DEB- 8, left m 1 of D. primaevus sp. nov. from La Piquera (debris); E. UCM-LPQ 1 - DER-DEB- 9, right m 1 of D. primaevus sp. nov. from La Piquera (debris); F. UCM-LPQ 1 - DER-DEB- 10, left m 1 of D. primaevus sp. nov. from La Piquera (debris); G. UCM-LPQ 1 - N 2 - DEB- 4, left m 1 of D. primaevus sp. nov. from La Piquera (level 2); H. UCM-LPQ 1 - N 1 - DEB- 2, left m 1 of D. primaevus sp. nov. from La Piquera (level 1); I. UCM-LPQ 1 - DER-DEB- 11, anterior portion of left m 1 of D. primaevus sp. nov. from La Piquera (debris); J. IPS- 116124, right m 1 of D. primaevus sp. nov. from Botardo D; K. IPS- 101292, right m 1 of D. primaevus sp. nov. from Sifón de Librilla 413; L. BA 1-2001 - R 11 - 1, right m 1 of D. julii from Baza- 1; M. UCM-LPQ 1 - DER-DEB- 7, right m 2 of D. primaevus sp. nov. from La Piquera (debris); N. UCM-LPQ 1 - DER-DEB- 12, right m 2 of D. primaevus sp. nov. from La Piquera (debris); O. UCM-LPQ 1 - DER-DEB- 13, left m 2 of D. primaevus sp. nov. from La Piquera (debris); P. UCM-LPQ 1 - N 1 - DEB- 6, left m 2 of D. primaevus sp. nov. from La Piquera (level 1); Q. UCM-LPQ 1 - N 1 - DEB- 7, left m 2 of D. primaevus sp. nov. from La Piquera (level 1); R. UCM-LPQ 1 - N 4 - DEB- 7, left m 2 of D. primaevus sp. nov. from La Piquera (level 4); S. IPS- 116138, right m 2 of D. primaevus sp. nov. from Botardo D; T. UCM-LPQ 1 - N 1 - DEB- 8, right m 3 of D. primaevus sp. nov. from La Piquera (level 1); U. UCM-LPQ 1 - N 5 B-DEB- 2, left m 3 of D. primaevus sp. nov. from La Piquera (level 5).	Figure 5. Lower dentition of the gerbils Debruijnimys primaevus sp. nov. from La Piquera (A – I, M – R, T, U; Early Pliocene, Duero Basin, Spain), Botardo D (J, S; Early Pliocene, Guadix-Baza Basin, Spain) and Sifón de Librilla 413 (K; Early Pliocene, Fortuna Basin, Spain), and Debruijnimys julii from Baza- 1 (L; Early Pliocene, Guadix-Baza Basin, Spain), in occlusal view. A. UCM-LPQ 1 - N 5 B-DEB- 1, right m 1, holotype of D. primaevus sp. nov. from La Piquera (level 5); B. UCM-LPQ 1 - N 4 - DEB- 6, left m 1 of D. primaevus sp. nov. from La Piquera (level 4); C. UCM-LPQ 1 - N 2 - DEB- 5, left m 1 of D. primaevus sp. nov. from La Piquera (level 2); D. UCM-LPQ 1 - DER-DEB- 8, left m 1 of D. primaevus sp. nov. from La Piquera (debris); E. UCM-LPQ 1 - DER-DEB- 9, right m 1 of D. primaevus sp. nov. from La Piquera (debris); F. UCM-LPQ 1 - DER-DEB- 10, left m 1 of D. primaevus sp. nov. from La Piquera (debris); G. UCM-LPQ 1 - N 2 - DEB- 4, left m 1 of D. primaevus sp. nov. from La Piquera (level 2); H. UCM-LPQ 1 - N 1 - DEB- 2, left m 1 of D. primaevus sp. nov. from La Piquera (level 1); I. UCM-LPQ 1 - DER-DEB- 11, anterior portion of left m 1 of D. primaevus sp. nov. from La Piquera (debris); J. IPS- 116124, right m 1 of D. primaevus sp. nov. from Botardo D; K. IPS- 101292, right m 1 of D. primaevus sp. nov. from Sifón de Librilla 413; L. BA 1-2001 - R 11 - 1, right m 1 of D. julii from Baza- 1; M. UCM-LPQ 1 - DER-DEB- 7, right m 2 of D. primaevus sp. nov. from La Piquera (debris); N. UCM-LPQ 1 - DER-DEB- 12, right m 2 of D. primaevus sp. nov. from La Piquera (debris); O. UCM-LPQ 1 - DER-DEB- 13, left m 2 of D. primaevus sp. nov. from La Piquera (debris); P. UCM-LPQ 1 - N 1 - DEB- 6, left m 2 of D. primaevus sp. nov. from La Piquera (level 1); Q. UCM-LPQ 1 - N 1 - DEB- 7, left m 2 of D. primaevus sp. nov. from La Piquera (level 1); R. UCM-LPQ 1 - N 4 - DEB- 7, left m 2 of D. primaevus sp. nov. from La Piquera (level 4); S. IPS- 116138, right m 2 of D. primaevus sp. nov. from Botardo D; T. UCM-LPQ 1 - N 1 - DEB- 8, right m 3 of D. primaevus sp. nov. from La Piquera (level 1); U. UCM-LPQ 1 - N 5 B-DEB- 2, left m 3 of D. primaevus sp. nov. from La Piquera (level 5).	2026-06-04	Agustí, Jordi;Piñero, Pedro;Blain, Hugues-Alexandre;Martín-Perea, David M.;Sevilla, Paloma;Laplana, Cesar		Zenodo	biologists	Agustí, Jordi;Piñero, Pedro;Blain, Hugues-Alexandre;Martín-Perea, David M.;Sevilla, Paloma;Laplana, Cesar			
40ECC7AE014C52939CDE8DB216EE5223.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1669857	https://doi.org/10.3897/fr.29.190187.figure6	Figure 6. Ranges of size variation (in mm) in the upper (M 1 and M 2) and lower (m 1 and m 2) teeth of Debruijnimys primaevus sp. nov. from La Piquera (LP, type locality; this paper), Puerto de la Cadena (PC; Piñero et al. (2017 a)), Botardo D (BO-D; Piñero and Agustí (2020)), Sifón de Librilla 413 (SIF- 413; Piñero and Agustí (2019)), La Bullana 2 B (LB- 2 B; Mansino et al. (2015)) and La Gloria 4 (LG- 4; Adrover et al. (1993)); Debruijnimys julii from Asta Regia 3 (AR- 3, type locality; Castillo and Agustí 1996), Gorafe 4 (GOR- 4; Martín-Suárez (1988)) and Baza 1 (BZ- 1; Piñero et al. (2017 b)); Debruijnimys davidi from Lissasfa (LSF, type locality; Geraads (1998)); Protatera almenarensis from Almenara Casablanca M (ACB-M, type locality; Agustí (1989 a)); Protatera algeriensis from Amama 2 (AMA- 2; Jaeger (1977)); and Protatera salvadori from Ses Fontanelles (SF, type locality; Agustí et al. (2026)). Mean sizes are marked by symbols inside the boxes.	Figure 6. Ranges of size variation (in mm) in the upper (M 1 and M 2) and lower (m 1 and m 2) teeth of Debruijnimys primaevus sp. nov. from La Piquera (LP, type locality; this paper), Puerto de la Cadena (PC; Piñero et al. (2017 a)), Botardo D (BO-D; Piñero and Agustí (2020)), Sifón de Librilla 413 (SIF- 413; Piñero and Agustí (2019)), La Bullana 2 B (LB- 2 B; Mansino et al. (2015)) and La Gloria 4 (LG- 4; Adrover et al. (1993)); Debruijnimys julii from Asta Regia 3 (AR- 3, type locality; Castillo and Agustí 1996), Gorafe 4 (GOR- 4; Martín-Suárez (1988)) and Baza 1 (BZ- 1; Piñero et al. (2017 b)); Debruijnimys davidi from Lissasfa (LSF, type locality; Geraads (1998)); Protatera almenarensis from Almenara Casablanca M (ACB-M, type locality; Agustí (1989 a)); Protatera algeriensis from Amama 2 (AMA- 2; Jaeger (1977)); and Protatera salvadori from Ses Fontanelles (SF, type locality; Agustí et al. (2026)). Mean sizes are marked by symbols inside the boxes.	2026-06-04	Agustí, Jordi;Piñero, Pedro;Blain, Hugues-Alexandre;Martín-Perea, David M.;Sevilla, Paloma;Laplana, Cesar		Zenodo	biologists	Agustí, Jordi;Piñero, Pedro;Blain, Hugues-Alexandre;Martín-Perea, David M.;Sevilla, Paloma;Laplana, Cesar			
40ECC7AE014C52939CDE8DB216EE5223.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://binary.pensoft.net/fig/1669858	https://doi.org/10.3897/fr.29.190187.figure7	Figure 7. Biostratigraphic distribution and evolution of Debruijnimys species and the Iberian Protatera, showing their chronological ranges. The localities marked with an asterisk are calibrated with palaeomagnetic data (Garcés et al. 1998, 2001; Martín-Suárez et al. 2000; Mansino et al. 2015; Piñero et al. 2017 a, 2018). Arrows indicate the two dispersal events of gerbils into the Iberian Peninsula. Abbreviations: ATNTS, Astronomically Tuned Neogene Time Scale; ELMA, European Land Mammal Ages; MN, Mammal Neogene.	Figure 7. Biostratigraphic distribution and evolution of Debruijnimys species and the Iberian Protatera, showing their chronological ranges. The localities marked with an asterisk are calibrated with palaeomagnetic data (Garcés et al. 1998, 2001; Martín-Suárez et al. 2000; Mansino et al. 2015; Piñero et al. 2017 a, 2018). Arrows indicate the two dispersal events of gerbils into the Iberian Peninsula. Abbreviations: ATNTS, Astronomically Tuned Neogene Time Scale; ELMA, European Land Mammal Ages; MN, Mammal Neogene.	2026-06-04	Agustí, Jordi;Piñero, Pedro;Blain, Hugues-Alexandre;Martín-Perea, David M.;Sevilla, Paloma;Laplana, Cesar		Zenodo	biologists	Agustí, Jordi;Piñero, Pedro;Blain, Hugues-Alexandre;Martín-Perea, David M.;Sevilla, Paloma;Laplana, Cesar			
