c3999648a950624126ebb24885672221 foss-rec_article_190187_en_1.pdf c635cb9c4ab4876cf30e4d21612c4c9f910cde11 foss-rec_article_190187_en_1.pdf 7042769a9a6e37f14a3356c5adabbbf5849519bb7175283fa7c3a7120ee88e01 foss-rec_article_190187_en_1.pdf Title: A new species of gerbil (Gerbillidae, Rodentia, Mammalia) from the Early Pliocene of the Iberian Peninsula and its bearing on faunal dispersals during the Messinian (Late Miocene) Subject: In this paper, the new gerbil species Debruijnimys primaevus sp. nov. is described from the Early Pliocene site of La Piquera in central Spain. This species differs from the type species Debruijnimys julii in its smaller dimensions and less developed longitudinal ridges in the lower molars. It can be derived from the Moroccan species Debruijnimys davidi by an increase in size and modification of the shape of the lower molars. Therefore, it cannot be related to the previously described Protatera almenarensis from other Late Miocene sites in Spain. It is proposed that the entry of the two gerbils into the Iberian Peninsula took place in two different phases during the Lago Mare stage of the Messinian. The first, involving the entry of Protatera almenarensis as well as other African and Asian elements, coincided with the first substage of the Lago Mare phase, between 5.546 and 5.352 Ma. The second took place during the late stage of the Lago Mare phase, between 5.338 and 5.333 Ma. After the Zanclean flood, members of the genus Debruijnimys persisted in several Pliocene sites of the Iberian Peninsula (D. primaevus sp. nov. and D. julii). Author: Jordi Agustí Creator: Adobe InDesign 21.1 (Windows) Producer: Adobe PDF Library 18.0 CreationDate: Thu Jun 4 09:57:08 2026 CEST ModDate: Thu Jun 4 09:57:16 2026 CEST Custom Metadata: no Metadata Stream: yes Tagged: yes UserProperties: no Suspects: no Form: none JavaScript: no Pages: 16 Encrypted: no Page size: 595.276 x 841.89 pts (A4) Page rot: 0 File size: 5143079 bytes Optimized: yes PDF version: 1.4 name type encoding emb sub uni object ID ------------------------------------ ----------------- ---------------- --- --- --- --------- BSXDTR+TimesNewRomanPS-ItalicMT CID TrueType Identity-H yes yes yes 3389 0 MTKMRV+SymbolMT CID TrueType Identity-H yes yes yes 3411 0 OYKITR+TimesNewRomanPSMT CID TrueType Identity-H yes yes yes 3412 0 IXLTND+NewsGothicStd Type 1C Custom yes yes yes 3392 0 MTKMRV+FFMarkPro-Bold Type 1C WinAnsi yes yes yes 3413 0 GSGCLH+NewsGothic-Oblique Type 1C WinAnsi yes yes yes 3416 0 YVNQDX+TimesNewRomanPSMT TrueType WinAnsi yes yes yes 3395 0 IXLTND+NewsGothicMT TrueType WinAnsi yes yes yes 3396 0 MTKMRV+TimesNewRomanPS-ItalicMT TrueType WinAnsi yes yes yes 3403 0 GSGCLH+TimesNewRomanPS-BoldMT TrueType WinAnsi yes yes yes 3407 0 RTTLJL+TimesNewRomanPS-BoldItalicMT TrueType WinAnsi yes yes yes 52 0 IXLTND+NewsGothicStd Type 1C WinAnsi yes yes yes 50 0 IXLTND+NewsGothicStd Type 1C WinAnsi yes yes yes 57 0 OEHYTR+TimesNewRomanPS-BoldMT CID TrueType Identity-H yes yes yes 60 0 Jhove (Rel. 1.28.0, 2023-05-18) Date: 2026-06-09 03:56:21 CEST RepresentationInformation: foss-rec_article_190187_en_1.pdf ReportingModule: PDF-hul, Rel. 1.12.4 (2023-03-16) LastModified: 2026-06-08 10:18:20 CEST Size: 5143079 Format: PDF Version: 1.4 Status: Well-Formed, but not valid SignatureMatches: PDF-hul ErrorMessage: Annotation object is not a dictionary ID: PDF-HUL-114 Offset: 5660 MIMEtype: application/pdf Profile: Linearized PDF PDFMetadata: Objects: 3454 FreeObjects: 3247 IncrementalUpdates: 1 DocumentCatalog: ViewerPreferences: HideToolbar: false HideMenubar: false HideWindowUI: false FitWindow: false CenterWindow: false DisplayDocTitle: false NonFullScreenPageMode: UseNone Direction: L2R ViewArea: CropBox ViewClip: CropBox PrintArea: CropBox PageClip: CropBox PageLayout: SinglePage PageMode: UseNone Language: en-US Outlines: Item: Title: A new species of gerbil (Gerbillidae, Rodentia, Mammalia) from the Early Pliocene of the Iberian Peninsula and its bearing on faunal dispersals during the Messinian (Late Miocene) Item: Title: Abstract Item: Title: Introduction Item: Title: Material and methods Item: Title: Systematic Palaeontology Children: Item: Title: Genus Debruijnimys Castillo & Agustí, 1996 Item: Title: Debruijnimys primaevus sp. nov. Item: Title: Discussion Item: Title: Conclusions Item: Title: Competing interests Item: Title: Author contributions Item: Title: Acknowledgements Item: Title: References Info: Title: A new species of gerbil (Gerbillidae, Rodentia, Mammalia) from the Early Pliocene of the Iberian Peninsula and its bearing on faunal dispersals during the Messinian (Late Miocene) Author: Jordi Agustí Subject: In this paper, the new gerbil species Debruijnimys primaevus sp. nov. is described from the Early Pliocene site of La Piquera in central Spain. This species differs from the type species Debruijnimys julii in its smaller dimensions and less developed longitudinal ridges in the lower molars. It can be derived from the Moroccan species Debruijnimys davidi by an increase in size and modification of the shape of the lower molars. Therefore, it cannot be related to the previously described Protatera almenarensis from other Late Miocene sites in Spain. It is proposed that the entry of the two gerbils into the Iberian Peninsula took place in two different phases during the Lago Mare stage of the Messinian. The first, involving the entry of Protatera almenarensis as well as other African and Asian elements, coincided with the first substage of the Lago Mare phase, between 5.546 and 5.352 Ma. The second took place during the late stage of the Lago Mare phase, between 5.338 and 5.333 Ma. After the Zanclean flood, members of the genus Debruijnimys persisted in several Pliocene sites of the Iberian Peninsula (D. primaevus sp. nov. and D. julii). Creator: Adobe InDesign 21.1 (Windows) Producer: Adobe PDF Library 18.0 CreationDate: Thu Jun 04 09:57:08 CEST 2026 ModDate: Thu Jun 04 09:57:16 CEST 2026 ID: 0xad9fe18ab3a0df478b2157c25bbea3b3, 0x2380d6614a55b443aa40fc926a40e4c4 Filters: FilterPipeline: FlateDecode FilterPipeline: DCTDecode Images: Image: NisoImageMetadata: FormatName: image/jpg CompressionScheme: JPEG ImageWidth: 1961 ImageHeight: 2428 ColorSpace: RGB BitsPerSample: 8 BitsPerSampleUnit: integer Intent: RelativeColorimetric Name: X Image: NisoImageMetadata: FormatName: image/png ImageWidth: 1961 ImageHeight: 1603 ColorSpace: black is zero BitsPerSample: 8 BitsPerSampleUnit: integer Filter: FlateDecode Intent: RelativeColorimetric Name: X Image: NisoImageMetadata: FormatName: image/jpg CompressionScheme: JPEG ImageWidth: 1961 ImageHeight: 2529 ColorSpace: black is zero BitsPerSample: 8 BitsPerSampleUnit: integer Intent: RelativeColorimetric Name: X Image: NisoImageMetadata: FormatName: image/jpg CompressionScheme: JPEG ImageWidth: 1961 ImageHeight: 1348 ColorSpace: black is zero BitsPerSample: 8 BitsPerSampleUnit: integer Intent: RelativeColorimetric Name: X Image: NisoImageMetadata: FormatName: image/jpg CompressionScheme: JPEG ImageWidth: 1961 ImageHeight: 2451 ColorSpace: black is zero BitsPerSample: 8 BitsPerSampleUnit: integer Intent: RelativeColorimetric Name: X Image: NisoImageMetadata: FormatName: image/png ImageWidth: 1961 ImageHeight: 1931 ColorSpace: RGB BitsPerSample: 8 BitsPerSampleUnit: integer Filter: FlateDecode Intent: RelativeColorimetric Name: X Image: NisoImageMetadata: FormatName: image/png ImageWidth: 1931 ImageHeight: 1881 ColorSpace: black is zero BitsPerSample: 8 BitsPerSampleUnit: integer Filter: FlateDecode Intent: RelativeColorimetric Name: X Fonts: Type0: Font: BaseFont: MTKMRV+SymbolMT Encoding: Identity-H ToUnicode: true Font: BaseFont: OYKITR+TimesNewRomanPSMT Encoding: Identity-H ToUnicode: true Font: BaseFont: OEHYTR+TimesNewRomanPS-BoldMT Encoding: Identity-H ToUnicode: true Font: BaseFont: BSXDTR+TimesNewRomanPS-ItalicMT Encoding: Identity-H ToUnicode: true Type1: Font: BaseFont: IXLTND+NewsGothicStd FontSubset: true FirstChar: 31 LastChar: 150 FontDescriptor: FontName: IXLTND+NewsGothicStd Flags: Nonsymbolic FontBBox: -187, -250, 1000, 1020 FontFile3: true EncodingDictionary: BaseEncoding: WinAnsiEncoding Differences: true ToUnicode: true Font: BaseFont: MTKMRV+FFMarkPro-Bold FontSubset: true FirstChar: 65 LastChar: 118 FontDescriptor: FontName: MTKMRV+FFMarkPro-Bold Flags: Nonsymbolic FontBBox: -680, -236, 1553, 1113 FontFile3: true Encoding: WinAnsiEncoding ToUnicode: true Font: BaseFont: GSGCLH+NewsGothic-Oblique FontSubset: true FirstChar: 32 LastChar: 237 FontDescriptor: FontName: GSGCLH+NewsGothic-Oblique Flags: Nonsymbolic, Italic FontBBox: -218, -250, 1165, 1020 FontFile3: true Encoding: WinAnsiEncoding ToUnicode: true TrueType: Font: BaseFont: YVNQDX+TimesNewRomanPSMT FontSubset: true FirstChar: 32 LastChar: 252 FontDescriptor: FontName: YVNQDX+TimesNewRomanPSMT Flags: Serif, Nonsymbolic FontBBox: -568, -307, 2046, 1040 FontFile2: true Encoding: WinAnsiEncoding ToUnicode: true Font: BaseFont: IXLTND+NewsGothicMT FontSubset: true FirstChar: 44 LastChar: 56 FontDescriptor: FontName: IXLTND+NewsGothicMT Flags: Nonsymbolic FontBBox: -155, -224, 1349, 942 FontFile2: true Encoding: WinAnsiEncoding ToUnicode: true Font: BaseFont: RTTLJL+TimesNewRomanPS-BoldItalicMT FontSubset: true FirstChar: 32 LastChar: 121 FontDescriptor: FontName: RTTLJL+TimesNewRomanPS-BoldItalicMT Flags: Serif, Nonsymbolic, Italic FontBBox: -547, -307, 1401, 1032 FontFile2: true Encoding: WinAnsiEncoding ToUnicode: true Font: BaseFont: MTKMRV+TimesNewRomanPS-ItalicMT FontSubset: true FirstChar: 32 LastChar: 232 FontDescriptor: FontName: MTKMRV+TimesNewRomanPS-ItalicMT Flags: Serif, Nonsymbolic, Italic FontBBox: -498, -307, 1333, 1023 FontFile2: true Encoding: WinAnsiEncoding ToUnicode: true Font: BaseFont: GSGCLH+TimesNewRomanPS-BoldMT FontSubset: true FirstChar: 32 LastChar: 237 FontDescriptor: FontName: GSGCLH+TimesNewRomanPS-BoldMT Flags: Serif, Nonsymbolic FontBBox: -558, -328, 2000, 1056 FontFile2: true Encoding: WinAnsiEncoding ToUnicode: true CIDFontType2: Font: BaseFont: MTKMRV+SymbolMT CIDSystemInfo: Registry: Adobe Registry: Identity Supplement: 0 FontDescriptor: FontName: MTKMRV+SymbolMT Flags: Symbolic FontBBox: 0, -220, 1113, 1005 FontFile2: true Font: BaseFont: BSXDTR+TimesNewRomanPS-ItalicMT CIDSystemInfo: Registry: Adobe Registry: Identity Supplement: 0 FontDescriptor: FontName: BSXDTR+TimesNewRomanPS-ItalicMT Flags: Serif, Symbolic, Italic FontBBox: -498, -307, 1333, 1023 FontFile2: true Font: BaseFont: OYKITR+TimesNewRomanPSMT CIDSystemInfo: Registry: Adobe Registry: Identity Supplement: 0 FontDescriptor: FontName: OYKITR+TimesNewRomanPSMT Flags: Serif, Symbolic FontBBox: -568, -307, 2046, 1040 FontFile2: true Font: BaseFont: OEHYTR+TimesNewRomanPS-BoldMT CIDSystemInfo: Registry: Adobe Registry: Identity Supplement: 0 FontDescriptor: FontName: OEHYTR+TimesNewRomanPS-BoldMT Flags: Serif, Symbolic FontBBox: -558, -328, 2000, 1056 FontFile2: true XMP: application/pdf A new species of gerbil (Gerbillidae, Rodentia, Mammalia) from the Early Pliocene of the Iberian Peninsula and its bearing on faunal dispersals during the Messinian (Late Miocene) In this paper, the new gerbil species Debruijnimys primaevus sp. nov. is described from the Early Pliocene site of La Piquera in central Spain. This species differs from the type species Debruijnimys julii in its smaller dimensions and less developed longitudinal ridges in the lower molars. It can be derived from the Moroccan species Debruijnimys davidi by an increase in size and modification of the shape of the lower molars. Therefore, it cannot be related to the previously described Protatera almenarensis from other Late Miocene sites in Spain. It is proposed that the entry of the two gerbils into the Iberian Peninsula took place in two different phases during the Lago Mare stage of the Messinian. The first, involving the entry of Protatera almenarensis as well as other African and Asian elements, coincided with the first substage of the Lago Mare phase, between 5.546 and 5.352 Ma. The second took place during the late stage of the Lago Mare phase, between 5.338 and 5.333 Ma. After the Zanclean flood, members of the genus Debruijnimys persisted in several Pliocene sites of the Iberian Peninsula (D. primaevus sp. nov. and D. julii). Jordi Agustí et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Jordi Agustí Pedro Piñero Hugues-Alexandre Blain David M. Martín-Perea Paloma Sevilla César Laplana Early Ruscinian Gerbillidae Messinian Salinity Crisis south-western Europe 2026-06-04T10:57:08+03:00 2026-06-04T10:57:16+03:00 2026-06-04T10:57:16+03:00 Adobe InDesign 21.1 (Windows) uuid:53b83c23-5f11-4e1d-b0ee-af5f0fd4fbd1 xmp.did:9F28E084BD8CE31184EC999A5D291C94 xmp.id:d3ce822a-a59a-c842-80ee-ff01b2a9a02c proof:pdf xmp.iid:879c875d-58db-a748-bd75-dcb1827c3004 xmp.did:4444FC51923DE811ADBCA6C78E600139 xmp.did:9F28E084BD8CE31184EC999A5D291C94 default converted from application/x-indesign to application/pdf Adobe InDesign 21.1 (Windows) / 2026-06-04T10:57:08+03:00 Adobe PDF Library 18.0 False Checksum: 64d8ebdb Type: CRC32 Checksum: c3999648a950624126ebb24885672221 Type: MD5 Checksum: c635cb9c4ab4876cf30e4d21612c4c9f910cde11 Type: SHA-1 Checksum: 7042769a9a6e37f14a3356c5adabbbf5849519bb7175283fa7c3a7120ee88e01 Type: SHA-256