Geologica Acta: an international earth science journal ISSN: 1695-6133 [email protected] Universitat de Barcelona España

GÓMEZ DE SOLER, B.; CAMPENY VALL-LLOSERA, G.; VAN DER MADE, J.; OMS, O.; AGUSTÍ, J.; SALA, R.; BLAIN, H.-A.; BURJACHS, F.; CLAUDE, J.; GARCÍA CATALÁN, S.; RIBA, D.; ROSILLO, R. A new key locality for the Pliocene vertebrate record of Europe: the Camp dels Ninots maar (NE Spain) Geologica Acta: an international earth science journal, vol. 10, núm. 1, 2012, pp. 1-17 Universitat de Barcelona Barcelona, España

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How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 .geologica-acta.com o c . a t c a 7 - 1 a - c 1 i g , o 2 l 1 o 0 e 2 g w. h w c w r 2 a 0 t M 7 a 1 , 0 e 1 0 n 0 i º 0 l N 0 n . o , 5 0 0 e 1 1 l . / b l 4 a o 4 V l 3 i 1 , a . a v A 0 t 1 c A : I a O c D i g o l o e G R. SALA B.GÓMEZDESOLERG.CAMPENYVALL-LLOSERA J.VAN DERMADE O.OMSJ.AGUSTÍ comparable to other remarkable maar sites suchDeposition as Messel,theatlake bottom seemsthe haveto Eocene taken place siteoxygenin depleted situated layers. inthisInway, Germany. Campdels Ninots is leprosa of them in anatomicaltheminof connection. rodentThe well as isolated remains. A minimum of five individuals of the chelonian helveticus recovered:threeindividuals of made it ideal for the preservation of fossils.delsNinots. TheparticularAt present,geological fiveconditions largethesite,ofwhich correspond mammal lacustrineto skeletons Asedimentation new in Pliocenemaar, anatomicalain Konservat-Lagerstätte connection have been in north-eastern Spain is described here for the first time. It is referred to as Camp thebiogeographic range ofsome vertebrate taxa, such as fortheCamp dels Ninots, near theMN15-MN16 transition. The Camp delNinots upfossil torecord theenables presentoneto extend time. The coexistence of KEYWORDS Avinguda de Catalunya 35, 43002 Tarragona, Spain. Gómez de Soler E-mail: Plaça Imperial Tarraco 1, 43005 Tarragona, Spain. J. Agustí E-mail: C/ Escorxadors/n,43003Tarragona, Spain.GómezdeSolerE-mail:[email protected] Campeny E-mail:[email protected] A newkeylocalityforthePliocenevertebraterecordofEurope: Burjachs E-mail:[email protected] GarcíaCatalánE-mail:[email protected] RibaE-mail:[email protected] ICREAResearchProfessor, InstitutCatalàdePaleoecologiaHumanaiEvolucióSocial,UniversitatRoviraVirgili (URV) cc64, 2placeEugèneBataillon,34095Montpellier, cedex5,France.J.Claude E-mail:[email protected] 5

2 tothe Iberian Peninsula. and Edifici C,CampusdelaUAB08193,Cerdanyoladel Vallès, Spain.O.OmsE-mail:[email protected] 1 Agustí E-mail:[email protected]@urv.cat BlainE-mail: [email protected] Agustí E-mail:[email protected]@urv.cat BlainE-mail: [email protected] Mammals. Chelonians. Amphibians.Pliocene.Konservat-Lagestätte. Maar. Carrer RamonLlull,15,17455CaldesdeMalavella,Spain.R.Rosillo E-mail:[email protected] H.-A. BLAIN Pelophylax José G.Abascal2,28006Madrid,Spain.J.Van derMadeE-mail:[email protected] DepartamentodePaleobiología,MuseoNacionalCienciasNaturales,CSIC García CatalánE-mail:[email protected] RibaE-mail:[email protected] 3

the CampdelsNinotsmaar(NESpain) 1 1 cf. DepartamentdeGeologia.UniversitatAutònomaBarcelona(UAB) InstitutCatalàdePaleoecologiaHumanaiEvolucióSocial(IPHES), 1 2 4 2 perezi Alephistigneresi F. BURJACHS

Taphonomicevidences ofthe skeletal remains indicates minimal (if any) weathering. ÀreadePrehistòria,UniversitatRoviraiVirgili (URV) ISE-M,UMR5554CNRS,UniversitédeMontpellier2 2 6 and freshwater fishes ( CentredeRecerquesArqueològiques Stephanorhinus jeanvireti 7

5 Apodemus atavus 1 , one ofone , S B A 2 J. CLAUDE Stephanorhinusjeanvireti 1 T C A R T Stephanorhinusjeanvireti Leuciscus [email protected] 2

6 , the amphibiansthe, cf. and [email protected] S. GARCÍA CATALÁN ?)complete the vertebrate assemblage uncovered Alephis tigneresi Mauremys leprosa . F. Burjachs E-mail: 3

and one ofoneand , Campeny E-mail: Tapirusarvernensis

1 Pleurodeles suggests an age of about 3.2Ma 2 have been recovered, some 4 D. RIBA [email protected] Tapirusarvernenis

sp., [email protected] 1 Lissotriton 2 or 5 R. ROSILLO Mauremys 1

2

, as , aff. 1 7

10 12 11 14 13 16 15 18 17 20 19 22 21 24 23 26 25 28 27 30 29 32 31 34 33 36 35 38 37 40 39 42 41 44 43 46 45 48 47 50 49 52 51 54 53 55

2 1 4 3 6 5 8 7 ARTICLE IN PRESS 9 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 . l a t e R E L O S E D Z E M Ó G . B ) 2 2 1 0 0 7 2 1 ( 0 0 7 0 1 0 - 0 1 . 5 , 0 ) 1 1 / ( 4 0 4 1 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G Catalan Coastal Ranges (Fig. 1A), and its morphology is morphology its and 1A), (Fig. the Ranges Coastal of Catalan rocks Paleozoic by bounded is basin This 1990). al., et (Pous Depression Selva La the in Girona of city the The site is located in the town of Caldes de Malavella near with singular shapes resembling little “puppets” (“ninots”). menilites of presence the from name its derives 2008) al., et Oms 2008; al., et Soler de Gómez 2007; al., et Catalán GEOLOGICAL SETTING its ageandpalaeoenvironmental setting. concerning information contribute to and fauna vertebrate the study of this new site, to provide information about the for obtained results palaeontological and geological first other vertebrates. The aim of this paper is to summarize the this of context project the yielded articulated skeletons of large mammals and in Ninots dels Camp at excavations Subsequent Peninsula. Iberian the of NE the archaeological in deposits and palaeontological fluvial the and volcanic,lacustrine Plio-Pleistocene of records of study the (Tarragona) Virgili i includes that project research Rovira multidisciplinary a initiated Universitat the of EvolucióSocial i Humana Paleoecologia de Català Institut the 2003 In maar. a as basin the interpreted first who (1999), al. et Vehí of prospections the during found were bones bovid was the which to bone, assigned fragmentary a was Ninots dels from Camp described fossil first The (1946). Sabarís Solé and was later followed by Bataller (1933), Llopis Lladó (1943) of lacustrine origin and Quaternary age. This interpretation locality this at sediments Vidalthe by considered who (1882) out carried were area this in studies first The Ninots. dels Camp of site maar Pliocene new the of time first the environmental changein termsofvegetation, climate,etc. found in such settings, provide detailed archives of palaeo- and macroscopic pollen (both remains). usually sediments, laminated of successions The flora and etc.) sizes (insects, several of vertebrates (fishes, amphibians, reptiles, mammals, etc.), invertebrates including found, is biota 2000). Typically,al., et whole the of largerepresentation a Mertz 1994; al., et Neuffer 1993; Büchel, 1977; (Franzen, respectively 45Ma, and 50Ma dated (Germany), Eckfeld in the case of the Eocene Fossil-Lagerstätten of Messel and as such fossils, preserved well exceptionally for potential great a have they Therefore, 2006). Kaulfuss, and (Lutz lakes meromictic as waters bottom anoxic have may and records. Such palaeo-environmental lakes are relatively deep, detailed protected from major erosion of preservation INTRODUCTION The palaeontological site of Camp dels Ninots (García Ninots dels Camp of site palaeontological The for account detailed a give we paper, present the In the for settings ideal are volcanos, in lakes Maars, Vcne 18) Several 1985). (Vicente, Leptobos FIGURE 1 Vehí et al.,2005). from (modified crater the of extension and area the from geology tailed De - B) Basin. Selva La LSB: Iberia. NE from units geological main the NW-SE. It is formed as a result of the distensivetectonics the of result a as formed NW-SE.is It oriented faults of sets two by controlled B C E a A P r

c T B e A R l L o O A n r Y o Geology of the Camp dels Ninots Volcano. A) Location within g S T c A g s C a a N B o a A r r r a l e d a l a a S l n u n m l v v v u d v d e e e I v e i s s a l l r i R n a s s a a

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B e y PALEO- PLIOCENE a a & 2 r n ZOIC y s e a 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 also present. Regarding clay minerals, montmorillonite is montmorillonite minerals, clay Regarding present. also sediment, quartz is this the dominant mineral, albite and microcline In being diatoms. benthic with clays laminated greenish by up built is subunit this (particularly detail, In leaves). remains tree plant of amount large findings, a including palaeontological most and skeletons mammal origin recent of deposits draping presentdaygeomorphology. wash slope of consists 4 unit Finally,industry. lithic of exclusively consisting remains, Quaternary age. In this unit, we found some archaeological of clays laminated reddish of 1meter around of consists 3 2.2, and 2.1 distinctive opal mineralizations subunitsare found (see Fig. 2). Unit In carbonates. no and sandstones isolated, includes 2.3 Subunit 2.4. and 2.2 2.1, subunits in found are ankerite) and dolomite include (that Carbonates broadly built up by greenish laminated clays with diatoms. is 2 Unit diatomites. and sandstones clays, greyish are 1 unit of sediments Lowermost 2). (Fig. maar the of parts units local four that canbepartiallyfoundinsmallertrenchesdugother observed we 4), to (1 top to base From Ninots. dels Camp the the for described section stratigraphic we reference There, 3). 2, (Figs. sector Argilera excavationCan the at metres 8 of one the is section -thick during several excavations, but the only available, relatively classified asasoft-substratemaarlake (Lorenz,2000). be can volcano this features, substrate maar.to According this of beds lacustrine the from come herein reported data as strata crater. the filling newsediments lake palaeontological The Ninots dels Camp the identified and described by identified was first who volcano (1999) etal. This Vehí 1B). (Fig. area latter this in found are volcano Ninots dels Camp the and 1976) Donville, 5Ma, at (dated units flood volcano Dalmai Sant de (Vilobí d’Onyar) Crosa or around the Caldes de in Malavella. Both as basalt such margins, basin the around all found are et Volcanicrocks 1999). Vehí al., in references and non-explosive summary (see and phases explosiveactivity both with cones volcanic and floods basaltic olivine-rich is of record volcanism a with Selva alkaline La The 1964). (Guardia, Iberian Peninsula the of NE the in BP years 10,000 to 14Ma from active was which 1983), al., et (Araña VolcanicComplex mainly during the Pliocene (Donville, 1976) as part of the Catalan place took activity Volcanic 2005). al., et (Vehí activitygeothermal marginsthe at Selvathe of Depression Neogene and thermalism shows that there is still important 1994). Tassone1999; al., al., et et Roca 1996; Roca, in presented is setting geotectonic (detailed Quaternary and the Neogene of most for Mediterranean western the affected that . l a t e R E L O S E D Z E M Ó G . B ) 2 2 1 0 0 7 2 1 ( 0 0 7 0 1 0 - 0 1 . 5 , 0 ) 1 1 / ( 4 0 4 1 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G uui 23 s h oe ht otis l articulated all contains that one the is 2.3 Subunit been studied has maar the of infill sedimentary The the during occurred episodes volcanic Several FIGURE 2 crater infillatCan Argilerasection(seelocationinFig. 3). at Camp dels Ninots would correspond to lithozone D in D lithozone to clays correspond would Ninots dels Camp at and silts studied materials the scheme, this laminated to According increase. while and debris decrease, rock wall pyroclasts top to base 2006; From al., others). et among Linder 2003; al., et (Pirrung lithozonation or succession stratigraphic vertical typical a in arranged right). 2, Fig. (see levels stratigraphic distinct three mammals in found are large of skeletons Articulated remains. fossil no with grains) piroclast of up built (basically sandstone fine-medium a by capped is unit the and sandstones and silts in richer are laminites upper be The unit. this to in present likely are laminae thousand a around that suggest fact that lamination may be difficult to discern, local counts the Despite found. be also can illite and one dominant the 3 4 2

opal 1 (mostly opal) Laterally extendedmineralization menilites greenish laminated clays with diatoms are sediments maar that observed generally is It DESCRIPTIVE reddish laminated slope wash and wittishcolor and diatomites mineralizations mineralizations greyish clays, UNITS light coloured sandstones interval with interval with sandstones interval with interval with and rounded Small (lessthan1cm) and elongated Large (morethan1cm) excavation code clays

small opal large opal laminites Stratigraphic succession and units of the Camp dels Ninots dels Camp the of units and succession Stratigraphic isolated 2.1 2.2 2.3 2.4 C4 C3 C2 C1 B A D 1m 2 0 3 4 7 8 5 6 hs ffmcvc c m f vf st sh dwkpkgrn pck wck md SECTION GENERAL Camp delsNinotsmaarsite(Spain) 10 cm vertebrate PALAEONTOLOGICAL

remains LOCATION OF carbonates sandstones and Whittish mudstones, and sandstones greyish mudstones Green andgreen- (mostly opal) Mineralizations and sandstones Greyish mudstones Reddish mudstones deformation Soft-sediment DETAILED

hs ffm f vf st sh LEVELS Isolated Articulated 3

10 12 11 14 13 16 15 18 17 20 19 22 21 24 23 26 25 28 27 30 29 32 31 34 33 36 35 38 37 40 39 42 41 44 43 46 45 48 47 50 49 52 51 54 53 55

2 1 4 3 6 5 8 7 ARTICLE IN PRESS 9 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10

ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1

. l a t e R E L O S E D Z E M Ó G . B METHODOLOGY OF EXCAVATION AND SAMPLING palustrine settingssuchaslasHigueruelas. in found is it as desiccation lake of evidence no observe we Additionally,2000). Lorenz, example, for (see setting proximal a in deposition indicate border, volcano the to the studied strata, together with the marginal position close the of infill lacustrine of volcano.lithozonation) stated (and features general The the of parts may deeper sediments in present such be that suggest (1999) al. et Vehí by data although 2), (Fig. site and section studied the for case the not is This 1992a). (Wuttke, sites palaeontological to anoxic under formed common conditions (such as sapropels) and are they may even be strata related settings lake Maar 2009). (Martín-Serrano, Higueruelas Las or 2005) Sachse, in references (see Eckfeld 2007), Micklich, and outstanding Some Gruber in references (see Messel literature. from those are examples the from well-known are flow deposits”. debris and turbidites some with clays and silt as “laminated defined was which (2003), al. et Pirrung of sense the the limits of the lake and the fossiliferous levels (Fig. 3). (Fig. levels fossiliferous the and lake the of limits the of field ownership, where we made different pits searching limits the by defined sector), Butano Can and Sector,Sector Cateura Pol Can Sector, Argilera (Can sectors four of parts small excavated have we date, present the to Up fieldwork. the of organization the for sectors several into divided was area This 1B). (Fig. respectively metres, 400 and 650 over of diameters minimum and maximum with

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I er AA D I 483.705 100m D I D BO I D I D Lissotriton ) cf. ?), (Leuciscus fishes cyprinid are: far so identified taxa The 4). (Fig. mammals and reptiles amphibians, fishes, freshwater of remains includes record PALAEONTOLOGY i Evolució Social at Tarragona. collections of the Institut Català de Paleoecologia Humana to theprotocoldeveloped byBurjachsetal.(2003). according to Girard and Renault-Miskovsky Beaulieu (1969), and as and acetolysis of elimination Goeury the by modified slightly by (1979), proposed by protocol Sector) the Butano using and Argilera Can Pol, (Can sectors residues. Pollen was extracted from samples from different at the from microfossils microscope pick to used was magnification binocular low A meshes. 2 millimeter with 0.5 sieves and superposed using samples, of sediment screen-washing the conventional the fossils by recovered Microvertebrate were analysis. pollen and vertebrate technical systems(López-Polínetal.,2009). leavesthe of some recovereddifferentwere with blocks in and remains others). Woodfruit among and 1996; Barron, 1988; plates Vicente, 1987; Siria, monographic de (Sanz classification case, its for this in or manuals, different using then and 1999) al., et (Ash description good first a with laboratory, the in identification taxonomical later a for field the in documented photographically were fossils made was cast a connexion, anatomical in (Gómez- Social Evolució preservedwell of remains case the In 2009). al., et Merino i Humana Català Paleoecologia Institut the de of laboratory the in excavation its for later protection polyurethane with blocks were in individually out out taken taken be articulated not could connection, that in elements found were skeletons When removal. their for impregnated were specimens damaged most The register). of number and square excavationunit, pit, sector, year, -CN-, initials by site the of (name code register their to corresponds that grid reference the using which were drawn and their position in space was recorded started with the delimitation and excavation of the remains, fossils the of recovery The (UTM). Mercator Transverse Universal the through positioned were occurrences fossil and Pits thick. 1m approximately are strata fossiliferous from 4m² to 100m² ranged for those pits size with fossil their evidences. The and meters 5 or 4 about was pits the of thenthe and fossil strata were abackhoe manually excavated. by The maximum depth opened first were pits These ers, lpi tigneresi , Alephis leprosa, p o o, h Cm dl Nnt vrert fossil vertebrate Ninots dels Camp the now, to Up the in stored provisionally are remains the All Sediment samples were taken from each level for small aff. helveticus, Pelophylax Camp delsNinotsmaarsite(Spain) tpaohns jeanvireti, Stephanorhinus cf. perezi, Pleurodeles Plant situ. in Mauremys sp., 4 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 . l a t e R E L O S E D Z E M Ó G . B ) 2 2 1 0 0 7 2 1 ( 0 0 7 0 1 0 - 0 1 . 5 , 0 ) 1 1 / ( 4 0 4 1 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G FIGURE 4 present because of a fault in the field. D) Skeleton of same fault in the field. E) Skeleton of ? inanatomicalconnection recoveredduring2005.H)Articulated frogskeletonPelophylaxcf. perezi recoveredduring2008. A to C) Skeletons of Skeletons C) to A recovered during A) 2005 B) 2004 and C) 2006; the posterior and anterior limbs in B and C are not are C and B in limbs anterior and posterior the 2006; C) and 2004 B) 2005 A) during recovered tigneresi Alephis Tapirus arvernensis recovered during the 2008. F) Stephanorhinus jeanvireti recovered during 2006; the anterior limbs are not present because of the Mauremys leprosa recovered during 2008. G) Newt Camp delsNinotsmaarsite(Spain)

Lissotriton 5

10 12 11 14 13 16 15 18 17 20 19 22 21 24 23 26 25 28 27 30 29 32 31 34 33 36 35 38 37 40 39 42 41 44 43 46 45 48 47 50 49 52 51 54 53 55

2 1 4 3 6 5 8 7 ARTICLE IN PRESS 9 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 and fruitimpressionswerefound(Fig.5E,F). lauroid with flora wood some addition, In 5). (Fig. of evergreentrees of leaves type subtropical a indicate clays, lacustrine in plant impressions as large preserved are hand, which fossils, other the On microzooremains). and spores fungi protists, Spirogyra, Botryococcus, Rivularia, Typha-Sparganiumvariousand taxa; NPP’s (Gloeotrichia, Cistaceae, Quercus, Olea/Phillyrea, Arbutus, Erica, Calluna, Ephedra whereas on the posterior half there are two or three smaller prezygapophyses, of basis the at vertebra the of side each on view,opens lateral foramen largeIn a Spain). (Madrid, Naturales Ciencias de Nacional Museo the at collection reptile and amphibian the from coming waltl Pleurodeles characteristic has been seen on some modern specimens of a Such (1997). Haller-Probst and (2003) Bailon and Ruiz Barroso- and by described as process process costal ventral costal the to not dorsal the of linked base the are to prezygapophyses posteriorly the material fossil arch. neural our vaulted On highly a and spine neural thin and high long, a Vertebraewith elongated, opisthocoelous, are sented by a few presacral vertebrae (centrum length >4mm). Argilera Sector. Pit7/8.K24’. Caudata Amphibia Pinus,Picea, Abies of presence the determine to possible been has it Nevertheless, microremains. abundant provided not Until now, palynological analysis of all excavated areas has macroremains. and palynology microremains, on focusing and silicaprecipitation. dissolution calcite to leading 2003), al., et Marini 2000, al., et (Varekamp waters basic slightly to acid generally have lakes Maar dissolution. selective as such processes abiotic sediments by caused bias lake taphonomical strong a in reflect may fact unusual intensive relatively and This sieving. surface excavated significant the despite found was bivalvesetc.) ostracods, (gastropods, organism few isolated materials found in C1 and D units (see Fig. 2). a for except C2 unit in localized been have remains fossil the cyprinids. All of forms three or two least at recognize to allowed data preliminar study, but under still are fishes and arvernenis Tapirus . l a t e R E L O S E D Z E M Ó G . B ) 2 2 1 0 0 7 2 1 ( 0 0 7 0 1 0 - 0 1 . 5 , 0 ) 1 1 / ( 4 0 4 1 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G ecito ad oprsn A ag nw i repre- is newt large A comparison. and Description Can “CN”08. Code: vertebrae. isolated 5 Material. cf. Pleurodeles sp. primarily underway, are studies Palaeobotanical invertebrate of shell single a not that remarkable is It Artemisia, Asphodelus, Poaceae, Cyperaceae and Cupressaceae,, Freshwater atavus. Apodemus Betula, Corylus/Myrica, Taxodiaceae, , the presence of abundant aquatic vegetation aquatic abundant with of generally presence the waters, still and shallow prefers species the period, aquatic its During 2004). al., et (García-París well represented in areas with a humid and moderate climate Lissotriton. to attributed vertebrae isolated the in with concordant be to seems (reversed vertebrae presacral its of premaxillae morphology The 2006). fused possibly Pleurodeles, and frontosquamosal a arch family of presence the the of with characteristics Salamandridae derived some shows and 4G) (Fig. view latero-dorsal in exposed is 6centimeters) of Béon1inFrance(RageandBailon,2005). knownis which helveticus, Miocene early late the since L. newt palmate extant the of that with well matches fossils foramina are large. The general morphology and size of the subcentral The 0.47. around is height height/vertebra arch neural spine and forked a well vaulted posteriorly neural and arch. The thin ratio neural high, long, a with elongated, genus the of characteristic are but found, was unfortunately skeleton it could not be preserved (Fig. excavation4G). articulated Vertebrae 2005 quasi-complete the a During sediment. the of washing about 2mm),afemurandhumerusrecovered by sieving- Camp dels Ninots by 4 presacral vertebrae (centrum length to referred 2. C2unit.R46’. Sector. Pit7/8.C2unit’. possibly anarticulatedspecimen. point ispermanent,itremainsinthewater allyearlong. water a if and aquatic mainly newtis This currents. strong or moderate with and streams avoids generally and 2.5 meters 10 between comprised depth a with environments 2004). al., et (García-París climate dry and warm under mainly Current a northernmost extension of the genus during the Pliocene. The presence de of Rambla of Pliocene LowerValdecebro III, Teruel, Spain (Sanchíz, 1977; or Estes, 1981). Miocene Upper the genus the record, fossil the genus to attribution an with concordant is morphology vertebrae Fossil foramina. postero-lateral h plae newt palmate The around length (total skeleton articulated The newt small-sized A comparison. and Description Pit Sector. Butano “CN”07. Code: material. Isolated - Argilera Can “CN”05. Code: specimen. Articulated - and femur,humerus, a vertebrae,a isolated 4 Material. Lissotriton aff. helveticus(Razoumosky, 1789) preferently frequents large water large frequents preferently Pleurodeleswaltl ie i a ra vrey f habitats of variety great a in lives Pleurodeles aff. Lissotriton and Tylototriton Pleurodeles in Camp dels Ninots suggests s currently is helveticus Lissotriton Camp delsNinotsmaarsite(Spain) helveticus s nw since known is Pleurodeles : opisthocoelous, Lissotriton: Fot t al., et Frost Chioglossa; s ersne in represented is In Pleurodeles. and a dense a and 6 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 . l a t e R E L O S E D Z E M Ó G . B y ivn-ahn o te eiet dcmn the document sediment, genus the the of representative a of presence of sieving-washing by recovered bones isolated few a as well as to 4H) (Fig. 2008 2005 of excavations the during recovered specimens 2. C2unit.S46’. Argilera Can “CN”08. Sector. Pit7/8.C2unit.M23,32’. 1’, J26, unit. C2 Argilera 7/8. Can Pit “CN”08. Sector. 1’, I18, unit. C2 7/8. Pit ArgileraSector. Can “CN”05. 1’, N23, unit. C2 7/8. Pit Sector. humerus andamandible. Anura and ispresentinthewater fromNovember toMay. newts Iberian the of aquatic less the is It millimeters. 700 found is today in areas with newta mean annual precipitation higher than this Catalonia, In cover. vegetal lakeside ) 2 2 1 0 0 7 2 1 ( 0 0 7 0 1 0 - 0 1 . 5 , 0 ) 1 1 / ( 4 0 4 1 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G FIGURE 5 ecito ad oprsn Svrl articulated Several comparison. and Description Pit Sector. Butano “CN”06. Code: material. -Isolated ArgileraCan “CN”05. Codes: specimens. isolated -Articulated an and specimens articulated 4 Material. Pelophylax cf.perezi (Seoane,1885) A andB)Surfacewith plantsfossils(lauroidleaves).C) Quecus drymeja.D)Daphnogenepolymorpha . E)woodimprint.F)Alnusfruit. . On Pelophylax. l, 95. oee, h ha wdhsotvn length to width/snout-vent closer is ratio head the However, 1995). al., et (Crochet vomers the of position relative the and ratio length length/snout-vent tibia the ratio, length snout-vent Pelophylaxperezi establish a link between these fossils and the extant species than and genus 130°) in shaft (approximately ilial apophysis sacral the open on more crest dorsal higher recognized: es ios a rpeet h eris ctto fr this for citation earliest the represent may Ninots dels Camp of Pliocene the in occurrence its Therefore, 2005). bones at least since the late Pliocene (Sanchíz, 1998; Blain, fragmentary and isolated from doubt with mentioned is it representativewhere Peninsula, Iberian the in genus the of only the currently is (P.perezi) frog marsh southern The fossilization. during compression little a of artifact an be ern. etrs f h genus the of Features bearing. teeth and maxilla premaxilla omosternum, ossified sternum with firmisternous ribs, of absence lacking, the urostyle of processes transverse articulation, sacro-urostylar urostyle that bears cylindrical sacral apophysis, bicondylar the with unfused sacrum arch, neural non-imbricate and short with column, vertebral diplasiocoelous observed: be familythe of characteristics specimens, can articulated the Sm dsrt caatrsis emt to permit characteristics discrete Some Rana. than P.ridibunda (Seoane, 1885), as the fourth toe length/ toe fourth the as 1885), (Seoane, Camp delsNinotsmaarsite(Spain) Pelophylax but this may this butP. perezi

a as be also can 7

10 12 11 14 13 16 15 18 17 20 19 22 21 24 23 26 25 28 27 30 29 32 31 34 33 36 35 38 37 40 39 42 41 44 43 46 45 48 47 50 49 52 51 54 53 55

2 1 4 3 6 5 8 7 ARTICLE IN PRESS 9 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 . l a t e R E L O S E D Z E M Ó G . B ) 2 2 1 0 0 7 2 1 ( 0 0 7 0 1 0 - 0 1 . 5 , 0 ) 1 1 / ( 4 0 4 1 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G than in cervical are elongated characters rather the that and are scute, found anal more than often humeral shorter in Ninots, the dels Camp anteriorlyvertebral In wide. is very fourthandvertebral scuteratherare rectangular firsttheand gaudryi,M. are diagnosticare genustheof pectoral and gular scutes, and the weakly developedrathertheshort bridge, entoplastron the buttresses partially covered by cases, irregular somein scutes, neural anal short series, the and, wide rather the scute, marginal second thereaching pentagonalfourth neural plate. Thewidefirst vertebral scute specimenoneexhibits heptagonal a thirdneural a plateand predominantlyandwideare anteriorly shortsided,however Neurals 2004). Tong, and (Claude family geoemydid the seventhperipheral plates andallowassign third tothesespecimens into ducts musk and plate, pygal long thanwider posteriormargin. bonearrangement,The analnotch,thethe smooth a with rather flat, and shape oval in shells areThe specimenbeing a subadult, while smallerlarger the 25cm,specimens to are 12 adults. from vary lengthsestimated shell sub-connected. individualsor five least present. The are At appendicular and skull bones. The material is often connectedrecorded by partial and more complete shells, as wellcalled Mediterranean as axial, speciesTurtle is PondThis 4F). (Fig. speciesthe referred to be chelonians can that by Ninots dels Camp at Pit 7/8,C2unit,N25,3; T23, 1;M22,2-6’. Sector, Pit7/8,C2unit, L24,1’ (Fig.4F). Argilera Can “CN”05, 1’, M22, unit, C2 7/8, Pit Sector, isolated material: Testudines and Arano, 1997;García-París etal.,2004). (Llorente water permanent of absence the by limited be to seems only which of presence the environments, aquatic Mauremysleprosa, sulcusand by the shape of its vertebral scute. Asin the living newspecimens found bytheposition ofthehumero-pectoral Mauremysportisi therefore,it differs from subadults of smallerindividual (subadult) doesnotshow lateral keels and Ninotsalsodifferfrom Pelophylax the Straight of Gibraltar opened (García-París et al., 2004). supposed to have split off from data, molecular to according although, species, ecito ad oprsn Rpie ae represented are Reptiles comparison. and Description -Isolated material. Code: “CN”05, Can Argilera Sector, Argilera Can “CN”05, Codes: skeletons. -Articulated Material. Two complete articulated skeletons and some Mauremys leprosa (Schweigger, 1812) M.

caspica Mauremys leprosa perezi gularsslightlyarepinched posteriorly, thirdthe fromthePliocene Italyof differs fromthe or

M s seis ht ie i ms sunny most in lives that species a is M. rivulata . rivulata, M. rivulataM. Mauremys and (Schweigger, so- the 1812), . Specimens from Camp dels P. saharica at 5.4Ma, when by theirlargerbysize. The M.caspica M.caspica (Claudeal.,2007).et and and P.perezi is M. leprosa

thefossil rivulata water streams,ponds,lakes oroxbows. living andthe shorter marginals and is therefore more similar to the gaudryi material of type the differs from Spainspecimen from intraspecific than smaller variation. fossil However, be the Perpignanveryissimilar to recognized on the basis of morphology and metrics. In a In metrics. and morphology of basis the on be recognized can which taxa than names more probably are There 2003). al., et Radulescu 1929; Piveteau, and Arambourg 1991; al., et Michaux 1984; Morales, 1981; (Gromolard, Europe Eastern from two and Westernfrom named were genus species the transferred the and Bovini to basal related closely be to genera both believed (2006) al. et Montoya but opinion, similar a had (1992) Geraads Boselaphini. the to latter the and Bovini the to belong to placed was which within “cordieri”then , until species the for Alephis Piveteau (1929), while Gromolard (1981) named the genus Parabos andAlephis. genera the of those to similar are Ninots dels Camp from Leptobos, or larger. All characters studied in the specimens of ranges upper the in are bones limb The Bison. and Bos than columns Leptobos, interlobular smaller have molars the and in directed backwards and are are not theymuch curved. keel, This is unlike anterior an and section oval an have They orbit. the behind little a originate cores horn The Europe. from known bovid species non-bovine any single than a larger bovid, of of 4A-C) (Figs. found were skeletons M25, 1-38;N23,1-7;N24,1-44;N25,1-27’ (Fig.4A). 1-35; M24, 1-31; M23, unit, C2 7/8, Pit Sector, Argilera 1-8; O48,1-3’ (Fig4C). “CN”06, 4B), N47, 1-16; P48, 1-21; O47, ‘(Fig. unit, C2 2, Pit Sector, Butano 1-12 L21, 1-45; K20, 1-17; M20, 1-44; L20, 2; and 1 K19, unit, C2 7 /8, Sector,ArgileraPit and Sudre,1991 but in Artiodactyla Mammalia and Bos Parabos,

h genus The articulated Three comparison. and Description Can “CN”05, Code: skeleton. articulated -Complete Can “CN”04, Codes: skeletons. articulated -Partial Material. Three articulatedskeletons: Alephis tigneresi Alephis resembles . leprosa.M. . In addition to the two genera, five species five genera, two the to addition In Alephis. by itsnarrowerby first vertebral, the narrower cervical, Bos . The cheek teeth are smaller high crown high smaller are teeth cheek The Bison. and Alephis M. leprosaM. and a nmd y rmor and Arambourg by named was Parabos , and claimed the former genus former the claimed and Parabos, Today, . The limb bones are robust, like robust, are bones limb The Bison. Michaux, Aguilar, Calvet, Duvernois Calvet,Michaux, Aguilar, . , but not as robust as in as robust as not but Leptobos, M. Mauremys leprosa M. leprosaM. Camp delsNinotsmaarsite(Spain)

gaudryi from the Pliocenethefromof and differencesand may to the P.to cordieri ie i calm inlives M. 8 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 humidity, temperatureandseasonality. variablesbeing important the with kind, some of optimum climatic a reflects probably most Neogene. This whole the of Tapiridae the of southernmost the of one and Europe of record southernmost the is Ninots tapirs favorableof climate. and habitats dels Camp at record The of distribution distribution the reflecting presumably time, over geographic changed and abundance The Stefanovic,2006). and (Made year the of part during food increased of because Europe seasonality, which limited the in availability of their preferred believed extinct is went It they forests. that humid to adapted are and and leaves fruits principally eat Tapirs similar. been have may tapir,livingMalayan Eisenmann (1994). and Guérin and Fejfar(1964) by described as 1828 Jobert, species the fits metrically specimen from Camp dels Ninots fits morphologically and and canines. These are all characteristics of characteristics all are These canines. and incisors of set full a and premolars, molarized teeth, cheek robust three and metatarsals manus in the the pes, in high placed metacarpals reduced nasals, robust lophodont with four structure foot mesaxonic presents: 4E) (Fig. found dpain o pn adcps But landscapes. open to adaptation 1991. tigneresi of material type morphology the to similar such particularly is has and Ninots dels Camp from material The more. out flare which cores, horn flattened more and longer has material younger geologically the sense, broad G20, 1-53,H20,1-44’ (Fig.4E). 1-3; F19, unit, C2 7/8, Pit Sector, Argilera Can “CN”08, Perissodactyla that werenotvery dryoropen. landscapes to adapted were existed they that Bovinisuggests elsewhere, evolved more when Europe in they persisted that well fact the of and lack grazing, to the adaptations developed but change, environmental to related probably is record fossil the in appearance to Their grazing. adaptations other lacked and crowned high less were . l a t e R E L O S E D Z E M Ó G . B ) 2 2 1 0 0 7 2 1 ( 0 0 7 0 1 0 - 0 1 . 5 , 0 ) 1 1 / ( 4 0 4 1 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G aeil Pril riuae seeo. oe “CN”06, Code: skeleton. articulated Partial Material. Stephanorhinus cf.jeanvireti (Guérin,1972) Tapirus arvernensis Tapirus oii ed o e pcaie gaes wih s an is which grazers, specialised be to tend Bovini ecito ad oprsn Te complete The comparison. and Description Code: skeleton. articulated complete one Material. Tapirus arvernensisCroizetandJobert,1828 Michaux, Aguilar, Calvet, Duvernois and Sudre, and Duvernois Calvet, Aguilar,Michaux, and thus, its ecology Tapirusits thus, and indicus is probably closely related to the to related closely probably is Tapirusand Croizet arvernensis and Parabos T.SW in arvernensis TapirusThe . skeleton Alephis Alephis al., 1991;Lordkipanidze etal.,2007). (Pasquier, Hungary et 1990; Aguilar Storch, and Fejfar 1990; and Bachelet, 1974; Germany France, throughout Georgia, to Spain Southern from Europe, of early Pleistocene and Pliocene the in distributed widely species a is species. this of population archaic an to belongs it that suggests M1 upper the of stephanodonty of degree low The t9. the from separated well small, is t12 The t4. the of wall posterior the to connected being shape, ovaland largein is t7 The established. well and large is t9 spur.evidenceposterior of and t6 between connection The any without rounded, is t3 The t3. and t2 to respect with presents a large and rounded t1, which is placed posteriorly oas f h murid the of molars scarcely represented at Camp dels Ninots by two first upper Code: “CN”08,Can Argilera Sector, Pit7/8,C2unit’. Sector, Pit7/8,C2unit,K20’. without teeth. Rodentia suggests that it preferred the same habitats that as suggestsjeanviretitapirs. crowns the on cementum Occiput shape, the low crowned overhangingteeth have occiputs (Zeuner, 1934). and to tend grazersthe lack of abundant aremorebrachydont (theremaining species). additionIn the grazers ( on materialfrom Vialette (Guérin,1980;table115). Stephanorhinus jeanvireti (Guérin, 1972), which is defined and than larger species, large a to belongs Ninots dels Camp from material The 1993). al., in them placed these species b) in 1982a, (1980, Guérin incisors. the of vestiges Europe, but not to the species “megarhinus”, which retains to belongs it of species the of one that suggesting incisors, reduced or of large indication any show not The does rhinoceros. symphysis anterior a mandibular of without presence the but indicates mandible, bones limb and skull partial with 1-7; M46,1-11,N46,1-4’ (Fig.4D). N45, 1-50; M45, 1-5; L45, unit, C2 2, Pit Sector, Butano ecito ad oprsn Sal aml are mammals Small comparison. and Description teeth. without maxilla a and (M1) molar upper -First Can Argilera “CN”04, Code: (M1). molar upper -First amaxilla and (M1) molars upper first Two Material. Apodemus atavusHeller, 1936 Present day rhinoceroses are either grazers or browsers. The comparison. and Description , and similar in size to the species the to size in similar and miguelcrusafonti, S. Ceratotherium simum (Heissig, 1996, 1999; Fortelius et Fortelius 1999; 1996, (Heissig, Stephanorhinus wasbrowser.a Itsknown geographical distribution Dicerorhinus, but it is current usage to place that are known from known are that Stephanorhinus Te pe M1 upper The atavus. Apodemus ) are hypsodont and the browsers Camp delsNinotsmaarsite(Spain) tpaohns etruscus Stephanorhinus n riuae skeleton articulated An

Apodemus atavus Apodemus Stephanorhinus 9

10 12 11 14 13 16 15 18 17 20 19 22 21 24 23 26 25 28 27 30 29 32 31 34 33 36 35 38 37 40 39 42 41 44 43 46 45 48 47 50 49 52 51 54 53 55

2 1 4 3 6 5 8 7 ARTICLE IN PRESS 9 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 . l a t e R E L O S E D Z E M Ó G . B ) 2 2 1 0 0 7 2 1 ( 0 0 7 0 1 0 - 0 1 . 5 , 0 ) 1 1 / ( 4 0 4 1 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G (or Equus (2003; table 1), Malusteni, Capeni and Iaras (Iaras 3?) have al. et 1992). Radulesco al., to et According (Bruijn omited were placed localities these were later while Beresti 1975), (Mein, MN16 and in Malusteni tables MN the In earlier 1). (Iaras boundary Gilbert-Gauss the above just MN15 (Malusteni, Capeni, Varghis) and the base of MN16 Age and correlation DISCUSSION However, by replaced is also including apparently and marks quotation the distribution oftaxa. the shows dels 6 Figure Camp Ninots. of age the least discuss are at can we or before which discussed mentioned be to localities, need and these here, relevance from of taxa of stratigraphic or distribution temporal accepted the and localities of positions units. accepted MN the between the contradictions of are There boundaries the for ages precise very gave (2001) al. et Agustí transitions, gradual of idea the to Contrary gradual. are MN units between transitions the that ended stated is they it though if even as boundaries, MN at given abruptly latter, generally the are In genera 1992). of al., ranges reference et the (Bruijn of given are lists localities faunal and distribution genera, unit, important MN of each of localities tables principal publication, with as later a well In as 1975). (Mein, occurrences taxa last typical and first the and localities principal the unit, MN each for provided, that chart a of reference the to fauna. fauna their of resemblance overall the of basis the on unit MN fossil an in other placed be should and localities locality reference a by but boundaries, their by defined not are units MN but taxon, specified a of occurrence last or first the by defined are that boundaries have bio-zones Normal 2001). al., the et Agustí 1992; al., et for mammals, on European and North African Neogene (Mein, 1975; Bruijn based scale, biostratigraphical with so thatthereisnofundamentforMN16age. of mixing, problems while in Sofia with only locality a is Kislang MN16. in placed and Kislang Gromolard 2003). cited (1981) al., et Radulesco 1981; (Gromolard, (1992), and al. et Bruijn MN15 in placed is which 1990), Soria, and (with “Parabos” (1992) al. et Bruijn to According a form units (MN) Neogene mammals The Radulesco et al. (2003) placed the Romanian localities Romanian the placed (2003) al. et Radulesco “Parabos”athanasiui The original publication on MN units consisted consisted units MN on publication original The ), while the latter the while stenonis), Plesippus/Allohippus s led ctd rm an (Pérez Layna from cited already is Leptobos Parabos and from Sofia, a possible a Sofia, Parabosfrom from Capenis, Virghis and Iaras, all Iaras, and VirghisCapenis, from Alephis t h M1-6 transition. MN15-16 the at Leptobos Alephis are still cited from MN16 Parabos and in the top of MN14 (Beresti), MN14 of top the in ages of relevant localities and relevantlocalities of ages Anancus are cited, from Alephis Alephis) ée ad oi, 90 ad ilry (N6 Villalta, (MN16; Villaroya1952; Agustí andOms,2001). and 1990) Soria, and Pérez (MN15; Layna of sites known well the between situated therefore is Ninots dels Camp of record extraordinary the largePliocene succession, contextSpanish mammal the of the In 2.5Ma. around dated while was transition 3.2Ma, MN16-17 the around events, Mammoth between and boundary Kaena MN15-16 the the situated (2001) al. et Agustí later. slightly or palced transition, be MN15-16 the should around Ninots dels Camp that suggests This indications. age precise less give species of other the while appearance of the last appearance and first MN16, the in possibly between or MN15 Parabos/Alephis, comprised thus is or than jeanviretirather Stephanorhinus , in MN16(Bruijnetal.,1992). placed generally is that locality (Fejfar,a Hajnácka 1964), of locality the in present also is It 2006). Stefanovic, and which belong probably to MN13 (Made et al., 2006; Made localities 1996), Heissig, 1980; (Guérin, (=Autrey) Gray 1996), this species is found in Baccinello V3 and Arc-près- Heissig, 1982a; 1980, (Guérin, MN14-15 be to considered lcd in placed of range the Although Middle earliest 1999). Heissig, after here assignation (generic Pleistocene the till Villafranchian earliest the from and MN16 or Villafranchianearliest the from Ruscinian or faunal unit MN15, late while the from known is miguelcrusafonti Stephanorhinus have overlapped. may ranges locality,temporal one their in neverfound are later of than co-occurrence the suggest Parabos lists (or faunal the that and bovids these replaced Westernin Europe which Leptobos, problem this has localities these solve of none that to note should we butnow, impossible is It 2010). Mateos, and (Made Europe Western in than Europe SE in earlier considerably appearing mammals of examples are there Quaternary,and Neogene the of all during hand, other the of range stratigraphic the Europe, of parts other in those to similar so that the first appearances of MN17 MN17, in localities three these placed we if and hand, one the 17 On 2001). or al., et Agustí MN16 1992; al., et (Bruijn in respectively appeared parts they other in Europe while of MN16, and MN15 in Romania in This would imply that imply would This . 5, neither 556, p. locality has also . The age of Camp dels Ninots dels Camp of age The megarhinus. Dihoplus? h rioeo fo Cm dl Nnt i probably is Ninots dels Camp from rhinoceros The codn t Géi (90 18a b te species the b) 1982a, (1980; Guérin to According would be extended upward. On upward. extended be Alephis/Paraboswould Leptobos. Though or Dicerorhinus ) with taxa that appeared considerably appeared that taxa with Alephis) Equus Eucladoceros (though in the text on Iaras, n MN16, in jeanvireti Stephanorhinus nor Dihoplus? and Equus r mentioned). are Eucladoceros Parabos/Alephis and Camp delsNinotsmaarsite(Spain) i generally is Stephanorhinus) Equus and (previously megarhinus appeared Eucladoceros S. jeanvireti S. miguelcrusafonti S. Eucladoceros are S. etruscus S. Leptobos is known 10 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 Palaeobiogeography last phase. this to belongs Ninots dels Camp from record Stefanovic, The 2006). and (Made ago 2.5Ma around extinct going Late the till and ago again4Ma peaking and Pliocene, 3 some between lasted species This appeared. arvernensis species the when abundance, in peak new a was there Miocene, latest the During 1988). Gómez-Alba, of known from Spain from Camallera II (Tapirus arvernensis Late tapir small Miocene rare the but Europe, in extinct went Tapirus VallesianCrisis, the During 2006). Stefanovic, and Made II Ponsic Can (Vallés and 1982; Pairo, Crusafont and (Golpe-Posse Penedés) Llobateres Can and Urgell) de (Seu species the where Spain, reaching abundantwidespread, more and were tapirs time, this At strong. very not was and seasonality widespread were environments closed humid when the Vallesianand time latest Aragonian a The 2006). were largeand middle four Stefanovic,and (Made species sized to three by represented is it where Europe, into Miocene genus The probably originated “tapir-vacuum”). in Asia the and dispersed during called the Middle is time America–this from as well (as Europe from Tapiridaedisappeared transition Miocene Early–Middle 1988). the Ginsburg, Around and (Cerdeño (Cuenca) and Valquemado 7A) Fig. (Zaragoza; Aragón are de they Cetina from Spain, known in and Miocene earliest and Oligocene the reflecting distribution oftheirfavored habitat. presumably greatly, fluctuated has space and time in abundance and distribution limited seasonality.Their of degree and low a with adapted environments closed are and humid to tapirs toes, lateral large with legged short being and folivores and frugivores Being 7). (Fig. world the of part this Tapiridaethe in of distribution of limits the to close chronologically and geographically habitats. were different. Probably, bovids tolerated more open habitats or dry optimal their that suggests This Italy). central in the while and the concentrations of the localities differ (in SE France, Turkey) and Greece Italy, southern Spain, southern wider (e.g., be to seems distribution geographic its However, of that to comparable is group this of occurrences of number remains). and distribution poor temporal with The localities some of exclusion the to lead might revision–revision (without literature the from trace other could we with occurrences all compared shows we 7B Figure if groups. record fossil the in rare be . l a t e R E L O S E D Z E M Ó G . B ) 2 2 1 0 0 7 2 1 ( 0 0 7 0 1 0 - 0 1 . 5 , 0 ) 1 1 / ( 4 0 4 1 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G ois f h tp of type the of Bovids the during Europe in present already were Tapiridae The record in Camp dels Ninots is interestingly in both is found in the localities El Firal El localities the in found is priscus Tapirus oaiis r concentrated are localities arvernensis Tapirus perd wih is which appeared, pannonicus Tapiriscus and Parabos aiu arvernensis . Tapirus em to seems Alephis Tapirus Tapirus the geographical distribution of distribution geographical the Unlike Spain. into range known the extend finds new The localities. few but from known is (without species literature The revision). the in cited occurrences other well the as as localities these shows 7C Figure Italy. northern in seven and France eastern and central in jeanviretifour , Mediterranean duringthePliocene. the of north the to species this of range the considerably from Perpignan, the record from Camp dels Ninots extends from distinct as species this considers Pleistocene. one the If in was Europe in species this of oldest record the now until but 2000), Broin, de Lapparent (de Algeria of (Ruscinian) Pliocene early the from known is It Africa. Northern to Peninsula Iberian the from extends may have preferredasimilarhabitat. it that suggesting tapirs, of distribution the of ranges the knowndistribution of FIGURE 6 graphic rangeasa resultofthegivenoccurrences. strati- the highlight lines solid The assignments. correct to or species synonymize to made was attempt No squares. open by indicated erally gen - are bone or tooth a of fragment a just on based (1981) Gromolard indicate a certain degree of doubt (“cf.”, “aff.” or “?”). Assignments by squares open and presence indicate squares Solid left. the on 1992) al., et Bruijn 1975; (Mein, units MN and (Ma) years of millions in Age 2006). Stefanovic, and Made 1984; Morales, 1997; Mazo, 2003; al., Tapirus (largely based on Gromolard, 1981; Guérin, 1980; Radulescu et earliest the and Guérin (1980) listed 13 localities with localities 13 listed (1980) Guérin The current geographical range of range geographical current The h tmoa dsrbto o te Bovidae the of distribution temporal The , selected Villafranchian Rhinocerotidae and Rhinocerotidae Villafranchian selected Leptobos, S. jeanviretiS. extendbeyondnot does Camp delsNinotsmaarsite(Spain) Parabosand Mauremys leprosa Mauremys Stephanorhinus Parabos,

, the Alephis, M. gaudryi M.

Alephis 11

10 12 11 14 13 16 15 18 17 20 19 22 21 24 23 26 25 28 27 30 29 32 31 34 33 36 35 38 37 40 39 42 41 44 43 46 45 48 47 50 49 52 51 54 53 55

2 1 4 3 6 5 8 7 ARTICLE IN PRESS 9 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 . l a t e R E L O S E D Z E M Ó G . B ) 2 2 1 0 0 7 2 1 ( 0 0 7 0 1 0 - 0 1 . 5 , 0 ) 1 1 / ( 4 0 4 1 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G Taphonomy uui 23 itra wt ioae snsoe) I i yet is levelsingle a to belong (what fossils such whether unclear It sandstones). isolated with (interval 2.3 subunit particular in and diatoms) with clays laminated (greenish 2 unit from are fossils other as well as skeletons complete different levels of the sedimentary succession, though most from and 3) suchas Fig. (see Sector Butano the areas and Can Argilera distant from are finds the and rule, the be (and fossil record in general) at Camp dels Ninots seems to on taphonomicandpreservation processes. hypotheses preliminary assist can observations geological and palaeontological some but Ninots, dels Camp at out Horizontal scattering of both large and small vertebrates carried been yet not have studies taphonomic Detailed FIGURE 7 A) Pietro, Pradalbino; 73)NovaVieska. San Monte 72) Capannoli; Valdarno, Lower 71) (Sj); Sienna Se), (Sj, 68) Desnes–Vincent; 67) Hambach; mais; 62) Karaboroun; 63) Esme-Manissa; 64) Layna; 65) Ptole- Schelde; 66) 61) Lozenets; 60) Beresti; 59) Bessarabia; South 58) Kislang; 53) Pont-de-Gail; 54) Montecarlo; 55) Val di Pugna; 56) Gravitelli; 57) Collonges; de Túnel 52) Jassans-Riottier; 51) Laurent-des-Arbres; St. 50) Puimasson; 49) Celleneuve; 48) Palais; Saint Ille-sur-Têt;47) 46) Moro; del Venta 45) Júcar; del Alcalá 44) Alcoy; 43) Rey; 42) del Arenas (P); Vallay P), (Tp, Gray 41) Camallera; 40) I; Ponsic Lloba- Can Can teres, 39) Firal; El 38) Valquemado; 37) Aragón; de Cetina 36) Ninots; dels Camp 35) Musselievo; 34) Sülzheim; 33) Stavropol; 32) (T,(T,1 Malusteni; Ilieni Iaras 31) A), 30) A); Capeni; 29) (T); Fagului Araci-Fântâna ), A (T,Varghis 28) Hrabarsko; 27) Zivojno; 26) Djonai; 25) Sütö; 24) Hajnácka; 23) Ivanovce; 22) Brebir; 21) Sostanje; 20) Spoleto; 19) Pietro; San Castel Montoro, Nera 18) Monticchiello; 17) V3; Baccinello 16) Barbara; Santa Gavile, Valdarno: Upper 15) sino; Vignola;12) Fosciata; Ca- Pieve 14) Glosina; di Sasso Livergnana, 13) Barga, Garfagnana: 11) Ponzano-Magra; Sarzanello, Magra: Valdi 10) Coupet (T), Vialette (T, Sj); 7) Montpellier; 8) Perpignan; 9) Villafranca; Trévoux5) Parabos); (T, (T,Étouaires 6) T); ( Merle Mont D), Le Se), Sj, (Tapirus, Autrey (Tapirus ), Changny 4) Wölfersheim; 3) Maalbeek; 2) Crag; Red 1) Legend: 1980). Guérin, (after text the in mentioned oses rhinocer other with localities of selection a Hooijer,and 1994; 1981) and Mors, 2008; Holec, 1996; Bianucci et al., 2001; Campanino et al., Stephanorhinus jeanvireti (after Guérin, 1980; additions from Lacombat cu et al., 2003; Spassov, 2005; Cuscani Politi, 1979; Dallan, 1988). C) Alephis (afterGromolard,1981;Morales,1984;Mazo,1997;Radules- B) 2005). Spassov, 1980; Guérin, from: additions 2006; with locality youngest sites suchasthatofMessel(Wuttke, of 1992a,b). absence the and tissues bioturbation. This setting resembles that soft of other maar lake of fossilization the by evidenced depleted as hypolimnion), oxygen with stratified, in (being waters place taken have to likely periods) sites such as Messel (Wuttke, 1992a). the lake. This type of occurrence is also found in other maar ofbottom thescavenged, undamaged nottosank werebut bottomofthe lake soon after death. Large mammal the corpses to sank mammals large that indicate to seems above, This,together with the sedimentological observations made arrangement.parallel a in and side their on lyingposition to follow the same pattern: limbs are in a completely indicating relaxingminimal 1978), (if (Behrensmeyer,any) weathering. 0” In all cases, stage body “weatheringposition seems fine out corpsetransportbycurrents. within ruling mudstones), laminated found (mostly sediments are grained skeletons Large in carcass completeness. role significant a have to (slumping, seem etc.) processes faulting, post-depositional but connexion, anatomical full are in skeletons complete bones as found vertebrate generally Large preservation). enhanced of period a or events several (indicating subunit the within spread event)are mortality or mass single a indicate could , the Spanish localities with Tapiridae, and the and Tapiridae, with localities Spanish the arvernensis, Tapirus eoiin t h lk bto i (t es fr some for least (at is bottom lake the at Deposition to belongs always preservation bone mammal Large Maps showing the distribution of taxa discussed in the text. the in discussed taxa of distribution the showing Maps (after Van der Made and Stefanovic, and Made der Van (after Tapiriscus Camp delsNinotsmaarsite(Spain) Dusino; 69) Roatto; 70) Montopoli 70) Roatto; 69) Dusino; and Parabos 12 - 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 ACKNOWLEDGMENTS its as well as better, site taphonomy and its palaeoenvironment during the the Pliocene. understand and get know to to us enable palaeobotanical to information and geological and palaeontological material better and more others. (Eocene, Messel among Spain) (Pliocene, or Higueruelas Las and Eckfeld Germany) as such sites lagerstätten maar significant other also understand better but to contribute will age, this for area the of ecosystem the study to stability ofthedistribution oftheseenvironments. of degree some suggesting times, environments) several humid Spain reached (closed habitat preferred their of presumably and tapirs of distribution of limits The tapirs. of species Miocene several of record southernmost the to the southernmost occurrence in SW Europe and is also close previously known onlyinthePleistocene. were which species, these of Europe in record oldest the constitute they as Ninots, dels Camp Pelophylaxin perezi units MN15and16,anageofabout3.2Ma. large mammal association is suggestive The of the transition of fossils. invertebrate no but remains plant numerous (Leuciscus fishes freshwater and perezi e utr i ijn d Cmnccó e a eeaia de Generalitat la de Comunicació de Mitjans i Cultura de the Townfrom Malavella,de Departament Caldes the of Council contributions from benefited has paper This appreciated. highly The are Gibert de M. manuscript. J. and Delclòs X. Cabrera, L. by work editorial the improve to revision contributed the strongly of process result a as out carried changes and comments The fieldwork. the in participated have who those all as well as and Repsol YPF, who gave permission to excavate on their fields, the tapir the Alephis tigneresi , the rhinoceros the new dataonvertebrate fossils. on emphasis particular putting Ninots, dels Camp the age, new a of study the CONCLUSIONS sp. and sp. turtle the . l a t e R E L O S E D Z E M Ó G . B ) 2 2 1 0 0 7 2 1 ( 0 0 7 0 1 0 - 0 1 . 5 , 0 ) 1 1 / ( 4 0 4 1 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G We hope that future excavation campaigns will deliverwill excavationcampaigns future We that hope This exceptional site not only provides a unique chance represents Ninots Tapirusdels Camp from arvernensis of presence the is Noteworthy The vertebrate taxa identified until now are the bovid the are now until identified taxa vertebrate The from obtained results first the present we paper this In We thank the Montalat, Esteve Miquel and Cateura families Cateura and Miquel Esteve Montalat, the Wethank aff. Lissotriton , the rodent Tapirusthe arvernenis, to et, cf. newts, two leprosa, Mauremys site of site Konservat-Lagerstätte , the frog the helveticus, Stephanorhinus jeanvireti, and leprosa Mauremys Apodemus atavus, Apodemus ?). There are also are There ?). Pelophylax Pleurodeles Pliocene cf. Bianucci, G., Mazza, P., Merola, D., Sarti G., Cascella, A., A., Cascella, G., Sarti D., Merola, P., Mazza, G., Bianucci, Behrensmeyer, A.K., 1978. Taphonomic and ecologic information aguas las de geológicas Condiciones 1933. J.R., Bataller, Barroso-Ruiz, C., Bailon, S., 2003. Los anfibios y los reptiles del paleoecológico análisis y tafonómico Estudio 1996. E., Barrón, Bachelet, B., 1990. Muridae et Arvicolidae S., P.,Wing, Wilf, K., Jonson, L.J., Hickey, B., Ellis, A., Ash, L., Cacho, García C., Escorza, Martín A., V.,Aparicio, Araña, de Pontien Vertébrésdu Les 1929. J., Piveteau, C., Arambourg, O., Oms, W., Krijgsman, M., Garcés, L., Cabrera, J., Agustí, hipparionine last the of age the On 2001. O., Oms, J., Agustí, J.P., Faillat, M., Calvet, B., Bachelet, J., J.P.,Aguilar, Michaux, REFERENCES of thisarticle. revision for Duke Charles to thanks Our Atapuerca. Fundación from grant a has Riba D. Naturales. Ciencias de Nacional Museo the of material reptile and amphibian the to access for Fernández González J.E. thanks Blain H.-A. (Gencat). SGR2009-324 and Inovación),e Ciencia de Ministerio (DGI, CGL2009-12703-C03 Catalunya and the projects CGL2008-03881, CGL2009-7896 and biostratigraphical data and paleoecological observations. observations. paleoecological and data biostratigraphical planckton di Val calcareous of of light the in Italy) (Tuscany, assemblage Pugna mammal Pliocene early The 2001. from boneweathering.Paleobiology, 4,150-162. del SeminariodeBarcelona,8,84pp. Geología de Laboratorio Barcelona, Cataluña. de minerales Cultura, Arqueología Monografías,267-278. de de Consejería Junta Andalucía, Zafarraya. de Boquete del In: Barroso-Ruiz, C. (ed.). El Pleistoceno superior de la Cueva Zafarraya. de Boquete del Cueva la de superior Pleistoceno de Complutense Universidad Madrid, 773pp. Madrid, Thesis. Doctoral de la macro y microflora miocena de la cuenca de la Cerdaña. Montpellier, 180pp. de Université Montpellier, Thesis. Doctoral évolution, biochronologie. systématique, France: la de sud du Pliocène du n ctgrzto o dctldnu ad net-veined and Institution, 65pp. dicotyledonous Smithsonian Washington,angiosperms. of monocotyledoneous categorization and description architecture–morphological leaf of Manual 1999. y petrológicos estructurales, caracteres neógeno-cuaternario geodinámicos. Acta GeológicaHispánica,18,1-17. volcanismo Cataluña: El de 1983. J., X., Albert, G., Gassiot, Ferrara, F., Barberi, R., Vaquer, R., Ortíz, Salonique. Annales dePaléontologie, 18(2-3), 59-139,3-14. 52, 247-260. Reviews, Science Earth art. the of State WesternEurope. of Parés, J.M., 2001. A calibrated mammal scale for the Neogene de Rendus Comptes Paris, Europe. l’Académie desSciences,332,291-297. western in faunas Mio-Pliocène en Roussillon. Palaeovertebrata, 20(4), 147-174. limite la de prochesrongeurs de faunesnouvelles Les 1991. Camp delsNinotsmaarsite(Spain)

(Rodentia, mammalia) 13

10 12 11 14 13 16 15 18 17 20 19 22 21 24 23 26 25 28 27 30 29 32 31 34 33 36 35 38 37 40 39 42 41 44 43 46 45 48 47 50 49 52 51 54 53 55

2 1 4 3 6 5 8 7 ARTICLE IN PRESS 9 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 . l a t e R E L O S E D Z E M Ó G . B ) 2 2 1 0 0 7 2 1 ( 0 0 7 0 1 0 - 0 1 . 5 , 0 ) 1 1 / ( 4 0 4 1 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G Donville, B., 1976. Géologie néogène de la Catalogne orientale. Catalogne la de néogène Géologie 1976. B., Donville, aln L, 98 Rtoaet di Ritrovamento 1988. L., Dallan, dalle provineiente Parabos di Cranio 1979. P., Politi, Cuscani 1995. H., Tunner, A., Ohler, A., Dubois, P.-A., Crochet, Late Suteethorn, V.,Tong,the J., H., Turtlesfrom Claude, 2007. Turtles Testudinoid Eocene Early 2004. H., Tong J., Claude (Perissodactyla, TapiridaeLes 1988. L., Ginsburg, E., Cerdeño, D., Ormezzano, A., Mottura, M.G., Forno, F., Campanino, Metodología 2003. M.J., Iriarte, J.A., F.,López-Sáez, Burjachs, Lecture Germany. Westeifel, the of Maars 1993. G., Büchel, V., Fahlbusch, G., Daxner-Höck, R., Daams, de, H. Bruijn, Paléoherpétofaune la de Contribution 2005. H.-A., Blain, 2(4), 177-210. minières, et géologiques recherches de Bureau du Bulletin 95, A, 203-219. serie Memorie di Pisa, Toscaza in Società Residente Naturali Della Scienze Tai Pisano. Monte il e Albano Monte el tra dell’area continentali depositi dui stratigrafiche considerazioni e (Lucca) Montecarlo du lacustre serie della (Pelophylax) 265-278. 11, Siena, in (Toscana).Fisiocritici dei della Accademia Atti Siena di pressi nei Pugna di Val di plioceniche formazioni a naturelle, 17(1-2),11-30. d’Histoire national of Muséum du description Bulletin Paris, taxon. and new diagnoses morphological Anura): (Amphibia, klepton associated their and 1885 pereziSeoane, 305-316. (Thailand). Basin 178(4), 1, Série France, de géologique Krabi société la de Bulletin the of Oligocene Eocene-Early modern Testudinoidea. Oryctos,5,3-45. of Origin the on Comments with France, Saint-Papoul, from Inférieur Miocène du et Européens. l’Oligocène Annales dePaléontologie, 74(2), 71-96. de Mammalia) 12(2), 439-499. Naturali, Scienze di Regionale Museo Dusino. del Bolletino Torino, from specimen the of Revision Italy. NW d’Asti, Villafranca near Roatto from Mammalia) (Rhinocerotidae, 1994. B., Sala, y objetivos de Catalunya, 11-18. arqueobotánica: en La d’Arqueologia Museu Barcelona, metodológicas. propuestas (eds.). muestras R. de Piqué, recogida R., Buxó, In: arqueopalinológica. Notes inEarthSciences,49,1-13. on Newsletters 1990. Reisensburg Stratigraphy, 26(2/3),65-118. mammals, fossil on group working Telles RCMNS the N., of Report Schmidt-Kittler, 1992. M., Antunes, A.J., Meulen, der Van D.F., Mayhew,E., Heizmann, J., P.,Morales, Mein, L., Ginsburg, d’Histoire naturelle, Institut de Paléontologie Humaine, 402pp. national Muséum Paris, Thesis. Doctoral d’Espagne. Pléistocène moyen au supérieur Pliocène du paysage du et climat l’évolutionduconnaissance de la àSquamata) &(Amphibia 107(3), Stratigrafia, e 425-438. Paleontologia di Italiana Rivista als 1771, Pallas, ridibunda Géi) 1972 (Guérin) jeanvireti Sterphanorhinus el argille nelle lyrix Alephis (Pelophylax) Rana Rana Gómez-Alba, J.A.S., 1988. Guía de Campo de los Fósiles de España Riba, S., García, O., Oms, G., Campeny, B., Soler, de Gómez Caracterización 1982. M., Crusafont-Paíro, J.M., Golpe-Posse, Goeury, C., Beaulieu, J.-L. de, 1979. À propos de la concentration techniques Nouvelles 1969. J., Renault‑Miskovsky, M., Girard, Bovini tribe the of analysis Phylogenetic 1992. D., Geraads, 2004. P., Herrero, A., Montori, M., García-París, D., Riba, B., Soler, de Gómez Campeny,G., S., Catalán, García Frost, D.R., Grant, T., Faivovich, J., Bain, R.H., Haas, A., Haddad, 50 vor Krokodile-Messel und Urpferdchen 1977. J.L., Franzen, (ober-ruscinische) pliozäne Eine 1990. G., Storch, O., Fejfar, Fejfar, O., 1964. The Lower-Villafranchian vertebrates from Hajnácka 1993. B., Sala, P., Mazza, M., Fortelius, In: Wellnhofer,Caudata. Gymnophiona, 1981. R., Estes, P. (ed.). y de Europa. Barcelona, Ediciones Omega S.A., 925pp. Jornades IX L’Escala, Selva). d’Arqueologia de lesComarquesGironines,13-23. 2007 la i Ninots. Malavella, dels de 2006 Camp (Caldes del El 2008. arqueopaleontològiques R., Intervencions Sala, R., Rosillo, D., Penedés. Acta Geológica Hispánica,17(1-2),95-101. de minéraux. PollenetSpores,XXI(1-2),239-251. du pollen à l’aide de la liqueur de Thoulet dans les sédiments Quaternaire, 4,275-284. du l’Étude pour française l’Association de Bulletin Rhône). du (Istres‑Bouches Cornille l’Abri de final Quaternaire sédiments du trois à appliqués Palynologie en préparation de Society, 104,193-207. Linnean he of (Mammalia: Journal Zoological Artiodactyla). Ciencias Naturales,640pp. de Nacional Consejo Museo (CSIC), Madrid, Científicas Investigaciones 24. de Ibérica, Fauna (eds.). J. J., Tamplado, Serrano, E., MacPherson, A., Guerra, J., Gosálbez, X., Bellés, J., Alba-Tercedor, M.A., Ramos, In: Lissamphibia. Corneli (Cercs),69-82. Sant Paleontología. en Investigadores Jóvenes de Encuentro III del Actas (eds.). B. Vila, J., Marmi, M., Furió, I., Vilar, Casanovas- S., Almécija, In: Girona). Malavella, de (Caldes Ninots dels Camp del Selva:yacimiento la el de depresión la 2007. Aportaciones preliminares al registro paleontológico de the American MuseumofNaturalHistory, 297,1-370. of Bulletin life. of tree Wheeler,amphibian W.C., The 2006. D.M., Green, J.D., Lynch, R.A., Nussbaum, R.C., Drewes, S.C., Raxworthy, C.J., Campbell, J.A., Blotto, B.L., Moler, P., C.F.B., De Sá, R.O., Channing, A., Wilkinson, M., Donnellan, Millionen Jahren.KleineSenckenberg-Reihe, 7,1-36. I. Rheinhessen. 71(1-2), 139-184. letahea, Senckenbergiana Gudersheim, Rodentia. Mammalia, Nagetiere: aus Kleinsäugerfauna near Filákovo in southern Slovakia. Rozpravy, 30, 1-115. European Palaeontographia Italica,40,63-155. western the of revision a with Pleistocene, of Rhinocerotidae) (Mammalia: Fischer Verlag, 15,115pp. Gustav München, 2. Part Paläoherpetologie, der Handbuch Kaup en el Mioceno superior del Vallés-del superior Mioceno el en Kaup Tapiruspriscus Camp delsNinotsmaarsite(Spain) S. etruscus S. (Falconer, 1868). (Falconer, Stephanorhinus Amphibia, 14 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 Hooijer, D.A., 1981. The first rhinocerotid of the Pretiglian “black from fauna mammal large Plio-Pleistocene A 1996. P., Holec, G., Rössner, In: Rhinocerotidae. Family 1999. K., Heissig, Heissig, K., 1996. The stratigraphical range of fossil rhinoceroses von Salamandridenmaterial Neues 1997. M.S., Haller-Probst, (Mammalia, tapirs Les 1994. V., Eisenmann, C., Guérin, (Mammalia, Rhinocerotidae Les 1982b. C., Guérin, Pléistocène du biozonation Première 1982a. C., Guérin, Perissodactyla) (Mammalia, rhinocéros Les 1980. C., Guérin, la de volcans des l’étude à Contribution 1964. P., Guardia, Gruber, G., Micklich, N., 2007. Messel. Treasures of the Eocene. (Mammalia, Bovidae grands Les 1981. C., Gromolard, Gómez-Merino, G., Ibáñez, N., Campeny, G., Gómez de Soler, B., . l a t e R E L O S E D Z E M Ó G . B ) 2 2 1 0 0 7 2 1 ( 0 0 7 0 1 0 - 0 1 . 5 , 0 ) 1 1 / ( 4 0 4 1 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G Zoology, 31(2),472-475. of Journal Netherlands Netherlands. the from fauna bones” Cracoviensia, 39(1),219-222. Zoologica Acta Slovakia. south Vieska, Nova and Strekov München, Verlag Dr. FriedrichPfeil,175-188. Europe. of mammals land Miocene The (eds.). K. Heissig, New Faunas.York, ColumbiaUniversity Press,339-347. Mammal Neogene Eurasian Western of evolution The H.W.Fahlbusch,(eds.). V., Mittmann, R.L., Bernor, In: in the late Neogene of Europe and the Eastern Mediterranean. 53-73. 37, Mitteilungen, Geologie Historische für und Palaeontologie Staatssammlung Bayerische Fundorten. süddeutschen Géobios, 27,113-127. occidentale. d’Europe Supérieur Miocene du Perissodactyla) actuelles: 15(4), 599-605. espèces Geobios, phylogénétiques. aux relations et évolutives tendances comparés occidentale d’Europe Perissodactyla) du Miocène terminal au Pléistocène Supérieur occidentale. d’Europe Geobios, 15(4),593-598. Supérieur Pléistocène au terminal l’étude Miocène du Perissodactyla) (Mammalia, Rhinocerotidae des de biostratigraphique résultat principal Européen, Documents desLaboratoiresdeGéologie,79(1-3),1185pp. Lyon, actuelles. espèces les Europe avec Comparaison en occidentale. Supérieur Pléistocène au terminal Miocène du París, 56pp. Mém. du Diplôme d’Études Supérieures. París, Université de coulées. leurs de paléomagnétisme du et Gerone de Province Darmstadt, HessichesLandesmuseum159pp. occidentale, Université Claude Bernard-Lyon I,271pp. Villeurbanne, d’Europe Thesis. Doctoral paléoécologie. Pliocène évolution, systématique, Paléoneurologie, Biostratigraphie, du Artiodactyla) de Barcelona.Memòriaespecialnúm4,201-210. Universitatla de Publicacions de Autònoma Servei Sabadell, (eds.). Gelabert Celià L. and Cortés Marigó J. Molina, Val S. Fernández, Cancelo C. In: Collections. History Natural for Solutions Global Finding Workshop: Consevation “I at Presented Girona). Malavella, de (Caldes Ninots dels Camp del paleontológico yacimiento del bóvido de cráneo un de 2009. Actuaciones de conservación-restauración y almacenaje atnSrao A, ea, . Grí-ots A, aá, L., Galán, A., García-Cortés, J., Vegas, A., Martín-Serrano, biogeographic Longstanding 2010. I., Mateos, der,van J. Made, Miocene Late P.,2006. Montoya, J., Morales, der, van J. Made, Made, J. van der, Stefanovic, I., 2006. A small tapir from the Turolian Lutz, H., Kaulfuss, U., 2006. A dynamic model for the meromictic Terravolcanoes. maar-diatreme of Formation V.,2000. Lorenz, Léón, de Ponce A., Vekua, T., Jashashvili, D., Lordkipanidze, Llorente, G.A., Arano, B., 1997. B., Arano, G.A., Llorente, Llopis Lladó, N., 1943. Estudio hidrogeológico de los alrededores de López-Polín, L., Roubach, S., Gómez de Soler, B., Campeny, G., 2010. Linder, H.,Gabriel,G.,Götze,H-J.,Kaeppler, R.,Suhr, P., 2006. the from Chelonians African 2000. de, F. Broin, de Lapparent of occurrence northernmost The 2008. Mörs, T. F., Lacombat, ap d Cltaa ocnc il (eta San SW Spain, (Central Europe). Sedimentary Geology, 222,52-63. the Field Volcanic in Calatrava lakes de maar Campo of setting Morphotectonic 2009. J.L., García-Lobón, A., Pérez-González, A., Granda, F.M., P.I., Ibarra Rubio, S., Martín-Alfageme, J.L., Gallardo-Millán, doi 10.1016/j.quaint.2009.11.015. 195-200. 1223-1224, International, Quaternary Europe. into and Africa of out Homo early of dispersal the and patterns 228-246. 238, Palaeoecology, Palaeoclimatology, Palaeogeography, context. wider a in record mammal Spanish the in turnover Neues Monatsheften, Jahrbuch fürGeologieundPaläontologie, 240(2),207-240. tapirs. Neogene the of stratigraphy and biogeography the on discussion a and (Bosnia) Kreka of 433-450. 157(3), Geowissenschaften, für Gesellschaft Deutschen der Zeitschrift Germany). Eocene, (Middle Maar Eckfeld lake Nostra, 6,284-291. from evidence early HomofromDmanisi,Georgia. Nature,449,305-310. Postcranial 2007. L., Rook, M., Nioradze, A., Mouskhelishvili, B., Martínez-Navarro, G., Kiladze, R., Kahlke, J., Agustí, M., Bukhsianidze, Tappen,M., O., Oms, R., Ferring, G.P.,H., Pontzer,Rightmire, C., Zollikofer, M., Monografias Revista EspañoladeHerpetología,3,164-166. de los anfibios y reptiles de España. Universidad de Granada, Biogeografia y DistribuciónJ.P. (eds.). Martínez-Rica, J.M., Caldas de Malavella (Gerona). Speleon, II, 103-164. Memòria especial núm 4, 233-239. Servei de Publicacions de la Universitat AutònomaMolina, deJ. Marigó Barcelona.Cortés and L. Celià Gelabert (eds.). Sabadell, for Natural History Collections”. In: C. CanceloPresented Fernández,at “I Consevation S. Val Workshop: Malavella, Spain). Girona, de (Caldes site Ninots Finding dels Camp Global Solutions Prelimimaryconsolidation tests of fossil plant imprints form the für Geowissenschaften, 157(3),355-372. Gesellschaft Deutschen der Zeitschrift Saxony). (East region Geophysical and geological investigation in the Upper Lusatia fossil chelonians.Palaeontologia Africana, 36,43-82. African the of catalog preliminary A Present. the to Jurassic und Paläontologie, Abhandlungen, 249(2),157-165. Geologie für Jahrbuch Neues Perissodactyla). (Mammalia, rhinoceros Pliocene Late rare the

Camp delsNinotsmaarsite(Spain) . In: Pleguezuelos, In: perezi. Rana

Stephanorhinus jeanvireti Stephanorhinus 15

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2 1 4 3 6 5 8 7 ARTICLE IN PRESS 9 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 . l a t e R E L O S E D Z E M Ó G . B ) 2 2 1 0 0 7 2 1 ( 0 0 7 0 1 0 - 0 1 . 5 , 0 ) 1 1 / ( 4 0 4 1 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G oa E, 96 L eouin ednmc d l Cuenca la de geodinámica evolución La 1996. E., Roca, reptiles squamate and 2005. Amphibians S., Bailon, J.-C., Rage, E., Stiuca, A., Petculescu, P.M, Samson, C., Estudio Radulescu, 1990. P., Badiella, L., Sugrañes, Solé J., Pous, Pirrung, M., Fischer, C., Büchel, G., Gaupp, R., Lutz, H., Neuffer, de comunidades las de Análisis 1990. D., Soria, B., Pérez, de sous-genre d´un évolutive Dynamique 1974. L., Pasquier, Agustí, R., Sala, G., Campeny, B., Soler, de Gómez O., Oms, Fossillagerstätte 1994. (eds.), H., Lutz,Neuffer, Gruber, G., F.O, de macrofauna su Moro: del Venta 1984. J., Morales, J., Made, der Van M.T, Alberdi, L., Ginsburg, P., Montoya, Sudre, M.P., Duvernois, M., Calvet, J.P, Aguilar, J., Michaux, Lutz, F.O., Neuffer, J.L., Franzen, C.C., Swisher, D.F., Mertz, Néogène du biozonation de Proposition 1975. P., Mein, Mazo, A.V., 1997. El yacimiento Rusciniense de Alcalá del Júcar The 2003. G., Saldi, M., Zuccolini, Vetuschi L., Marini, la actualidad. Acta GeológicaHispánica, 29(1),3-25. Catalano-Balear y áreas adyacentes desde el Mesozoico hasta Gers, southwesternFrance).Geodiversitas, 27(3),413-441. from the late early Miocene (MN 4) of Béon 1 (Montréal-du- Paleontología, de 1,549-558. Coloquios Romania. of mammals large Pliocene 2003. Acta (Girona). Selva La Geológica Hispánica,25(4),261-269. de depresión la de geoeléctrico the Eifelarea(Germany). Terra Nova, 15,125-132. in deposits crater maar of sucession Lithofacies F.-O.,2003. Calera La y (Soria) Layna (Teruel). Paleontología yEvolució, 23,231-238. de Plioceno del mamíferos actuelles Université deMontpellier, 176pp. et occidentale. fósiles populations d’Europe de dentaires caràcteres Muridae geology, Abstract CD-ROM, 69p. 33rd historical and paleontology to contributions General HPF-01 6-14, Symposia (IGC). August Congress Geological World Oslo, International Catalonia). (Pliocene, dels Ninots Camp el of site maar the at record fossil the a volcano: inside Trapped 2008. F., Burjachs, J., Made, der Van J., Eckfelder Maar. Mainz, Mainzer Naturwiss Mioceno Arch, BeiheftComplutense deMadrid,340pp. del ,16, 222pp. continental Universidad la de Editorial Madrid, mediterraneo, Terminal biostratigrafía y mamíferos, in biostratigraphy. Geodiversitas, 28(1),137-173. of locality Pliocene early importance their Alcoyand (Spain) Morales, J., Soria, M.D., 2006. Fossil large mammals from the Pliocène duRoussillon.Geobios,24(6),735-745. 1991. J., Naturwissenschaften, 87,270-274. Eifel, Germany: a calibration mark for the Eocene time scale. site, fossil maar Eckfeld the of dating Numerical 2000. H., Trabajossobre Neógeno-Cuaternario,4,112p. Madrid, mammifères. des partir à méditerranéen (Albacete). EstudiosGeológicos,53,275-286. Volcanology andGeothermalResearch,121,83-98. of Journal waters. lake volcanic of distribution pH bimodal lpi tigneresi Alephis . td boérqe des biométrique Etude (Sylvaemus). Apodemus otrl hss Montpellier, Thesis. Doctoral o. p, n oié ova du nouveau Bovidé un sp., nov. Wuttke, M., 1992a. Death and burial of the vertebrates. In: Schaal, fauna Villalta,la de J.F.conocimiento Contribuciónal 1952. de., de termal estación la de geológico Estudio 1882. Ll.M., Vidal, (Barcelona). de Montjuïc fòssil flora La 1988. J., Vicente, Vicente, J., 1985. Troballa d’un Troballa1985. Vicente,J., Relationship 2005. R., Linares, C., Roqué, J., Bach, M., Vehí, edifici Un 1999. L., Pallí, C., Roqué, A., Pujadas, M., Vehí, Volcanic2000. G.L.Jr.Rowe, G.B., Pasternack,Varekamp, J.C., Tassone, A., Roca, E., Muñoz, J.A., Cabrera, L., Canals, M., 1994. Spassov, N., 2005. Brief review of the Pliocene ungulate fauna of los de hidrogeológicas Características 1946. L., Sabarís, Solé floras las de conocimiento el para Datos Siria, 1987. de A., Sanz Handbuch (ed.). P.Wellnhofer,In: Salentia. 1998. B., Sanchíz, Cuaternario y Neogeno del anfibios Nuevos 1977. B., Sanchíz, deposit flow mass fossiliferous remarkable A 2005. M., Sachse, Oligocene 1999. M., Marzo, L., Cabrera, M., Sans, E., Roca, life andoftheEarth. Oxford,ClarendonPress,263-275. of history the into insight An Messel. Ziegler,W.(eds.). S., 64, España, de 3-203. Minero y Geológico Instituto del Boletín (Logroño). Villaroya de Plioceno del fósiles mamíferos de Mapa GeológicodeEspaña,9,65-91. del Comisión la de Boletín (Gerona). Malavella de Caldas del Natura la de d’Estudis Barcelonès Nord,1-93. Institut Natural, d’Història Societat Gramenet, de Coloma Santa 1. monogràfica, Sèrie d’Estudis delaNaturadelBarcelonèsNord,1(1),86-88. Centre del Butlletí Natural, d’Història Societat Selva). (La 60(6), 331-338. Mediterranean margin. Journal of soil and water conservation, wetlands: The case of the Prats de Sant Sebastià, northwestern of environmentalgenesis and ground-water thermal between Volcà del Camp dels Ninots. Quaderns de la Selva, 11, 45-72. el Girona): Selva, (La Malavella de Caldes a inèdit volcànic of Journal constraints Chemical Volcanology II. andGeothermalResearch,97,161-179. systematics lake Hispánica, 29(2-4), 3-37. Geológica Acta Cenozoico. el durante continental catalán margen del septentrional sector del Evolución Bulgaria. Quaternaire,2,201-212. Anales delInstitutodeEstudiosGerundenses,1,236-269. Picants. Fonts llamados Gerona de carbónicos manantiales 303. 295- 21, Evolució, y Paleontología Cataluña. de pliocénicas Pfeil, 275pp. der Paläoherpetologie, tome 4. München, Verlag Dr. Friedrich de Madrid,863pp. española. Doctoral Thesis. Madrid, Universidad Complutense de Europa. Origen, desarollo y relaciones de la batrachofauna 51, 173-184. Facies, considerations. palaeoecological and taphonomical, (Germany)-sedimentological, Maar Eckfeld Eocene the in 233. 209- 315, Tectonophysics, Mediterranean). (northwestern margin Catalan central the of evolution Miocene Middle to Camp delsNinotsmaarsite(Spain) Leptobos a Caldes de Malavella de Caldes a 16 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 55 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 ARTICLE IN PRESS 9 8 7 6 5 4 3 2 1 Wuttke,M., 1992b. Conservation-dissolution transformation. On the . l a t e R E L O S E D Z E M Ó G . B ) 2 2 1 0 0 7 2 1 ( 0 0 7 0 1 0 - 0 1 . 5 , 0 ) 1 1 / ( 4 0 4 1 3 1 , . a 0 t 1 c A : I a O c D i g o l o e G and of the Earth. Oxford, Clarendon Press, 263-275. S.,Ziegler, W. (eds.). Messel. insightAn intothe history lifeof behaviourbiogenicof materials during fossilization. In:Schaal, enr F, 94 De eihne zice Shdlom und Schädelform zwischen Beziehungen Die 1934. F., Zeuner, published OnlineJuly2011. revision acceptedDecember2010; Manuscript receivedApril2010; eeses bi e rzne ud osln Nashörnern. fossilen und Berichte derNaturforschendeGesellschaft,34,21-80. rezenten den bei Lebensweise

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