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New Dryolestidan Mammal from the Hauterivian–Barremian Transition of the Iberian Peninsula

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New Dryolestidan Mammal from the Hauterivian–Barremian Transition of the Iberian Peninsula New dryolestidan mammal from the Hauterivian–Barremian transition of the Iberian Peninsula GLORIA CUENCA−BESCÓS, AINARA BADIOLA, JOSÉ I. CANUDO, JOSÉ M. GASCA, and MIGUEL MORENO−AZANZA Cuenca−Bescós, G., Badiola, A., Canudo, J.I., Gasca, J.M., and Moreno−Azanza, M. 2011. New dryolestidan mammal from the Hauterivian–Barremian transition of the Iberian Peninsula. Acta Palaeontologica Polonica 56 (2): 257–267. Crusafontia amoae sp. nov. (Dryolestida, Stem Cladotheria) is represented by two isolated upper molars (M4 or M5 and M6 or M7) from the terminal Hauterivian–basal Barremian (Early Cretaceous) of the El Castellar Formation (Galve, Spain). The molars have a deep ectoflexus, a distinct metacone, a continuous metacrista, and an antero−lingually placed paracone. They differ from the molars of the other species of the genus, Crusafontia cuencana, by their larger size, by their outsized parastyle, by the pointed lingual slope of the paracone, their more symmetrical appearance with a deep ectoflexus in occlusal view, and the well−developed metacone. Revision of three isolated teeth previously attributed to Crusafontia cuencana suggests that instead of being upper premolars belonging to Crusafontia cuencana they in fact be− long to Pocamus pepelui, of the stem cladotherian superorder Zatheria, probably “peramuran”. As such, the stem Cladotheria record from the Early Cretaceous of the Iberian Peninsula is composed of two dryolestids (Crusafontia amoae and Crusafontia cuencana) and one zatherian (Pocamus pepelui). Key words: Mammalia, Cladotheria, Dryolestida, Crusafontia, systematics, Cretaceous, Iberian Peninsula. Gloria Cuenca−Bescós [[email protected]], Ainara Badiola [[email protected]], José I. Canudo [[email protected]], José M. Gasca [[email protected]], and Miguel Moreno−Azanza [[email protected]], Grupo Aragosaurus−IUCA (http://www.aragosaurus.com), Paleontología, Facultad de Ciencias, C/ Pedro Cerbuna 12, Universidad de Zaragoza, 50009 Zaragoza, Spain. Received 29 December 2009, accepted 3 August 2010, available online 14 September 2010. Introduction South America, and possibly Australia, and in the Late Creta− ceous of Argentina and possibly North America (Henkel and The Stem clade Cladotheria are important and diversified Krebs 1969; Freeman 1979; Prothero 1981; Bonaparte 1990, Mesozoic mammals that comprise several groups of pre− 1994, 1996, 2002; Krebs 1993; Canudo and Cuenca−Bescós tribosphenic mammals, the dryolestoids, the amphitheriids, 1996; Martin 1999; Sigogneau−Russell 1999; Kielan−Jawo− Vincelestes, and the zatherians, including Zatheria and “per− rowska et al. 2004). The Early Cretaceous Stem Cladotheria amurans” (Kielan−Jaworowska et al. 2004). The Stem Clado− from the Iberian Peninsula are represented by Dryolestida and theria represent the intermediate grade between the plesio− peramurans from the provinces of Teruel (Galve) and Cuenca morphic “symmetrodontans”, with a reversed−triangle molar (Uña) in the Iberian Range in central and north−eastern Spain pattern like Spalacotherium and their kin, and the apomor− (Henkel and Krebs 1969; Krebs 1993; Canudo and Cuenca− phic tribosphenic mammals, with the triangle−molar pattern Bescós 1996; Martin 1998) (Fig. 1). of metatherians and eutherians (Kielan−Jaworowska et al. More than a hundred Late Jurassic (Tithonian) and Early 2004). The molars of Stem Cladotheria share with sym− Cretaceous (Hauterivian–Barremian) vertebrate−fossil−bea− metrodontans the reversed−triangle molar pattern, though the ring beds are present in the Galve syncline, around the vil− upper molars are wider than the lower ones, having a third lage of Galve (Teruel), which are included in four forma− (lingual) root and the labial stylar cusps being more elabo− tions of the Galve sub−basin (Ruiz−Omeñaca et al. 2004), rate. The last upper premolars may be molariform in some that forms part of the Maestrazgo Basin (Fig. 1). The known groups, though they have only two roots. stem cladotherians from Galve are from two localities of the The stem cladotherians are known from the Middle Juras− El Castellar Formation and one from the Camarillas Forma− sic on (Kirtlington, Forest Marble Formation in Britain; Gui− tion. The upper layers of the El Castellar Formation are ter− marota Mine in Portugal; Morrison Formation in the USA). minal Hauterivian−basal Barremian in age. The fossil mam− They survive throughout the Cretaceous: in the Early Creta− malian remains were found in two different localities of the ceous of Europe (Spain, Britain) and Morocco, Mongolia, El Castellar Formation: Galve Th, and Pelejón 2. The third Acta Palaeontol. Pol. 56 (2): 257–267, 2011 doi:10.4202/app.2009.0157 258 ACTA PALAEONTOLOGICA POLONICA 56 (2), 2011 (Krebs 1985) in the upper levels of the El Castellar Forma− tion, terminal Hauterivian–basal Barremian in age (Canudo and Cuenca−Bescós 1996). Near the Galve syncline in the Iberian Maestrazgo Basin N province of Teruel, other Mesozoic mammals have been de− Range Calanda scribed (Cuenca−Bescós and Canudo 2003; Badiola et al. Oliete Josa Ol Pe 2008). Montalbán Herbers Fault Recently, a new dryolestid upper tooth has been found in Pa Tortosa GALVE Aliaga Mo the El Castellar Formation. In this paper, it is assigned to a Tertiary Morella Fig. 2 Miravete Fault Mesozoic Ga Turmell Fault Zone new species of Crusafontia. The aim of this paper is twofold, Paleozoic Vinaròs Sa on the one hand a description of this new species of Dryo− lestida from the Hauterivian–Barremian transition of the (Umbral Zone) (JavalambreTERUEL High) Mora de Rubielos Galve area, and on the other hand a revision of the premolars CUENCA Serranía de Pg of stem cladotherians, especially those assigned to Crusa− Cuenca Basin Mediterranean Sea fontia cuencana from several localities of the Early Creta− 20 km CASTELLÓN ceous of Galve. MAMMAL MAMMAL AGE FORMATION LEVELS LEVELS Institutional abbreviation.—MPZ, Museo de Paleontología de la Universidad de Zaragoza. LA HUÉRGUINA UPPER Other abbreviation.—CC 2, Cuesta Corrales 2. BARREMIAN LOWER CAMARILLAS EL CASTELLAR Geographic and geological setting UPPER HAUTERIVIAN U A (CUENCA) GALVE (TERUEL) The localities of Galve are situated in the geographic domain Serranía de Cuenca Basin Maestrazgo Basin of the Central Iberian Range, in the province of Teruel in the sub-basin Galve sub-basin NE of Spain (Fig. 1). The new mammalian tooth was found in the locality of Cuesta Corrales 2. The site is 1.5 km east of the Fig. 1. Geographical and geological setting of Galve (Teruel, Spain). A.Sim− plified geological map of the Iberian Peninsula. B. Palaeogeographic sub− village of Galve, on the eastern flank of the Galve syncline basins within the Maestrazgo Basin and active faults during Early Cretaceous (Fig. 2). sedimentation, modified from Salas et al. (2001), and the palaeogeographi− The Galve syncline is situated to the west of the Galve cal—and geographical—relationship with the Uña area (Cuenca, Spain). sub−basin, which together with six other sub−basins com− C. Litho− and chronostratigraphy of the Mesozoic mammal record from the prises the Maestrazgo Basin (Salas et al. 2001; Liesa et al. areas of Galve and Uña. Abbreviations: Ga, Galve; Mo, Morella; Ol, Oliete; 2006). The Galve sub−basin is NNW−SSE oriented, and is Pa, Las Parras; Pe, Perelló; Pg, Peñagolosa; Sa, Salzedella. 40 km long and 20 km wide (Fig. 1B). It was formed as a consequence of the Jurassic–Cretaceous extensional defor− mammalian site of Galve is Poca, in the upper layers of mation of the Iberian rift (Liesa et al. 2006). In the Galve the Camarillas Formation. The mammals described are the sub−basin, marine, continental and transitional sediments dryolestid Crusafontia cuencana Henkel and Krebs, 1969 outcrop from the Kimmeridgian (Late Jurassic) to the early at the sites of Galve Th and Pelejón 2, and the peramurid Aptian (Díaz Molina and Yébenes 1987). Sites of Mesozoic Pocamus pepelui Canudo and Cuenca−Bescós, 1996 at the vertebrates from the Galve syncline are distributed through− site of Poca. The species Crusafontia cuencana was first out the stratigraphic sequence. The geological units that are discovered and described by Henkel and Krebs (1969) in richest in vertebrates are the Villar del Arzobispo Forma− the late Barremian site of Uña in the province of Cuenca (La tion (late Tithonian–early Berriasian), the El Castellar For− Huérguina Limestone Formation, the Serranía de Cuenca mation (terminal Hauterivian–basal Barremian), and the Basin; see Buscalioni et al. 2008 for a bibliography). Be− Camarillas Formation (early Barremian) (Sanz et al. 1987; tween the terminal Hauterivian–basal Barremian of the El Estes and Sanchiz 1982; Díaz Molina and Yébenes 1987; Castellar and Camarillas formations, and the late Barremian Ruiz−Omeñaca et al. 2004; Canudo et al. 2006; Badiola et of the La Huérguina Formation, there is a span of around al. 2009) (Figs. 1C, 2). 2Ma. The El Castellar Formation represents the first syn−rift Multituberculate and “symmetrodontan” mammals have unit included in the Wealden facies of the Galve sub−basin. It also been described from the Galve area (Crusafont−Pairó lies above an unconformity on the Villar del Arzobispo For− and Adrover 1966; Crusafont and Gibert 1976; Hahn and mation, and there is a stratigraphic gap that extends from Hahn 1992, 2002; Canudo and Cuenca−Bescós 1996; Ba− the Berriasian to the upper Hauterivian (Díaz Molina and diola et al. 2009, 2011). The only species of Early Cretaceous Yébenes 1987; Liesa et al. 2006; Meléndez et al. 2009). The Iberian “symmetrodontan”
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