MastiaRevista del Museo Arqueológico Municipal de Cartagena

Geología y Paleontología de Cueva Victoria

2012-2014 Segunda ÉpocaNúmeros 11-12-13

L. Gibert y C. Ferràndez-Cañadell(Editores Científicos)

MastiaRevista del Museo Arqueológico

Municipal de Cartagena

«Enrique Escudero de Castro»

Segunda Época

Cartagena, 2015

Números 11-12-13 / Años 2012-2014

AYUNTAMIENTODE CARTAGENA

Mastia

CONSEJO DE REDACCIÓN

Director, Miguel Martín Camino

Secretario, Dr. Miguel Martínez Andreu

Museo Arqueológico Municipal de Cartagena

«Enrique Escudero de Castro»

CONSEJO ASESOR

Prof. Dr. Lorenzo Abad (Universidad de Alicante)

Prof. Dr. Juan Manuel Abascal (Universidad de Alicante)

Prof. Dr. José Miguel Noguera Celdrán (Universidad de Murcia)

Prof. Dr. Sebastián F. Ramallo Asensio (Universidad de Murcia)

Prof. Dr. Jaime Vizcaíno Sánchez (Universidad de Murcia)

Carlos García Cano, Manuel Lechuga Galindo (Dirección General de Bienes Culturales, CARM)

Dr. Cayetano Tornel Cobacho (Archivo Municipal de Cartagena)

CORRESPONDENCIA E INTERCAMBIO

Museo Arqueológico Municipal de Cartagena «Enrique Escudero de Castro»

C/ Ramón y Cajal, nº 45 · 30205 Cartagena

Telf.: 968 128 967/128 968 · e-mail: museoarqueologico@ayto-cartagena.es

ISSN: 1579-3303Depósito Legal: MU-798-2002

© De esta edición: Museo Arqueológico Municipal de Cartagena «Enrique Escudero de Castro»© De los textos: Sus autores© De las ilustraciones: Sus autores© Imagen de la cubierta: Excavación en Cueva Victoria.

Gestión editorial: Gráficas Álamo, S.L. graficasalamo@gmail.com www.graficasalamo.com

1: Excavación en Cueva Victoria (Andamio Superior A), 20 de julio de 2010.2: Tercer molar inferior izquierdo de Theropithecus (CV-MC-400), vista oclusal.3: Cuarto premolar inferior izquierdo de Theropithecus (CV-T2), vistas bucal y lingual.4: Falange intermedia del quinto dedo de la mano derecha de Homo sp. (CV-0), vista dorsal.(Fotos: Carles Ferràndez-Cañadell).

Portada (Explicación)

Índice

Prólogo 9Prologue

EMILIANO AGUIRRE

Presentación 11Foreword

L. GIBERT y C. FERRÀNDEZ-CAÑADELL

Introducción. Cueva Victoria, un yacimiento de vertebrados del Pleistoceno Inferior 17Introduction. Cueva Victoria, an early Pleistocene vertebrate site

C. FERRÀNDEZ-CAÑADELL y L. GIBERT

Historia de la minería de Cueva Victoria 47Mining history of Cueva Victoria

M. A. PÉREZ DE PERCEVAL, J. I., MANTECA MARTÍNEZ y M.A. LÓPEZ-MORELL

Las mineralizaciones ferro-manganesíferas de la mina-cueva Victoria 59y su contexto geológico

Fe-Mn mineralizations of the mine-cave Victoria and their geological context

J. I. MANTECA y R. PIÑA

Microscopía electrónica de las mineralizaciones cársticas de óxidos de hierro y 75manganeso de Cueva Victoria (Cartagena, Murcia)

Electron microscopy of the karstic mineralizations of Fe and Mn oxydes of Cueva Victoria (Cartagena, Murcia)

D. ARTIAGA, L. GIBERT y J. GARCÍA-VEIGAS

Edad del yacimiento de Cueva Victoria y su relación con otros yacimientos de la 85Península Ibérica

Age of Cueva Victoria site and its relationship with other sites in the Iberian peninsula

L. GIBERT L. y G. SCOTT

230Th/U-dating of the Cueva Victoria flowstone sequence: Preliminary results and 101palaeoclimatic implications

Datación mediante 230Th/U de la secuencia de espeleotemas de Cueva Victoria: Resultados preliminares e implicaciones

paleoclimáticas

A. BUDSKY, D. SCHOLZ, L. GIBERT y R. MERTZ-KRAUS

Reconstrucción y génesis del karst de Cueva Victoria 111Reconstruction and genesis of the Cueva Victoria karst

A. ROS y J. L. LLAMUSÍ

Modelización tridimensional mediante escáner 3D y tomografía eléctrica de alta 127resolución, en Cueva Victoria I

Three-dimentional modelization by means of 3D Scanner and High-Resolution Electric Tomography in Cueva Victoria I

A. ESPÍN DE GEA, A. GIL ABELLÁN y M. REYES URQUIZA

Contexto sedimentario y tafonomía de Cueva Victoria 139Sedimentary context and taphonomy of Cueva Victoria

C. FERRÀNDEZ-CAÑADELL

Génesis de una acumulación osífera excepcional en Cueva Victoria (Cartagena, 163Murcia, España)

Genesis on an exceptional bone accumulation at Cueva Victoria (Cartagena, Murcia, Spain)

J. VILÀ-VINYET, Í. SORIGUERA-GELLIDA y C. FERRÀNDEZ-CAÑADELL

Anfibios y escamosos de Cueva Victoria 175Amphibians and squamate reptiles from Cueva Victoria

H. A. BLAIN

Las tortugas del yacimiento del Pleistoceno inferior de Cueva Victoria (Murcia, España) 199Turtles from the early Pleistocene site of Cueva Victoria (Murcia, Spain)

A. PÉREZ-GARCÍA, I. BONETA, X. MURELAGA, C. FERRÀNDEZ-CAÑADELL y L. GIBERT

A brief review of the Spanish archaic Pleistocene arhizodont voles 207Breve revisión de los topillos arrizodontos arcaicos de España

R. A. MARTIN

Estado de conocimiento de los Insectívoros (Soricidae, Erinaceidae) de Cueva Victoria 227The Insectivores (Soricidae, Erinaceidae) from Cueva Victoria: state of the art

M. FURIÓ

The Lower Pleistocene Bats from Cueva Victoria 239Los murciélagos del Pleistoceno inferior de Cueva Victoria

P. SEVILLA

Aves del Pleistoceno inferior de Cueva Victoria (costa sudoriental mediterránea de 253la península Ibérica)

Aves from the early Pleistocene of Cueva Victoria (southeastern mediterranean coast of the Iberian peninsula)

A. SÁNCHEZ MARCO

The latest Early Pleistocene giant deer Megaloceros novocarthaginiensis n. sp. and the 269fallow deer Dama cf. vallonnetensis from Cueva Victoria (Murcia, Spain)

El ciervo gigante Megaloceros novocarthaginiensis n. sp. y el gamo Dama cf. vallonnetensis del Pleistoceno inferior de

Cueva Victoria (Murcia, Spain)

J. VAN DER MADE

Estudio de los caballos del yacimiento de Cueva Victoria, Pleistoceno Inferior (Murcia) 325Study of the horses from Cueva Victoria, early Pleistocene (Murcia)

M. T. ALBERDI y P. PIÑERO

The rhinoceros Stephanorhinus aff. etruscus from the latest Early Pleistocene of Cueva 359Victoria (Murcia, Spain)

El rinoceronte Stephanorhinus aff. etruscus del final del Pleistoceno inferior de Cueva Victoria (Murcia, España)

J. VAN DER MADE

Elephant remains from Cueva Victoria 385Fósiles de elefante de Cueva Victoria

M. R. PALOMBO y M. T. ALBERDI.

Canid remains from Cueva Victoria. Specific attribution and biochronological implications 393Fósiles de cánidos de Cueva Victoria. Asignación específica e implicaciones biocronológicas

M. BOUDADI-MALIGNE

Úrsidos, hiénidos y félidos del Pleistoceno inferior de Cueva Victoria (Cartagena, Murcia) 401Early Pleistocene ursids, hyaenids and felids from Cueva Victoria (Cartagena, Murcia)

J. MADURELL-MALAPEIRA, J. MORALES, V. VINUESA y A. BOSCAINI

Los primates de Cueva Victoria 433Primates from Cueva Victoria

F. RIBOT, C. FERRÀNDEZ-CAÑADELL y L. GIBERT

Grupos pendientes de estudio o revisión 453Groups needing study or revision

C. FERRÀNDEZ-CAÑADELL

Preparación de restos fósiles de Cueva Victoria, Cartagena 463Preparation of fossil remains from Cueva Victoria, Cartagena

A. GALLARDO

9

Por las montañas quebradas, que llegan a la costa mediterránea entre Alicante y Cartagena, se puede ver una historia kárstica compleja, además de los restos de antiguas minas de galena argentífera en torno a La Unión. También se ob-servan en esta región algunas series sedimentarias, incluso en complicadas cavidades abiertas al exterior como es el caso de Cueva Victoria.

Cueva Victoria fue estudiada por José Gibert Clols, desde primeros de 1980 hasta su prematura muerte en el 2007. José Gibert fue un eminente científico y una gran y ejemplar persona. Insigne en una ciencia particularmente difícil, como es la Paleoantropología, ciencia que estudia las particularidades del ser humano y su evolución a través de hallaz-gos en residuos sedimentarios de remotos tiempos prehistóricos,

Cueva Victoria es una cavidad en la que se conservan parte de los sedimentos que la rellenaron y de los que se infieren sucesivos cambios climáticos y ambientales. Algunos de estos sedimentos contienen fósiles que ilustran más estas condiciones, además de la evolución de grupos biológicos. En Cueva Victoria se han podido estudiar muchos fósiles de vertebrados grandes y pequeños, algunos de ellos muy singulares como un primate del género de los “gelada”, Theropithecus.

Tales restos fósiles se encuentran en puntos muy diversos de Cueva Victoria, pero en un mismo repetido material sedi-mentario: una brecha fosilífera que presenta fósiles de vertebrados entre pequeños cantos o detritus rocosos, todo ello en ocasiones muy cementado y duro. Esta brecha se encuentra pegada en partes de la actual pared y techos de la cue-va, también en forma de bloques caídos por la actividad minera que se desarrolló en la cueva durante parte del siglo XX.

Lo más atractivo de este yacimiento fue una falange 2ª de la mano derecha (CV-0). Fue preciso examinar su distinción de la de otros primates, sobre todo del gelada Theropithecus, bien representado en Cueva Victoria y que tiene una talla parecida aunque algo más pequeña que la de los humanos. Fue José Gibert quien estudió en detalle no sólo esa falange sino otras de humanos y primates no humanos, asignándola a los primeros, con fundamento, conclusión que fue reafirmada con nuevas técnicas por otros especialistas, como los doctores Pérez Claros y Palmqvist, de la Universi-dad de Málaga. Su antigüedad fue una de las cosas más discutidas habiéndose demostrado recientemente una edad próxima al millón de años.

Esta monografía está dedicada a la memoria del Dr. José Gibert Clols quien dirigió las investigaciones en este yacimiento durante veintitrés años. El volumen nos ofrece veinticinco capítulos sobre Cueva Victoria que nos permitirán conocer y aprender mucho más sobre la Paleontología y Geología de este yacimiento emblemático. Vale la pena leer los trabajos que siguen, aunque no es pena saber más sino tiempo bien empleado, y mucho mejor cuando podáis ir por Cartagena y que os guíen en una visita a Cueva Victoria.

Emiliano AguirreReal Academia de Ciencias Exactas, Físicas y Naturales

PrólogoPrologue

11

Cueva Victoria es un yacimiento kárstico con vertebrados fósiles del Pleistoceno Inferior. Fue excavado inicialmente no como un yacimiento fosilífero, sino como mina de manganeso, incluyendo métodos tan expeditivos como el uso de ex-plosivos. Los mineros explotaron las mineralizaciones de hierro y manganeso, pero Cueva Victoria también es conocida por especialistas y coleccionistas, por la presencia de otros minerales como baritina, rodocrosita, romanechita, goethita, hollandita, calcofanita, coronadita, etc. A pesar de que la acción minera excavó alrededor del 80 % de los sedimentos fosilíferos, dejando sólo testimonios de la brecha en techo y paredes, Cueva Victoria ha suministrado miles de restos fósiles que han revelado una diversidad extraordinaria. Con las contribuciones de este volumen monográfico, la lista de especies de vertebrados identificadas en Cueva Victoria se acerca al centenar, algo extraordinario en un yacimiento. Cueva Victoria es el único yacimiento en Europa con restos fósiles del cercopitécido africano Theropithecus oswaldi, pariente cercano del babuino actual gelada. La presencia de esta especie africana en el sureste de la península ibérica aporta datos para entender los modelos de dispersión de mamíferos en el Pleistoceno. Por último, los restos fósiles de Cueva Victoria incluyen una falange humana, lo que la convierten en uno de los pocos yacimientos europeos con restos humanos del Pleistoceno Inferior.

Cueva Victoria fue dada a conocer a la comunidad científica en 1970 por Arturo Valenzuela, quien la presentó en el I Congreso Nacional de Espeología como un karst fósil, destacando sus minerales, pero describiendo también los restos de vertebrados fósiles. A finales de los 70 y principios de los 80, Joan Pons investigó su fauna fósil, en colaboración con miembros del Institut de Paleontologia de Sabadell, publicando una serie de trabajos sobre carnívoros fósiles. En estos años se presenta públicamente el primer resto humano, una falange, junto con una serie de supuestas industrias líticas sobre hueso que despiertan un interés añadido al yacimiento. En 1984 se inician campañas de excavación con cierta regularidad, dirigidas por el Dr. José Gibert, que año a año van incrementando la colección de vertebrados fósiles. En los años 1985 a 1999 se publican varios estudios sobre la fauna de Cueva Victoria, interpretaciones de su edad, estudios anatómicos de la falange humana y el descubrimiento de Theropithecus. También se publican nuevos mode-los sobre la dispersión de mamíferos en el Pleistoceno inferior que destacan la importancia del estrecho de Gibraltar como ruta alternativa a la dispersión de África a Europa, sustentados por la fauna fósil de Cueva Victoria y también de los yacimientos de Orce, situados a unos escasos 150 km. A partir de 2008, gracias a la financiación de la Consejería de Cultura, el Consorcio Sierra Minera y el Ayuntamiento de Cartagena, las excavaciones dan un salto cualitativo, ya que se instala un andamio con el que se puede acceder a la parte superior de la brecha de relleno, la más rica en fósiles, pero situada a varios metros del suelo. El andamio permite por primera vez un trabajo completo y detallado, iniciándose una excavación sistemática y metodológica, cartografiando los fósiles para obtener también información tafonómica. A partir de ese momento se añaden piezas importantes a la colección situadas en un contexto estratigráfico y tafonómico, entre ellas nuevos restos de Theropithecus, que se publican en el Journal of Human Evolution. Gracias al andamio se puede también muestrear la pared a diferentes niveles estratigráficos para llevar a cabo un estudio paleomagnético, así como realizar dataciones radiométricas en el espeleotema superior. Los resultados permiten refinar la edad de la

Luís Gibert Beotas y Carles Ferràndez Cañadell

PresentaciónForeword

12

asociación fósil, situándola entre 850.000 y 900.000 años, coincidiendo con la primera gran caída del nivel del mar que tiene lugar en el Cuaternario, hecho que refuerza las hipótesis de una dispersión de fauna de África a Europa a través de Gibraltar. A partir de 2009 se invita a paleontólogos especialistas en diversos grupos de vertebrados fósiles, así como a geólogos de distintas disciplinas, a visitar la cueva y a participar en el estudio del yacimiento y su fauna. De esta colaboración surge una serie de estudios que amplían notablemente el conocimiento de la asociación de vertebrados fósiles de Cueva Victoria, así como de la formación y la edad del yacimiento. Este volumen reúne los trabajos fruto de esta colaboración y pretende ser una actualización del conocimiento sobre Cueva Victoria en los diversos ámbitos de la geología y la paleontología.

Esta monografía está dividida en dos partes, en una primera parte se tratan temas de la geología de Cueva Victoria: la historia de las labores mineras (M. A. Pérez de Perceval, J. I. Manteca y M. A. López-Morell), las mineralizaciones de hierro y manganeso (J. I. Manteca y R. Piña; D. Artiaga, L. Gibert y J. García-Veigas); la datación de los espeleotemas y su interpretación paleoclimática (A. Budsky, D. Scholz, L. Gibert y R. Mertz); la espeología (A. Ros y J. L. Llamusí); la edad del yacimiento a partir de datos paleomagnéticos (L. Gibert y G. R. Scott), y los estudios geofísicos para modelizar tridimensionalmente la cueva y para descubrir nuevas cavidades (A. Espín de Gea, A. Gil Abellán y M. Reyes Urquiza).

A continuación, dos capítulos enlazan la geología con la paleontología, con estudios sobre la formación del yacimiento y de las acumulaciones de restos fósiles (C. Ferràndez-Cañadell, J. Vilà Vinyet e Í. Soriguera). Los siguientes capítulos están dedicados a los diferentes grupos fósiles. Se estudian los anfibios y reptiles (H.-A- Blain; A. Pérez-García, I. Boneta, X. Murelaga, C. Ferràndez-Cañadell y L. Gibert), los arvicólidos (R. A. Martin), los quirópteros (P. Sevilla), los insectívoros (M. Furió), las aves (A. Sánchez Marco), los cérvidos (J. Van der Made), los caballos (M. T. Alberdi y P. Piñero), los rino-cerontes (J. Van der Made), los elefantes (M. R. Palombo y M. T. Alberdi), los cánidos (M. Boudadi-Maligne), los úrsidos, hiénidos y félidos (J. Madurell-Malapeira, J. Morales, V. Vinuesa y A. Boscaini), los primates (F. Ribot, C. Ferràndez-Caña-dell y L. Gibert), y se acaba con un repaso a los grupos pendientes de estudio o revisión (C. Ferràndez-Cañadell) y un trabajo sobre la preparación y restauración de los restos fósiles (A. Gallardo).

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AGRADECIMIEnToS

Agradecemos, en primer lugar, a todos los autores su esfuerzo y dedicación para aportar capítulos de calidad a esta monografía y les pedimos disculpas por el retraso sufrido en la publicación. En segundo lugar, agradecemos a todas aquellas personas e instituciones que han colaborado de forma directa o indirecta para que esta monografía sea una realidad: a todo el personal del Museo Arqueológico de Cartagena y especialmente a María Comas Gabarrón, Directora del Museo Arqueológico Municipal Enrique Escudero de Castro durante los últimos años y ahora Directora General de Bienes Culturales; a Miguel Martínez Andreu, quien siempre nos mostró su apoyo, tanto en su etapa de Director del Museo Arqueológico como en la de investigador, y a Miquel Martín Camino, investigador del Museo de Arqueológico de Cartagena y miembro del consejo de redacción de MASTIA, que nos ha prestado su ayuda en la etapa de edición de este volumen. Nuestra sincera gratitud al Ayuntamiento de Cartagena, especialmente a Pilar Barreiro Álvarez, alcaldesa de Cartagena; a los concejales del Ayuntamiento de Cartagena que se han implicado en el proyecto de Cueva Victoria, María Rosario Montero Rodríguez, Nicolás Ángel Bernal y Carolina Beatriz Palazón. Expresamos nuestro agradecimiento a los técnicos y responsables de la Dirección General de Bienes Culturales, Miguel San Nicolás del Toro, Manuel Lechu-ga Galindo, Jefe de Servicio de Museos y Exposiciones y especialmente a Gregorio Romero Sánchez, paleontólogo y técnico del Servicio de Patrimonio, por animarnos desde el primer momento en esta iniciativa.

A los miembros del Centro de Estudios de la Naturaleza y el Mar de Cartagena (CENM), nuestra más sincera gratitud a Andrés Ros y José Luis Llamusí, que nos han apoyado y dado asesoramiento técnico sobre cuestiones de seguridad en la cavidad y han colaborado de forma muy activa en las diferentes jornadas de puertas abiertas celebradas en los últimos años. Nuestra especial agradecimiento a Ignacio Manteca Martínez de la Universidad Politécnica de Cartagena y compañeros de Departamento de Ingeniería Minera, Geológica y Cartográfica por su interés y apoyo en todos los aspectos geológicos y patrimoniales de Cueva Victoria, así como a Mariano Mateo y los miembros de la Asociación de Vecinos del Llano del Beal, por su ayuda y apoyo al proyecto de investigación. También a todos los colegas y voluntarios que han participado de forma altruista en las excavaciones a lo largo de estos años, especialmente a Alfredo Iglesias, Julià Gonzàlez, Florentina Sánchez, Fernando González y a nuestras compañeras Emma La Salle y María Lería por su ayuda y paciencia durante tanto tiempo. A Pepa Beotas, Patxu Gibert y Blanca Gibert por ayudarnos y compartir tantas campañas en Cueva Victoria.

Finalmente, queremos dar las gracias a todas aquellas instituciones que han apoyado las investigaciones de Cueva Vic-toria en estos últimos 30 años: Consejería de Cultura de la Región de Murcia, Ayuntamiento de Cartagena, Universidad de Barcelona, Universidad Politécnica de Cartagena, EarthWatch Institute y Diputación de Barcelona.

Este trabajo es una contribución al Grup de Recerca Consolidat 2014 SGR 251 Geologia Sedimentària de la Generalitat de Catalunya y al Programa Ramón y Cajal del Ministerio de Economía y Competitividad del Gobierno de España.

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DEDICAToRIA

“Success is not final, failure is not fatal: it is the courage to continue that counts” (El éxito no es definitivo, el fracaso no es fatídico. Lo que cuenta es el valor para continuar)

Winston Churchill

Dedicamos este volumen al Dr. José Gibert Clols, director de las investigaciones en Cueva Victoria desde 1984 hasta su prematura muerte en 2007. José Gibert es para nosotros un ejemplo de pasión por el conocimiento, tenacidad, ho-nestidad y profesionalidad. Realizó su última campaña en Cueva Victoria en verano de 2007, pero no la pudo terminar. Después de ser atendido en el Hospital de Cartagena ese verano fue finalmente ingresado en un hospital de Barcelona, delegando en nosotros la responsabilidad de continuar el trabajo y cerrar la campaña en la fecha prevista del 31 de septiembre, así lo hicimos. Moriría una semana después, el 7 de octubre de 2007, dejándonos un gran legado y una gran responsabilidad.

Cueva Victoria fue un lugar donde José Gibert trabajó con pocos recursos pero con mucha dedicación y libertad. Duran-te los 23 años que estuvo al frente de las investigaciones se sintió querido y apoyado por la sociedad civil, académica y administrativa del conjunto de la Región de Murcia. Los que tuvimos el privilegio de trabajar junto a él sabemos que fue una persona excepcional, con una gran vocación y calidad humana. A principios de los años ochenta, su trabajo y descubrimientos en el Sureste de la Península Ibérica, en Orce y Cueva Victoria, le permitieron establecer nuevas teorías que quebrantaban el viejo paradigma de la ocupación tardía de Europa por el Hombre. José Gibert propuso, de manera pionera, que la humanidad llegó a Europa cerca de un millón de años antes de lo establecido en aquel momento, proponiendo además que esa migración se hizo por Gibraltar en lugar de rodeando el Mediterráneo. Después de una euforia inicial generalizada, su trabajo fue duramente criticado de forma poco rigurosa. No obstante, la presencia de fauna africana en Cueva Victoria junto a homínidos avalan esa idea, y nuevos hallazgos en Orce y en otros yacimientos han supuesto que, 30 años después, nadie dude de que la ocupación de Europa fue muy temprana. Por otro lado, nuevos hallazgos y las mejoras en las técnicas de datación han determinado que las primeras evidencias de presencia humana en Europa con industria lítica de tipo olduvaiense y los primeros vestigios también en Europa de industria achelense se hallan en el sureste de la Península Ibérica (en Orce y en Cueva Negra del Río Quípar, Caravaca). Estos hechos, junto a la presencia de primate africano Theropithecus en Cueva Victoria, única en Europa, apoyan de manera más convincente la hipótesis de que durante el Pleistoceno inferior se dieron varias dispersiones desde África hacia Europa a través de Gibraltar.

Sin duda, José Gibert estaría hoy muy satisfecho no sólo por ver que sus ideas se van consolidando sino también por ver editado este volumen especial de MASTIA dedicado a Cueva Victoria, donde se integran y actualizan todos los re-sultados de las investigaciones realizadas en este lugar excepcional. Creemos que este volumen es parte de su legado pues sin su dedicación a Cueva Victoria, esta monografía no existiría.

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DR. JoSé GIBERT CLoLS (1941-2007)

La trayectoria profesional y figura humana de José Gibert Clols destacan desde muy pronto y en diferentes aspectos. Durante el bachillerato fue un estudiante brillante, obteniendo 23 matrículas de honor en el colegio de los Agustinos de Zaragoza. Su carrera universitaria en Ciencias Geológicas en la Universidad de Barcelona se vio truncada por la muerte de su padre a mitad de los estudios, teniéndose que responsabilizar de la familia y del negocio familiar. Aun así, consiguió Matrícula de Honor en Paleontología, disciplina que siempre le interesó especialmente. Una vez licenciado en 1968, inició su tesis doctoral, bajo la dirección del Dr. Miquel Crusafont, sobre los insectívoros fósiles de España. Consiguió una beca para realizar el doctorado de la Fundación Juan March, que le facilitó colaborar con centros ex-tranjeros, especialmente franceses y holandeses. De esta colaboración aprendió nuevas técnicas, que se aplicaron por primera vez en España en la investigación de micromamíferos y publicó varios estudios en revistas internacionales. En 1971 fue profesor ayudante de Paleontología Humana en la Universidad de Barcelona. Una vez doctorado en 1973, compaginó su labor investigadora en el Instituto de Paleontología de Sabadell con la docencia de enseñanza media, en la que alcanzó el grado de Catedrático de Ciencias Naturales. En 1976 vio la necesidad de desarrollar la investigación en paleontología del Cuaternario Ibérico. Para ello organizó, desde el Instituto de Paleontología, una campaña de pros-pección en la cuenca de Guadix-Baza en Granada, donde consideró que existía un gran potencial fosilífero. Después de planificar esa prospección por los sectores que juzgó con mayores posibilidades para la localización de yacimientos fosilíferos, descubrió el yacimiento de Venta Micena, probablemente el yacimiento del Pleistoceno Inferior europeo

José Gibert Clols en 2005

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más rico y extenso que se conoce. Durante 1982 organizó una campaña de excavaciones e identificó un fragmento de cráneo que clasificó como humano. Este hallazgo rompió el paradigma establecido, al proponer la presencia humana en el Sur de Europa cerca de un millón de años antes de lo establecido. Como todos los hallazgos revolucionarios, este fósil generó una polémica que se inició al morir el Dr. Crusafont, la mayor autoridad en paleontología de vertebrados en España y avalador de la humanidad del fósil.

José Gibert afrontó el problema basándose en el poder resolutivo del método científico y enfocándolo desde una pers-pectiva pluridisciplinar, estableciendo colaboraciones con distintos especialistas, incluyendo científicos en el innovador campo de la bioquímica aplicada a la paleontología. Los resultados fueron concluyentes, al detectarse, en laboratorios de España y Estados Unidos, proteínas humanas en los fósiles cuestionados y encontrar, en cráneos humanos infanti-les actuales, los caracteres anatómicos cuestionados en el cráneo fósil. De forma paralela, fueron identificados nuevos fósiles humanos, así como industrias líticas, que aportaron evidencias complementarias de la presencia de homínidos en el Pleistoceno inferior de Orce. El descubrimiento de la falange de Cueva Victoria en 1984 por Juan Pons supuso un apoyo importante a la teoría de una ocupación humana antigua de la Península y la asociación de ese fósil con primates africanos avaló la idea de una dispersión por Gibraltar. Entre 1986 y 1993, José Gibert publicó y divulgó los resultados de estas investigaciones por todo el mundo, dando a conocer Orce y Cueva Victoria a la comunidad científica internacional. Este ejercicio le permitió organizar un Congreso Internacional de Paleontología Humana en Orce en 1995, en el que participaron más de 300 especialistas de 18 países y que incluyó una visita a Cueva Victoria, generándose un debate fructífero sobre las vías de colonización y las edades de las primeras ocupaciones humanas en Europa. Orce y Cueva Victoria pasaron a ser lugares de referencia en el mundo de la paleontología humana. Habían pasado 13 años desde el descubrimiento y los datos y la comunidad científica le daba al fin la razón. A partir de ese momento álgido, su carrera en Orce entra la etapa más difícil, al ser excluido de la excavación e investigación de los yacimientos por él descubiertos. Sin embargo, lejos de abandonar Orce, José Gibert se interesó por otras localidades fosilíferas de la zona, como Barranco del Paso y Fuentenueva-1, estableciendo nuevas colaboraciones que le permitieron resolver la edad del conjunto de yacimientos de Orce. Al mismo tiempo, intensificó sus investigaciones en Cueva Victoria hasta el momento que fueron interrumpidas por su prematura muerte.

El Dr. José Gibert publicó 181 artículos (52 de ellos en revistas internacionales), 2 libros y ha sido editor o coeditor de 6 monografías. La hipótesis de que la presencia humana más antigua de Europa se sitúa en el Sur de la Península Ibérica hace 1,3 millones de años fue provocadora y revolucionaria en 1982, pero gracias a sus investigaciones y perseverancia ha sido suficientemente demostrada y está plenamente establecida y aceptada en la actualidad.

Durante su carrera, el Dr. José Gibert Clols recibió los siguientes premios y distinciones por su trabajo:

• 1983 Premio de la Generalitat de Catalunya a la innovación pedagógica en Ciencias Naturales.• 1985 Premio al Vallesano del año, modalidad Ciencia.• 1986 Concesión por el Excmo. Ayuntamiento de Orce del título “Hijo Adoptivo”• 1998 Premio Narciso Monturiol a la Investigación Científica (Colectivo al Inst. Crusafont) de la Generalitat de Ca-

talunya.• 2000 Insignia de Oro del Colegio de Ingenieros Técnicos de Minas de Cartagena.• 2001 Cartagenero del siglo XX, Excmo. Ayuntamiento de Cartagena.• 2005 Medalla Narciso Munturiol al Mérito Científico y Técnico concedida, a título personal, por la Generalitat de

Catalunya.• 2007 Insignia de Plata del Colegio de Ingenieros de Minas de Cartagena.• 2007 Premio nacional El Vallenc (Ayuntamiento de Valls), modalidad Ciencia.• 2010 Medalla de la Vila a título póstumo, Castellar del Vallés.• 2013 El ayuntamiento de Mora d’Ebre le dedica la Semana Cultural.• 2014 Medalla de Oro de la provincia de Granada, Diputación de Granada.

MASTIA 11-12-13, 2012-14, PP. 207-225ISSN: 1579-3303

Robert A. Martin*

A brief review of the Spanish archaic Pleistocene arhizodont voles

Breve revisión de los topillos arrizodontos arcaicos de España

* Department of Biological Sciences, Murray State University, kY, USA. Robert.martin@murraystate.edu

AbstractThe recent recognition of Chaline’s vole as the monotypic member of the new Pleistocene genus Victoriamys, appar-ently descended from a Mimomys ancestor distinct from that which gave rise to other voles with arhizodont (rootless) molars, naturally leads to the speculation that other Spanish Pleistocene arhizodont voles may have originated from distinct Mimomys ancestors. This hypothesis is tentatively accepted for the Spanish Pleistocene genera Microtus, Arvi-cola, Tibericola, Iberomys and Victoriamys. A new phylogenetic scenario is provided for Arvicola within which only two species, A. sapidus and A. terrestris, are recognized. The presence of an m1 morphotype inseparable from that of I. brecciensis and other m1 morphotypes from Huéscar not normally associated with Iberomys is puzzling and allows for a number of competing hypotheses regarding systematics of the Huescar-1 sample and origin of Iberomys in Spain.

Key WordsPleistocene, arvicolid, vole, Microtus, Allophaiomys, Iberomys.

ResumenLa reciente caracterización del topillo de Chaline como miembro monotípico del nuevo género pleistoceno Victoriamys, que aparentemente desciende de un antecesor Mimomys distinto al que dio origen a otros arvicolinos con molares arrizodontos (sin raíces), lleva a especular sobre el origen de otros especies arrizodontas del Pleistoceno español pue-dan haberse originado de distintas especies ancestrales de Minomys. Esta hipótesis se acepta de manera provisional para los géneros españoles del Pleistoceno Microtus, Arvicola, Tibericola, Iberomys y Victoriamys. Se presenta un nuevo escenario filogenético para Arvicola, en el cual sólo se reconocen dos especies, A. sapidus and A. terrestris. La presencia de un morfotipo de m1 indiferenciable del de I. brecciensis y otros morfotipos de m1 de Huéscar normalmente no asociados a Iberomys es desconcertante y da pie a varias hipótesis alternativas respecto a la sistemática de la muestra de Huéscar-1 y el origen de Iberomys en España.

Palabras ClavePleistoceno, arvicólidos, topillos, Microtus, Allophaiomys, Iberomys.

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InTRoDUCTIon

One might think that the Pyrenees Mountains in north-ern Spain would present a formidable barrier to the dis-persal of small mammals into the Iberian Peninsula, but gauging by the Spanish fossil record of arvicolid rodents this does not seem to be the case. Many species from a variety of arvicolid genera have been recorded from Pli-ocene and Pleistocene deposits (Cuenca-Bescós et al., 2010a). Nevertheless, there appears to have been a sig-nificant difference in arvicolid diversity between northern and southern Spain during the Pleistocene that may be unrelated to sampling bias. This difference is particularly pronounced when we compare arvicolid species diver-sity from the Atapuerca sequence of north-central Spain to that of the Baza Basin and Cueva Victoria in the south (Agustí et al., 2010; Cuenca-Bescós et al., 2010a; Mar-tin, 2012a). It is difficult to explain this dichotomy by taphonomic bias, and it may be that the difference in diversity is related to environmental zonation during the Pleistocene, with more habitat complexity to the north

than the south, perhaps mediated by climatic patterns that mantained a more arid, Mediterranean climate to the south. Despite these differences, the combination of arvicolid records from the rich Atapuerca sequence with those of the Baza Basin, where a number of critical fossil assemblages are stratigraphically superposed and supplemented by a detailed paleomagnetic (pmag) his-tory (Gibert et al., 2007; Cuenca-Bescós et al., 2010a; Martin, 2012a), allows the construction of an arvicolid biochronological Pleistocene sequence in Spain that is unparalleled for western Europe (Fig. 1).

Until recently, one of the most contentious issues in the establishment of the Spanish arvicolid sequence was the phylogenetic and taxonomic position of a va-riety of small and middle-sized Pleistocene voles with arhizodont (rootless) molars. Initially, most of these species were referred to the taxon Allophaiomys (kor-mos, 1933), considered either as a genus or subgenus of Microtus. Thus, we had A. pliocaenicus, A. chalinei and A. vandermeuleni. In the Levant, we also had the

Fig. 1. Arvicolid rodent biochronology for some Spanish Pleistocene localities. Abbreviations: AZ = arvicolid zones; BC = Barranco de los Conejos; BP = Barranco del Paso; VM = Venta Micena; BL = Barranco León D (=5); FN 3 = Fuentenueva 3; LRU = Trinchera Elefante, Lower Red Unit, Atapuerca sequence; LQ = Loma Quemada; Hu1 = Huéscar 1; CN = Cueva Negra; CV = Cueva Victoria; D5 = level D5, Cal Guardiola; TD3-8a = Trinchera Dolina, levels 3-8a, Atapuerca sequence; TD8b = Trinchera Dolina, level 8b, Atapuerca sequence; CB1 = Cullar Baza 1; TD10 == Trinchera Dolina, level 10, Atapuerca sequence; o = assumed present but not recorded; x = present; cf = compares favorably with, aff = has affinities with.

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troublesome middle Pleistocene Microtus jordanica from ‘Ubeidiya, originally questionably referred by Haas (1966) to Arvicola and later named as a distinct sub-genus of Microtus, Tibericola, by von koenigswald et al. (1992). Recently, Spanish investigators also recog-nized an extinct early Pleistocene species of Arvicola, A. jacobaeus, perhaps descended from Mimomys savini (Cuenca-Bescós et al., 2010b). With the exception of Nadachowski (1991), throughout the early 1990s Eu-ropean arvicolid specialists continued to think of small Pleistocene voles with relatively simple, rootless molars except Arvicola as derived from a common Mimomys ancestor. But problems persisted with this interpretation, as the only potential ancestor for all these arhizodont species in a somewhat morphological and chronolog-ical transitional position was Microtus (Allophaiomys) deucalion, a diminutive species with an essentially Mi-crotus-like dental morphology, but with the negative enamel differentiation of Mimomys (van der Meulen, 1974). The three species M. jordanica, A. chalinei and A. vandermeuleni displayed considerably larger molars than M. deucalion and the average occlusal patterns of the former three species tended to be atypical for either M. deucalion or A. pliocaenicus. What did this imply?

Two investigators, Nadachowski (1991) and Ünay et al. (2001) at least partially understood the problem. Nad-achowski recognized the similarity in dental pattern of “Microtus” jordanica and the extant Chionomys, and suggested an origin of Chionomys through “M.” jordan-ica, separate from other Microtus-like living voles. Thus, in one scenario, Chionomys was not directly descended from either Microtus deucalion or M. pliocaenicus, but from a ghost lineage leading from another species of Mimomys. Likewise, Ünay et al. (2001) realized that “Al-lophaiomys” chalinei, “A.” vandermeuleni and A. jordan-ica were likely distinct from Microtus, and they referred the three species to Tibericola, treated as a distinct genus, descended from M. deucalion. They may have been wrong about “T.” chalinei, but they were on the right track. More recently, Gloria Cuenca-Bescós (Cuen-ca-Bescós et al, 1995 and other papers), by recognizing Iberomys as a distinct genus, also realized the possible polyphyletic origin of some Microtus-like arvicolids. In this essay I will briefly review the archaic Spanish Pleis-tocene voles with rootless molars in the context of a dynamic period in which a number of Mimomys species were experimenting with arhizodonty.

One of the most significant difficulties in separating early generic clades of arhizodont species is identifying syn-apomorphies. How, for example, does one separate the earliest Arvicola from the earliest Microtus, especially since early Arvicola (A. jacobaeus) displays a simple m1 and negative enamel differentiation like M. deucalion? This difficulty led Ruiz Bustos (1999) to develop an elaborate dental evolutionary scheme in which he pro-posed that M. deucalion was a small Arvicola, ancestral to extant Arvicola through Allophaiomys chalinei, which he considered also as an Arvicola. The recent identifica-tion of A. chalinei as the distinct genus Victoriamys (see below) negates Ruiz-Bustos’s ingenious hypothesis, but it also demonstrates how similar these taxa are in dental morphology. The answer to this problem seems to be that we need to branch out to molars other than the m1 and to mandibular anatomy. For instance, as noted by Gibert et al. (2007) and Martin (2012a), all known species of Arvicola have a complex m3 (with distinct T4) often mimicking the morphology of m2. Although this morphology may also be seen in some Mimomys species and Victoriamys, it is not seen in Microtus, in which T4 is usually reduced or absent. As shown by the position of the mandibular foramen in Victoriamys (Mar-tin, 2012a), helpful characters are also found on the mandible. We will need all of these characters and more to tease out relationships among the early Pleistocene voles.

TERMInoLoGY AnD ABBREVIATIonS

Arvicolid molar occlusal morphology (Fig. 2) follows van der Meulen (1973). Lower molars are designat-ed by lower case, uppers by upper case letters; acc = anteroconid complex, acd = anteroconid of first lower molar (m1). Anterior and posterior enamel edges of triangles, as seen in occlusal morphology under the dis-secting microscope, may display distinct differences in thickness (be differentiated). The term negative enamel differentiation applies if the posterior (trailing) edges in the lowers are thicker than the anterior (leading) edges (opposite in the uppers- anterior edges thicker). Most Mimomys dentitions display negative differentiation. Positive enamel differentiation describes the condi-tion in which the posterior edges are thinner than the anterior edges in the lowers (opposite in the uppers). Most Microtus species have positive differentiation. A few extinct and extant arhizodont voles display the un-differentiated condition, in which anterior and posterior

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edges are of equal thickness. As shown by Martin et al. (2006), molars that appear undifferentiated under the light or dissecting microscope may actually be slightly differentiated if edges are measured under the electron microscope. Rhizodont molars possess roots; arhizodont molars are rootless and evergrowing. Hypsodont = high-crowned, hypselodont = high-cowned and rootless.

Allocation of fossil samples to appropriate taxa requires a consistent methodology and a stable and comprehen-sible taxonomic platform. I agree with modern phyloge-netic systematists that a classification should represent phylogeny and I further agree that the search for synapo-morphies, or shared derived characters, is fundamental to that process. In this approach, an evolutionary grade

Fig. 2. Molar occlusal surface terminology of Ondatra zibethicus (after Hibbard [1950] and van der Meulen [1973]).AC(D) = anterior cap or anteroconid, ACC = anteroconid complex (all structures on m1 anterior to T3), AL = anterior lobe (loop), BRA = buccal re-entrant angle (fold), BSA = buccal salient angle, LRA = lingual re-entrant angle, LSA = lingual salient angle, PC = posterior cap, PL = posterior lobe (loop), TTC = trigonid-talonid complex. The re-entrant angles are sometimes referred to as synclines, and the salient angles as anticlines.

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seems to be untenable, representing as it does simply a stage in morphological evolution that all related taxa passed through on their way to expressing their crown position endpoint morphologies (character sets). Thus utility and validity of the genus Allophaiomys, coined by kormos (1933) for extinct European voles with a sim-ple m1 with three triangles and a variable anteroconid, becomes debatable, as the simple m1 is, in cladistic par-lance, a symplesiomorphy and of negligible taxonom-ic value. The Allophaiomys concept was examined by Martin and Tesakov (1998), and they contrasted two alternatives in which Allophaiomys was considered: 1) a valid taxon, representing as it did during the ear-ly Pleistocene a set of genetically closely related spe-cies, regardless what clades they would later generate and 2) an invalid taxonomic wastebasket, defined by a set of dental symplesiomorphies and our inability to recognize the correct relationships of its members. Pure cladistic protocol demands we follow 2) and reject Al-lophaiomys, but the biological reality is that a species is both a member of a clade in its own time and, if it is subsequently shown to be ancestral to a later clade, a member of the clade to which it is ancestral. In a sense, it is a reflection of a temporal paradox, the old problem of the time traveler who cannot, theoretically, be in two places simultaneously. Despite my earlier conclusions to the contrary, Allophaiomys seems like a reasonable sub-genus of early Pleistocene Microtus, reflecting a number of genetically closely related species (during their time) with simple molars that cannot be clearly allocated to an extant subgenus.

SPECIES REVIEW

The archaic Spanish Pleistocene arhizodont voles are reviewed below. Generic level features are considered first, followed by characters, localities, temporal range, and brief remarks for each species.

Genus MICROTUS Schrank, 1798

The most recent summary of the early evolution of Eu-ropean Microtus is that of Maul et al. (2007). These authors presented a careful evaluation correlating the appearance of various dental morphologies with pale-omagnetic information. They bypassed providing a dis-tinctive dental definition for Microtus, and even noted (2007, p. 245) that the most primitive Microtus m1 re-sembles that of “a small Arvicola.” The early members of

Microtus from western Europe, such as M. deucalion, M. pliocaenicus, M. nutiensis, M. burgondiae and M. lavo-cati are small, and it seems likely be that these species originated from a small Mimomys. In addition to relative-ly small size, Microtus is recognized in western Europe by the following dental characters: 1) molars rootless, 2) crown cementum abundant; 3) relatively thin enamel, 4) m3 simple (reduced T4), 5) mandibular foramen not a thin slit on posterior edge of mesial side of ascending ramus (tends to be more centrally located). The latter feature is usually coupled with a relatively long lower incisor that displaces the mandibular foramen mesially. Negative enamel differentiation is found in the extinct Microtus deucalion (van der Meulen, 1974), the extinct M. pliocaenicus (Hír, 1998) and extant M. umbrosus and occasionally in extant M. guatemalensis (Martin, 1987). The enamel may be undifferentiated in the living M. oaxacensis (Martin, 1987). Most other extinct and extant species of Microtus display positive differentiation (Nadachowski and Zagorodnyuk, 1996). Consequently, while the majority of Microtus species possess positive enamel differentiation, if species with negative differen-tiation are to be included within Microtus, then we can-not use positive enamel differentiation as a character to define the genus. The most we can say, which appears to be well substantiated in the literature (Maul et al., 2007), is that most Microtus have positively differenti-ated molars and are descended from an ancestor such as M. deucalion with negative differentiation. From this concept we must also conclude that extant species such as M. umbrosus, M. guatemalensis and M. oaxacensis probably are not descended from M. pliocaenicus, the latter which usually includes a large proportion of molars with positive differentiation (Hír, 1998; Martin, 1998). Although the North American pine vole, Pitymys pineto-rum, has positively differentiated enamel, its early Pleis-tocene relative P. cumberlandensis (van der Meulen, 1978) had thick, undifferentiated enamel and P. pin-etorum must, by definition, also have evolved from an early Pleistocene ancestor other than M. pliocaenicus. For additonal reasons, Martin (2012b) followed van der Meulen (1978) in considering the North American pine voles as the distinct genus Pitymys.

With the identification of new Microtus (Allophaiomys) species from Siberia such as M. (A.) tchumakovi and M. (A.) reservatus (Zazhigin, 1998), we face the distinct possibility that the various recognized extinct and extant subgenera of Microtus originated from more than one

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species of Microtus with negatively differentiated or un-differentiated enamel. Some Microtus subgenera may have also arisen at larger size than those in western Eu-rope, as M. reservatus has a mean m1 length of 2.91 mm (2.35-3.50 observed range). If these species origi-nated from the same basal ancestral Mimomys species then Microtus can still be considered monophyletic, but with the recognition of Iberomys and Victoriamys, that concept must be constantly tested.

MICROTUS (ALLOPHAIOMYS) PLIOCAENICUS (kormos 1933)

Dental and mandibular characters: size small (mean length m1 <2.85 mm); samples often include various combinations of negatively differentiated, positively dif-ferentiated or undifferentiated enamel; m1 with three triangles and a simple acd, T4-5 usually widely confluent with each other and the anterior loop; T6-7 rarely de-veloped on m1; M3 simple (T4 usually confluent with posterior loop); enamel relatively thin.

Localities: Cañada de Murcia (Sesé, 1989); Venta Mi-cena (Ruiz Bustos, 1993).

Age in Spain: ~1.4 Ma (this study, Fig. 1).

Remarks: Relatively simple first lower molars at the Mi-crotus pliocaenicus grade have been reported from a number of early Pleistocene localities in Spain, but most are advanced in morphology beyond M. plioacenicus from the type locality, Betfia 2 in Roumania (Hir, 1998).

MICROTUS (ALLOPHAIOMYS) LAVOCATI (Laplana and Cuenca-Bescós, 2000)

Dental and mandibular characters: size small (mean length m1 <2.85 mm); enamel differentiation positive; T4 often closed from posterior loop in M3; enamel rel-atively thin. Differs from M. pliocaenicus in the following m1 characters: 1) constriction of dentine isthmus be-tween T5 and the anteroconid resulting in confluent T4-5, 2) tendency of internal enamel borders of BRA3 and LRA4 to form elongated neck connecting T4-5 to acd, 3) occasional appearance of incipient T6.

Localities: Lower Red Unit of Trinchera Elefante (TEL-RU), Atapuerca (Laplana and Cuenca-Bescós, 2000); Cal Guardiola (Vallparadís), Layer D3 (Minwer-Barakat

et al., 2011); El Chaparral (López-García et al., 2012); Barranco León (Agustí and Madurell, 2005; Gibert et al., 2007), Fuentenueva 3 (Agustí and Madurell, 2005).

Age in Spain: ~1.3 (1.2 Ma; cosmogenic nuclide date for TELRU; Carbonell et al., 2008) - ~ 0.79 Ma (López-García et al., 2012).

Remarks: In its small size, relatively thin, positively dif-ferentiated enamel, and relatively simple m1 and M3, M. lavocati shows close relationship to, and likely deriva-tion from, M. pliocaenicus. The recent addition of both Microtus burgondiae and M. nutiensis (both reported as Allophaiomys) to the rodent contingent from TELRU at Atapuerca (Cuenca-Bescós et al. 2010a, b, 2013) introduces a new taxonomic/phylogenetic conundrum to the history of Spanish early Pleistocene arhizodont arvicolids.

M. burgondiae was first named as Microtus (Suranomys) malei burgondiae by Chaline (1972) for some m1s from the French Pleistocene locality Les Valerots. Chaline (1972) also segregated out other m1s from the same site and named them M. (Allophaiomys) pliocaenicus nutiensis. No arvicolid experts today would use these names for the specimens in question, but the names have been transformed into species epithets describ-ing various samples of European Pleistocene Microtus m1s somewhat advanced in complexity from the type series of M. pliocaenicus from Betfia 2 in Roumania (Hir, 1998). For instance, Sesé (1989) tentatively referred a specimen from Puerto Lobo, a site in the Baza Basin, to M. (A.) cf burgondiae. Agustí and Madurell (2005) later referred the material from Puerto Lobo as well as that from Fuentenueva 3, Baranco León 5 and Loma Que-mada in the Baza Basin to Allophaiomys lavocati. Gibert et al. (2007) questioned the reference of the Fuent-enueva 3 Microtus to A. lavocati, and allocated the ma-terial to M. cf pliocaenicus. Agustí and Madurell (2005) also referred some Microtus specimens from other Baza Basin localities such as Venta Micena, Cañada de Murcia and Fuente Nueva 2 to Allophaiomys ruffoi, a species named from the Italian Cava Sud by Pasa (1947). Gib-ert et al. (2007) later conservatively referred the Venta Micena specimens to M. cf pliocaenicus.

There are so many questions with all this material it will take a Ph. D. thesis to sort it out. For instance, how many species are actually present in the Les Valerots Micro-

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tus material? There are specimens illustrated by Chaline (1972; e.g., illustrations 2, 3 and 13 in fig. 24) allo-cated to M. (Allophaiomys) pliocaenicus nutiensis that include M. pliocaenicus, M. lavocati and M. (Stenocra-nius) hintoni/gregaloides morphotypes. The specimens Chaline illustrates as M. (Suranomys) malei burgondiae (Chaline, 1972, fig. 24; illustrations 14-24) are morpho-logically similar, with T4-5 confluent, T6 undeveloped, and an incipient T7, but they bear little resemblance to the much more complicated specimens referred to the same taxon in Fig. 25, many of which would fit easily into a number of Terricola species.

The identification of M. burgondiae, M. nutiensis and M. ruffoi from various Spanish sites certainly shows that m1 morphotypes advanced over those of typical M. pliocae-nicus appeared in Spain during the early Pleistocene, but what taxonomic name should be affixed to them? Is sympatry of three closely related species of early Mi-crotus from TELRU likely, or are the Microtus m1 mor-photypes from that site variants of the single species M. lavocati? Not having studied most of the material, I have no answer to the questions raised. The m1 characters of M. lavocati as initially described seem distinct, but if they were chosen from a wider expression of variation that also encompasses one of Chaline’s (1972) taxa, then Chaline’s name could have priority. However, as noted above, the Les Valerots samples need further evaluation in order to finally determine the number and nature of the Microtus present in that site. For now, as the Fuent-enueva 3 and Barranco León D m1s are advanced over those of typical M. pliocaenicus, a temporary residency in M. lavocati seems appropriate until all the systemtic details can be unwound.

Genus ARVICOLA Lacépède, 1799

Arvicola is a geographically widespread genus during the Pleistocene, with representatives reported from western Europe through the Ukraine (Rekovets and Nadachowski, 1995; Maul et al., 2000; Cuenca-Bescós et al., 2010b). It is interesting that this genus has so long been considered separate from early Microtus (Al-lophaiomys) because the m1 occlusal pattern of most Arvicola species is similar to that of archaic Microtus. I suspect the reason was predominantly because: 1) of large size of fossil Arvicola molars, 2) the earliest fossil Arvicola and some modern populations of A. sapidus retain negative enamel differentiation, and 3) the almost

identical size and dental pattern to Arvicola is seen in the extinct European Mimomys savini, long considered the ancestor for Arvicola. For many years, the fossil re-cords of Mimomys savini and the earliest Arvicola were consecutive, and it was generally accepted that at least one population of M. savini evolved into Arvicola (A. cantianus = A. mosbachensis) during the middle Pleis-tocene (Maul et al., 2000). We now understand that this evolutionary hypothesis is wrong, as an early Pleistocene species of Arvicola, A. jacobaeus, is now known (Cuen-ca-Bescós et al., 2010b). In the Atapuerca sequence of north-central Spain, Arvicola jacobaeus actually appears before Mimomys savini, though the latter species occurs considerably earlier than the first Arvicola in the Baza Basin of southern Spain (Gibert et al., 2007). Mimomys savini still makes an ideal ancestor for Arvicola, but the details remain to be determined.

Generic features of Arvicola include the following: 1) size large (mean m1 length ranges from 3.25 - 4.09 mm [Maul, 2000]), 2) relatively thick enamel, 3) ear-liest species with negative enamel differentiation, later may be undifferentiated or positively differentiated, 4) m1 simple, with three closed triangles and a simple acd, 5) m3 complex (T4 well developed), 6) mandibular foramen centrally located on mesial side of ascending ramus.

ARVICOLA JACOBAEUS (Cuenca-Bescós and others, 2010b)

Dental and mandibular characters: Position of the mandibular foramen on A. jacobaeus is unknown, but it is centrally located on the mesial side of the ramus in extant A. sapidus (Cuenca-Bescós, pers. com.). Based on the table of characters given in Cuenca-Bescós et al. (2010b), the dental morphology of A. jacobaeus is very similar to that of A. sapidus, so much so that rec-ognition of A. jacobaeus as a distinct species is a matter of personal taste. The distinctions of A. jacobaeus from A. sapidus include the following: 1) in a few M3s of A. jacobaeus there is a constriction at the base of the molar, an indication that a root might have formed late in development, whereas this has not been reported in A. sapidus, 2) average size of the m1 is larger in A. sapi-dus, 3) the M3 of A. jacobaeus is, on average, simpler than that of A. sapidus, with some of the specimens dis-playing two closed triangles rather than three, the latter which is typical for A. sapidus. The enamel in A. jacobae-

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us is negatively differentiated, as in Mimomys savini and Arvicola sapidus, and the SDQ values (a reflection of extent of negative vs positive differentiation) are virtually identical in A. jacobaeus and A. sapidus (133 and 123-132, respectively).

Localities: Lower Red Unit of Trinchera Elefante (TEL-RU), Atapuerca Hills, Burgos (Cuenca-Bescós et al., 2010b).

Age: ~1.2 Ma (cosmogenic nuclide date; Carbonell et al., 2008).

Remarks: The various hypotheses available in the Eu-ropean literature for the origination of Arvicola and sub-sequent speciation of A. sapidus (with negative enamel differentiation) and A. terrestris (with positive differenti-ation) include at least two extinct, intermediate species, treated either as A. cantianus or A. mosbachensis in western Europe (Maul, 2000) and A. chosaricus in the Ukraine (Rekovets and Nadachowski, 1995). With the recent description of A. jacobaeus, we now have three extinct species. Evolutionary scenarios for the origin of the extant water voles include either one or two line-ages, including at least one scenario in which Arvicola would be polyphyletic, with A. sapidus and A. terrestris derived from different Mimomys ancestors (van kolf-schoten, 1993). If one accepts the philosophical po-sition that speciation is a branching process and that evolution will most often follow the most parsimonious route, then the phylogenetic interpretation of European Arvicola samples becomes considerably easier. In Fig. 3 I propose a simple scenario for western European water voles, namely that the earliest Arvicola (A. jacobaeus) is the earliest A. sapidus, and that all subsequent samples in which negative enamel differentiation dominates are also A. sapidus. Samples dominated by positive enam-el differentiation can be considered A. terrestris, and those intermediate represent the speciation event from A. sapidus to A. terrestris in progress. As discussed by Cuenca-Bescós et al. (2010b) the origin of A. terrestris from A. sapidus is supported by karyotypic information (A. sapidus, 2n =40; A. terrestris, 2n = 36). This scenar-io is also supported by the recent evaluation of middle Pleistocene European Arvicola samples by Escude et al. (2008) concluding that only one species (their A. can-tiana) was recognizable and that enamel differentiation coefficients (SDQ values) from these samples could not be used to order them chronologically. One would hope

that there would eventually be a series of dental charac-ters rather than just the SDQ values to distinguish fossil samples of A. sapidus and A. terrestris.

Genus TIBERICOLA von koenigswald, Fejfar and Tch-ernov, 1992

The genotype for Tibericola was Hass’s (1966) “Arvico-la (?)” jordanica from ‘Ubeidiya in Israel. It was restudied by von koenigswald et al. (1992) and referred to the new Tibericola, considered as a subgenus of Microtus. Ünay et al. (2001) named a new species of Tibericola, T. sakayensis, from Değirmendere in Turkey, and raised Tibericola to generic status, specifically referring two Spanish species previously considered as Allophaiomys, A. vandermeuleni and A. chalinei, to Tibericola. New material of T. jordanica was reported by Goren-Inbar et al. (2000) from Gesher Benot Ya’aqov in Israel. Tiberi-cola is here tentatively considered a distinct genus (but see below) based on the following characters: 1) large size (mean m1 length 3.0-3.5 mm), 2) enamel relative-ly thick and undifferentiated, 3) relatively deep penetra-tion of LRA3 on m1, isolating T4 from T5 on m1 and relatively deep penetration of LRA4 in advanced species, nearly isolating T5 from the acd.

TIBERICOLA VANDERMEULENI (Agustí, 1991)

Dental and mandibular characters: Specimens of T. vandermeuleni from Barranco de Los Conejos in the Baza Basin of southern Spain illustrated by Agustí (1991) display the features of an early member of the genus: 1) large size (mean m1 length 3.11 mm), 2) enamel relatively thick and undifferentiated, 3) relatively deep penetration of LRA3, isolating T4 from T5 on m1.

Localities: Barranco de Los Conejos, Baza Basin, south-ern Spain (Agustí, 1991); Barranco del Paso (Martin, 2012a).

Age: ~1.5 Ma (Gibert et al., 2007).

Remarks: T. vandermeuleni appears to be in the center of a cline in a set of characters distinguishing primitive through advanced (derived) members of the genus, as noted by Ünay et al. (2001). T. sakaryaensis displays the most underived occlusal character states, with a rel-atively simple m1 in which BRA3 ranges from shallow to moderately deep, thus expressing a three to near four

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closed triangle morphology, with more specimens with LRA3 shallow than deep. In nine m1s of T. vandermeu-leni reported by Agustí (1991), LRA3 averaged consist-ently deeper than in T. sakaryaensis, more definitively isolating T4 from T5. This trend continues in T. jordan-ica, with deep penetration of both LRA3 and LRA4 in most specimens, sometimes nearly isolating T5 from the anteroconid.

Nadachowski (1991) noted the similarity in dental features between T. jordanica and extant Chionomys, and in one of a number of phylogenetic scenarios he proposed a connection with T. jordanica and derivation from an unknown Mimomys ancestor separate from the Mimomys that gave rise to Microtus. In his fig. 24, Na-dachowski (1991) refers the ‘Ubeidiya arvicolid to “(?) Chionomys” jordanica, which would be arguable but

Fig. 3. Proposed phylogenetic scenario for Arvicola. Ma = millions of years ago. The horizontal line representing the evolution of rooted to rootless molars applies only to the change from Mimomys to Arvicola; M. savini molars retain roots, although in some populations roots appear to form late in development.

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reasonable if the phylogenetic connection is substanti-ated. Ünay et al. (2001) preferred Microtus deucalion as an ancestor for Tibericola, but another phylogenetic scenario seems more likely.

Tibericola species are found in early Pleistocene Spanish deposits with an arvicolid that will probably eventually be shown to be a Microtus similar to M. deucalion (current-ly recognized as Mimomys oswaldoreigi; Agustí et al., 1993) (Figs. 1, 4). M. deucalion retains negative enamel differentiation, small size and a simple m1 morphology, while Tibericola species display the derived conditions of undifferentiated enamel and a deep LRA3 on m1. Although contemporaneity in the early Pleistocene does not rule out a possible ancestor-descendant relationship between M. deucalion and T. vandermeuleni, M. deu-calion is not found in earlier sites in depositional basins from southern Spain, all of which contain only Mimomys (Agustí, 1990; Opdyke et al., 1997; Minwer-Barakat et al., 2008). Consequently, it seems likely that Tibericola vandermeuleni originated from an early species of Mi-momys other than the one ancestral to Microtus.

A nagging concern I have is that Tibericola may be the equivalent of Allophaiomys, a trashbasket for most-ly larger voles with undifferentiated enamel and a tendency to develop closed T4, all of which are like-ly symplesiomorphic for voles going from an ancestor with negative enamel differentiation to a larger Microtus with at least four closed triangles on m1. Conceivably, Tibericola jordanica, T. sakaryensis and T. vandermeu-leni are unrelated, each having evolved from a separate

M. deucalion-like or Mimomys ancestor. There is also a distinct resemblace between the m1 morphology of T. vandermeuleni and V. chalinei (Fig. 4), though the acd in most specimens of V. chalinei appears simpler than that of T. vandermeuleni despite its later occurrence (in what would be the first arvicolid lineage to display a sim-plification of m1 pattern through time, the Barranco de los Conejos and Barranco del Paso T. vandermeuleni could be the earliest Victoriamys). Based on both size and published dental features, M. (Allophaiomys) reser-vatus (Zazhigin, 1998) could be referred to Tibericola. Yet more problems to be solved.

Genus VICTORIAMYS Martin, 2012

Originally described as Allophaiomys chalinei by Alcal-de et al. (1981) from Cueva Victoria, this species from Spain and Italy was subsequently referred to the new genus Victoriamys (Martin, 2012a) based on the unique position of the mandibular foramen as a thin slit along the posterior edge of the mesial side of the ascending ramus. As there is only one species recognized, the de-tailed list of characters for the genus is given below.

VICTORIAMYS CHALINEI (Alcalde, Agustí and Villalta, 1981)

Dental and mandibular characters: size large (mean m1 length >3.0 mm<3.5 mm); mandibular foramen formed as a thin slit along posterior edge of mesial side of ascending ramus; m1 with 3-5 triangles, BRA3 and LRA4 often symmetrically deep, isolating confluent T4-5

Fig. 4. Lower first molars of Iberomys brecciensis (No. 195), Iberomys huescarensis (Nos. 193, 194 (possibly), 257) and Microtus sp. (Nos. 192, 256) from Huéscar 1 (Photos courtesy of C. Sesé and P. Peláez-Campomanes).

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from a simple acd; BRA3 on m1 usually shallow and not provergent; m3 complex, with well-developed T4; M3 with well-developed BRA3 and LRA4, the latter of-ten curving back into the anterior cap; enamel mostly undifferentiated, with occasional positively differentiated edges.

Localities: Cueva Victoria; Almenara (Casablanca) 3 (Al-calde et al., 1981); Trinchera Dolina 3-6 (Atapuerca) (Cuenca-Bescós et al., 2010a); Cal Guardiola Layer D5 (Vallparadís) (Minwer-Barakat et al., 2011); Cueva Ne-gra del Estrecho del Río Quípar (Walker et al., 2006); Bagur 2 (Laplana, 1999); Huetor Tájar 1 and 8, Tojaire 1 (García-Alix et al., 2009); El Chapparal (López-García et al., 2012).

Age in Spain: 0.98-0.79 Ma, in magnetically reversed sediments between the Jaramillo and Brunhes chrons (Martin, 2012a).

Remarks: This species (as Allophaiomys chalinei) was also reported from Italy, allocated to the Farneta Faunal Unit at 1.4 Ma (Masini and Sala, 2007), but there is no independent substantiation for that date, which was based on biostratigraphic considerations, including the taxonomy of V. chalinei considered initially as a relatively primitive member of the genus Allophaiomys. Neverthe-less, as a suitable ancestor for V. chalinei is not known in Spain, it may be that Chaline’s vole is a Pleistocene immigrant, with a longer history elsewhere in southern Europe.

The phylogenetic relationship of Victoriamys to Tiber-icola is uncertain because neither the morphology of m3 nor the position of the mandibular foramen on the mandible has been described for Tibericola. Both taxa display large size and undifferentiated enamel, but these are expected symplesiomorphies for any arhizo-dont vole descended from a large Mimomys. Tibericola vandermeuleni seems to be restricted in Spain to the early Pleistocene, in localities between the Olduvai and Jaramillo subchrons (Martin, 2012a). By that time the defining occlusal m1 pattern of Tibericola had appeared, with deep LRA3 practically closing T4 from T3 (Agustí, 1991). Victoriamys, on the other hand, is only known from post-Jaramillo age sediments, probably between 0.98-0.78 Ma, and yet many m1s display a less de-rived pattern than in T. vandermeuleni, with only three closed triangles and a relatively simple anteror loop, as

in Arvicola (Agustí, 1991; Martin, 2012a). The availa-ble evidence suggests independent origins of Tibericola and Victoriamys from distinct Mimomys ancestors, but as noted above, other hypotheses need to be tested as well.

As noted in Martin (2012a), the extant Iberomys cabre-rae also has a short lower incisor with a mandibular for-mamen placed on the posterior edge of the ascending ramus. This foramen opens dorsally, rather than pos-teriorly as in Victoriamys, but the short lower incisor may indicate a relationship at a higher taxonomic level despite considerable differences in dental morphology. On the other hand, the short lower incisor may be ple-siomorphic for a number of unrelated, small ancestral Mimomys species and the similarity in mandibular struc-ture could have been derived independently.

Genus IBEROMYS (Chaline, 1972)

Iberomys was originally described as a new subgenus of Microtus by Chaline (1972), restricted to the Iberian peninsula. Iberomys is currently considered a distinct genus by Spanish paleontologists, including the extinct I. huescarensis (Ruiz Bustos, 1988), I. brecciensis (or I. mediterraneus; Cuenca-Bescós et al, 2014) and the ex-tant I. cabrerae (= I. dentatus). Mazo et al. (1985) iden-tified two arhizodont voles from Huéscar 1 as Microtus brecciensis and Microtus (Pitymys) gregaloides. Ruiz Bustos renamed some of the Huéscar 1 M. gregaloides specimens as the new species Pitymys huescarensis. Cuenca-Bescós et al. (2014) developed an evolutionary scenario deriving I. brecciensis and I. cabrerae from “P.” huescarensis, now treated as Iberomys huescarensis. The Microtus brecciensis specimens reported and illus-trated by Mazo et al. (1985) were considered to repre-sent variants in the I. huescarensis population. In their model, I. brecciensis appears first in deposits of Brunhes age, such as levels TD 8b, 10 and 11 in Gran Dolina from the Atapuerca hills (Cuenca-Bescós et al., 2010a) and Cullar-Baza 1 in the Baza Basin (Sesé, 1989).

Recently, I examined the arvicolid specimens from Huéscar 1 at the National Museum of Natural History in Madrid. There are six m1s, of which two (HU-195, HU-256) were referred to Microtus brecciensis and four (HU-192, HU-193, HU-194, HU-256) to Microtus gre-galoides by Mazo et al (1985) (Fig. 4). In the caption to Fig. 3 of Mazo et al (1985) HU-256 was accidentally

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numbered HU-195 (Sesé, pers. com.). Thus, in the in-dividual illustrations of Fig. 3, Nos. 8 and 12 of Mazo et al (1985) bear the same number. When Ruiz Bustos (1988) defined Pitymys huescarensis, he included as the holotype HU-193, and as paratypes HU-194 and HU-195. Ruiz Bustos likely meant to include Nos. 10, 11 and 12 in Fig. 3 of Mazo et al (1985) rather than Nos. 8, 11 and 12, but from the list of paratypes in the species description, that cannot be determined. Conse-quently, at least for the basis of defining P. huescarensis, both m1s identified as HU-195 must be ignored. That leaves us with HU-192, 193, 194, 256 and 257. The holotype right m1, HU-193, demonstrates a general-ized five triangle pattern with T4-5 confluent and closed from the anterior loop by deep penetrance of BRA3 and LRA4. Lingual triangles are not overly developed rela-tive to the labial triangles as they are in I. brecciensis and I. cabrerae. HU-257, another right m1, exhibits the same basic pattern as HU-193, but lingual triangles 3 and 5 appear larger relative to the labial triangles, and the anterior loop displays the somewhat anteriorly-pos-teriorly flattening characteristic of later Iberomys species. LRA3 and LRA4 are also distinctly provergent in HU-257, a condition that is also characteristic of I. brecciensis. Together, HU-193 and HU-257 could be accepted as an ancestral m1 morphology for later Iberomys, con-firming the validity of I. huescarensis as envisioned by Cuenca-Bescós et al. (2014). However, other m1s in the Huéscar 1 sample could create problems for the overall evolutionary scenario and the timing of events as proposed by Cuenca-Bescós et al. (2014). HU-195 demonstrates all the characters of Iberomys breccien-sis, including distinct asymmetry of lingual and labi-al triangles as a result of deep and provergent lingual reentrant folds and closure of T4-5 by deep penetrance of LRA3. It is unlikely that the morphology of HU-195 represents part of the variation of I. huescarensis as discussed above and represented by HU-193 and HU-257, although HU-256 appears intermediate in some ways, with a relatively primitive anterior loop combined with deep lingual reentrants that close T4-5. HU-192 and HU-194 are also enigmatic. Posterior to the anterior loop they closely resemble HU-193. However, the ante-rior loop morphology is a bit unusual for either I. brec-ciensis or I. cabrerae. HU-192 displays what is usually referred to in the literature as an “arvaloid” morphology and HU-194 exhibits a “gregaloid” morphology (Reko-vets and Nadachowski, 1995). Both usually indicate an evolutionary trajectory towards lineages other than Iber-

omys. Because of these observations and the limited sample size, a number of hypotheses are possible, as follows: 1) the sample is homogeneous and the varia-tion expressed is of one species, I. huescarensis; 2) two species are present, I. huescarensis and I. brecciensis; 3) three or four species are present, represented by I. huescarensis, I. brecciensis and one or two additional species of the arvaloid and gregaloid groups. If I. brec-ciensis is present and distinct from I. huescarensis, then even if I. brecciensis originated from I. huescarensis it was at an earlier time than represented by Huéscar 1 (at ~0.90 Ma) and much earlier than envisioned by Cuenca-Bescós et al. (2014). However, this contradic-tion could be removed if the I. brecciensis specimens are intrusive. Recently, Dumo et al (2014) suggested that exposures in the Huéscar area included a mixture of early and middle Pleistocene sediments, in part be-cause their optically stimulated luminescence (OSL) dates at Huéscar were considerably younger (0.42-0.57 Ma) than the pre-Brunhes dates preferred by other in-vestigators based on faunal associations. If they did not sample only sediments younger than those at Huéscar 1, their results allow for the I. brecciensis specimen(s) from Huéscar 1 to be intrusive from younger sediments and not, therefore, contemporaneous at Huéscar 1 with I. huescarensis. If this is the case, the evolutionary sce-nario proposed by Cuenca-Bescós et al. (2014) remains intact. As noted by Dumo et al (2014), the hypothesis of a pre-Brunhes age for Huéscar 1 was based in part on the assumption by Gibert et al (2007) that Huéscar 1 lies below Puerto Lobo in the Baza Basin, in paleomag-netically reversed sediments. This needs to be tested. If the sediments at or above Huéscar 1 are reversed, then the OSL dates are too young, or possibly represent only later sediments. If the sediments display normal paleomagnetism, then the OSL dates likely apply to the level with the Huéscar 1 arvicolids, and the evolutionary scenario for Iberomys of Cuenca-Bescós et al. (2014), while perhaps still intact in essence, would need to be modified to represent the new dating, and the associa-tion of I. brecciensis with I. huescarensis would not be as unusual as it is at 0.90 Ma. The paleomagnetic data should also be augmented, if posible, by new sampling for microverterbates through the Huéscar exposures.

IBEROMYS HUESCARENSIS (Ruiz Bustos, 1988)

= Microtus (Pitymys) gregaloides Mazo et al, 1985= Pitymys huescarensis Ruiz Bustos, 1988= Iberomys huescarensis Cuenca-Bescós et al, 1995

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Dental and mandibular characters: size small (mean m1 length = 2.63 mm; Mazo et al [1985]), enamel positively differentiated. Anterior loop simple, with T6 absent or poorly developed. T7 moderately developed. The anterior loop may show some anterior-posterior flat-tening as in later Iberomys, and if HU-194 belongs with this species the anterior loop morphology may mimic the shape of simple M. gregaloides. T4-5 are confluent and closed from the anterior loop by deep penetrance of LRA4 and BRA3. The anterior enamel border of T4 tends to slope backwards, rather than horizontally or slightly anteriorly as in M. gregalis and M. gregaloides.

Localities: Huéscar 1 (Ruiz Bustos, 1988); Gran Do-lina, Atapuerca, levels TD3-4, TD6, TD8a (Cuenca-Bes-cós et al.,1995); Cal Guardiola level D5 and Vallparadís level EVT7 (Minwer-Barakat et al., 2011); Cueva Negra (Walker et al., 2006); El Chaparral (López-García et al, 2012).

Age in Spain: post-Jaramillo and pre-Brunhes sedi-ments, from about 0.98-0.79 Ma (Cuenca-Bescós et al., 2010a; Martin, 2012; López-García et al., 2012).

Remarks: In my early Pleistocene Spanish arvicolid biochronology (Martin, 2012a, fig. 5) I included Iber-omys huescarensis (as Microtus huescarensis) in the Lower Red Unit of Trinchera Elefante (TELRU) from the Atapuerca sequence following Carbonell et al. (2008). Subsequently, Cuenca-Bescós et al. (2010a, 2013) did not include reference to I. huescarensis from TELRU, list-ing only A. burgondiae and A. nutiensis. The specimens now referred to A. burgondiae and A. nutiensis from TELRU may represent part of the variation in Microtus (Allophaiomys) lavocati (see above).

IBEROMYS BRECCIENSIS (Giebel, 1847)

Dental and mandibular characters: Identifiable as Iberomys by considerable triangle assymetry, with lin-gual triangles extended as a result of deep penetrance of lingual reentrant folds; T4-5 closed; LRA4 often pene-trates deeply into the anterior loop. Differs from I. cabre-rae in more provergent than horizontal lingual reentrants and more rounded triangle apices.

Localities: Huéscar 1 and levels TD8B at Gran Dolina in the Atapuerca hills (Cuenca-Bescós et al, 2014); Cullar Baza 1 (Sesé, 1989) and other middle to late Pleisto-

cene sites in Spain (e.g., Cova del Llentiiscle; Montaña-na, 2010).

Age in Spain: (if not intrusive at Huéscar 1) late Matuy-ama through Brunhes, from approximately 0.90- ? Ma.

Remarks: According to Cuenca-Bescós et al (2014) the proper name for I. brecciensis is I. mediterraneus, a species named by Chaline (1967). Until I consider their argument in detail, for historical continuity I will retain the name I. brecciensis for this vole.

DISCUSSIon

Late Pliocene and early Pleistocene deposits throughout Eurasia include many members of the successful ge-nus Mimomys, considered at this time as monophyletic, derived from a Eurasian species of Promimomys (van der Weerd, 1979; Repenning et al., 1990). The reality, of course, may be considerably more complex, with a variety of Mimomys and Mimomys-like species origi-nating from various Promimomys species as the latter taxon spread across Eurasia into North America (Martin, 2010). Nevertheless, for convenience of discussion we will here consider Mimomys monophyletic. In the late Pliocene and early Pleistocene various Mimomys spe-cies lost the enamel atolls on m1 and M3, reduced the Mimomys-kante (ridge) on m1, and began experiment-ing with hypselodonty, resulting in a series of species in which roots formed very late in development. Some of these, segregated by mean length of the m1, include: 1) < 3.0 mm: M. blanci (van der Meulen, 1973) , M. pu-sillus (Méhely, 1914), M. oswaldoreigi (Agustí, 1993), M. tornensis (Jánossy and van der Meulen, 1975), M. tigliensis (Tesakov, 1998) and M. haplodentatus (Savinov and Tutkova, 1987); 2) > 3.0 mm: M. savini (Hinton, 1910). As noted by Tesakov (1998), M. torn-ensis, M. haplodentatus and M. tigliensis form a group of sibling species recognized as the Mimomys subge-nus Tcharinomys, but it is also possible they represent geographic variation in the single species M. tornensis. Tesakov (1998) also pointed out that more study of the Barranco de los Conejos M. oswaldoreigi is neces-sary in order to rule out the accidental combination of arhizodent first lower molars of an early Microtus near M. deucalion with a rooted M3 from a small Mimomys.

Species of Arvicola, Tibericola and Victoriamys appear to have originated at relatively large size (average m1

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Fig. 5. First appearances of various archaic Spanish arvicolid arhizodont voles and Mimomys savini. A = Victoriamys chalinei, Cueva Victoria (from Martin, 2012); B = Microtus lavocati, Atapuerca, Trinchera Elefante, Lower Red Unit (from LaPlana and Cuenca-Bescós, 2000); C = Iberomys huescarensis, holotype m1; M3 unknown), Huéscar 1 (from Mazo et al., 1985); D = Arvicola jacobaeus, Atapuerca, Trinchera Elefante, Lower Red Unit (illustrations are of A. mosbachensis from Mosbach, Germany; from Maul et al. 2000); E = Mimomys savini, Fuentenueva 3 (m1s from Agustí and Madurell, 2005; M3 from Untermassfeld, Germany [Maul, 2001]); F = Microtus pliocaenicus, Venta Micena (illustrations of type specimens from Betfia 2, Roumania; from Hir, 1998); G = Mimomys oswaldoreigi or Microtus cf deucalion m1s and a small Mimomys M3, Barranco de los Conejos (from Agustí et al., 1993); H = Tibericola vandermeuleni, Barranco de los Conejos (from Agustí, 1991). The time-line is in millions of years. Figures not to scale.

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length > 3.0 mm), and the only dentally advanced early Pleistocene Mimomys from western Europe in the appropriate size range to fill an ancestral position is Mimomys savini. Position of the mandibular foramen has not been published for either Mimomys savini or Tibericola, but it is distinct in Victoriamys, formed as a thin slit on the posterior edge of the mesial side of the ascending ramus. The Mimomys ancestor for Victori-amys should express this morphology. Arvicola has the more standard, central position of the foramen, and if it is descended from Mimomys savini, the latter should also display this condition. My suspicion is that there is a number of sibling species masquerading under the name of M. savini, and that a more comprehensive anal-ysis of fossil material referred to this and related species that have subsequently been synonymized with M. sa-vini (such as M. newtoni, M. milleri, M. majori and M. intermedius) will serve to identify potential ancestors for the larger early Pleistocene arhizodont voles.

Another feature of some phylogenetic value is form of the m3, either complex or simple. The complex con-dition is seen in Mimomys savini, M. tigliensis (some specimens; Tesakov, 1998), all extant Arvicola, Victo-riamys chalinei, and a few other, generally earlier, Mi-momys, such as M. pliocaenicus, M. coelodus and M. ostramosensis (Martin, 2012a). While the Mimomys ancestors for Arvicola and Victoriamys clearly possessed a complex m3, the ancestor of Microtus may not have, as neither M. deucalion nor extant Microtus species dis-play this condition. Consequently, we must look care-fully among the various small Mimomys for the salient combination of features that can pinpoint the species ancestral to Microtus. The starting place would seem to be a thorough morphological and chronological evalua-tion of early Pleistocene small Mimomys, as the limits of dental and mandibular features and chronological rang-es among the species are presently undefined.

The recent report by López-García et al. (2012) of the arvicolid community from El Chapparal, an early Pleis-tocene site from the Betic Cordillera of southwestern Spain, shows that we are still in the learning stages of understanding the history of arvicolid dispersal and di-versity on the Iberican Peninsula. They reported a com-bination of Allophaiomys lavocati, Victoriamys chalinei, Iberomys huescarensis, Pliomys episcopalis and Micro-tus (Terricola) arvalidens from El Chaparral. Neither M. arvalidens nor V. chalinei had previously been reported

synchronic with A. lavocati. If not a mixed assemblage, I agree with their assessment that this could represent a late occurrence of A. lavocati, probably in an unusual southern ecological setting. Consequently, I have not in-cluded El Chaparral in Fig. 1, as I would not, based on its arvicolids, know exactly where to put it. For now, I agree with López-García et al. (2012) that El Chaparral is likely post-Jaramillo and pre-Brunhes in age.

In conclusion, a review of the Spanish archaic Pleisto-cene arhizodont voles Microtus, Arvicola, Iberomys, Ti-bericola and Victoriamys (Fig. 5) suggests they emerged during the early Pleistocene, possibly from separate Mi-momys ancestors as the latter genus experimented with hypselodonty in a number of lineages. Microtus deu-calion originated from a small Mimomys with a simple m3. Arvicola sapidus (= A. jacobaeus = A. cantianus [in part]) originated from Mimomys savini and A. terrestris (= A. mosbachensis [in part]) originated from A. sapi-dus, probably during the late Pleistocene, the latter step recognized by a switch from negative to positive enam-el differentiation. Victoriamys and Tibericola (if distinct) originated from separate large, currently unrecognized, Mimomys speces. The ancestor for Victoriamys should have a short lower incisor and a complex m3. The denti-tion and mandible of Tibericola are insufficiently known to speculate on features of its ancestor other than large size, though a Mimomys ancestor might show an occa-sional tendency of deep penetration of LRA3 on m1.

The possible record of Iberomys brecciensis from Hués-car 1 is troublesome and allows a number of evolution-ary and dispersal scenarios.

ACknoWLEDGMEnTS

I thank L. Gibert and G. Scott for inviting me to partici-pate in their ongoing geological investigations in Spain and C. Ferràndez Cañadell for help in providing refer-ences and cataloguing specimens. G. Cuenca-Bescós kindly provided photographs of mandibular anatomy and occlusal morphology in extant Arvicola sapidus and Iberomys cabrerae. I am particularly grateful for the hos-pitality of Carmen Sesé and Pablo Peláez-Campomanes while studying the Huéscar material in their care. A. Tesakov provided helpful insights and information that greatly improved the manuscript.

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LITERATURE CITED

AGUSTÍ, J. 1990. The plio-pleistocene arvicolid succes-sion in southern and eastern Spain; pp. 21-32 in O. Fejfar and W.-D. Heinrich (eds.), International Sympo-sium on the Evolution, Phylogeny and Biostratigraphy of Arvicolids (Rodentia, Mammalia). Geological Survey, Prague.

AGUSTÍ, J. 1991. The Allophaiomys complex in south-ern Europe. Geobios 25: 133-144.

AGUSTÍ, J., CASTILLO, C., GALOBART, A., 1993. Hetero-chronic evolution in the late Pliocene-early Pleistocene arvicolids of the Mediterrranean area. Quaternary Inter-national 19: 51-56.

AGUSTÍ, J., MADURELL, J., 2005. Los arvicólidos (Muroidea, Rodentia, Mammalia) del Pleistoceno inferi-or de Barranco León y Fuente Nueva 3 (Orce, Granada). Monografías de Arqueología. Junta de Andalucía, Sevilla, pp. 137-145.

AGUSTÍ, J., BLAIN, H.-A., FURIÓ, M., DE MARFÁ, R., SAN-TOS-CUBEDO, A. 2010. The early Pleistocene small mammal succession from the Orce región (Guadix-Ba-za Basin, SE Spain) and its bearing on the first human occupation of Europe. Quaternary International 223-224:162-169.

ALCALDE, G., AGUSTÍ, J., VILLALTA, F. J. 1981. Un nuevo Allophaiomys (Arvicolidae, Rodentia, Mammalia) en el Pleistoceno inferior del sur de España. Acta Geológica Hispánica 16: 203–205.

CARBONELL, E., BERMÚDEZ DE CASTRO, J. M., PARÉS, J. M., PÉREZ-GONZÁLEZ, A., CUENCA-BESCÓS, G., OLLÉ, A., MOSQUERA, M., HUGUET, R., MADE, J. VAN DER, RO-SAS, A., SALA, R., VALLVERDÚ, J., GARCÍA, N., GRANGER, D. E., MARTINON-TORRES, M., RODRÍGUEZ, STOCk, G. M., VERGÉS, J. M., ALLUÉ, E., BURJACHS, F., CÁCERES, I., CANALS, A., BENITO, A., CHALINE, J. 1967. Microtus mediterraneus, n. sp., nouvel Arvicolidé (Rodentia) du Pléistocène moyen de France. Comptes Rendues de la Academie des Sciences (Série D) 265: 900-903.

CUENCA-BESCÓS, G., CANUDO, J. I., LAPLANA, C., 1995. Los Arvicólidos (Rodentia, Mammalia) de los niveles in-feriores de Gran Dolina (Pleistoceno Inferior, Atapuerca,

Burgos, España). Revista Española de Paleontología 10: 202-218.

CUENCA-BESCÓS, G., ROFES, J., LÓPEZ-GARCÍA, J. M., BLAIN, H.-A., DE MARFA, R. J., GALINDO-PELLICENA, M. A., BENNÁSAR-SERRA, M. L., MELERO-RUBIO, M., ARSUAGA, J. L., BERMÚDEZ DE CASTRO, J. M., CAR-BONELL, E., 2010a. Biochronology of Spanish Quater-nary small vertebrate faunas. Quaternary International 212: 109-119.

CUENCA-BESCÓS, G., AGUSTÍ, J., LIRA, M., RUBIO, M., ROFES, J., 2010b. A new species of water vole from the early Pleistocene of southern Europe. Acta Palaeontolog-ica Polonica 55: 565-580.

CUENCA-BESCÓS, G., ROFES, J., LÓPEZ-GARCÍA, J. M., BLAIN, H.-A., DE MARFA, R. J., RABAL-GARCÉS, R., SAU-QUÉ, V., ARSUAGA, J. L., BERMÚDEZ DE CASTRO, J. M., CARBONELL, E., 2013. The small mammals of Sima del Elefante (Atapuerca, Spain) and the first entrance of Homo in western Europe. Quaternary International 295: 28-35.

CUENCA-BESCÓS, G., LÓPEZ-GARCÍA, J. M., GALIN-DO-PELLICENA, M. A., GARCÍA-PEREA, R., GISBERT, J., ROFES, J., VENTURA, J., 2014. Pleistocene history of Iberomys, an endangered endemic rodent from south-western Europe. Integrative Zoology 9: 481-497.

DÍEZ, C., LOZANO, M., MATEOS, A., NAVAZO, M., RODRÍGUEZ, J., ROSELL, J., ARSUAGA, J. L., 2008. The first hominin of Europe. Nature 452: 465–470.

ESCUDÉ, E., MONTUIRE, S., DESCLAUX, E., QUÉRÉ, J.-P., RENVOISÉ, E., JEANNET, M., 2008. Reappraisal of ‘chronospecies’ and the use of Arvicola for biochronol-ogy. Journal of Archaeological Science 35: 1867-1879.

GARCIA-ALIX, A., MINWER-BARAkAT, R., MARTÍN SU-AREZ, E., FREUDENTHAL, M., 2009. Small mammals from the Early Pleistocene of the Granada Basin, south-ern Spain. Quaternary Research 72: 265–274.

GIBERT, L., SCOTT, G., MARTIN, R., GIBERT, J., 2007. The early to middle Pleistocene boundary in the Baza basin (Spain). Quaternary Science Reviews 26: 2067-2089.

223

A BRIEF REVIEW OF THE SPANISH ARCHAIC PLEISTOCENE ARHIZODONT VOLES

GOREN-INBAR, N., FEIBEL, C. S., VEROSUB, k. L., MEL-AMED, Y., kISLEV, M. E., TCHERNOV, E., SARRAGUSTI, I., 2000. Pleistocene milestones on the Out-of-Africa corridor at Gosher Benot Ya’aqov, Israel. Science 289: 944-947.

HAAS, G., 1966. On the vertebrate fauna of the Lower Pleistocene site ‘Ubeidiya. Israel Academy of Science and Humanities: 1-68.

HINTON, M. A. C., 1910. A preliminary account of the British fossil voles and lemmings with some remarks on the pleistocene climate and geography. Proceedings of the Geological Association 21: 489-507.

HIR, J., 1998. The Allophaiomys type-material in the Hungarian collections. Paludicola 2: 28-36.

JÁNOSSY, D., VAN DER MEULEN, A. J., 1975.On Mi-momys (Rodentia) from Osztramos-3, north Hungary. Proceedings konikl. Nederl. Akademie van Wetenschap-pen 78: 381-391.

kOENIGSWALD, W. VON, FEJFAR, O., TCHERNOV, E., 1992. Revision einiger alt- und mittelpleistozäner Arvi-coliden (Rodentia, Mammalia) aus dem östlichen Mit-telmeergebiet (‘Ubeidiya, Jerusalem and kalymnos-Xi). Neues Jahrbuch für Geologie und Paläontologie Ab-handlung, 1-23.

kOLFSCHOTEN, T. VAN, 1993. On the origin of the mid-dle Pleistocene larger voles. Quaternary International 19: 47-50.

kORMOS, T., 1933. Neue Wühlmäuse aus dem Ober-pliozän von Püspökfürdö. Neue Jahrbuch fur Minerolo-gie Geologie Paleontologie 69: 323-346.

kRETZOI, M., 1941. Die unterpleistozane saugetierfauna von Betfia bei Nagyarad. Sonderdruck aus Földtani kö-zlön 71: 7-12.

LAPLANA, C., 1999. Presencia de Microtus (Allo-phaiomys) chalinei Alcalde, Agustí y Villalta, 1981 (Arvi-colidae, Rodentia) en el yacimiento de Bagur-2 (Pleisto-ceno inferior, Girona, España). Treballs del Museum de Geologia de Barcelona 8: 25–32.

LAPLANA, C., CUENCA-BESCÓS, G., 2000. Ua nueva es-pecie de Microtus (Allophaiomys)(Arvicolidae, Rodentia, Mammmalia) en el Pleistoceno inferior de la Sierra de Atapuerca (Burgos, España). Revista Española Paleon-tología 15: 77-87.

LÓPEZ-GARCÍA, J. M., CUENCA-BESCÓS, G., BLAIN, H.-A., CÁCERES, I., GARCÍA, N., VAN DER MADE, J., GUTIÉR-REZ, J. M., SANTIAGO, A., PACHECO, F. G., 2012. Bio-chronological data inferred from the early Pleistocene Arvicolinae (Mammalia, Rodentia) of the El Chaparral site (Sierra del Chaparral, Cádiz, southwestern Spain). Journal of Vertebrate Paleontology 32: 1149-1156.

MARTIN, R. A., 1987. Notes on the classification of some North American fossil Microtus. Journal of Vertebrate Paleontology 7: 270-283.

MARTIN, R. A., 1998. Time’s arrow and the evolutionary position of Orthriomys and Herpetomys. Paludicola 2: 70-73.

MARTIN, R. A., 2010. The North American Promimomys immigration event. Paludicola 8: 14-21.

MARTIN, R. A., 2012a. Victoriamys, a new generic name for Chaline’s vole from western Europe. Geobios 45: 445-450

MARTIN, R. A., 2012b. Further notes on the Port kenne-dy Cave arvicolid rodents. Paludicola 9: 13-20.

MARTIN, R. A., TESAkOV, A., 1998. Introductory remarks: does Allophaiomys exist? Paludicola 2: 1-7.

Martin, R. A., Crockett, C. P., Marcolini, F., 2006. Variation of the schmelzmuster and other enamel characters in molars of the primitive Pliocene vole Ogmodontomys from kansas. Journal of Mammalian Evolution 13: 223-241.

MASINI, F., SALA, B., 2007. Large and small mammmal distribution patterns and chronostratigraphic boundaries from the Late Pliocene to the Middle Pleistocene of the Italian Peninsula. Quaternary International 160: 43-56.

MAUL., L., 2001. Die kleinsäugerreste (Insectivora, Lag-omorpha, Rodentia) aus dem Unterpleistozän von Un-

224

ROBERT A. MARTIN

termassfeld. Monographien des Römish-Germanischen Zantralmuseums Mainz 3: 783-887.

MAUL, L., REkOVETS, L., HEINRICH, W.-D., kELLER, T., STORCH, G., 2000. Arvicola mosbachensis (Schmidt-gen 1911) of Mosbach 2: a basic sample for the ear-ly evolution of the genus and reference for further bi-ostratigraphical studies. Senckenbergiana Lethaea 80: 129-147.

MAUL, L., HEINRICH, W.-D., PAREFITT, S. A., PAUNES-CU, A.-C., 2007. Comment on the correlation between magnetostratigraphy and the evolution of Microtus (Ar-vicolidae, Rodentia, Mammalia) during the Early and ear-ly Middle Pleistocene. Cour. Forsch. Inst. Senckenberg 259: 243-263.

MAZO, A.V., SESÉ, C., RUIZ BUSTOS, A., PEÑA, J. A. 1985. Geología y paleontología de los yacimientos Plio-Pleisto-cenos de Huéscar (Depression de Guadix-Baza, Grana-da). Estudios Geologicos 41:467-493.

MÉHELY, L., 1914. Fibrinae Hungariae. Die ternären und quartären wurzelzähnigen Wühlmšäuse Ungarns. Annals Natural History Museum Hungary 12: 155-243.

MEULEN, A. J. VAN DER., 1973. Middle Pleistocene smaller mammals from the Monte Peglia, (Orvieto, Italy) with special reference to the phylogeny of Microtus (Ar-vicolidae, Rodentia). Quaternaria 17: 1-144.

MEULEN, A. J. VAN DER., 1974. On Microtus (Allo-phaiomys) deucalion (kretzoi, 1969), (Arvicolidae, Ro-dentia), from the Upper Villányian (lower Pleistocene) of Villány-5, S. Hungary. Proceedings konikl. Nederl. Akade-mie van Wetenschappen 77: 260-266.

MEULEN, A. J. VAN DER., 1978. Microtus and Pitymys from Cumberland Cave, Maryland, with a comparison of some New and Old World species. Annals of the Carne-gie Museum 47: 101-145.

MINWER-BARAkAT, R., GARCÍA-ALIX, A., MARTÍN-SUÁREZ, E., FREUDENTHAL, M., 2008. The laterst Rus-cinian and early Villanyian Arvicolinae from southern Spain re-examinend: biostratigraphical implications. Journal of Vertebrate Paleontology 28: 841-850.

MINWER-BARAkAT, R., MADURELL-MALAPEIRA, J., ALBA, D. M., AURELL-GARRIDO, J., DE ESTEBAN-TRIVIGNO, S., MOYÀ-SOLÀ, S., 2011. Pleistocene rodents from the Torrent de Vallparadís section (Terrassa, northeastern Spain) and biochronological implications. Journal of Ver-tebrate Paleontology 31: 849-865.

MONTAÑANA, S. 2010. Puntulizaciones sobre la fauna del Pleistoceno inferior/medio de la Cova del Llentiscle (Vilamarxant, Valencia). Archivo de Prehisoria Levantina 28:11-45.

NADACHOWSkI, A., 1991. Systematics, geographic var-iation, and evolution of snow voles (Chionomys) based on dental characters. Acta Theriologica 36: 1-45.

NADACHOWSkI, A, ZAGORODNYUk, I., 1996. Recent Allophaiomys-like species in the Palaearctic: Pleistocene relicts or a return to an initial type. Acta Zoologica Cra-coviensia 39: 387-394.

OPDYkE, N., MEIN, P., LINDSAY, E., PEREZ-GONZALES, A., MOISSENET, E., NORTON, V. I., 1997. Continental deposits, magnetostratigraphy and vertebrate paleontol-ogy, late Neogene of eastern Spain. Palaeogeography, Palaeoclimatology, Palaeoecology 133: 129-148.

PASA, A., 1947. I mammiferi di alcune antiche brecce veronesi. Memoirs Museo Civico di Storia Naturele di Verona 1: 1-111.

REPENNING, C. A., FEJFAR, O., HEINRICH, W.-D., 1990. Arvicolid rodent biochronology in the northern hem-isphere; pp. 385-418 in O. Fejfar and W.-D. Heinrich (eds.), International Symposium on the Evolution, Phy-logeny and Biostratigraphy of Arvicolids (Rodentia, Mammalia). Geological Survey, Prague.

REkOVETS, L., NADACHOWSkI, A., 1995. Pleistocene voles of the Ukraine. Paleontologia i Evolució 28-29: 145-245.

RUIZ BUSTOS, A., 1988. Estudio sobre los arvicolidos cuaternarios. Paleomammalia 2: 1-89.

RUIZ BUSTOS, A. 1993. New data on lower Pleistocene arvicolids. The Venta Micena, Betfia-IX and Villany-5 sites. Communicaciones de las IX Jornados de Paleont-ologia: 60-64.

225

A BRIEF REVIEW OF THE SPANISH ARCHAIC PLEISTOCENE ARHIZODONT VOLES

RUIZ BUSTOS, A., 1999. The role of the water voles (Ar-vicola, Rodentia) in the Quaternary. Estudios Geológicos 55: 89-95.

SAVINOV, P. F., TUTkOVA, L. A., 1987. Pliocene mam-mals from the Aktogay locality; pp. 99-102 in kozham-kulova, B.S., P. F. Savinov, L. A. Tjutkova and T. k. Pak. Materials on the History of Fauna and Flora of kazakh-stan, 9: 82-120, Alma-Ata [in Russian].

SESÉ, C., 1989. Micromamíferos del Mioceno, Plioceno y Pleistoceno de la cuenca de Guadix-Baza (Granada); pp. 187-215 in M. T. Alberdi and F. Bonadonna (eds.), Geología y Paleontología de la Cuenca de Guadix-Ba-za. Museo Nacional de Ciencias Naturales, Madrid.

TESAkOV, A., 1998. Voles of the Tegelen fauna. Me-dedelingen Nederlands Instituut voor Toegepaste Ge-owetenschappen 60: 71-134.

ÜNAY, E., EMRE, O., ERkAL, T., kEçER, M., 2001. The rodent fauna from the Adapazari pull-apart basin (NW Anatolia): its bearings on the age of the North Anatolian fault. Geodinamica Acta 14: 169-175.

WALkER, M. J., ESTRELLA, T. R., GARCÍA, J. S. C., JIMÉN-EZ, M. A. M., SCHWENNINGER, J.-L., MARTÍNEZ, M. L., JIMÉNEZ, A. L., DEL TORO, M. S. N., HILLS, M. D., WALk-LING, T., 2006. Cueva Negra del Estrecho del Río Quípar (Murcia, southest Spain): an Acheulean and Levalloi-so-Mousteroid assemblage of Paleolithic artifacts exca-vated in middle Pleistocene faunal context with hominin skeletal remains. Eurasian Prehistory 4: 2-42.

WEERD, A. VAN DER., 1979. Early Ruscinian rodents and lagomorphs (Mammalia) from the lignites near Ptole-mais (Macedonia, Greece). Proceedings konikl. Nederl. Akademie van Wetenschappen 82: 127-170.

ZAZHIGIN, V., 1998. Taxonomy and evolution of the ge-nus Allophaiomys (Microtinae: Rodentia: Mammalia) in Siberia. Paludicola 2: 126-129.

PrólogoEmiliano Aguirre

PresentaciónL. Gibert y C. Ferràndez-Cañadell

Introducción. Cueva Victoria, un yacimiento de vertebrados del Pleistoceno InferiorC. Ferràndez-Cañadell y L. Gibert

Historia de la minería de Cueva VictoriaM. A. Pérez de Perceval, J. I. Manteca Martínez y M.A. López-Morell

Las mineralizaciones ferro-manganesíferas de la mina-cueva Victoria y su contexto geológicoJ. I. Manteca y R. Piña

Microscopía electrónica de las mineralizaciones cársticas de óxidos de hierro y manganeso de Cueva Victoria (Cartagena, Murcia)D. Artiaga, L. Gibert y J. García-Veigas

Edad del yacimiento de Cueva Victoria y su relación con otros yacimientos de la Península IbéricaL. Gibert L. y G. Scott230Th/U-dating of the Cueva Victoria flowstone sequence: Preliminary results and palaeoclimatic implicationsA. Budsky, D. Scholz, L. Gibert y R. Mertz-kraus

Reconstrucción y génesis del karst de Cueva VictoriaA. Ros y J. L. Llamusí

Modelización tridimensional mediante escáner 3D y tomografía eléctrica de alta resolución, en Cueva Victoria IA. Espín de Gea, A. Gil Abellán y M. Reyes Urquiza

Contexto sedimentario y tafonomía de Cueva VictoriaC. Ferràndez-Cañadell

Génesis de una acumulación osífera excepcional en Cueva Victoria (Cartagena, Murcia, España)J. Vilà-Vinyet, Í. Soriguera-Gellida y C. Ferràndez-Cañadell

Anfibios y escamosos de Cueva VictoriaH. A. Blain

Las tortugas del yacimiento del Pleistoceno inferior de Cueva Victoria (Murcia, España)A. Pérez-García, I. Boneta, X. Murelaga, C. Ferràndez-Cañadell y L. Gibert

A brief review of the Spanish archaic Pleistocene arhizodont volesR. A. Martin

Estado de conocimiento de los Insectívoros (Soricidae, Erinaceidae) de Cueva VictoriaM. Furió

The Lower Pleistocene Bats from Cueva VictoriaP. Sevilla

Aves del Pleistoceno inferior de Cueva Victoria (costa sudoriental mediterránea de la península Ibérica)A. Sánchez Marco

The latest Early Pleistocene giant deer Megaloceros novocarthaginiensis n. sp. and the fallow deer Dama cf. vallonnetensis from Cueva Victoria (Murcia, Spain)J. van der Made

Estudio de los caballos del yacimiento de Cueva Victoria, Pleistoceno Inferior (Murcia)M. T. Alberdi y P. Piñero

The rhinoceros Stephanorhinus aff. etruscus from the latest Early Pleistocene of Cueva Victoria (Murcia, Spain)J. van der Made

Elephant remains from Cueva VictoriaM. R. Palombo y M. T. Alberdi

Canid remains from Cueva Victoria. Specific attribution and biochronological implicationsM. Boudadi-Maligne

Úrsidos, hiénidos y félidos del Pleistoceno inferior de Cueva Victoria (Cartagena, Murcia)J. Madurell-Malapeira, J. Morales, V. Vinuesa y A. Boscaini

Los primates de Cueva VictoriaF. Ribot, C. Ferràndez-Cañadell y L. Gibert

Grupos pendientes de estudio o revisiónC. Ferràndez-Cañadell

Preparación de restos fósiles de Cueva Victoria, CartagenaA. Gallardo

ARTÍCULOS

AYUNTAMIENTODE CARTAGENA