Articles

FIGURE 1. Geographical and geological setting of the Fonelas SSC-3 site. 1, location of the Guadix and Baza Basins in the southeastern Iberian Peninsula; 2, digital elevation model (DEM) (3D) of the Guadix and Baza Basins, and the high ground surrounding them. The position of Fonelas SCC-3 is indicated along with other important Early Pleistocene sites in the Guadix area?? (FP-1: Fonelas P-1; M-9: Mencal-9) and the three most important in the Baza area?? (VM: Venta Micena; BL-5: Barranco León-5; FN-3: Fuente Nueva-3); 3, DEM of the Guadix and Baza Basins showing their Early Pleistocene fossil sites (those listed above along with others, as yet undescribed in the literature, found during the Fonelas Project), plus the area's Paleozoic and Mesozoic ridges, corridors and associated Neogene basins. The current landscape model for the Fardes River Valley includes large expanses of badlands that increase the number outcrops of fossiliferous units and paleontological sites.

 figure1

FIGURE 2. Stratigraphic profiles for Fonelas SCC-1 (which contain FSCC-3) and Fonelas P-1. Magnetostratigraphic information for each is provided (modified after Arribas et al., 2009; Pla-Pueyo et al., 2011). FSCC-3 is located at the lower part of the reverse Chron C2r.2r; its age is 2.4-2.5 Ma (Pla-Pueyo et al. 2011).

figure2 

FIGURE 3. Fragment of the right maxilla of Anancus arvernensis mencalensis nov. ssp. (Fonelas SCC-3-001, holotype). 1, labial view; 2, occlusal view. Scale bar equals 10 cm.

figure3 

FIGURE 4. Occlusal view of the right M2 of Anancus arvernensis mencalensis nov. ssp. (Fonelas SCC-3-001, holotype) showing the abbreviations employed in the description of the dental structures. Scale bar equals 5 cm.

figure4 

FIGURE 5. Occlusal view of the right M3 of Anancus arvernensis mencalensis nov. ssp.  (Fonelas SCC-3-001, holotype) showing the abbreviations employed in the description of the dental structures. Scale bar equals 5 cm.

figure5 

FIGURE 6. Comparative occlusal view of the M3 of Anancus arvernensis and Anancus osiris. 1, Anancus sp. cf. Anancus arvernensis turoliensis from Molino de Manuel, Alfacar (from Bergounioux and Crouzel [1958]); 2, Anancus arvernensis arvernensis from Baza-1 (after Ros [2010]); 3, Anancus arvernensis mencalensis from Fonelas SCC-3, described in this paper; 4, Anancus arvernensis chilhiacensis from Chilhac (from Boeuf [1992]); 5, Anancus osiris from Gizeh (from Arambourg [1945]). Red arrows show the rectangular shape of the distal part of the tooth in A. arvernensis mencalensis nov. ssp. Note the resemblance with the shape of the Miocene Anancus from Alfacar. Specimens not to scale.

 figure6

FIGURE 7. Scatter plot of M3 measurements (in mm) from Western European Anancus (plus A. osiris from North Africa). Note: no metric data are available in the bibliography for the M3 of A. arvernensis from Las Higueruelas (specimens held at the Museo Geominero do not allow the complete visualization of M3).

figure7

 

TABLE 1. Measurements of the teeth and their features (in mm) from the maxilla fragment FSCC-3-001 (holotype) classified as Anancus arvernensis mencalensis nov ssp.

 
 

L

W 1

W2

W3

W4

W5

W6

WI

H

M2

120.18

70.43

73.15

69.11

66.38

45.26

-

60.87

-

M3

166

69.61

74.05

74.06

70.37

68.67

64.84

44.61

64.7

 

TABLE 2. Comparison of M3 from Fonelas SCC-3-001 and other specimens of Anancus arvernensis from Europe.

 

Fossil Site

Anancoidy

No. lophs in M3

Cement

Age

Reference

Fonelas SCC-3

absent

6

present

2.5-2.4 Ma

This work

Dorkovo

pronounced

5-6

absent

4.5 Ma

Metz-Muller, 1995

Baza-1

pronounced

5

absent

~4 Ma

Ros, 2010

Las Higueruelas

pronounced

5

absent

3.3 Ma

Own data

Hajná?ka

pronounced

5

traces

Villafranchian

Lupták, 1997

Chilhac

weak

5

present

2.3-2.2 Ma

Boeuf, 1992

 

TABLE 3. Comparison of dental features in Fonelas SCC-3-001 and other European and African anancine gomphotheres.

 

Species

Anancoidy in upper molars

No. lophs in M2

No. lophs in M3

Enamel folding

Age

Reference

A. arvernensis mencalensis

weak

4

6

none-coarse

Early Pleistocene

This work

A. arvernensis turoliensis

weak

4

5-6

none

Late Miocene

Own data

A. arvernensis arvernensis

pronounced

4-5

5-6

coarse-moderate

Pliocene

Own data

A. arvernensis falconeri

pronounced

4

5

coarse

Early Pleistocene

Guérin, 2004

A. arvernensis chilhiacensis

weak

4

5

coarse

Early Pleistocene

Boeuf, 1992

A. kenyensis

weak

4

5-6

none-coarse

Late Miocene-Early Pliocene

Hautier et al., 2009

Sanders et al., 2010

A. petrocchii

weak

5

unknown

unknown

Late Miocene

Sanders, 2011

Sanders, 2007, 2008

A. capensis

pronounced

4

6-7

moderate-strong

Late Miocene or Early Pliocene

Sanders, 2007

A. ultimus

moderate-pronounced

5

6-7

moderate-strong

Early-mid Pliocene

Sanders, 2011

A. osiris

weak

4

6

none-coarse

Late Pliocene-Early Pleistocene

Arambourg, 1945

 
Print Email

The last Iberian gomphothere (Mammalia, Proboscidea): Anancus arvernensis mencalensis nov. ssp. from the earliest Pleistocene of the Guadix Basin (Granada, Spain)

Guiomar Garrido and Alfonso Arribas

Plain Language Abstract

The discovery in 2006 of a 2.4-2.5 million year-old gomphothere jaw fragment in sedimentary layers of the Guadix Basin (Granada, Spain) provided information on the dental anatomy of the last Iberian Anancus. Detailed morphological analysis showed the upper third molar to possess a combination of primitive and derived features. These primitive features include six lophs with coarse folding of the enamel, indiscernible anancoidy (the cones face each other), and a rectangular distal outline. Among the more derived features are the presence of cement in the tooth-valleys and the reduced overall size of the molar. This mosaic of features, which has never before been reported, allows a new chronosubspecies of the European gomphothere to be proposed - Anancus arvernensis mencalensis. This subspecies is the most recent Anancus known for the Iberian Peninsula.

Resumen en Español

El último gonfoterio (Mammalia, Proboscidea) ibérico: Anancus arvernensis mencalensis nov. ssp. del Pleistoceno basal de la cuenca de Guadix (Granada, España)

En este trabajo se describe un fragmento de maxilar perteneciente a Anancus arvernensis localizado en el yacimiento de Fonelas SCC-3 (Cuenca de Guadix, Granada, España), del Pleistoceno basal (c.a. 2.5-2.4 Ma). El fósil, que conserva M2 y M3 en sus alveolos correspondientes, es atribuido a una nueva cronosubespecie basada en la peculiar combinación de caracteres observada en el tercer molar, el cual muestra un patrón estructural primitivo junto con un mosaico de caracteres derivados. Entre los primeros encontramos un una condición masiva y hexalofodonta, con un contorno distal subcuadrangular, un esmalte apenas plegado y ausencia de anancoidía. Sin embargo, esta condición primitiva se encuentra combinada con una serie de condiciones derivadas típicas de los últimos representantes del linaje, como es el pequeño tamaño de los molares, o la presencia de agregados de cemento que cubren los valles del diente. Esta combinación nos permite determinar una nueva subespecie, Anancus arvernensis mencalensis nov. ssp., que constituye el representante más moderno del género Anancus conocido hasta el momento en la península Ibérica (MNQ 17a). Esta nueva subespecie representaría un estadio evolutivo intermedio entre Anancus arvernensis arvernensis y Anancus arvernensis chilhiacensis, el último representante conocido del linaje europeo. Por lo tanto, A. arvernensis mencalensis permite reconocer una clina temporal entre los mastodontes durante el límite Plio-Pleistoceno e identifica importantes cambios en la configuración del tercer molar superior. Estos cambios anatómicos podrían considerarse como una respuesta a la aridificación climática acaecida hace unos 2.5 Ma y que tuvo como resultado un cambio en las comunidades vegetales.

PALABRAS CLAVE: Proboscidea; Anancus; nueva cronosubespecie; Pleistoceno basal; cuenca de Guadix; Península Ibérica

Traducción: Author

Résumé en Français

Le dernier gomphothère ibérique (mammifères, proboscidiens): Anancus arvernensis mencalensis nov. ssp . du tout début du Pléistocène du bassin de Guadix (Grenade, Espagne)

Ce travail décrit une nouvelle découverte de Anancus arvernensis ̶ un fragment de maxillaire qui préserve la M2 et la M3 ̶ du tout début du Pléistocène (environ 2.5 à 2.4 Ma) du site Fonelas SCC-3 (Cuenca de Guadix, Grenade, Espagne). Ce fossile est attribué à une nouvelle sous-espèce chronologiques basés sur la combinaison des caractéristiques anatomiques représentés sur la M3 : un motif anatomique primitif, en plus de caractéristiques dérivées. Les traits primitifs comprennent la condition hexalophodonte, un contour distale massif et rectangulaire, le pliage de l'émail discret, et un anancoidé indiscernable. Les caractéristiques dérivées ̶ vallées des dents couvertes par du ciment et la petite taille de la dent ̶ sont typiques des derniers représentants de la lignée. Cette mosaïque de caractéristiques permet la proposition d'une nouvelle sous-espèce chronologiques: Anancus arvernensis mencalensis nov. ssp . Celle-ci est la plus jeune représentante du genre Anancus connu de la péninsule ibérique ( MNQ 17a ), et représente un stade évolutif intermédiaire entre Anancus arvernensis arvernensis et Anancus arvernensis chilhiacensis, la dernier représentante connue de la lignée européenne. A. arvernensis mencalensis fait donc partie d'un déclin temporel pour la configuration de la M3 au cours de la transition Plio-Pléistocène. Ces changements anatomiques ont pu se produire en réponse à l' aridification qui a commencé vers 2,5 Ma, qui a produit des changements dans la composition des communautés végétales .

MOTS-CLÉS: proboscidiens; Anancus; nouvelles sous-espèces chronologiques; Pléistocène; bassin de Guadix, Péninsule ibérique

Translator: Kenny J. Travouillon

Deutsche Zusammenfassung

Das letzte Iberische Gomphotherium (Mammalia, Proboscidea): Anancus arvernensis mencalensis nov. ssp. aus dem frühesten Pleistozän des Guadix Beckens (Granada, Spanien)

Diese Arbeit beschreibt einen neuen Fund von Anancus arvernensis ̶ ein Maxillafragment mit erhaltenem M2 und M3 – aus dem frühesten Pleistozän (ca. 2.5-2.4 Ma) aus der Fonelas SCC-3 Fundstelle (Cuenca de Guadix, Granada, Spanien). Dieses Fossil wird basierend auf einer Kombination von anatomischen Merkmalen des M3 einer neuen Chronounterart zugeschrieben: ein primitives anatomisches Muster plus abgeleitete Merkmale. Die primitiven Merkmale beinhalten eine hexalophodonte Kondition, eine massive, rechteckige distale Kontur, unauffällige Zahnschmelzfaltung und eine kaum wahrnehmbare Anacoidität. Die abgeleiteten Merkmale - Zahngruben die mit Zement bedeckt sind und eine geringe Gesamtgröße des Zahns – sind typisch für die letzten Vertreter dieser Linie. Dieses Merkmalsmosaik erlaubt das Aufstellen einer neuen Chronounterart: Anancus arvernensis mencalensis nov. ssp. Dies wäre der jüngste Vertreter der Gattung Anancus auf der Iberischen Halbinsel (MNQ 17a). Er repräsentiert eine evolutionäre Zwischenstufe zwischen Anancus arvernensis arvernensis und Anancus arvernensis chilhiacensis, dem letzten bekannten Vertreter der Europäischen Linie. A. arvernensis mencalensis ist daher ein Teil der temporalen Kline in der Konfiguration des M3 über die Plio-Pleistozän Transition. Diese anatomischen Veränderungen traten möglicherweise als Reaktion auf die Aridifikation auf, die vor ca. 2.5 Millionen Jahren einsetzte und die zu Veränderungen in der Zusammensetzung der Pflanzengemeinschaften führte.

SCHLÜSSELWÖRTER: Proboscidea; Anancus; neue Chronounterart; Frühes Pleistozän; Guadix Becken; Iberische Halbinsel

Translator: Eva Gebauer

Arabic

387 arab

Translator: Ashraf M.T. Elewa

 

 

arribas

Alfonso Arribas Herrera is a paleontologist whose research covers taphonomy, taxonomy, biostratigraphy and the paleobiogeography of large Quaternary mammals. Alfonso holds a degree in Geology and a Ph.D. in Paleontology from Madrid’s Complutense University. He works for the Geological Survey of Spain, and is head of the Estación Paleontológica Valle del Río Fardes in Fonelas (Guadix Basin, Granada). He has a deep concern for the conservation of Spanish wildlife, especially the Iberian wolf.

divider

 

garridoGuiomar Garrido Álvarez-Coto is a paleontologist whose research covers the taxonomy of large Quaternary mammals. Guiomar holds a degree in Biological Sciences (Zoology) and a Ph.D. in Paleontology from Madrid’s Complutense University. She works for the Geological Survey of Spain and has been part of the Fonelas Project since 2001. In 2006 she completed her doctoral thesis on the systematic paleontology of large mammals from the Fonelas P-1 Early Pleistocene fossil site (Guadix Basin, Granada, Spain). Guiomar is involved in the conservation of the Iberian wolf (Canis lupus signatus), a species endangered by a disastrous management plan that allows annual killings because of conflicts with farmers.

 

Authors shown with Spanish wolves (Canis lupus signatus), an endangered subspecies in Spain.