(Mammalia, Proboscidea): Anancus Arvernensis Mencalensis Nov

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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

ABSTRACT

This work describes a new finding of Anancus arvernensis ̶ a maxilla fragment that preserves M2 and M3 ̶ from the earliest Pleistocene (c.a. 2.5-2.4 Ma) at the Fon- elas SCC-3 site (Cuenca de Guadix, Granada, Spain). This fossil is attributed to a new chronosubspecies based on the combination of anatomical features shown by M3: a primitive anatomical pattern plus derived features. The primitive features include the hexalophodont condition, a massive, rectangular distal outline, inconspicuous enamel folding, and indiscernible anancoidy. The derived features ̶ tooth-valleys covered by cement and the small overall size of the tooth ̶ are typical of the last representatives of the lineage. This mosaic of features allows a new chronosubspecies to be proposed: Anancus arvernensis mencalensis nov. ssp. This would be the youngest representative of the genus Anancus known for the Iberian Peninsula (MNQ 17a), and represents an intermediate evolutionary stage between Anancus arvernensis arvernensis and Anan- cus arvernensis chilhiacensis, the last known representative of the European lineage. A. arvernensis mencalensis therefore forms part of a temporal cline in the configuration of M3 over the Plio-Pleistocene transition. These anatomical changes could have occurred as a response to the aridification that began around 2.5 Ma, which led to changes in the composition of plant communities.

Guiomar Garrido. Estación Paleontológica Valle del Río Fardes, Instituto Geológico y Minero de España (IGME), Ríos Rosas 23, 28003 Madrid, Spain [email protected] Alfonso Arribas. Estación Paleontológica Valle del Río Fardes, Instituto Geológico y Minero de España (IGME), Ríos Rosas 23, 28003 Madrid, Spain [email protected]

KEYWORDS: Proboscidea; Anancus; new chronosubspecies; Early Pleistocene; Guadix Basin; Iberian Peninsula

PE Article Number: 17.1.4A Copyright: Society for Vertebrate Paleontology January 2014 Submission: 20 March 2013. Acceptance: 17 December 2013

Garrido, Guiomar and Arribas, Alfonso. 2014. The last Iberian gomphothere (Mammalia, Proboscidea): Anancus arvernensis mencalensis nov. ssp. from the earliest Pleistocene of the Guadix Basin (Granada, Spain). Palaeontologia Electronica Vol. 17, Issue 1;4A; 16p; palaeo-electronica.org/content/2014/639-last-iberian-mastodon http://zoobank.org/2E37A18F-DD62-4589-82A9-07F7F9BC965E GARRIDO AND ARRIBAS: LAST IBERIAN MASTODON

INTRODUCTION contrast, for the same chronological interval, different species of the genus Anancus are accepted, Anancine gomphotheres (Anancus Aymard [in both in Africa and Asia. Currently, five African spe- Dorlhac, 1855]) originated in Eurasia. They proba- cies of Anancus have been identified (Sanders, bly derived from Tetralophodon Falconer, 1857, 2011): Anancus kenyensis (MacInnes, 1942), displacing it during the late Miocene to become dis- Anancus ultimus Sanders, 2011, Anancus capen- tributed throughout the Old World until the early sis Sanders, 2007, Anancus petrocchii Coppens, Pleistocene (ca. -2.3 Ma). The type species is 1965, and Anancus osiris Arambourg, 1945. Five Anancus arvernensis (Croizet and Jobert, 1828), Asian species of Anancus are also recognized: the only one presently accepted to have lived in the Anancus sinensis (Hopwood, 1935) and Anancus west of the European continent. A. arvernensis cuneatus (Teilhard de Chardin and Trassaert, was fairly common in European ecosystems during 1937) from China, two species based on materials Pliocene. recovered in the foothills of the Siwaliks in India, In certain European Villafranchian deposits, i.e., Anancus sivalensis (Cautley, 1836) and Anan- Anancus fossils are found along with other species cus perimensis (Falconer and Cautley, 1846), and of Proboscidea. For example, they are associated Anancus kazachstanensis (Aubekerova, 1974) with Mammut borsoni (Hays, 1834) in Perrier-Etou- from Kazakhstan. aires and Vialette (MNQ 16a), and with Mammuthus meridionalis (Nesti, 1825) in Saint GEOLOGICAL AND CHRONOLOGICAL Vallier (MNQ 17a; Viret, 1954; Guérin, 2004), SETTING Costa San Giacomo (MNQ 17b; Palombo and Valli, 2003) and Chilhac (MNQ 17b; Boeuf, 1992). Mam- The paleontological site of Fonelas Solana del muthus Brookes, 1828 appeared in Western Cortijo Conejo 3 (Fonelas SCC-3) was discovered Europe around 3 m.y.a. and for the moment no Ibe- during the systematic surveying campaign of 2006, rian site has yet provided any evidence of coexis- undertaken as part of the Fonelas Project. The site tence between Anancus and Mammuthus. is located in the Guadix Basin (Granada, Spain) Anancus eventually become extinct, while Mam- near the town of Fonelas (UTM coordinates muthus meridionalis expanded with great success 482407, 4141795) (Figure 1.1-1.3). across the continent. The environment of the time Fonelas SCC-3 is part of Unit V (Early Pleisto- seemed to have been optimal for this new lineage, cene) of the basin fill (Pla-Pueyo et al., 2011), and for which chronological variation across Eurasia the fossilized remains of Proboscidea found in it fall has been described (Lister et al., 2005). into a layer of conglomerates that form part of a In the Iberian Peninsula, fossils of A. arver- high-energy fluvial channel facies within the Axial nensis have been recovered from many sites (see System (Viseras, 1991). Magnetostratigraphic Mazo and van der Made, 2012) between zones studies of this stratrigraphic column have shown a MNQ 12 (Concud, Teruel; Alcalá, 1994) and MNQ good correlation with GTS2004 (Lourens et al., 16-17 (Villarroya, La Rioja; Villalta, 1952). The 2004), locating Fonelas SCC-3 to the base of a most important is the site of Las Higueruelas (Ciu- reverse magnetozone (Matuyama Chron) where dad Real, MNQ 16; Mazo et al., 1980), where two successive subchrons with normal polarity are abundant fossil remains of this species have been recognized (Reunion C2r.1n and Olduvai C2n). recovered. Fonelas SCC-1 (MNQ 18; Guérin, 1990; Mein, In the Guadix Basin, Anancus was previously 1990) is located between these subchrons, near cited for Huélago-1 (MNQ 17; Alonso et al., 2002), the bottom of the Olduvai magnetozone. Therefore, where it may have coexisted with Mammuthus. Fonelas SCC-3 can be precisely assigned to the However, this cannot be verified due to the tapho- early Matuyama (pre-Reunion; Figure 2), i.e., it is nomic reworking of the Anancus fossil tooth frag- 2.4-2.5 million years old and therefore within the ments (A. Arribas, personal observation). MNQ 17a zone (according to Nomade et al., in Anancus appears in Africa near the end of the press). Miocene, coincident with the first appearance of elephants on that continent. It persisted there well MATERIAL AND METHODS into the Early Pleistocene. The fossil described in this work was collected To date, only A. arvernensis is fully recog- during the above-mentioned 2006 survey and is nized in the European context. It existed for several provisionally held at the headquarters of the Geo- million years and showed wide variation. As a con- logical Survey of Spain in Madrid’s Museo Geom- sequence distinct subspecies are recognized. In

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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. inero. The nomenclature and methods used in the MGM, Museo Geominero, Madrid, Spain; MN, anatomical description of the molar cusps were Mammal Neogene Zone; MNCN, Museo Nacional those proposed by Tassy (1986) (modified and de Ciencias Naturales, Madrid, Spain; MNQ, Mam- simplified for the upper teeth). Measurements were mal Neogene Quaternary Zone; m.y.a., million taken using a Mitutoyo calliper (precision 0.01 years ago; pr, pretrite principal tubercle; po, pos- mm). trite principal tubercle; pacc, pretrite anterior central conule; ppcc, pretrite posterior central conule; Abbreviations popcc, postrite posterior central conule; prm, pre- H, crown height; L, length; M, upper molar; m, trite mesoconule; pom, postrite mesoconule; ac, lower molar; Ma, mega-annum; me, mesoconule; anterior cingulum; pc, posterior cingulum; W,

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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).

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width; WI, width index (maximum width x 100 / with six lophs, inconspicuous enamel folding, indis- length). cernible anancoidy, a widened distal area forming a subrectangular outline, and an absent basal cingu- SYSTEMATIC PALEONTOLOGY lum. The derived features of this tooth include cementum deposits in the valleys between the Order PROBOSCIDEA Illiger, 1811 cusps and an overall small size, intermediate Superfamily ELEPHANTOIDEA Gray, 1821 between Anancus arvernensis arvernensis and Family GOMPHOTHERIIDAE Hay, 1922 Anancus arvernensis chilhiacensis. Subfamily ANANCINAE Hay, 1922 Measurements: See Table 1. Genus ANANCUS Aymard (in Dorlhac, 1855) Type species. Anancus arvernensis (Croizet and Description Jobert, 1828) The Anancus material recovered at Fonelas Genus diagnosis. Gomphotheriid with a high, SCC-3 consists of a fragment of hemimaxilla in short skull. Elevated dome. Enlarged tympanic which M2 and M3 are perfectly preserved in their bulla. Short mandible without tusks. Straight upper alveoli (Figure 3). These teeth show bunodont mor- tusks without enamel. Loss of premolars. Tetralo- phology and narrow valleys between the lophs. phodont intermediate molars (occasionally pentalo- The preserved maxillary bone fragment hardly phodont M2). Pretrite posterior central conule allows for its anatomy to be examined, although it reduced in upper molars. Reduction of the pretrite is possible to appreciate a palatine foramen that anterior central conule on the lower molars and opens beyond the mesial limit of the third molar. fusion with the mesoconelet. Alternation of the pre- The specimen was not considerably transported or trite and postrite half-loph(id)s (anancoidy) allows rolled before final deposition and burial, as indi- the establishment of an alternative contact of suc- cated by the absence of any abrasion signs on its cessive loph(id)s, especially for the lower interme- surface. diates and m3. The upper third molar has five or six M2 is a tetralophodont tooth, with moderately lophs and shows a trend towards simplification. worn structures, especially in the mesial region Cementum is normally absent, although it is pre- (Figures 3.2, 4), in which thick (6-8 mm) smoothly sent in later taxa (Tassy, 1986 [modified]; Hautier et folded enamel is visible. Despite the wear, an ante- al., 2009). rior cingulum with a straight mesial edge is clearly Anancus arvernensis mencalensis nov. ssp. seen. The cingulum has become fused to the prin- Figures 3-5 cipal tubercles due to wear. However, a mesio-lingual low conule appears separate from the first http://zoobank.org/B3BAE8B4-1E82-448A-893A- 4C814F7A1B8B pretrite principal tubercle (pr1). The outline of the mesio-distally flattened, postrite posterior central Derivatio nominis: From Cerro Mencal, a Jurassic conule (popcc1) can also be distinguished. The inselberg rising 1447 m above sea level, represent- second and third lophs show a pretrite anterior ing a paleogeographic high in the northern part of central conule (pacc2 and pacc3) fused to the pre- the Guadix Basin (Figure 1.3). This high is a geo- trite principal tubercle (pr2 and pr3), and a meso- logical and historical landmark for this area. Men- conule (me2 and me3) fused to the postrite cal comes from the Arabic for cemetery or transit, principal tubercle (po2 and po3). The fourth loph probably related to the presence of burial sites with shows the same structure as the previous two, traces of prehistoric dolmens, and to caves occu- although there is no mesoconule joined to the pos- pied during Paleolithic times. trite principal tubercle. Finally, a cingulum in the Holotype: Right maxilla fragment with M2 and M3 central region of the distal part of the tooth is subdi- (FSCC-3-001; Figure 3). Held at the Museo vided into three lobes, revealing the original exis- Geominero. tence of a large distal conule shifted to the lingual Type locality: Fonelas SCC-3 (Guadix Basin, region, and two minor conules in the distal area of Granada, Spain). Early Pleistocene (c.a. 2.5-2.4 the postrite principal tubercle of the fourth loph. m.y.a.; Pla-Pueyo et al., 2011). M3 is a hexalophodont tooth, with lophs Occurrence: Fonelas SCC-3 (Guadix Basin, slightly inclined towards the mesial region (Figure Granada, Spain). 3.1). There is no visible cingulum in the basal Diagnosis: Anancus with tetralophodont interme- region (neither labial nor lingual). The reduced diate upper molars. The upper third molar shows a wear shown by this tooth allowed a detailed exam- unique combination of primitive and derived fea- ination of its structures. The lophs are formed by tures. Among the former are a simple M3 crown

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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.

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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.

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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.

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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 1W2W3W4W5W6WIH 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

pairs of thick conical cusps known as principal mesio-labial region of the pretrite principal tubercle tubercles, the height and overall size of which is (pr4), the cusps of which reach the same height reduced towards the distal region. No anancoid (which equals that of ppcc3). The mesoconule pattern among the pretrite and postrite principal (prm4) is completely fused to the postrite principal tubercles is seen (Figure 5). The principal tuber- tubercle (po4); both reach the same height, which cles of one loph and those of the next are sepa- makes differentiation difficult. rated by deep valleys covered by cement The fifth loph consists of the two principal aggregates. Between the principal tubercles of tubercles which are substantially lower in height each loph lie a number of intermediate cusps that than the previous ones, and a pretrite mesoconule disappear towards the distal region of the tooth - (prm5) attached to pr5. In the mesio-labial area of except the central conule, which is observed in all po5 there is a small conical cusp occupying the lophs (and is more individualized in the three labial valley between po4 and po5. The degree of mesial lophs). The tooth therefore shows a com- development and differentiation of the M3 distal plex morphology in its mesial region that becomes cusps, including the entoflexus behind the fifth simpler towards its distal part. Mesially, there is a loph, suggests they should be understood as an well-developed anterior cingulum (ac), slightly independent sixth loph and not part of the posterior shifted to the mesio-labial region and connected to cingulum. Curiously, this potential sixth loph the pretrite anterior central conule of the first loph (including pr6 and po6) does not emerge directly (pacc1). from the labial and lingual sides of the tooth, but Between the pretrite (pr1) and postrite (po1) from irregular platforms with a rectangular outline. principal tubercles of the first loph lies a meso- In this region, the cement is less abundant, which conule subdivided into two completely fused allows examination of the deep valley between the conules (prm1 and pom1). In the valley that sepa- components of the fifth and sixth lophs. Finally, in rates the first and second lophs there is a low pop- the distal region of the tooth, three basal thicken- cc1, the top of which emerges from a large cement ings form the outline of the posterior cingulum. aggregate. In the central area of the second loph there is a large pretrite mesoconule (prm2), high COMPARISON enough to almost reach the principal tubercles. A The Fonelas SCC-3 hemimaxilla can be very compressed cusp is seen between this and attributed to Anancus according to the anatomical the postrite principal tubercle (po2), forming the features of the upper molars: bunodont morphol- postrite mesoconule (pom2). The third pretrite prin- ogy, a tetralophodont M2 and hexalophodont M3 cipal tubercle (pr3) appears with a well-developed with pretrite central conules (pacc and ppcc) (these and completely individualized pretrite anterior cen- are more developed than the postrite conules), and tral conule (pacc3) that almost reaches the height distal lophs with a reduced pretrite posterior central of the third pretrite principal tubercle. A lower pre- conule. At first glance, the most striking feature of trite posterior central conule (ppcc3) reaches the the specimen is the absence of anancoidy in M3, a height of the fourth pretrite principal tubercle (pr4). feature quite evident in almost all upper third Thick cement aggregates appear among these ele- molars of Anancus arvernensis. The absence of a ments that prevent the cusp bases from being basal cingulum also contrasts with other studied seen; only the latter’s apical region outcrops. The M3 specimens of A. arvernensis. A description of central area shows a single large mesoconule the Anancus species recognized for Old World (prm3) slightly shifted towards the postrite principal paleontological sites is provided below so that tubercle (po3). In height it reaches the principal comparisons can be made of the main anatomical tubercles of the third loph. The fourth loph is sub- dental features differentiating them from Fonelas stantially simpler than the previous ones, with a SCC-3 specimen (see Tables 2 and 3): pretrite anterior central conule (pacc4) fused to the

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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 Hajnáčka 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.

No. Anancoidy in No. lophs in Species lophs in Enamel folding Age Reference upper molars M3 M2

A. arvernensis weak 4 6 none-coarse Early Pleistocene This work mencalensis A. arvernensis weak 4 5-6 none Late Miocene Own data turoliensis A. arvernensis pronounced 4-5 5-6 coarse-moderate Pliocene Own data arvernensis A. arvernensis pronounced 4 5 coarse Early Pleistocene Guérin, 2004 falconeri A. arvernensis weak 4 5 coarse Early Pleistocene Boeuf, 1992 chilhiacensis A. kenyensis weak 4 5-6 none-coarse Late Miocene-Early Hautier et al., 2009 Pliocene 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 Sanders, 2007 Pliocene A. ultimus moderate- 5 6-7 moderate-strong Early-mid Pliocene Sanders, 2011 pronounced A. osiris weak 4 6 none-coarse Late Pliocene-Early Arambourg, 1945 Pleistocene

Anancus in Western Europe fied as Tetralophodon. This taxon is probably also present in Iberian sites such as Alfacar (Bergou- Anancus arvernensis is, for now, the only nioux and Crouzel, 1958). This subspecies has widely accepted species of this genus for Western massive third upper molars that show weak (nearly Europe. This Late Miocene-Early Pleistocene gom- absent) anancoidy. phothere had tetralophodont intermediate molars Anancus arvernensis arvernensis (Croizet and and upper third molars with five or six lophs. The Jobert, 1828). The most typical and abundant form enamel folding is coarse to fine, and the molar of the genus during the European Pliocene. The crown simple or complex and usually shows clear type series is found in the Pliocene of Perrier-les- anancoidy (especially so in m3). This taxon Etouaires (MN16). The upper third molars are nor- includes several recognized subspecies: mally pentalophodont, in which anancoidy is clearly Anancus arvernensis turoliensis Gaziry, 1997. visible. The distal part of this tooth narrows and The most primitive form of the species and typical there is always a well-developed basal cingulum. of Turolian sites such as Dorn-Dürkheim 1 (MN11- Anancus arvernensis falconeri Osborn, 1926. MN12) where it was first described. According to This subspecies is based on fossils from the Late Metz-Muller (2000), this material should be classi- Pliocene of Red Crag (England). It has also been

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cited for the Late Pliocene French site of Saint Val- Ma; Sanders et al., 2010) from eastern and central lier (Guérin, 2004), and is considered typical of the Africa; it has tetralophodont intermediate molars Middle Villafranchian and of the lowermost Late Vil- and upper third molars with five or six lophs and lafranchian. It has pentalophodont upper third usually unfolded, very thick (5-7 mm) dental molars, with anterior principal tubercles strongly enamel. The crown morphology is simple, with inclined in the forward direction. Anancoidy is con- massive lophs and low conelets; anancoidy is spicuous. Narrow valleys between the lophs con- weakly expressed. This species evolved into A. tain cement deposits. ultimus, an Early-Mid Pliocene species also from Anancus arvernensis chilhiacensis Boeuf, 1992. eastern and central Africa. In A. ultimus, the inter- This subspecies is based on specimens from the mediate molars are pentalophodont and upper Early Pleistocene deposits of Chilhac (MNQ17b), third molars have six or seven lophs. The enamel recently estimated at about 2.2 or 2.3 million years is coarsely to finely folded and anancoidy is clear. old (Boivin et al., 2010; Nomade et al., in press, A. capensis is a Latest Miocene-Early Pliocene respectively) - the last known record of this genus species recorded for Langebaanweg (South in Europe. Only pentalophodont upper third molars Africa). It has tetralophodont intermediate molars are observed and anancoidy is very weak. Cement and upper third molars with six or seven lophs. The is present in the valleys between the lophs. The M3 enamel shows moderate to strong folding and of this species is substantially smaller than in A. anancoidy is pronounced. A. petrocchi is a Late arvernensis arvernensis. Together, these features Miocene species recorded from As Sahabi (Libya), appear to be the result of the evolution of popula- with pentalophodont intermediate molars and third tions of A. arvernensis leading to a reduction in the molars with six lophs; the molar crowns are simple size of their third molars and the addition of cement and show weak anancoidy (Sanders, 2007, 2008). to them, among other skull and jaw features men- Finally, A. osiris is a Late Pliocene-Early Pleisto- tioned by Boeuf when she proposed A. arvernensis cene species from northern Africa (Egypt, Morocco chilhiacensis. This subspecies has recently been and Algeria). It has tetralophodont intermediate suggested a synonym of A. arvernensis falconeri molars, simple M3 crowns with six lophs, no (Metz-Muller, 2000). enamel folding, and shows weak anancoidy. Some Anancus arvernensis mencalensis. This paper authors consider A. kenyensis and A. osiris to be proposes the existence of this chronosubspecies of synonymous (Hautier et al., 2009). The similarities Anancus for the Early Pleistocene of the Iberian between these species were reported by Mackaye Peninsula. (2001). However, other authors of more recent publications continue to give validity to A. osiris Anancus in Eastern Europe (Sanders, 2011) as an independent species, a Despite the variability observed in the Anan- trend that it has been followed in the present work. cus samples from the Turolian to the Villafranchian of Eastern Europe, there is no real consensus on DISCUSSION the possibility of differentiating more than one spe- The dentition of the aforementioned species cies. The Russian species Anancus alexeevae Bai- of Anancus shows great variability in terms of the gusheva, 1971 does not seem to have been widely number of loph(id)s, a feature mentioned by many recognized in recent papers, although Titov (2008) other authors (Bergounioux and Crouzel, 1958; continues to maintain the subspecies Anancus arv- Metz-Muller, 1995, 1996; Hautier et al., 2009). The ernensis alexeevae for the Villafranchian form from intermediate molars of Anancus are normally Liventsovska (Russia). A systematic review of this tetralophodont, e.g., in A. arvernensis, A. group of Proboscidea is needed for Eastern kenyensis, A. capensis and A. osiris. In A. petroc- Europe. The outcome will depend on the complete- chii and A. ultimus, however, they are pentalopho- ness of the available fossil remains; the great vari- dont (Sanders, 2008, 2011). One specimen ability observed for the upper molars may lead to attributed to A. sivalensis from the Siwaliks, drawn distinctions between morphotypes. by Falconer and Cautley (1846), shows also a fifth Anancus in Africa loph in M2. The upper third molars of Anancus (Table 3) In Africa, several Anancus species have been can have five, six or seven lophs (Kalb and Froeh- differentiated, based on different combinations of lich, 1995). Thus, in the Eurasian species A. arv- cheek tooth features (Sanders, 2011). A. kenyensis ernensis, M3 is variable in terms of loph number; is a Late Miocene-Early Pliocene species (7.4-4.3 specimens with five or six are known. With regard

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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. Speci- mens not to scale.

to the African species, the M3 of A. kenyensis may ers, 2008). Supposedly, as indicated in the specific also have five or six lophs, while those of A. ultimus diagnosis, the molars of A. arvernensis are easily and A. capensis have six or seven (Hautier et al., recognized by their anancoid structure. Such is the 2009; Sanders, 2011). case for all the European specimens that have The number of lophs is therefore very vari- been attributed to this species (e.g., the Spanish able, and although most European specimens specimens from Las Higueruelas [MGM], Baza-1 have five in their upper third molar (Table 2), the [Ros, 2010] or Alcalá del Júcar [Mazo, 1997]), existence of an hexalophodont M3 in the Fonelas- although the lower dental elements generally show SCC-3 specimen does not by itself allow any defin- more marked anancoidy than the upper ones. itive classification to be made. However, it does Care must be taken when determining the allow the Metz-Muller’s (2000) assertion that there presence of anancoidy, especially in specimens is an absence of hexalophodont upper third molars showing advanced wear. The more worn a tooth is, in MNQ 17 to be rejected. the more marked anancoidy appears to be. This is The upper molars studied from other Euro- due to the fusion (observed by wear) of the pretrite pean sites and attributed to A. arvernensis, such as principal tubercles and the pretrite central conules those from Dorkovo (Bulgaria; Metz-Muller, 1995), immediately preceding them, and the union of the Baza-1 (Spain; Ros, 2010), Las Higueruelas postrite principal tubercles with the mesoconules. (Spain; our own data) and Hajnáčka (Slovakia; This leads to clearer alternation of the cusps than Lupták, 1997), show a clear anancoid pattern and would be seen in an unworn tooth. This is why the a pentalophodont M3 (except for some specimens Fonelas SCC-3 seems to show anancoidy in M2, from Dorkovo) with a thick cingulum that runs along yet there is no real anancoidy in either M2 or M3 the base of the crown ̶ traits not seen in the Fon- (see comparison in Figure 6). elas SCC-3 specimen. Certain anatomical similarities between A. Anancoidy ̶ the most important autapomorphic arvernensis and A. osiris should be noted (Figure trait included in the diagnosis of the genus ̶ is vari- 6). These were considered sister taxa by Tassy able in M3. African forms such as A. kenyensis and (1986). A. osiris was described in 1945 by Camile A. osiris barely show an anancoid pattern for their Arambourg based on a complete upper third molar third upper molars (Hautier et al., 2009). It is also recovered from a site in the deposits of Gizeh near very weak in A. petrocchii from As Sahabi (Sand- Cairo (Egypt), the age of which is uncertain. Sub-

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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).

sequently, A. osiris has been cited for other sites of de Manuel in Alfacar (Granada Basin), described North Africa, such as Ain Boucherit (Algeria; Aram- by Bergounioux and Crouzel (1958) and held at the bourg, 1970), with an age estimated at 2.32 Ma Madrid Natural History Museum (MNCN-12,959). (Sahnouni et al., 2002), and possibly in Ahl al This specimen might be classified as Anancus arv- Oughlam (Morocco), where Anancus sp. cf. A. ernensis turoliensis, although its revision is beyond osiris has been cited with an age of 2.5 Ma (Ger- the scope of this article. Hence, the absence of aads and Metz-Muller, 1999). In our opinion A. anancoidy observed in the Fonelas SCC-3 speci- osiris might be related to the last forms of A. arv- men is shared only by anancines recorded in Turo- ernensis (Figure 6, Table 3), but the former is sub- lian deposits, African Anancus and A. arvernensis stantially larger (Figure 7). Understanding the chilhiacensis from Chilhac (Table 3, Figure 6). phylogenetic relationships between Pleistocene It is important to note that the cement seen in species of Anancus depends on the discovery of the tooth-valleys of African Anancus has also been new samples that might allow a more in-depth reported in A. kenyensis and A. ultimus (Sanders, comparison. 2011). In European specimens, and in late repre- Some Spanish specimens in which the anan- sentatives of Anancus such as A. arvernensis chil- coid pattern is hardly noticeable have been classi- hiacensis and in the fossil described in the present fied as A. arvernensis. Such is the case of the only work, cementum appears in the valleys between anancine fossil (an upper third molar) from Molino

13 GARRIDO AND ARRIBAS: LAST IBERIAN MASTODON

the cusps. No cement is seen in A. arvernensis ACKNOWLEDGMENTS arvernensis. This study was funded by research projects Taking into account the number of lophs in the IGME 2005-009, IGME 2275.11 and by the Conse- upper molars, the absence of any basal cingulum jería de Cultura de la Junta de Andalucía. We or discernable anancoidy, the development of cen- express our gratitude to M. Kaiser and J. Roda, the tral cusps and a square shape for the distal part of owners of the land where the Fonelas SCC-3 site M3, together with the presence of cementum is located, for allowing us access throughout 2006. aggregates, the hemimaxilla recovered from Fon- M. McDonald, editor of the American Philosophical elas SCC-3 would appear to be that of a new chro- Society, kindly provided us with useful literature on nosubspecies. It is here proposed as Anancus fossil proboscideans. J. Pascual, of the IGME arvernensis mencalensis nov. ssp., the most recent library, provided us with invaluable help in finding Iberian gomphotherid known to date. documents. We also thank C. Bonnefon of the A trend towards reduced size in the upper Central Library of the Muséum National d'Histoire molars (Figure 7) between the Pliocene A. arvern- Naturelle (Paris) for her efforts in providing us with ensis arvernensis and the Early Pleistocene sub- useful literature. Dr. B. Sánchez-Chillón gave us species A. arvernensis mencalensis and A. access to comparative materials held at the arvernensis chilhiacensis should be noticed. Such MNCN, Madrid. We also thank Drs. M.P. Ferretti a reduction was previously indicated by Boeuf and W. Sanders for sharing with us their extensive (1992), who mentions that the jaw size of Anancus knowledge of fossil proboscideans. We would like arvernensis decreases in size eventually becoming to thank G.N. Markov for his comments on the pre- more "brevirostrine." Anancus arvernensis mencal- liminary manuscript. A. Burton, E. Díaz and I. ensis nov. ssp. can be understood as a native geo- Cotarelo provided us with linguistic assistance. graphical variation with its own molar characteristic Finally, we thank the valuable comments of an ̶ an adaptative solution to new southern Iberian anonymous referee which helped to improve the Quaternary environments as occurs in other Pleis- manuscript. tocene Euroasiatic Proboscideans (e.g., Mam- muthus; Lister et al., 2005), and probably indicating REFERENCES that an uncommon environment was developed in the Southern part of Iberia that would stimulate Alcalá, L. 1994. Macromamíferos Neógenos de la fosa local morphologic differentiation. de Alfambra-Teruel. Instituto de Estudios Turolenses, Museo Nacional de Ciencias Naturales, Teruel. CONCLUSIONS Alonso, M.A., Hoyos, M., and Alberdi, M.T. 2002. Tafonomía y ambiente sedimentario del yacimiento The recognition of Anancus arvernensis men- de Huélago. Estudios Geológicos, 58:11-25. calensis nov. ssp. in MNQ 17a, an Iberian form of Arambourg, C. 1945. Anancus osiris, un mastodonte intermediate-size between the typical Pliocene nouveau du Pliocène inférieur d'Egypte. Bulletin de European Anancus arvernensis arvernensis la Societé Géologique de France, 15:479-495. (MNQs 14-15-16) and the French Pleistocene Arambourg, C. 1970. Les Vertébrés du Pléistocène de Anancus arvernensis chilhiacensis (MNQ 17b), l’Afrique du Nord. Archives du Muséum National d’Histoire Naturelle, 10:1-127. supports the existence of a temporal cline among Arribas, A., Garrido, G., Viseras, C., Soria, J.M., Pla, S., these last gomphotheres of southwestern Europe. Solano, J.G., Garcés, M., Beamud, E., and Carrión, Anancus arvernensis mencalensis shows import- J.S. 2009. A Mammalian Lost World in Southwest ant changes in the upper third molar: the lophs Europe during the Late Pliocene. PLoS ONE, 4(9):1- face one another reducing anancoidy until its dis- 10. e7127. appearance, the distal part of M3 has a rectangular Aubekerova, P.A. 1974. A new species of mastodont outline, and cement aggregates are seen in the Anancus kazachstanicus Aubekerova from the loca- tooth valleys. The size of the upper third molar is tion of Esekartkan. Teriologiya, 2: 65-77. progressively reduced in the lineage A. arvernensis Aymard, A. 1855. Anancus Aymard. Anancus macroplus arvernensis-A. arvernensis mencalensis-A. arvern- Aymard. In Dorlhac, M.J, (ed.), Notice géologique sur ensis chilhiacensis. These changes could have le cratère de Coupet et sur son gisement de gemmes et d’ossements fossiles. Annales de la Societé occurred as a response to the aridification that d’Agriculture, Sciences, Arts et Commerce du Puy, began around 2.5 m.y.a., resulting in changes in 19:497-517. the composition of plant communities.

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Baigusheva, V.S. 1971. Fossil Terrestrial Fauna of the Hautier, L., Mackaye, H.T., Lihoreau, F., Tassy, P., Vig- Liventsovka Quarry (the Northeastern Azov Region). naud, P., and Brunet, M. 2009. New material of Anan- Materials on Anthropogene Faunas of the USSR, cus kenyensis (Proboscidea, mammalia) from Toros- 49:5-28. Menalla (Late Miocene, Chad): Contribution to the Bergonioux, F.M. and Crouzel, F. 1958. Les mastodontes systematics of African anancines. Journal of African d’Espagne. Estudios Geológicos, 14:223-365. Earth Sciences, 53:171-176. Boivin, P., Barbet, P., Boeuf, O., Devouard, B., Besson, Hay, O.P. 1922. Further observations of some extinct J.C., Hénot, J.M., Devidal, J.L., Constantin, C., and elephants. Proceedings of the Biological Society of Charles, L. 2010. Geological setting of the lower Washington, 35:97-101. Pleistocene fossil deposits of Chilhac (Haute-Loire, Hays, I. 1834. Descriptions of the specimens of interior France). Quaternary International, 223-224:107-115. maxillary bones of Mastodons in the Cabinet of the Boeuf, O. 1992. Anancus arvernensis chilhiacensis nov. American Philosophical Society, with remarks on the subsp. (Proboscidea, Mammalia), un mastodonte du genus Tetracaulodon. Transactions of the American Plio-Pléistocène de Haute-Loire, France. Geobios, Philosophical Society, N.S., IV:317-339. mém. sp. 14:179-188. Hopwood, A.T. 1935. Fossil Proboscidea from China. Brookes, J. 1828. A catalogue of the anatomical and Paleontologia Sinica, C9:1-108. zoological museum of Joshua Brookes, part 1. R. Illiger, C.D. 1811. Prodromus systematis mammalium et Taylor, London. avium additis terminis zoographicis uttriusque clas- Cautley, P. 1836. Note on the teeth of the Mastodon a sis. Salfeld, Berlin, 301 pp. dents etroites of the Siwalik Hills. Journal of the Asi- Kalb, J.E. and Froehlich, D.J. 1995. Interrelationships of atic Society of Bengal, 5:294-296. Late Neogene Elephantoids: new evidence from the Coppens, Y. 1965. Les Proboscidiens du Tchad, leur Middle Awash Valley, Afar, Ethiopia. Geobios, contribution à la chronologie du Quaternaire africain. 28:727-736. Actas del V Congreso panafricano de Prehistoria y Lister, A.M., Sher, A.V., van Essen, H., and Wei, G. 2005. de Estudio del Cuaternario:331-387. Museo Arque- The pattern and process of mammoth evolution in ológico de Santa Cruz de Tenerife (Canarias). Eurasia. Quaternary International, 126-128:49-64. Croizet, J.B. and Jobert, A. 1828. Recherches sur les Lourens, L., Hilgen, F.J., Shackleton, N.J., Laskar, J., ossements fossiles du département du Puy-de- and Wilson, D. 2004. The Neogene Period, p. 409- Dôme. Clermont-Ferrand, Paris. 440. In Gradstein, F.M., Ogg, J.G., and Smith, A.G. Falconer, H. 1857. On the species of mastodon and ele- (eds.), A Geological Time Scale 2004. Cambridge phant ocurring in the fossil state in Great Britain Part University Press. 1. Mastodon. Quarterly Journal of the geological Lupták, P. 1997. Upper jaw fragment of the Anancus arv- Society of London, 13:307-360. ernensis (Croizet & Jobert 1828) (Mastodontidae, Falconer, H. and Cautley, P.T. 1846. Fauna Antiqua Siva- Proboscidea, Mammalia) from the Villafranchian of lensis, Being the Fossil Zoology of the Sewalik Hills Hajnáčka, Slovakia. Geologica Carpathica, 48 in the North of India. Smith, Elder & Col. London. (5):341-343. Gaziry, A.W. 1997. Die Mastodonten (Proboscidea, MacInnes, D.G. 1942. Miocene and post-Miocene Pro- Mammalia) aus Dorn-Dürkheim 1 (Rheinhessen), p. boscidea from East Africa. Transactions of the Zoo- 73-115. In Franzen, J.L. (ed.). Die Säugetiere aus logical Society of London, 25:33-106. dem Turolium von Dorn-Dürkheim 1 (Rheinhessen, Mackaye, H.T. 2001. Les proboscidiens du Mio-Pliocène Deutschland). Courier Forschungsinstitut Sencken- du Tchad : Biodiversité, Biochronologie, Paléoécolo- berg, 197. gie et Paléobiogéographie. Unpublished PhD Thesis, Geraads, D. and Metz-Muller, F. 1999. Proboscidea University of Poitiers. (Mammalia) du Pliocène final d’Ahl al Oughlam Mazo, A.V. 1997. El yacimiento Rusciniense de Alcalá (Casablanca, Maroc). Neues Jahrbuch für Geologie del Júcar (Albacete). Taxonomía y bioestratigrafía. und Paläontologie, Monatshefte, 1:52-64. Estudios Geológicos, 53:275-286. Gray, J.E. 1821. On the natural arrangement of verte- Mazo, A.V. and van der Made, J. 2012. Iberian mast- brose animals. London Medical Repository, 15 odonts: Geographic and stratigraphic distribution. (1):296-310. Quaternary International, 255:239-256. Guérin C. 1990. Biozones or mammal units? Methods Mazo, A.V., Alberdi, M.T., and Bone, E. 1980. Le gise- and limits in biochronology, p. 119-130. In Lindsay, ment à Anancus arvernensis d’Alcolea de Calatrava E.H., Fahlbusch, V., and Mein, P. (ed s.), European (Ciudad Real) dans la Ruscinien (Pliocène) de la Neogene mammal chronology. Plenum. Mesete espagnole. Bulletin de la Société Belge de Guérin, C. 2004. Les Proboscidiens (Mammalia) du gise- Géologie, 89 (3):145-178. ment villafranchien moyen de Saint Vallier (Drôme, Mein, P. 1990. Updating of MN zones, p.73-90. In Lind- France). Geobios, 37, Supp. 1:306-317. say, E.H., Fahlbusch, V., and Mein, P. (eds.), Euro- pean Neogene mammal chronology. Plenum.

15 GARRIDO AND ARRIBAS: LAST IBERIAN MASTODON

Metz-Muller, F. 1995. Mise en évidence d’une variation Sahnouni, M., Hadjouis, D., Van der Made, J., Derradji, intra-spécifique des caractères dentaires chez Anan- A., Canals, A., Medig, M., Belahrech, H., Harichane, cus arvernensis (Proboscidea, Mammalia) du gise- Z., and Rabhi, M. 2002. Further research at the Old- ment de Dorkovo (Pliocène ancien de Bulgarie, owan site of Aïn Hanech, north-eastern Algeria. Jour- Biozone MN14). Geobios, 28 (6):737-743. nal of Human Evolution, 43:925-937. Metz-Muller, F. 1996. A mandible of Anancus arvernen- Sanders, W.J. 2007. Taxonomic review of fossil Probos- sis (Proboscidea, Mammalia, Pliocene) with pentalo- cidea (Mammalia) from Langebaanweg, South phodont M2’s – significance of the pentalophodont Africa. Transactions of the Royal Society of South grade in Anancus. Neues Jahrbuch für Geologie und Africa, 62 (1):1-16. Paläontologie, Monatshefte, 12:709-726. Sanders, W.J. 2008. Review of fossil Proboscidea from Metz-Muller, F. 2000. La population d'Anancus arvernen- the Late Miocene-Early Pliocene site of As Sahabi, sis (Proboscidea, Mammalia) du Pliocène de Libya. Garyounis Scientific Bulletin, Special Issue, Dorkovo (Bulgarie); étude des modalités évolutives 5:241-256. d'Anancus arvernensis et phylogénie du genre Anan- Sanders, W.J. 2011. Proboscidea, p. 233-262. In Harri- cus. Unpublished PhD Thesis, Muséum National son, T. (ed.), Paleontology and Geology of Laetoli: d'Histoire Naturelle, Paris. Human Evolution in Context. Volume 2: Fossil Nesti, F. 1825. Sulla nuova specie di elefante fossile del Hominins and the Associated Fauna. Springer Sci- Valdarno all’illustrisimo Sig. Dott. Prof. Ottaviano Tar- ence+Business Media. gioni Torzzetti (Lettera sopra alcune ossa fossili del Sanders, W.J., Gheerbrant, E., Harris, J.M., Saegusa, Valdarno non per anco descritte). Nuovo Giornale dei H., and Delmer, C. 2010. Proboscidea, p. 161-251. In Letterati II, 24:195-216. Werdelin, L. and Sanders, W.J. (eds.), Cenozoic Nomade, S., Pastre, J.F., Guillou, H., Faure, M., Guérin, Mammals of Africa. University of California Press. C., Delson, E., Debard, E., Voinchet, P., and Mes- Tassy, P. 1986. Nouveaux Elephantoidea (Mammalia) sager, E. In press. 40Ar/39Ar constraints on some dans le Miocène du Kenya. Cahiers de Paléontolo- French landmark Late Pliocene to Early Pleistocene gie, CNRS. large mammalian paleofauna: paleoenvironmental Teilhard de Chardin, P. and Trassaert, M. 1937. The Pro- and paleoecological implications. Quaternary Geo- boscideans of southeastern Shansi. Palaeontologia chronology. doi: 10.1016/j.quageo.2012.12.006. Sinica, C13:1-58. Osborn, M.F. 1926. Additional new genera and species Titov, V. 2008. Late Pliocene large mammals from North- of the mastodontoid Proboscidea. American Museum eastern Sea Azov Region. PhD Thesis, Rostov-on- Novitates, 238:1-16. Don: SSC RAS Publishing. Palombo, M.R. and Valli, A.M.F. 2003. Remarks on the Villalta, J.F. 1952. Contribución al conocimiento de la biochronology of mammalian faunal complexes from fauna de mamíferos fósiles del Plioceno de Villarroya the Pliocene to the Middle Pleistocene in France. (Logroño). Boletín del Instituto Geológico y Minero Geologica Romana, 37:145-163. de España, 64. Pla-Pueyo, S., Viseras, C., Soria, J.M., Tent-Manclús, Viret, J. 1954. Le loess à bancs durcis de Saint-Vallier J.E., and Arribas, A. 2011. A stratigraphic framework (Drôme) et sa faune de mammifères villafranchiens. for the Pliocene-Pleistocene continental sediments of Nouvelles Archives du Museum d'Histoire Naturelle the Guadix Basin (Betic Cordillera, S. Spain ). Qua- de Lyon, 4. ternary International, 243(1):16-32. Viseras, C. 1991. Estratigrafía y sedimentología del rel- Ros, S. 2010. Los Proboscídeos del Plio-Pleistoceno de leno aluvial de la Cuenca de Guadix (Cordilleras las Cuencas de Guadix-Baza y Granada. Unpub- Béticas). Unpublished PhD Thesis, Universidad de lished PhD Thesis, Universidad de Granada. Granada.