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Keywords:

  • Felidae;
  • Machairodontinae;
  • Miocene;
  • cranial anatomy;
  • Spain

Abstract

  1. Top of page
  2. Abstract
  3. Material and methods
  4. Systematic palaeontology
  5. Paramachaerodus maximiliani () ;
  6. Paramachaerodus orientalis () ;
  7. Promegantereon ogygia ()
  8. Phylogenetic analysis of Paramachaerodus and Promegantereon
  9. Conclusions
  10. References
  11. Supporting Information

Abstract:  A systematic revision of the sabre-toothed cat genus Paramachaerodus Pilgrim, 1913 is presented. Two species are recognized within Paramachaerodus, Pa. orientalis, and Pa. maximiliani, and the genus Promegantereon Kretzoi, 1938 is retrieved to include Promegantereon ogygia. Material from the Turolian Spanish localities of Crevillente-2 (MN 11, Alicante) and Las Casiones (MN 13, Teruel), which was previously assigned to Paramachaerodus, is now included in the tribe Metailurini. The exceptional discoveries at the Spanish Vallesian (MN 10, Madrid) fossil site of Batallones-1 have made it possible to characterize the dentition and cranial anatomy of a previously very poorly known machairodontine cat, formerly included in Paramachaerodus as Pa. ogygia, which now can be distinguished from Pa. orientalis and Pa. maximiliani by the following features: canines without crenulations, P3 with a marked disto-lingual expansion, P4 without ectostyle and with a well-developed protocone, M1 bucco-lingually elongated and double-rooted, m1 with a larger talonid, and primitive mandible morphology. Thus, the population from Batallones-1 constitutes a clearly different form from the genus Paramachaerodus, and we propose its inclusion in the genus Promegantereon Kretzoi, 1938, together with an upper canine from Crevillente-2 (MN 11), very similar to those from Batallones-1. In contrast, Pa. orientalis shows the following apomorphies: crenulated canines, P3 reduced in size and without disto-lingual expansion, P4 with a clear ectostyle as well as a reduced, backwardly displaced protocone and with a rounded and single-rooted M1. The species Pa. maximiliani is characterized by its slightly larger size, crenulated canines, very elongated P3 with a moderate disto-lingual expansion and P4 and M1 similar to those of Pa. orientalis. Paramachaerodus orientalis is recorded at Puente Minero (MN 11, Teruel), Concud (MN 12, Teruel), Crevillente-15, and Crevillente-16 (both MN 12, Alicante), and Paramachaerodus maximiliani in Venta del Moro (MN 13, Valencia). The available data suggest that Pr. ogygia was present in the Iberian Late Vallesian and Early Turolian faunas (MN 10 and MN 11) but disappeared after that age. Paramachaerodus was present in the faunas throughout the Turolian, with the species Pa. orientalis and Pa. maximiliani, this latter being probably part of the same immigration event that occurred in the Late Turolian and involved other mammal taxa such as camelids and ursids.

The upper Vallesian (Miocene) locality of Batallones-1 (MN 10, Madrid) has yielded one of the most complete samples of primitive sabre-toothed cats known in the fossil record, including up to 24 individuals of a medium-sized sabre-toothed cat previously referred by us (Salesa et al. 2005, 2006) to the species Paramachaerodus ogygia. Virtually, all bones of the skeleton are represented, and the skulls in particular present a unique opportunity for systematic revision of this hitherto poorly known species. As a result, we now propose that Pa. ogygia should be referred to the genus Promegantereon Kretzoi, 1938.

The systematics of the sabre-toothed felid genus Paramachaerodus have been very confusing for a long time, owing to the great number of species created on the basis of a scanty record, characters of poor diagnostic value and even changes in spelling. Paramachaerodus was created by Pilgrim (1913) to group several felid fossils from the Late Miocene localities of Pikermi (Greece), Maragheh (Iran), and Eppelsheim (Germany), which had been named, respectively, as Machairodus schlosseri Weithofer, 1888, Machairodus orientalis Kittl, 1887, and Felis ogygia Kaup, 1832, and some remains from the Dhok Pathan zone of Hasnot (Pakistan). Pilgrim did not include the species Machairodus hungaricus Kormos, 1911, but Kretzoi (1938) later did. In the same paper, Kretzoi created the genus Promegantereon for Felis ogygia. Surprisingly, Pilgrim (1915) included in the genus Paramachaerodus (under the name of Paramachaerodus cf. Paschlosseri) new material from the Dhok Pathan zone of Hasnot, India (now Pakistan) consisting of two hemimandibles, GSI-140 and GSI-141, despite the fact that these fossils differed markedly from each other and from all material previously assigned to this genus, as Matthew (1929) pointed out. Pilgrim did not designate a type species for the genus Paramachaerodus in the original description in 1913, but he did it later (Pilgrim 1931) and chose M. orientalis as the type. In 1929, Kretzoi had proposed the new genus Pontosmilus for the species Pa. ogygia, Pahungaricus, Paschlosseri, Paorientalis and Paindicus (for the mandibular fragment GSI-141), with Pa. orientalis as the type species and the genus Paramachaerodus restricted to a new species, Papilgrimi, which included only the specimen GSI-140 (Text-fig. 1A–C). However, this scheme was rejected by Pilgrim (1931) and his view has been adopted by subsequent authors, with the genus Pontosmilus being a synonymy of Paramachaerodus. Most recent authors have also tended to simplify the systematics of Paramachaerodus, recognizing only the two species Pa. ogygia (Kaup, 1832) and Pa. orientalis (Kittl, 1887) (Beaumont 1975, 1978; Morales and Soria 1977; Montoya 1994; Morlo 1997; Ginsburg 1999). We agree with others (Morales 1984; Alcalá 1994; McKenna and Bell 1997) in also recognizing the validity of the species Pa. maximiliani (Zdansky, 1924) from the Turolian of Shangyingou (China), a fossil site classically known as ‘Locality 12 of Zdansky’. This species, although morphologically similar to Pa. orientalis, is slightly larger than the latter and has more derived P3 and P4. Thus, in these more recent schemes, the species Pa. hungaricus and Pa. schlosseri are considered synonyms of Pa. orientalis. In our view, the species Pa. indicus should be transferred to the subfamily Felinae, owing to the absence of machairodont characters (such as the verticalization of the mandibular symphysis) and the presence of a very large p3 (inferred by the existence of two large alveoli), although at this stage we can offer no further opinion on its generic status. As for Paramachaerodus pilgrimi, the only one that Kretzoi (1929) recognized within this genus, we consider that the presence of a large p3 (equal in size to p4), the absence of a mesial cusp in both lower premolars, the absence of flattening in the lower canines and the markedly robust and thick mandibular corpus prevent its inclusion in either the genus Promegantereon or Paramachaerodus, although the machairodont affinities of its mandibular symphysis seem clear (Text-fig. 1A–C).

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Figure TEXT-FIG. 1..  Holotype of Paramachaerodus pilgrimi Kretzoi, 1929, GSI-140, a fragment of left hemimandible with broken lower canine, p3, p4 and partially broken m1, in A, occlusal, B, buccal, and C, lingual views.

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Based on this background, we present a review on the systematics of the genus Paramachaerodus and a detailed description of the fossils from Batallones-1, now included in the genus Promegantereon.

Material and methods

  1. Top of page
  2. Abstract
  3. Material and methods
  4. Systematic palaeontology
  5. Paramachaerodus maximiliani () ;
  6. Paramachaerodus orientalis () ;
  7. Promegantereon ogygia ()
  8. Phylogenetic analysis of Paramachaerodus and Promegantereon
  9. Conclusions
  10. References
  11. Supporting Information

The present revision is based on more than 14 partial or complete skulls, 15 mandibles and several isolated teeth of Promegantereon and Paramachaerodus collected from 7 Late Miocene fossil localities in Spain: Batallones-1 (MN 10, Madrid), Puente Minero (MN 11, Teruel), Crevillente-2 (MN 11, Alicante), Crevillente-15, and Crevillente-16 (MN 12, Alicante), Concud (MN 12, Teruel), and Venta del Moro (MN 13, Valencia) (Text-fig. 2). The abundant postcranial remains of Pr. ogygia from Batallones-1 are being studied in a set of separate papers and are not included in the present systematic revision because all previous taxonomic discussions have been based entirely on cranial and dental material. The measurements were taken with a digital calliper and are illustrated in Text-figure 3 and listed in Tables 1–3.

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Figure TEXT-FIG. 2..  Fossil localities discussed in the text. A, Map of Spain showing the location of fossil localities discussed in the text (BAT-1, Batallones-1; CD, Concud; CG, Cerro de la Garita; CR, Crevillente localities; PM, Puente Minero; VM, Venta del Moro). B, Stratigraphic distribution of the European localities discussed in the text. Data from Bruijn et al. (1992), Bernor (1986), Morales et al. (1992, 2000, 2004), and Montoya and Alberdi (1995).

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Figure TEXT-FIG. 3..  Explanation of the cranial, mandibular and dental measurements taken in this work (illustrations modified from Merriam and Stock 1932).

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Table 1.   Measurements in mm of the upper and lower dentition of Promegantereon ogygia from Batallones-1.
PieceMeasurementMax.Min.AverageSt. Dev.n
CMDL16.6313.3614.890.93028
BLW9.487.798.860.53121
CH40.1930.3935.772.81226
TL71.8762.0366.353.80110
P3MDL17.7311.4915.371.55220
BLW8.715.407.811.15212
CH8.436.767.610.48917
P4MDL27.4722.8325.151.19225
BLW13.4211.9612.640.50213
CH13.2110.2811.990.68821
cMDL11.889.4010.750.67927
BLW8.316.477.610.51526
CH23.9817.3720.931.78428
TL48.4240.3343.283.1337
p3MDL13.3610.4511.890.75536
BLW6.864.765.640.43828
CH7.775.126.210.59231
p4MDL18.7314.9516.460.90527
BLW8.216.677.340.38723
CH10.867.459.380.84122
m1MDL20.8917.5619.250.96822
PadL9.597.528.760.65319
PrdL11.269.2510.120.52521
BLW8.737.578.240.33421
PadH12.649.4610.630.78219
PrdH11.219.8010.490.46717
Table 2.   Measurements in mm of the skulls and mandibles of Promegantereon ogygia from Batallones-1.
MeasurementMax.Min.AverageSt. Dev.n
LAL73.8753.6064.955.24019
LPL148.51125.63137.807.0308
LR90.7970.3579.216.5118
LNB73.7061.0165.883.9878
LBO84.2360.5173.087.9379
LDT209.75174.20194.249.9979
LPPF124.66106.29115.056.2827
LNO143.85131.86137.858.4782
TFL112.29108.69110.621.8153
LP34.0330.6232.211.7163
LO36.9733.7435.951.4924
LB176.65153.05166.078.5859
LPA93.0373.3484.307.1607
LPP88.3378.6685.173.6157
LSD85.2073.2680.184.0296
BU33.2128.3831.231.53710
LCOR135.30111.54118.485.99515
CM174.9768.2071.631.98716
COM165.4553.7259.023.53218
LSY48.3641.4745.031.68323
HRM60.8150.7755.712.86414
MAM27.9425.1626.450.90118
LM132.29110.93122.116.10919
Table 3.   Measurements in mm of the upper and lower dentition of the different species of the genus Paramachaerodus and Promegantereon ogygia from Crevillente-2.
 PieceLocalityMDLBLWCHTL
  1. Localities: CON, Concud; CRE2, Crevillente-2; MAR, Maragheh, PM, Puente Minero; SHA, Shangyingou; VM, Venta del Moro (part of the measurements of Pa. orientalis are taken from Pilgrim 1931, and Kittl 1887, and those of Pa. maximiliani are from Zdansky 1924).

Paramachaerodus orientalis
 PM-575CPM16.718.06
 PM-576CPM16.358.62
 C-25CCON16.208.98
 C-26cCON11.637.3825.16
 CRA C-1P3CON12.806.466.92
 CRA C-2P4CON19.207.8710.75
 CG4Jp4CON19.438.8911.07
 NHMW 2007z0172/0001CMAR17.009.6041.44
 NHMW 2007z0172/0001P3MAR15.007.05
 NHMW 2007z0172/0001P4MAR28.0013.75
 M-8959cPIK14.009.50
 M-8959P3PIK14.00
 M-8959p4PIK19.00
 M-8959m1PIK22.009.50
 M-3829m1MAR23.0010.00
 M-3828P4MAR28.5014.00
Paramachaerodus maximiliani
 PMU-M69CSHA20.5011.00
 PMU-M69CSHA18.0011.00
 VM-38CSHA24.9310.47
 PMU-M69P3SHA16.307.20
 PMU-M69P4SHA29.0013.30
Promegantereon ogygia
 CR2-880CCCRE216.849.7343.4270.33

Institutional Abbreviations.  The institutions that house these materials and the corresponding abbreviations are as follows: Museo Nacional de Ciencias Naturales-CSIC (Madrid): Batallones-1 (B and BAT-1), Concud (CRA C, CG and C) and Venta del Moro (VM); Museu de Geología de la Universitat de València (Burjassot, Valencia): Crevillente-2 (CR2), Crevillente-15 and Crevillente-16 (CR). Fundación Conjunto Paleontológico de Teruel-Dinópolis: Las Casiones (KS), and Puente Minero (PM). We have used for comparisons material belonging to the collections of the Evolutionsmuseet of Uppsala Universitet, Sweden (PMU), Naturhistorisches Museum Vienna, Austria (NHMW), Hessisches Landesmuseum Darmstadt, Germany (HLMD-Din) and The Geological Survey of India, India (GSI).

Anatomical and measurement abbreviations

Dental abbreviations.  C, upper canine; c, lower canine; I1, first upper incisor; i1, first lower incisor; M1, first upper molar; m1, first lower molar; P1, first upper premolar; p1, first lower premolar. Dental measurement abbreviations: BLW, bucco-lingual width; CH, crown height; MDL, mesiodistal length; PadH, height of the paraconid; PadL, length of the paraconid; PrdH, height of the protoconid; PrdL, length of the protoconid; TL, total length of the piece (for canines).

Cranial measurement abbreviations.  BU, maximum length of the tympanic bullae; LAL, length from the rostral border of the lacrimal bone to the rostral border of the premaxilla; LB, length from the caudal border of the occipital to the rostral border of the premaxilla; LBO, dorsal length from the caudal border of the occipital to the rostral border of the sagittal crest; LDT, total dorsal length; LNB, length from the rostral border of the sagittal crest to the caudal border of the nasals; LNO, length from the caudal border of the nasals to the caudal border of the occipital; LO, length from the rostral border of the lacrimal bone to the postorbitary process of the zygomatic; LP, length from the rostral border of the lacrimal bone to the postorbitary process of frontal bone; LPA, length from the rostral border of the premaxilla to the caudal border of the palate; LPL, length from the caudal border of the occipital to the rostral border of the lacrimal bone; LPP, length from the caudal border of the palate to the caudal border of the foramen magnum; LPPF, length from the caudal border of the occipital to the rostral border of the postorbitary process of the frontal bone; LR, length from the caudal border of the nasals to the rostral border of the premaxilla; LSD, length of the upper dental series; TFL, length from the postorbitary process of the frontal bone to the caudal border of the occipital. Mandibular measurements abbreviations: CM1, length from the mesial border of the lower canine to the distal border of m1; COM1, length from the carnassial notch of m1 to the caudal border of the mandibular condyle; HRM, height of the mandibular ramus; LCOR, length from the mesial border of the incisors to the caudal border of the coronoid process; LM, length from the rostral border of the mandibular symphysis to the caudal border of the mandibular condyle; LSY, length of the lower jugal series; MAM, length from the dorsal border of the mandibular condyle to the ventral border of the angular process. General measurements abbreviations: Max., maximum; Min., minimum; St. Dev., standard deviation; n, number of individuals.

Cladistic analysis

The analysed characters and their states are described in the Appendix S1. The data matrix included 5 taxa and 25 unordered and unweighted cranial and dental characters (Table 4). It was compiled in MacClade 4.05 and run in PAUP 4.0b10 (Swofford 2002) using a branch and bound search, which produced 1 tree of 23 steeps (CI = 1.000; RI = 1.000). We used Proailurus lemanensis, from de Late Oligocene of Eurasia as outgroup, as this species is widely recognized as the earliest true felid (Véron 1995; Hunt 1998; Peigné 1999; Rothwell 2003). We also included Pseudaelurus quadridentatus, from the Middle Miocene of Sansan, as this form has been classically related to Pr. ogygia because of their similar dental morphology (Beaumont 1975).

Table 4.   Data matrix used in the phylogenetic analysis.
TaxaCharacters
12345678910111213141516171819202122232425
Promegantereon ogygia0101010000010101000111010
Paramachaerodus orientalis1111111010111111110111110
Paramachaerodus maximiliani111?110?0111111??1??11??1
Pseudaelurus quadridentatus?100000000000000000?11000
Proailurus lemanensis?000000000000000000000000

Systematic palaeontology

  1. Top of page
  2. Abstract
  3. Material and methods
  4. Systematic palaeontology
  5. Paramachaerodus maximiliani () ;
  6. Paramachaerodus orientalis () ;
  7. Promegantereon ogygia ()
  8. Phylogenetic analysis of Paramachaerodus and Promegantereon
  9. Conclusions
  10. References
  11. Supporting Information

Order CARNIVORA Bowdich, 1821 Suborder FELIFORMIA Kretzoi, 1945 Family FELIDAE Fischer, 1817 Subfamily MACHAIRODONTINAE Gill, 1872 Genus PARAMACHAERODUS Pilgrim, 1913

  • 1913 Paramachaerodus Pilgrim, p. 291.

  • 1924 Paramachairodus Zdansky, p. 130

  • 1929 Pontosmilus Kretzoi, p. 1298.

  • 1929 Protamphimachairodus Kretzoi, p. 1316.

Type species. Machairodus orientalis Kittl, 1887 from the Late Miocene locality of Maragheh (Iran, MN 12).

Remarks.  In his original publication Pilgrim (1913) did not assign a type species for the genus, although later (Pilgrim, 1931) he argued that his intention had been to designate Machairodus schlosseri. In that latter publication, he identified both Machairodus schlosseri and M. orientalis Kittl, 1887, based on material from Maragheh (Iran, MN12), as genosyntypes, from which he selected M. orientalis as the genolectotype while arguing that they were in any event synonymous.

The spelling of some genera and species names within the felid subfamily Machairodontinae has been confusingly variable, so that a comment on this matter is opportune. The genus Paramachaerodus was created by Pilgrim (1913), and this spelling has generally been adhered to as shown in the listings below. However, the spelling Paramachairodus (with an ‘i’) has also been used from time to time by authors such as Zdansky (1924) and more recently Beaumont (1975) with no formal proposal for the change and no explanation of it, and it was used by ourselves in earlier publications (Salesa et al. 2003, 2005, 2006) because it appeared to be the standard form. However, as no formal change has been recorded, and the variations appear to be arbitrary, the original spelling of Paramachaerodus must be retained and we do so in this publication. The genus Machairodus was created by Kaup (1832), and although this has been at times rendered as Machaerodus the original spelling is still in common use.

Emended diagnosis.  Machairodontinae with moderately enlarged and laterally flattened upper canines, and reduced lower canines; presence of crenulations in the borders of both upper and lower canines. I1 and I2 smaller than I3, which has a caniniform crown; absence of P1, p1, P2 and p2; P3 with a mesiodistally elongated crown, a small or absent mesial cusp, and a well-developed distal one; P4 with a straight buccal border, a small ectostyle, a well-developed parastyle and a relatively reduced protocone, mesiolingually oriented; relatively reduced M1; p3 clearly smaller than p4; m1 with a very much reduced talonid, composed of a simple crest. Mandibular symphysis without flange, but clearly verticalized, with a flat and rough rostral surface. Relatively wide nasal bones. Moderately developed sagittal crest.

Paramachaerodus maximiliani (Zdansky, 1924) Plate 1; Text-figures 4–5

  1. Top of page
  2. Abstract
  3. Material and methods
  4. Systematic palaeontology
  5. Paramachaerodus maximiliani () ;
  6. Paramachaerodus orientalis () ;
  7. Promegantereon ogygia ()
  8. Phylogenetic analysis of Paramachaerodus and Promegantereon
  9. Conclusions
  10. References
  11. Supporting Information
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Figure EXPLANATION OF PLATE 1.   Figs 1–2. Paramachaerodus maximiliani (Zdansky, 1924), Holotype, PMU-M69, partial skull from Shangyingou (China). 1, lateral view. 2, ventral view.

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Figure  TEXT-FIG. 4. . Paramachaerodus maximiliani (Zdansky, 1924) from Shangyingou (China). A, detailed view of the left upper dentition of PMU-M69, showing P3, P4 and M1. B, lingual view of right upper canine (PMU-M7266). C, lateral view of fragment of right hemimandible with fragmented p4 and m1 (PMU-M7260).

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Figure TEXT-FIG. 5..  VM-38, upper canine of Paramachaerodus maximiliani from Venta del Moro, in A, buccal, and B, lingual views.

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  •  1924 Machairodus maximiliani Zdansky, p. 120, pl. 27, figs 7–9; pl. 28, figs 3–5.

  •  1929 Protamphimachairodus maximiliani Kretzoi, p. 1316.

  •  1931 Paramachaerodus orientalis Pilgrim, p. 188, pl. 8, fig. 6.

  •  1945 Protamphimachairodus indicus Simpson, p. 189, pl. 8, fig. 7.

  •  1984 Paramachaerodus maximiliani Morales, p. 409.

  •  1997 Paramachaerodus maximiliani McKenna & Bell, p. 24.

  •  1999 Paramachaerodus orientalis Ginsburg, p. 192.

Holotype.  PMU-M69, partial skull from Shangyingou (Late Miocene, Locality 12 of Zdansky, Henan province, China) figured by Zdansky (1924, pl. 28, figs 3–5) and housed at the collections of the Evolutionsmuseet of Uppsala Universitet (Sweden).

Other material.  PMU-M7260, fragment of right hemimandible with fragmented p4 and m1 from Shangyingou figured by Zdansky (1924, pl. 27, figs 7–9) (Text-fig. 4C); PMU-M7266, fragment of right upper canine from Shangyingou figured by Zdansky (1924, pl. 28, fig. 5) (Text-fig. 4B); VM-38, fragment of left upper canine from Venta del Moro (MN 13, Valencia, Spain), described and figured by Morales (1984) (Text-fig. 5).

Emended diagnosis.  Skull with moderately developed sagittal crest. Relatively high zygomatic arches. Moderate postorbital process of frontal and very much reduced postorbital process of zygomatic. Relatively wide nasal bones, rectangular in shape. Premaxillae strongly projected rostrally, with a procumbent incisor arcade. I1 and I2 with basal cingulum and buccal and lingual protuberances. I3 markedly larger than I2 and I1, and I2 larger than I1. Crenulated upper canines. P3 with a moderate distal cusp and strong principal cusp; relatively elongated crown, showing a moderate disto-lingual expansion and a marked mesial constriction. P4 with a small ectostyle, a well-developed parastyle, large paracone and long metacone-metastyle; reduced protocone, located between the parastyle and the paracone and slightly antero-buccally oriented. P3 and P4 are aligned in occlusal view. p4 with high principal cuspid, well-developed distal cuspid, and strong distal cingulum; m1 with paraconid slightly lower than protoconid, mesial border of paraconid vertical, and absence of talonid.

Differential diagnosis. Paramachaerodus maximiliani can be distinguished from Pa. orientalis by its longer C-P3 diastema, the alignment of P3 and P4, the relatively smaller protocone on P4, and absence of talonid on m1; Pa. maximiliani can be distinguished from Promegantereon ogygia by its crenulated canines, procumbent incisor arcade, elongated P3, presence of parastyle on P4, straight buccal border of P4, protocone of P4 reduced and mesio-lingually oriented, presence of ectostyle on P4, relatively reduced M1, and absence of talonid on m1.

Spanish localities.  Venta del Moro (MN 13, Valencia).

Discussion.  This species is well known from the relatively good sample of Shangyingou (China), including an almost complete skull (PMU-M69). Its presence outside China is only recorded in Spain, in the Turolian locality of Venta del Moro, and represented only by one isolated upper canine. The fossil of Venta del Moro is slightly larger than the homologous piece from the Chinese locality, although their morphology is very similar: marked lateral flattening and presence of crenulations in both mesial and distal margins. Even though the upper canines of Pa. orientalis from Maragheh and Pikermi show this basic morphology, their size is much smaller, supporting the taxonomic validity of Pa. maximiliani. The felid sample from Venta del Moro also includes the tiger-sized sabre-toothed Amphimachairodus giganteus, with crenulated and flattened upper canines, but much larger than the specimen VM-38. The differences in size within the canine sample of Pa. maximiliani from China and Spain could be because of a marked sexual dimorphism, which would be higher than that of Promegantereon ogygia (Salesa et al. 2006) and closer to that in larger machairodonts such as Machairodus aphanistus (Antón et al. 2004; Salesa et al. 2006). The presence of Pa. maximiliani in these Chinese and Spanish localities poses an interesting paleogeographic question, because of the absence of fossils of this species in between those geographical areas. For the moment, there is no convincing explanation for this apparently vicariant distribution of Pa. maximiliani, although it is highly probable that this species were present throughout all Eurasia.

Paramachaerodus orientalis (Kittl, 1887) Plate 2; Text-figures 6–8

  1. Top of page
  2. Abstract
  3. Material and methods
  4. Systematic palaeontology
  5. Paramachaerodus maximiliani () ;
  6. Paramachaerodus orientalis () ;
  7. Promegantereon ogygia ()
  8. Phylogenetic analysis of Paramachaerodus and Promegantereon
  9. Conclusions
  10. References
  11. Supporting Information
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Figure EXPLANATION OF PLATE 2.   Figs 1–3. Paramachaerodus orientalis (Kittl, 1887), Holotype, NHMW 2007z0172/0001, partial skull from Upper Maragheh (Iran). 1, dorsal view. 2, anterior view. 3, ventral view.

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Figure TEXT-FIG. 6..  Detailed views of the holotype of Paramachaerodus orientalis (Kittl, 1887), NHMW 2007z0172/0001. A, buccal view of the right upper toothrow, showing I3, C, P3 and P4. B, occlusal view of the right upper toothrow, showing left I1, and right I1, I2, I3, C, P3, P4 and the alveolus of M1.

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Figure TEXT-FIG. 7..  Dentition of Paramachaerodus orientalis from Concud (A–F, I–O) and Puente Minero (G, H). A–C, right p4, CG4J, in A, occlusal, B, lingual, and C, buccal views. D–F, right p4, CRA C-2, in D, occlusal, E, lingual, and F, buccal views. G–H, left upper canine, PM-576, in G, buccal and H, lingual views. I–J, fragment of left lower canine, C-26, in I, buccal and J, lingual views. K–L, fragment of left upper canine, C-25, in K, buccal and L, lingual views. M–O, right p3, CRA C-1, in M, occlusal, N, lingual, and O, buccal views.

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Figure TEXT-FIG. 8..  Complete mandible, M8959, of Paramachaerodus orientalis from Pikermi. A, lingual view of left hemimandible. B, lingual view of right hemimandible. It is noticeable that the differences in the development of the mandibular symphysis that apparently exist among the two pieces are because of the presence of a hard matrix in this area, as both hemimandibles clearly belong to the same individual.

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  •  1887 Machairodus orientalis Kittl, p. 329, pl. 14, figs 15, pl. 15, figs 12.

  •  1888 Machairodus schlosseri Weithofer, p. 233, pl. 11, figs 17.

  •  1901 Felis orientalis Boule, p. 569.

  •  1901 Felis schlosseri Boule, p. 569.

  •  1911 Machairodus hungaricus Kormos, p. 182, pl. 17.

  •  1913 Paramachaerodus orientalis Pilgrim, p. 291.

  •  1929 Pontosmilus indicus Kretzoi, p. 1300.

  •  1929 Machaerodus sivalensis Matthew, p. 506.

  •  1930 Propontosmilus matthewi Kadic and Kretzoi, p. 48.

  •  1932 Paramachaerodus indicus Pilgrim, p. 189, pl. 8, fig. 7.

  •  1938 Paramachaerodus schlosseri Kretzoi, p. 108.

  •  1938 Paramachaerodus hungaricus Kretzoi, p. 108.

  •  1938 Propontosmilus matthewi Kretzoi, p. 108.

  •  1938 Pontosmilus orientalis Kretzoi, p. 109.

  •  1951 Paramachaerodus matthewi Kretzoi, p. 409.

  •  1952 Paramachaerodus schlosseri hungaricus Kretzoi, p. 24, pl. 1, fig. 1; pl. 2, fig. 4.

  •  1952 Pontosmilus orientalis Kretzoi, p. 24.

  •  1976 Megantereon orientalis Kurtén, p. 192.

  •  1994 Paramachairodus orientalis Alcalá, p. 150, pl. 8, figs k–n.

Holotype.  NHMW 2007z0172/0001, partial skull from the Late Miocene (Turolian, MN 11) of Upper Maragheh (Iran), housed in the Naturhistorisches Museum Vienna (Austria) (Pl. 2, Text-fig. 6).

Emended diagnosis.  Machairodontinae with crenulations in both upper and lower canines. Zygomatic arches as high as P. maximiliani. Relatively wide nasal bones. Premaxillae strongly projected rostrally, with a moderately procumbent incisor arcade. I3 markedly larger than I2 and I1, which are very similar in size. I1 and I2 with basal cingulum and buccal and lingual protuberances. Relatively short diastema between I3-C and C-P3. P3 with a well-developed principal cusp, without mesial cusp and with a small distal cusp; P3 crown laterally flattened and without disto-lingual expansion. P4 with a well-developed parastyle and a small ectostyle; with lingually oriented and reduced protocone, located between paracone and parastyle; long metacone-metastyle ridge. P3 and P4 are not aligned in occlusal view, the mesial border of P3 being lingually oriented. Rounded and single-rooted M1. Mandible with verticalized mandibular symphysis and relatively long diastema between c-p3; p3 smaller than p4, without mesial cuspid and with a very much reduced distal cuspid. p4 with developed mesial and distal cuspids, the former being smaller than the mesial one. Both p3 and p4 are wider distally than mesially. Lower carnassial with high paraconid and protoconid, the latter being higher than the former; there is a very small talonid, developed as a basal expansion of the crown, with no trace of cuspids on it.

Differential diagnosis. Paramachaerodus orientalis can be distinguished from Pa. maximiliani by its smaller size, relatively shorter muzzle, relatively smaller I3, relatively shorter P3, which lacks any disto-lingual expansion, and by the orientation of P3 in relation to P4, which is not aligned in occlusal view, the mesial border of P3 being lingually oriented, and the talonid of m1 slightly larger. Paramachaerodus orientalis can be distinguished from Promegantereon ogygia by its wider nasal bones, crenulated upper and lower canines, upper canines with higher degree of lateral compression, absence of disto-lingual expansion in P3, shorter P3, presence of ectostyle in P4, protocone of P4 relatively reduced and more distally placed, by the orientation of P3 in relation to P4, which is not aligned in occlusal view, the mesial border of P3 being lingually oriented, by the relatively reduced and single-rooted M1, absence of p2, by the presence of a mesio-buccal basal expansion on the crown of p4, by a reduced talonid in m1, by the more derived mandibular symphysis, which is highest and bears an incipient mandibular flange, and by the lower coronoid process.

Spanish localities.  Crevillente-15 and Crevillente-16 (MN 12, Alicante), Puente Minero (MN 11, Teruel), and Concud (MN 12, Teruel).

Material.  Puente Minero: PM-575 and PM-576 (upper canines belonging to the same individual) (Text-fig. 7G–H); Concud: CRA C-1 (fragment of right hemimandible with p3) (Text-fig. 7M–O); CRA C-2 (right p4, probably belonging to the same individual as CRA C-1) (Text-fig. 7D–F); CRA C-3 (fragment of left P3); CG-4J (right p4) (Text-fig. 7A–C) and C 25 (fragment of upper canine) (Text-fig. 7K–L).

Description.  The material from Puente Minero is described in Alcaláet al. (1991), and Alcalá (1994); the material from Concud is described in Morales and Soria (1977); and the material from Crevillente-15 and Crevillente-16 is described in Montoya and Alberdi (1995).

Discussion.  The temporal range of this species in Spain is the same as in the rest of Eurasia, that is, MN11 and MN12. The dental and cranial remains of Pa. orientalis and Pa. maximiliani show enough differences to be considered as two different species, although the latter has sometimes been cited as a synonym of the former. The dental, cranial, and mandibular morphology of both forms shares a derived state in relation to Promegantereon ogygia, supporting the existence of these two lineages of medium-sized sabre-toothed cats in the Late Miocene of Spain. For instance, the mandibular morphology of Pa. orientalis is more derived than that of Pr. ogygia, as the material from Pikermi shows, with a verticalized and very high mandibular symphysis (Text-fig. 8A–B). The large canine from Venta del Moro indicates a relative increase in body size within the lineage Pa. orientalis-Pa. maximiliani, but the known populations of Pa. orientalis show a relatively homogenous size, clearly different from the fossil of Venta del Moro, with the form from Pikermi being very similar in size to the form from Concud.

Genus PROMEGANTEREON Kretzoi, 1938

Type species. Felis ogygia Kaup, 1832 from the Late Miocene locality of Eppelsheim (MN 9, Germany).

Promegantereon ogygia (Kaup, 1832) Text-figures 9–13

  1. Top of page
  2. Abstract
  3. Material and methods
  4. Systematic palaeontology
  5. Paramachaerodus maximiliani () ;
  6. Paramachaerodus orientalis () ;
  7. Promegantereon ogygia ()
  8. Phylogenetic analysis of Paramachaerodus and Promegantereon
  9. Conclusions
  10. References
  11. Supporting Information
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Figure TEXT-FIG. 9..  Lingual view of HLMD-Din 1141, holotype of Promegantereon ogygia, from Eppelsheim, a fragment of right hemimandible with the lower canine, p3 and p4.

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Figure TEXT-FIG. 10..  Skulls of Promegantereon ogygia from Batallones-1. A, C, B-847, skull with articulated mandible and atlas in A, right and C, dorsal views. B, left view of B-1377, skull with articulated mandible. D, right view of BAT-1′06 F6-57, skull with articulated mandible. E, ventral view of the skull B-3434. F, ventral view of the skull B-7022.

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Figure TEXT-FIG. 11..  Mandibles of Promegantereon ogygia from Batallones-1. A–C, right hemimandible BAT-1′01 E5-17 in A, buccal, B, lingual, and C, occlusal views. D–F, right hemimandible B-462 in D, buccal, E, lingual, and F, occlusal views. G–H, complete mandible B-1676 in G, right buccal and H, occlusal views.

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Figure TEXT-FIG. 12..  Dentition of Promegantereon ogygia from Batalllones-1. A–B, left upper canine, B-3116, in A, buccal, and B, lingual views. C–D, left upper canine, B-566, in C, buccal, and D, lingual views. E–G, left I3, B-2490, in E, buccal, F, lingual, and G, distal views. H–J, left i3, B-1981, in H, buccal, I, lingual, and J, distal views. K–L, right lower canine, B-1007, in K, lingual, and L, buccal views. M–N, right lower canine, B-1471, in M, lingual, and N, buccal views. O–Q, right P3, BAT-1′02 D7-72, in O, occlusal, P, lingual, and Q, buccal views.

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Figure TEXT-FIG. 13..  Dentition of Promegantereon ogygia from Batallones-1. A–C, detail of the dentition of the hemimandible B-462, showing the alveolus for p2, p3, p4 and m1 in A, occlusal, B, buccal, and C, lingual views. D–E, fragment of left maxilla, B-3570, showing P3, P4 and M1 in D, buccal, and E, occlusal views. F–G, fragment of right maxilla, B-6051, showing the tip of the erupting upper canine, D3 and a partially erupted P4 in F, buccal, and G, occlusal views.

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  •  1832 Felis ogygia Kaup, p. 156, pl. 2, figs 6–8.

  •  1832 Felis antediluviana Kaup, p. 157, pl. 2, figs 9–12.

  •  1842 Felis Ogygea Blainville, pp. 144–145, pl. 16.

  •  1869 Felis pardus eppelsheimensis Köppen, p. 141.

  •  1879 Machaerodus ogygius Cope, p. 171.

  •  1888 Machairodus ogygia Weithofer, p. 233.

  •  1929 Pontosmilus ogygius; Kretzoi, p. 1299.

  •  1931 Paramachaerodus ogygia; Pilgrim, p. 140.

  •  1935 Paramachaerodus ogygius; Haupt, p. 39.

  •  1935 Neofelis (?) antediluviana; Haupt, p. 39.

  •  1938 Promegantereon ogygia; Kretzoi, p. 108.

  •  1975 Paramachairodus ogygia; de Beaumont, p. 394.

  •  2003 Paramachairodus ogygia; Salesa et al., p. 605.

Holotype.  HLMD-Din 1141, fragment of right hemimandible with lower canine, p3 and p4 from Eppelsheim (MN 9, Germany; Kaup 1832), housed in the Hessisches Landesmuseum Darmstadt (Germany) (Text-fig. 9).

Remarks.  The species name ogygia, initially created by Kaup (1832) for the species Felis ogygia, has at times been rendered as ogygius (Cope 1879; Kretzoi 1929) and even ogygea by Blainville (1842), but the original spelling of Kaup is the one in common use and is retained here.

Diagnosis.  Machairodontinae of the size of a puma, Puma concolor. Dentition without crenulations. Premaxilla only slightly projected rostrally, with a nonprocumbent incisor arcade. Upper canines with a moderately inflated root, with smooth and laterally flattened crown; both mesial and distal borders of the crown show a soft ridge, more marked on the mesial side, which continues lingually at the base of the crown. P1, p1, and P2 are absent. The P3 is wide, with a marked disto-lingual expansion; it has a small distal cusp and a minute or absent mesial cusp. P4 without ectostyle and with a well-developed protocone, placed between the parastyle and paracone and buccally oriented. Lower canines very much reduced in comparison with the upper ones; p2 vestigial but present; p3 smaller than p4, without mesial cuspid and with a very much reduced distal cuspid; p4 with developed mesial and distal cuspids, the former located lingually and the latter buccally. Both p3 and p4 are wider distally than mesially. Lower carnassial with high paraconid and protoconid, the latter being higher than the former; the m1 shows a small talonid, and a tiny metaconid above it. Mandible with moderately verticalized symphysis and high coronoid process.

Localities.  Batallones-1 (MN 10, Madrid) and Crevillente-2 (MN 11, Alicante).

Material.  Batallones-1: 16 complete or partial skulls (BAT-1′06 F6-57, B-6051, B-3434, B-4869, B-5197, B-3848, B-7022, B-4236, B-5406, B-7021, B-4766 (1), B-4054, B-1377, B-847, B-4322 and B-4778), 16 complete or incomplete mandibles (BAT-1′01 E5-17, B-5198, B-5269, B-732, B-2376, B-4208, B-1676, B-462, B-3848, B-751, B-2270, B-134, B-5406, B-7042, B-1377, B-4322 and B-4778), 20 upper canines (B-566, B-5270 (1), B-568, B-3116, B-271, B-7022, B-4322, B-5197, B-4869, B-1377, B-847, B-3961, B-3693, B-4640, B-4418, B-1, B-241, B-4236, B-5406 and B-4778), 11 P3 (B-7022, B-7021, B-3570, B-4322, B-3848, B-5406, B-1377, B-847, B-3434, B-5197 y B-4778), 13 P4 (B-1959, B-7022, B-7021, B-3570, B-3848, B-4766 (1), B-5406, B-1377, B-847, B-4322, B-3434, B-5197 and B-4778), 17 lower canines (B-96, B-1524, B-1007, B-1471, B-1117, B-601, B-4552, B-3848, B-134, B-5406, B-7042, B-1377, B-847, B-432 (2), B-1676, B-5198 and B-4778), 14 p3 (B-7042, B-2270, B-751, B-3848, B-134, B-5406, B-1377, B-847, B-4322, B-2376, B-4208, B-462, B-1676 and B-5198), 13 p4 (B-1961, B-751, B-7042, B-3848, B-3574, B-134, B-5406, B-4322, B-2376, B-4208, B-462, B-1676 and B-5198) and 11 m1 (B-2270, B-751, B-7042, B-3848, B-134, B-2673, B-1676, B-2376, B-4208, B-462 and B-5198), 1 D3 (B-6051), and several M1 and incisors.

Crevillente-2: one left upper canine, CR2-880.

Description

Cranium.  The skull of Promegantereon ogygia (Text-fig. 10A–F) is similar in size to that of an extant puma (Puma concolor). The nasals are relatively narrow, sub-rectangular in shape, with rostral and caudal margins of similar width and slightly curved caudally. The rostral border of the premaxilla is slightly curved rostrally, producing a U-shaped arcade, although less procumbent than that of Pa. orientalis. There is a short diastema, approximately 12 mm long, between I3 and the upper canine. The postorbitary process of the frontal is sharp, rough, and slightly laterally projected. The lambdoidal and sagittal crests are strong and very well developed. The frontal and sagittal crests join at the level of the postorbitary process of the frontal. The infraorbitary foramen is large and placed at the level of P3. The lacrimal foramen is placed in the inner face of the rostral border of the orbit. There is a very robust nuchal crest. The sphenopalatine foramen is smaller than that of a pantherin; it is placed in the orbital wall of the palatine bone, close to the suture maxillo-palatine. The posterior palatine foramen is much smaller than the sphenopalatine foramen, and it is located slightly rostrally to the latter. The optic foramen and the orbital fissure are placed as in a pantherin cat, so that the optic foramen is located slightly above and rostrally to the orbital fissure. The rotundum foramen is also developed as in pantherins, being located on the dorsal border of the caudal end of the pterygoid process. The oval foramen opens on the basisphenoid bone, between the pterygoid process and the postglenoid process. The zygomatic arch is much more robust than that of a similarly sized pantherin, and higher dorso-ventrally; its dorsal border, which delimits the ventral margin of the orbit, is straight. The postorbitary process of the zygomatic is vestigial, but it is always present. A central ridge, which runs from the level of the condyloid foramen to the aperture of the Eustachian tube, is frequently apparent in the basioccipital. The condyloid foramen is clearly separated from the posterior lacerum foramen and thus there is not a common depression for both foramina. The tympanic bullae are less inflated than those of a pantherin; they are developed from the paraoccipital process to the caudal border of postglenoid process, without making contact with the latter. The caudal ectotympanic penetrates rostrally beside the medial margin of the ectotympanic. The external auditory meatus is a single opening, rostro-laterally oriented. The stylomastoid foramen and the tympano-hyal depression are very close to each other. Both are placed in a more rostral position than in pantherins, that is, at the level of the central part of the bulla. The paraoccipital process is reduced in size when compared with that of pantherins, being dorsally displaced in comparison with its position in the latter. The mastoid process is well developed and projected ventrally to the level of the ventral border of the tympanic bulla. The pterygoid process has a rostrally curved caudal border. The palate is triangular-shaped, with a nasopharyngeal fossa delimitated by the walls of the pterygoid bone. The palatine fissurae are placed close to the rostral border of palate.

Upper incisors.  I1 has a laterally flattened crown, without cingulum, but with a smooth basal tuberosity on the lingual side. I2 is slightly larger and less compressed than I1; its lingual basal border has a light crest. I3 (Text-fig. 12E–G) is much larger than the other incisors; its crown is sharp and caniniform; its mesial surface is convex and smooth, whereas the distal one is concave and bears a light cingulum along its lingual border; on the buccal surface, there is a marked ridge from the base to the tip of the crown.

Upper canines.  The crown of the upper canines is laterally flattened and curves backwards (Text-fig. 12A–D). There are no crenulations on its margins. Both buccal and lingual surfaces are smooth, although the mesio-lingual border bears a marked crest developed from the base to the middle of the crown.

P3.  This tooth (Text-fig. 12O–Q) is placed parallel to the maxilla margin, with its mesial half slightly displaced lingually. The crown is mesio-distally lengthened, with a straight buccal border, and a marked disto-lingual protuberance, showing a marked constriction at the level of the main cusp; there is no cingulum, except on the distal border. There is no mesial cusp, whereas the distal and central cusps are strong with the latter being much higher. Just posterior to the distal cusp, there is a very much reduced distal tubercle.

P4.  The upper carnassial has a strong protocone, placed between the parastyle and paracone and buccally oriented (Text-fig. 13D–E). The parastyle is well developed, and there is no trace of ectostyle. The metacone-metastyle edge is slightly shorter than the parastyle, with a marked notch separating both structures. There is no cingulum. P3 and P4 are aligned in occlusal view.

M1.  This tooth (Text-fig. 13E) is very much reduced, although it has two roots; it has a bucco-lingually lengthened crown, which bears a light ridge developed from the mesio-buccal to the lingual borders.

D3.  The paracone (Text-fig. 13F–G) is well developed. The parastyle is very low, and in its mesial border there is a marked and buccally displaced ectostyle. The metacone is also low and is followed by a shortened metastyle. The protocone is strong and placed at the level of the metacone, which means displaced posteriorly in relation to the P4 protocone.

Mandible.  The mandibular corpus has two mental foramina, the anterior one being placed at the level of the postcanine diastema, and the posterior one, which is smaller, at the level of p3 (Text-fig. 11A–H). Nevertheless, the size of these foramina is very variable within the Batallones-1 sample and thus probably lacks taxonomic value. There may be accessory foramina, very small and placed around the mental foramina. The mandibular symphysis has a verticalized anterior border, and there is an almost straight angle between the ventral border of the symphysis and the mandibular corpus. In anterior view, the symphysis is high and rectangular-shaped; its lateral margins develop a light mental ridge, more marked in its ventral half. The masseteric fossa is deep and its anterior border reaches the level of the m1 protoconid. The coronoid process is lower than that of pantherins; its posterior margin shows a smooth backwards curvature and does not surpass the level of the anterior margin of the mandibular condyle. The angular process is postero-ventrally oriented, with a rough lateral border; its medial surface has an antero-posteriorly oriented ridge that produces a marked groove. The mandibular foramen is placed close to the ventral border, behind the level of the anterior margin of the coronoid process.

Lower incisors.  i1 has a laterally flattened crown, more than that of I1 (Text-fig. 12H–J). i2 is less compressed than i1; both i1 and i2 have morphologically very simple crowns. The crown of i3 is triangular-shaped, although less sharp than that of I3; the mesial face is smooth and convex, whereas the distal one is concave and has a light tuberosity in the disto-lingual border of its base; i3 is larger than i1 and i2, but the difference in size is less than that seen between the upper incisors.

Lower canines.  The crown is clearly smaller than that of the upper canine; it shows a smooth lateral compression, and it is curved backwards (Text-fig. 12K–N). The lingual face is concave and has a longitudinal ridge, whereas the buccal face is smooth and slightly convex.

p2.  This tooth is very much reduced and single-rooted, but it is present in 54% of the available sample of hemimandibles. Its crown is very small, low and has an elliptical shape that is mesiodistally elongated.

p3.  The crown is triangularly shaped, with a low central cuspid and a reduced distal one (Text-fig. 13A–C). There is no mesial cuspid. Behind the distal cuspid, there is a smooth posterior cingulum. In occlusal view, the mesial sharpening of the crown can be seen. This piece is not aligned with p4 and m1, its mesial part being lingually oriented.

p4.  The crown is triangularly shaped, with a high central cuspid and small mesial and distal cuspids (Text-fig. 13A–C); both mesial and distal cuspids are separated from the central one by marked notches. The crown has a marked posterior cingulum around the distal cuspid. As in p3, the crown of p4 is sharpened mesially.

m1.  The protoconid is higher than the paraconid, and they are separated by a marked notch (Text-fig. 13A–C). The talonid is reduced, but retains a tiny metaconid. There is no trace of a cingulum.

Discussion.  With the description of the new material from Batallones-1, the separation of Promegantereon ogygia from those species included in the genus Paramachaerodus is clear. In 1913, when Pilgrim introduced the genus, the species Pr. ogygia was solely known from some fragments of mandible with lower dentition, and with no available information on its upper dentition and skull morphology. Nevertheless, because Pa. orientalis, Pa. schlosseri and Pr. ogygia shared a reduced p3, all these forms were placed by Pilgrim within the same genus and separated from the Metailurini, the other group of similarly sized sabre-toothed cats. More recently, the excavations at Batallones-1 have yielded a huge amount of new anatomical information on Pr. ogygia and show it to be more primitive than the forms from Pikermi and Maragheh and also from the Metailurini. In the process, the Batallones-1 felid has become one of the best known of the machairodontines, and a complete characterization of its dental, cranial and mandibular anatomy has finally been possible. These new data clearly indicate its separation from Paramachaerodus and the need for another genus, Promegantereon, to accommodate that separation. Text-figure 14 shows the phylogenetic relationships discussed here in the form of a cladogram, characters used in which are given in the Appendix S1.

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Figure TEXT-FIG. 14..  Cladogram showing the phylogenetic relationships of the machairodontine genera Promegantereon and Paramachaerodus, and the primitive felids Pseudaelurus quadridentatus and Proailurus lemanensis. The numbers indicate which characters support each clade.

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The temporal range of Pr. ogygia in Eurasia is from MN 9 to MN 11, although in Spain this species is recorded in MN 10 (Batallones-1) and MN 11 (Crevillente-2). The holotype from Eppelsheim, a fragment of left hemimandible with lower canine, p3 and p4, is smaller than most of the homologous pieces from Batallones-1, but there is one hemimandible from this latter locality with p3-m1 (BAT-1′01 E5-17) that is even smaller than the holotype. There are also some small differences between the holotype and the Batallones-1 sample: the p3 from Eppelsheim is aligned with p4, whereas in the Spanish sample it is lingually oriented, although the whole area of the alveolus of p3 in the holotype is broken and glued and thus its orientation may require re-assessment; the p3 from the holotype seems to be less reduced in comparison with p4; the p4 in the holotype shows a much less developed posterior cingulum; and finally, the mandibular corpus is lower in the Batallones-1 specimens. Nevertheless, considering the different age of the two samples and the state of preservation of the holotype, we regard all these differences as being within the expected range of intraspecific variation, and we do not find reasons to separate the forms into different species.

Phylogenetic analysis of Paramachaerodus and Promegantereon

  1. Top of page
  2. Abstract
  3. Material and methods
  4. Systematic palaeontology
  5. Paramachaerodus maximiliani () ;
  6. Paramachaerodus orientalis () ;
  7. Promegantereon ogygia ()
  8. Phylogenetic analysis of Paramachaerodus and Promegantereon
  9. Conclusions
  10. References
  11. Supporting Information

The cladogram (Text-fig. 14) shows a clade including Pr. ogygia and the two species of Paramachaerodus supported by 6 synapomorphies, whereas that including the two species of Paramachaerodus is supported by 7 synapomorhies. A clade including Pseudaelurus quadrientatus, Pr. ogygia, Pa. orientalis and Pa. maximiliani is supported by three synapomorphies. The cladogram shows that the genus Paramachaerodus is more derived than the genus Promegantereon, which do not show any autapomorphy.

Conclusions

  1. Top of page
  2. Abstract
  3. Material and methods
  4. Systematic palaeontology
  5. Paramachaerodus maximiliani () ;
  6. Paramachaerodus orientalis () ;
  7. Promegantereon ogygia ()
  8. Phylogenetic analysis of Paramachaerodus and Promegantereon
  9. Conclusions
  10. References
  11. Supporting Information

During the Late Miocene, two different lineages of medium-sized sabre-toothed cats coexisted in the Iberian Peninsula. One of them, including Paramachaerodus orientalis and Pa. maximiliani, shows a more derived dentition, with crenulations in the canines, lengthening of P3, a more trenchant P4 (reduction of protocone and development of ectostyle), markedly reduced talonid in m1 and a more derived mandible that has lost p2. The other lineage, consisting at this moment of a single species, Promegantereon ogygia, retained primitive characters, such as the absence of crenulations, relatively short P3, a P4 with strong protocone and without ectostyle, a metaconid on the talonid of m1, and a relatively high coronoid process, and was present at Batallones-1 and Crevillente-2. Both lineages coexisted throughout the biozone MN 11, but after this there is no record of Promegantereon.

In parallel with the development of these lineages, another group of sabre-toothed felids, the Metailurini, underwent a rapid radiation from MN 11 onwards, leading to the appearance of several genera such as Metailurus, Therailurus, Fortunictis, Dinofelis and Stenailurus. The Metailurini also developed laterally flattened canines, although these were less compressed and shorter than those of the Smilodontini (Promegantereon, Paramachaerodus, Megantereon and Smilodon) and Homotherini (Machairodus, Amphimachairodus, Homotherium and Xenosmilus), in addition to a more slender postcranial skeleton (Roussiakis et al. 2006). The two species of Paramachaerodus became extinct in the Turolian (Late Miocene) and were replaced by the more derived and jaguar-sized Megantereon. The disappearance of both Promegantereon and Paramachaerodus may be associated with the progressive aridification that occurred during the Vallesian and Turolian (Fortelius et al. 2002), which led to the predominance of savannas over wooded habitats. In this context, it is noticeable that the metailurines had slender proportions in comparison with those of Pr. ogygia, which is the only species for which the postcranial skeleton is well known. These different proportions could be pointing towards an ecological segregation, with the metailurines occupying open spaces (Roussiakis et al. 2006) while Pr. ogygia favoured more forested areas, which might have prevented any competition between both groups of primitive sabre-toothed felids. This interpretation is also supported by differences in the development of the nasal bones, which were narrower in Pr. ogygia than in Pa. orientalis and Metailurus major. This also results in a narrower nasal cavity in the former, which itself may be related to the likely differences in the habitat. The body proportions of the Late Miocene–Pliocene genus Megantereon were very similar to those of Pr. ogygia, which probably indicates similar habitat requirements, but while the cranial and skeletal features related to the machairodont type of killing were just perceptible in Pr. ogygia (Salesa et al. 2005, 2006) they were much more developed in Megantereon and thus suggest a significant difference in hunting success between the genera.

The spreading of grasslands at the expense of forest at the end of the Vallesian (Agustíet al. 1999; Fortelius et al. 2002) surely benefited the metailurines, helping in their radiation and restricting the species diversity of the forest-dwelling sabre-toothed felids. The postcranial elements of the largest species of Paramachaerodus, Pa. maximiliani, are virtually unknown, making it impossible to assess the body proportions of this species; its size was intermediate between that of the earlier species of the genus and that of Megantereon, and this could have meant that the niche occupied by Pa. maximiliani became progressively narrower, leading to the disappearance of the species.

Acknowledgements.  This study is part of the research projects CGL2005-03900/BTE, CGL2004-02094/BTE, CGL2008-05813-C02-01, CGL2008-00034/BTE (Dirección General de Investigación-MEC and MCI), CNRS-PIC 2009137 and the Research Group CAM-UCM 910607. M.J.S. holds a ‘Ramón y Cajal’ contract from the Ministerio de Educación y Ciencia in the MNCN-CSIC. We thank the Comunidad Autónoma de Madrid (Dirección General de Patrimonio Histórico) and Gobierno de Aragón (Dirección General de Patrimonio Cultural) for their continuous funding support and research permissions, and A.T. thanks the British Council for travel funding. The National Geographic Society provided additional support (Grant 6964-01). We also thank Dr Stéphane Peigné (Muséum national d’Histoire naturelle, Paris), Dr Ursula Göhlich (curator for Vertebrate Palaeontology, Naturhistorisches Museum Wien), Alice Schumacher (photographer, Naturhistorisches Museum Wien), and Dr Oliver Sandrock (curator of the department of Palaeontology, Hessisches Landesmuseum, Darmstadt) for providing us with the images of the holotypes of Paramachaerodus maximiliani (S.P.), Pa. orientalis (U.G. and A.S) and Promegantereon ogygia (O.S.). We thank Dr María Consuelo Sendino for providing us with the images of Pa. orientalis from Pikermi, housed in the Natural History Museum of London, and Dr Andrew Currant (curator, Natural History Museum of London) for allowing us the access to that material. We also thank Dr Vijay Prakash Mishra (Director, Palaeontology Division, Northern Region, Geological Survey of India at Lucknow) for providing us with images of Pa. pilgrimi, and Dr Lars Werdelin (senior curator) and Maria Björnsdotter, both from Evolutionsmuseet of Uppsala Universitet, Sweden, for the access to the material of Pa. maximiliani from China.

Editor. Adriana López-Arbarello

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  2. Abstract
  3. Material and methods
  4. Systematic palaeontology
  5. Paramachaerodus maximiliani () ;
  6. Paramachaerodus orientalis () ;
  7. Promegantereon ogygia ()
  8. Phylogenetic analysis of Paramachaerodus and Promegantereon
  9. Conclusions
  10. References
  11. Supporting Information
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Supporting Information

  1. Top of page
  2. Abstract
  3. Material and methods
  4. Systematic palaeontology
  5. Paramachaerodus maximiliani () ;
  6. Paramachaerodus orientalis () ;
  7. Promegantereon ogygia ()
  8. Phylogenetic analysis of Paramachaerodus and Promegantereon
  9. Conclusions
  10. References
  11. Supporting Information

Appendix S1. Character analysis.

FilenameFormatSizeDescription
PALA_1013_sm_APPENDIX-Salesa.doc28KSupporting info item

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