The type species of the suid genus
We here report the re-discovery of the collection of suids examined by Lartet (1851) and select a lectotype for the species
There are two taxa of bunodont tetraconodonts at Villefranche d'Astarac and Simorre, one of which represents a new genus, which is defined and described.
Por designación original, la especie tipo del género de suidos
En este trabajo se reporta el re-descubrimiento de la colección de suidos examinados por Lartet (1851) y se selecciona un lectotipo para la especie
Hay dos taxones de tetraconodontos bunodontos en Villefranche d’ Astarac y Simorre, uno de los cuales representa un nuevo género, que se define y describe.
As was the custom during the mid-19th century, authors generally did not nominate holotypes for the species that they named, and Lartet (
The absence of a holotype for
Lartet's (
In 1925, Pilgrim erected the genus
Vitry (
A cast of the teeth of the suid mandible from Le Fousseret (MHNT PAL 2012.0.206) housed in Toulouse which was donated to the University of Bologna, Italy, was mentioned by Stehlin (
Van der Made (
Van der Made (
Recent valorisation of the collections at the Muséum d'Histoire Naturelle, Toulouse, has led to the rediscovery of several specimens labelled on the fossils themselves with the words “
The aim of this paper is to describe and interpret the re-discovered suid fossils from Villefranche d'Astarac and to select a lectotype for the species
The fossils described herein are curated at the Muséum d'Histoire Naturelle, Toulouse, and the Muséum National d'Histoire Naturelle, Paris. An un-catalogued cast of a left mandible, from Le Fousseret, the original of which is housed at the Natural History Museum Toulouse (MHNT PAL.2012.0.206) is in the university collection in Bologna, Italy. Although the latter cast is quite poor, it is historically an important specimen because it contributed to the formation of Stehlin's (
Measurements of fossils were made with sliding calipers to the nearest tenth of a mm. Images were taken with a Sony Cybershot camera, and enhanced with Photoshop Elements 3.
The nomenclature of cusps, grooves and other structures in the teeth of suids follows the systems of Hünermann (
It is noted that Stehlin (
In the metric analyses provided in this paper, the breadth variation is given to the left of the bivariate plot (a thin vertical line for each tooth position with the range of breadths represented by a thick line), and the length variation beneath the bivariate plot (a horizontal line with the range of lengths represented by a thick line). This way of presenting the data provides visual and metric access to the proportional differences that exist between different tooth positions, an important source of information concerning tetraconodont teeth, because the inter-dental proportions help define the various taxa. Traditional bivariate plots of each tooth position on its own fail to distinguish several of the taxa, which can have very similar molar dimensions, for instance, but very different premolar dimensions. Likewise, traditional univariate plots fail to distinguish some taxa, but when combined with bivariate plots the differences emerge. Whilst this new way of presenting the data appears at first glance to be complicated, its advantages outweigh its inconveniences. The same measurement data are presented in the bivariate plots (in which tooth positions are labelled) and in the univariate plots. It is stressed that the combination employed in this paper yields a more coherent assessment of dental variation than either the bivariate or univariate plots on their own. Once the reference bivariate plots have been prepared, fossils from other localities can be compared with them, in this paper presented by letters of the alphabet in the univariate plots.
BSPG – Bayerische Staatsammlung für Paläontologie und Historische Geologie, München
DPZ – Department of Palaeontology, Zaragosa
GIU – Geology Institute, University of Belgrade
GPIT – Geologisches und Palaeontologisches Institut Tübingen
HLMD – Hessisches Landesmuseum, Darmstadt
IGGML – Institut für Geowissenschaften/Geologie de Montanuniversität, Leoben
IPS – Institut Català de Paleontoloia Miquel Crusafont, Sabadell
IPUW – Institut für Paläontologie, Universität Wien
LMJG – Landesmuseum Joanneum Graz
MHNT – Muséum d'Histoire Naturelle, Toulouse
MNHL – Museum of Natural History, Lisbon
MNHN – Muséum National d'Histoire Naturelle, Paris
MSIM – Museo San Isidro, Madrid
NHMB – Natural History Museum, Basel
NHMM – Naturhistorisches Museum, Mainz
NHMW – Naturhistorisches Museum, Wien
SMNS – Staatliches Museum für Naturkunde in Stuttgart
Tetraconodont suids have been found at many localities in Europe (
Location of selected European and North African tetraconodont localities. 1) Malartic, 2) Sansan, 3) Simorre, 4) Villefranche d'Astarac, 5) Bonnefond, 6) Le Fousseret, 7) Saint Gaudens (Valentine), 8) Lublé, Hommes, 9) Channay, Pontlevoy, 10) La Grive-Saint-Alban, 11) Fonte do Pinheiro, Pero Filho, Azambujeira, 12) Carpetana, Puente de Vallecas, Alhambra, Somosaguas, 13) Montejo de la Vega, 14) El Buste, La Ciesma, 15) St Quirze, 16) Can Llobateres, Can Ponsic, Santiga, 17) Mira, 18) Bou Hanifia (Dublineau), 19) Beglia, 20) Charmoille, 21) La Chaux-de-Fonds, 22) Gau-Weinheim (Wiesberg), 23) Esselborn, Eppelsheim, 24) Steinheim, 25) Breitenbrunn, 26) Urlau, Tutzing, 27) Pischelsberg, 28) Göriach, 29) Au, 30) Przeworno, 31) Klein Hadersdorf, Atzelsdorf, 32) Rosenthal 33) Neudorf, 34) Pitten, 35) Rudabanya, 36) Elgg, 37) Lucane, 38) Mala Miliva, 39) Nuri Yamut, 40) Pasalar, 41) Bâlâ, 42) Sahabi, 43) Wadi Natrun.
List of European Tetraconodont localities and their estimated correlations to the MN Zonation
Locality | Correlation this paper | Locality | Correlation this paper |
---|---|---|---|
Alhambra | MN 5-6 | Lucane | MN 5 |
Anwil | MN 7/8 | Mala Miliva | MN 5 |
Atzelsdorf | MN 9 | Malartic | MN 7/8 |
Au | MN 5-6 | Mira | MN 9 |
Azambujeira | MN 8-9 | Montejo de la Vega | MN 5 |
Bonnefond | MN 8-9 | Neudorf | MN 6 |
Breitenbrunn | MN 7/8 | Pasalar | MN 6 (or 5) |
Can Llobateres | MN 9 | Pero Filho | MN 6 |
Can Ponsic | MN 9 | Pischelsberg | MN 6 |
Channay | MN 5 | Pitten | MN 6 |
Charmoille | MN 9 | Pontlevoy | MN 5 |
El Buste | MN 7/8 or 9 | Przeworno | MN 6 or 7/8 |
Elgg | MN 5 or 6 or 7 | Puente de Vallecas | MN 5 |
Eppelsheim | MN 8 + 9 + 10 | Rosenthal | MN 5-6 |
Esselborn | MN 9 | Rudabanya | MN 9 |
Fonte do Pinheiro | MN 9 | Sansan | MN 6 niveau répère |
Gau Weinheim | MN 8 + 9 | Santiga | MN 9 |
Göriach | MN 6 | Simorre | MN 7/8 |
Hommes | MN 5 | Somosaguas | MN 5 |
Klein Hadersdorf | MN 7 | Steinheim | MN 7 |
La Chaux-de-Fonds | MN 7/8 | St-Gaudens (Valentine) | MN 7/8 |
La Ciesma | MN 9 | St Quirze | MN 7/8 |
La Grive-Saint-Alban | MN 7/8 | Tutzing | MN 6 |
Le Fousseret | MN 7/8 | Urlau | MN 7/8 |
Lublé | MN 5 | Villefranche d'Astarac | MN 7/8 |
Family Suidae Gray,
Tetraconodont suids in which the anterior premolars (p/1, p/2, P1/, P2/) are not vertically offset in the jaw with respect to the p/3 and P3/, crowns of anterior premolars slightly sectorial without marked mesial and distal buccal expansion and with quite tall, pointed main cusp, anterior premolars markedly smaller than P3/ and P4/; occlusal surface of talonid of m/3 slightly twisted with respect to the surface of the anterior lophids, distal root of m/3 slightly inclined buccally, talonid cusp positioned slightly buccal of the midline of the tooth; M3/ with posterior accessory cusplet slightly to the lingual side of the midline of the crown, largely posterior to the hypocone; dentine-enamel junction of the molars have relatively low relief (dentine exposure occurs with medium wear); lower male canines scrofic in section with shallow lingual and buccal longitudinal gutters and a prominent longitudinal ridge of cementum on the enamel-free distal surface.
Austria (Ilz, St Oswald): France (Bonnefond, Castelnau d'Arbieu, Channay (Faluns), Doué-la-Fontaine, Gers, Hommes, La Grive-St-Alban, Le Fousseret, Lublé, Meigné-le-Compte, Noyant-le-Plain, Pontigné, Pontlevoy, Simorre): Germany (Gunzberg, Hammerschmiede, Hinterauerbach bei Wartenburg, Kleineisenbach, Neufeld, Obere Susswassermolasse, Pischelsberg, Tutzing, Urlau, Wartenberg bei Erding): Poland (Przeworno): Portugal (Fonte de Pinheiro): Spain (Alhambra, Carpetana, El Buste): Switzerland (Braunkohle Ostschweiz, Elgg, Hohrain-Waldi, Schlatt am Kohlfirst).
partim
Species of
MN 6, Villefranche d'Astarac, France.
Austria (Ilz, St Oswald): France (Castelnau d'Arbieu, Channay (Faluns), Doué-la-Fontaine, Gers, La Grive-St-Alban, Le Fousseret, Lublé, Pontlevoy, Simorre): Germany (Hammerschmiede, Hinterauerbach bei Wartenburg, Kleinei-senbach, Neufeld, Pischelsberg, Urlau, Wartenberg bei Erding): Poland (Przeworno): Spain (Alhambra, Carpetana): Switzerland (Braunkohle Ostschweiz, Elgg).
The jaw fragment, MHNT PAL 2011.0.84.1.2, contains p/2 and p/3. The p/2 is a slender, sectorial tooth considerably smaller than the p/3. It has thin enamel, visible distally where flakes of enamel have broken off the tooth. The apex of the tooth is unworn and pointed, the mesial accessory cusplet low and exposing dentine, there is no cingulum, and the enamel is wrinkled. The cervix of the tooth is at the same level as that of the p/3 (ie there is no vertical offset between these two teeth, an important point considering the other specimens from the locality). The roots are almost vertical and not particularly enlarged.
The p/3 is a large tooth, heavily worn, with thin enamel. The mesial accessory cusplet is lightly worn and has cingular extensions buccally and lingually. The precristid is sharp and prominent, and the enamel wrinkled. The posterior accessory cusplet is low, small and with cingular folds buccally and lingually.
The p/4 is heavily worn with coarsely wrinkled enamel, a low mesial accessory cusplet from which rounded cingula depart buccally and lingually. The posterior accessory cusplet is worn, but was low and had cingular folds buccally and lingually. On the buccal side there is a cingular swelling, but not forming a true cingulum, and on the lingual side there is no sign of a cingulum. The anterior root is strongly developed.
The m/1 is very deeply worn. It has a large basal pillar in the buccal end of the median transverse valley.
The m/2 is heavily worn, showing four main cusps and well developed anterior, median and posterior accessory cusplets in the midline of the tooth. There is a large but low basal pillar in the buccal end of the median transverse valley. The beaded anterior cingulum is mesio-distally broad but fades out on the buccal and lingual surfaces of the crown. The enamel is wrinkled.
The m/3 is in heavy to medium wear with dentine exposed on the mesial cusp pair. The median accessory cusplet is compressed mesio-distally but broad bucco-lingually. There is a clear buccal cingulum. The talonid complex is positioned slightly buccally to the midline of the crown. The talonid is slightly twisted with respect to the two anterior lophids and this is reflected in the orientation of the distal root, which leans slightly laterally. This is important to note because in other specimens from Villefranche d'Astarac, the talonid of the m/3 is not twisted and the distal root is more vertically oriented in the jaw.
MNHN SML 76 is an unworn germ of a left P3/, which shows an apex that is subdivided into two points, a weak disto-lingual shelf bordered by cingular margin, a well developed mesial cusplet flanked either side by short cingular folds, and the enamel is wrinkled. Light wear would eradicate the evidence for the bifid apex.
The P4/, MNHN SML 37 possesses its roots, of which there are three, a large circular lingual one, and two subequal slightly compressed buccal ones.
The only M1/s in the collection are fragmentary and worn.
M2/, MNHN SML 57 has four main cusps at the corners of the tooth and prominent mesial, median and distal accessory cusplets in the midline. The mesial and distal cingula are beaded and do not reach the lingual and buccal surfaces of the crown. Both ends of the median transverse valley show low basal pillars which form shallow foveae between them and the bigger median accessory cusplet. There is quite marked buccal and lingual crown flare.
M3/, SML 36 shows a well developed buccal cingulum, above all on the buccal side of the paracone fading out distally. The crown narrows distally due to the small structures around the hypocone and the better developed hypoconule (talon complex). The four main cusps recall the same parts of the M2/ except that the second loph is narrower than the first one. Both specimens have a small pointed basal pillar in the buccal end of the median transverse valley, and a bigger, flatter pillar in the lingual end of the valley.
MNHN SML 72 is a right lower canine with a scrofic section. The lingual surface is much broader than the buccal surface. The distal surface is devoid of enamel and neither does it have a ridge of cementum.
MNHN SML 81 is interpreted to be left d/3 because of its tall triangular profile in lateral view. It has a small disto-lingual swelling which has almost been eradicated by wear. The precristid and mesial accessory cusplet are also heavily affected by wear. There are the bases of two roots.
MNHN SML 80 is a better preserved p/3 from Simorre, but it is broken lingually and mesially. The enamel is wrinkled, the main cusp is tall (27.2 mm) with prominent pre- and post-cristids, and a small development of a lingual cristid descending from the apex, but soon fading into the surface of the tooth. The distal cusplet is tall, and shows a lingual fold.
MNHN SML 39 is a left p/4 in medium wear. The mesial accessory cusplet is strong, with cingular folds either side of it, but which fade out on the lateral and lingual sides of the tooth. The distal accessory cusplet is strong and taller than the mesial one, and has broader folds either side, with depressions buccally and lingually separating it from the main cusp. The enamel is wrinkled and the tooth has three roots, a strong one mesially, and two distally.
MNHN SML 74 is a left m/3 in light wear lacking its roots. It shows a well formed beaded cingulum on the buccal surface of the protoconid which fades out distally before reaching the hypoconid. The talonid complex is positioned slightly to the buccal side of the midline of the crown. In other respects the tooth is similar to the specimen from Malartic described below.
The following description is based on the Toulouse (MHNT PAL 2012.0.206) and Paris mandibles (MNHN HGP 16) from Le Fousseret which represent a single individual. The wear pattern on the teeth is very close in the two specimens, and each of them has a green label stuck to it with the mention “
The left and right mandibles from Le Fousseret are poorly preserved and lack most of their bases. In the left mandible a small section of the ventral margin is preserved beneath the m/2, and here the jaw is 55 mm deep measured from the gingival level. The breadth of the jaw measured at the same place is 26.5 mm. The ascending ramus sweeps upwards starting at the level of the front loph of m/3, and thus in lateral view hides the rearmost part of the m/3. There is a mental foramen beneath the front of the p/4 at about half the height of the jaw.
The p/3 is lightly worn, with dentine showing down the postcristid of the main cusp, which is tall and located above the gap between the roots. There is a marked pre-cristid which ends basally in a prominent accessory cusplet flanked by cingular folds. Distally there is a small distal tubercle supported by tall cingular folds buccally and lingually, such that it is about half the total height of the tooth. The enamel is wrinkled, and there is no sign of an interstitial contact facet caused by abrasion with the p/2.
The p/4 is a robust tooth supported by two stout roots. It is moderately worn, exposing dentine from the apex of the main cusp down the post-cristid and onto the distal accessory cusplet. The latter structure is flanked by broad cingular folds. Anteriorly there is a precristid which terminates in a mesial accessory cusp and cingular folds. There is a prominent interstitial contact facet caused by the p/3.
The m/1 is deeply worn, exposing much dentine on all four main cusps as well as the three accessory cusplets arranged mesially, centrally and distally in the midline of the tooth. There is a well developed basal pillar in the buccal end of the median transverse valley.
The m/2 is worn to the stage where small dentine lakes have formed on the four main cusps and the median accessory cusplet. The mesial cingulum extends slightly onto the buccal surface where it shows three beads. The basal pillar in the buccal end of the median transverse valley is likewise formed of three beads, now worn down. Distally, there are cingular cusplets either side of the distal accessory cusplet.
The m/3 is constructed along the same lines as the m/2 except that behind the hypoconulid it possesses a stout talonid as tall as the main cusps, with strong beaded cingular folds extending mesially onto the buccal and lingual sides. The talonid is more or less in the midline of the crown. Because this tooth is less worn than the m/2, it shows remnants of the Fürchen and the coarse enamel wrinkles that typify tetraconodont molars. The cingulum is well developed at the buccal end of the median transverse valley, and on the mesial half of the protoconid.
Non
Large species of
Lectotype HLMD cast Fr3, right M3/. (The current whereabouts of the original fossil are unknown).
MN 8–9, Bonnefond, Haute Garonne, France.
France (Bonnefond); Germany (Gau-Weinheim; Obere Süsswassermolasse (Wartenberg bei Erding)); Portugal (Fonte de Pinheiro); Spain (El Buste, ?La Ciesma).
Wartenberg bei Erding (Obere Süsswassermolasse): BSPG 1952.I.279, left M3/
Large specimens of
The molars of
Bivariate and univariate plots of the upper teeth (top) and lower teeth (bottom) of
A suid jaw from the fluvial deposits of the Palaeo-Tagus at Fonte do Pinheiro, Portugal, was described by Roman (
Van der Made (
The question concerning the validity of the species
The mandible from El Buste possesses a deep sublingual fossa beneath the molar row, shallowing beneath the p/4 before joining the genial fossa. The symphysis, which is not fused in this sub-adult individual (p/4 in the process of erupting) extends back as far as the rear of p/2. The front of the jaw is broken, but shows parts of the alveoli for the central and second incisors, the small root (probably a female individual) and a two-rooted p/1. There is a short diastema between the p/1 and the p/2. There are two mental foramina, one beneath the front root of p/2, the other in the upper third of the jaw beneath the p/4.
It should be noted that the lectotype upper third molar from Bonnefond (27,5×20 mm) is considerably smaller than the M3/ in the type specimen of
Golpe-Posse (
Azanza (
It is concluded that the tetraconodont mandibles from Fonte do Pinheiro and El Buste belong to
Simorre: MNHN Si 322, right I3/; MNHN SML 34, right upper canine; SML 75, left M2/, SML 79, left M2/; SML 323, left lower canine; SML 269, right m/1; SML 73, left m/2; SML 1144, left m/2; SML 77, right m/2.
Some isolated or heavily worn tetraconodont teeth from Simorre and Rajegats cannot be attributed with certainty to a species. They are identified as
An isolated upper incisor, MNHN Si 322, is interpreted to be a right I3/ It has a broad distal crest descending from the apex, and a sharp lingual crest leading distally that closes off an elongated distal fovea. The protocrista ends in a low fold directed lingually.
The upper canine MNHN SML 34 is bucco-lingually compressed, is triangular in buccal view and has a root with a prominent lingual groove. There is a cementum deposit buccally. The posterior crista is sharp, and flanked by a zone on the buccal surface of the tooth which is devoid of enamel. The protocrista is sharp at its base but has been worn away towards the apex by contact with the lower canine.
The two M2/s from Simorre (MNHN SML 75, 79) vary slightly in dimensions and wear, but are morphologically similar to each other.
MNHN SML 323 is a deeply worn left central lower incisor, showing a prominent lingual crest and strong mesio-distal compression.
The only m/1s in the sample are broken, MNHN SML 269 being the front half of an unworn tooth showing the post-cristid of the metaconid detached apically to form a separate small cusplet, and Fürchen 2 and 3 meeting to carve off a small accessory cusp between the metaconid and the median accessory cusplet.
Three second lower molars in the collection (MNHN SML 73, 1144, 77) can be described together. SML 73 is unworn, shows wrinkled enamel, the detached post-cristid of the metaconid, and the accessory cusp between the metaconid and the median accessory cusplet. The mesial cingulum extends onto the buccal surface where it forms a series of beads before fading out distally. There is a cingular remnant in the buccal end of the median transverse valley but the lingual end of the valley is open. The hypoconulid is robust and flanked by two low cingular crests that lead mesio-lingually and mesio-buccally to close off two small foveae.
HLMD Fr 3, cast of the lectotype right M3/ of
Austria (Au bei Aufkenz, Göriach): France (Auverse, Doué-la-Fontaine, Faluns, Lasse, Malartic, Simorre (Rajegats), Villefranche d'Astarac): Germany (Eppelsheim, Gau Weinheim, Laichingen): India (Siwaliks); Pakistan (Siwaliks): Serbia (Mala Miliva): Spain (Puente de Vallecas, Somosaguas): Switzerland (Anwil): Turkey (Pasalar)
Species of
MNCN SOM N Cata 2 2008–83, left mandible containing p/2-m/3.
MN 5, Somosaguas, Madrid, Spain.
Spain (Somosaguas), Serbia (Mala Miliva)
The species name is for the Universidad Complutense, Madrid, in the grounds of which is situated Somosaguas where the type material was excavated.
SOM CATA 2'08–83 is an adult left mandible containing p/2 to m/3 in light wear. The jaw is deep and rather narrow (48,5 mm deep beneath m/1, 21 mm broad at m/1) with a deeply excavated sub-lingual fossa that extends anteriorly almost to the rear of the symphysis. The genial fossa is at the level of the back of the p/2. There are two mental foramina, one beneath the gap between p/2-p/3, the other beneath the rear of p/3, both are about mid-height of the jaw. There is a superior transverse torus slightly overhanging the genial fossa.
The p/2 has two roots that are parallel to each other and steeply inserted in the jaw. The crown has a long posterior projection of the crown which is slightly lower than the cervix of the p/3. The main cusp is positioned slightly mesial to the gap between the two roots, making the pre-cristid shorter than the post-cristid. There is a well formed mesial accessory cusplet with light cingular folds buccally and lingually. The post-cristid bifurcates about half way from the apex of the main cusp towards the distal margin of the crown, sending a low ridge linguo-distally, which makes the rear of the tooth broader than the front. The disto-buccal surface of the tooth is slightly swollen.
The p/3 is much larger than the p/2, its mesial accessory cusplet being taller than the entire p/2 in front of it and the main cusp about twice the height of the p/2. The crown is posed on two stout roots that slant slightly anteriorly. The main cusp is located above the rear root, which makes the pre-cristid longer than the post-cristid. The mesial accessory cusp is about half the height of the tooth, and has cingular folds lingually and buccally that border off low depressions behind them. The posterior accessory cusplet is taller than the mesial one, and is more voluminous with thick cingular folds extending buccally and lingually. The buccal enamel is heavily wrinkled and the enamel extends further rootwards buccally than lingually, especially on the distal root.
The p/4 is shorter than the p/3 but has approximately the same distal breadth. Its main cusp is also slightly lower than that of the p/3. The mesial accessory cusplet is about half the height of the tooth. The buccal cingular fold is strong and vertically oriented, but the lingual one is weak. The distal accessory cusplet is better individualised in the p/4 than in the p/3, and its cingular folds emerge from its rear edge. Narrow but quite deep grooves separate the distal accessory cusplet from the main cusp, especially marked on the buccal side of the crown. The enamel extends further rootwards buccally than lingually especially on the distal root.
The m/1 is damaged by cracking and displacement of the lingual half relative to the buccal half.
The m/2 is also cracked but the two parts are close together. The crown shows the usual four main cusps arranged in two lophids, with well formed mesial, central and distal accessory cusplets disposed along the mid-line of the tooth. The distal accessory cusp is large, the central one bucco-lingually broad, but mesiodistally compressed, and the mesial cusplet is small. The mesial cingulum is sharp and beaded and there are tiny cingular remnants in the buccal end of the median transverse valley.
The m/3 is lightly worn. The two anterior lophids are close in size and morphology to the m/2, but there is a prominent talonid complex comprised of the distal accessory cusplet accompanied distally by a large talonid cusp flanked by large cingular cusplets lingually and buccally. The talonid cusp is slightly to the buccal side of the midline of the crown. The buccal cingulum is well developed on the front half of the anterior lophid but fades out distally, but does leave a small remnant in the buccal end of the median transverse valley. In distal view the distal root of the m/3 is observed to slant slightly towards the buccal side of the jaw.
MNCN Som-N-83 the isolated lower left third molar is heavily worn, but it is possible to observe that the talonid is located slighty buccal to the midline of the tooth, and that its occlusal surface appears not to be twisted with respect to the surface of the two anterior lophids.
The upper third molar from Somosaguas (Van der Made & Salesa,
As was already noted by Van der Made & Salesa (
Petronijevic (
partim
Species of
IPS 1868, left maxilla containing P4/-M3/ (Golpe-Posse,
DPZ 2001, El Buste right mandible of
MN 5, Puente de Vallecas, Spain.
Austria (Au bei Aufkenz, Göriach): France (Auverse, Doué-la-Fontaine, Faluns, Lasse, Malartic, Simorre (Rajegats), Villefranche d'Astarac): Germany (Eppelsheim, Gau Weinheim, Laichingen): Spain (Puente de Vallecas): Switzerland (Anwil): Turkey (Pasalar).
Puente de Vallecas
Some of the fossils from Puente de Vallecas were briefly described by Golpe-Posse (
MSIM 2421 is a right mandible with p/4-m/3. The p/4 is a large, inflated tooth with the Innenhugel so close to the protoconid that with slight wear it has fused with it. The m/3 MSIM 290, has a centrally positioned talonid.
Villefranche d'Astarac
MHNT PAL 2012.0.197 is a left mandible fragment containing p/2-p/3 and an isolated p/4 which has interstitial wear facets that indicate that it belongs to the same individual. The p/2 is lower crowned than the p/3 and it is is offset vertically with respect to the p/3 such that its crown is aligned with the root of the p/3. This results in a distinct step in the occlusal surface of the tooth row. The main cusp of the p/2 is positioned above the gap between the two roots, thus in the anterior third of the tooth. The mesial accessory cusplet forms a low point from which low cingular folds depart buccally and lingually. The postcristid is long and bifurcates distally, sending a low rounded crest buccally. There is a slight bend in the cervix of the tooth above the gap between the two roots on the buccal and lingual sides. The roots are stout, the posterior one being enlarged, the mesial one smaller and both slightly inclined mesially.
The p/3 is a taller tooth with stout roots and the main cusp in the middle of the tooth. The mesial accessory cusplet is low and bordered by buccal and lingual cingular crests. The precristid is low and rounded due to light wear. The postcristid is worn away, with enamel exposed as far as the posterior accessory cusplet. This cusplet is low and is bordered by low, rounded cingular folds. In buccal and lingual views the cervix is observed to rise crownwards to a marked degree above the gap between the two roots. The enamel extends further towards the root on the buccal side than it does on the lingual one. The enamel is wrinkled. The two stout roots are inclined mesially.
Metric comparison of the upper (top) and lower cheek teeth (bottom) of
The p/4 is heavily worn, and shows a prominent posterior accessory cusplet with buccal and cingular folds walling off lingual and buccal depressions between the main cusp and the posterior accessory cusplet. The cervix is bent upwards above the gap between the two roots. The enamel extends further rootwards on the buccal side than the lingual one. The roots are stout.
The left m/1 (MHNT PAL 2012.0.195) lacks the disto-lingual cusp (entoconid). The crown is in light to medium wear, and shows four bunodont main cusps, and strongly developed median and posterior accessory cusplets. The anterior accessory cusplet is weak, as is the mesial cingulum. There is a prominent basal pillar in the buccal end of the median transverse valley.
MNCN SOM N Cata 2 2008–83, holotype left mandible of
MHNT PAL 2010.84.3 is a left m/3 in a fragment of mandible. The tooth is deeply worn exposing dentine in all the main cusps and the median and posterior accessory cusplets. The wear pattern so produced shows almost circular dentine islands separated from each other by thick enamel walls. There is a basal pillar in the buccal end of the median transverse valley. The talonid consists of a low cusp positioned slightly to the buccal side of the midline of the crown, directly behind the hypoconulid (posterior accessory cusplet) and it has well developed cingula running anteriorly on the buccal and lingual sides.
Isolated teeth of
The right maxilla in the Lartet collection (MHNT PAL 2012.0.196) contains the P2/ and P3/, and there is an isolated P4/ which belongs to the same individual judging from the compatible interstitial wear facets of the teeth. The P2/ is a low crowned tooth which is inserted in the maxilla well below the cervix of the P3/, which means that there is a prominent step or offset in the occlusal surface of the tooth row. There is almost no bend in the cervix on the lingual side, but a slight bend is present on the buccal side of the tooth. P2/ has two stout roots which are inclined slightly anteriorly, with the main cusp positioned above the gap between the two roots. The crown is moderately worn, with dentine exposed apically and in the distal cusplet complex. The mesial accessory cusplet is low, with cingular crests buccally and lingually. The distal accessory cusplet is deeply worn as is the disto-buccal end of the postcrista.
MNCN SMCN 3, stereo occlusal view of left m/3 of
The P3/ is a large tooth, triangular in occlusal outline. It has three roots, one stout one anteriorly, and two posterior ones. The tooth is deeply worn, which has eradicated most of the morphology, with the anterior wear facet extending onto the root of the tooth, but it is possible to make out that there was a mesial accessory cusplet bordered by buccal and lingual cingular folds. Above the gap between the mesial and distal roots the cervix bends rootwards to a slight extent on both the buccal and lingual sides of the tooth.
Isolated upper teeth of
The P4/ is heavily worn, but it is possible to deduce that it had two main cusps (one buccal, the other lingual) with pre- and post-crista leading from the buccal cusp towards the midline of the tooth anteriorly and posteriorly respectively. There are strong mesial and distal cingula.
Bivariate and univariate plots of the lower cheek teeth of
The right D4/ (MHNT PAL 2012.0.199) has a slightly trapezoidal occlusal outline, with four main cusps and weak mesial, median and distal accessory cusplets. The enamel is heavily wrinkled. Wear has exposed dentine at the apices of the four main cusps.
IPS 1868, left maxilla containing P4/-M3/, holotype of
The isolated right M2/ (MHNT PAL 2012.0.198) is deeply worn. It shows four main cusps with a strong median accessory cusplet, a weaker anterior cusplet and a weak posterior cusplet. It has strong lingual flare, but the buccal surface of the tooth is almost vertical.
MHNT PAL 2012.0.194 is a damaged right maxilla set in a plinth of plaster-of-paris. It contains the P4/ and M2/ in the bone, but the M3/ is isolated behind. The roots of the P3/ and M1/ are partly preserved. The P3/ has three roots, the M1/ has four. The P4/ is moderately worn, showing large dentine islands at the apices of both the buccal and the lingual cusps, surrounded by thick enamel walls. The mesial and distal cingula are well developed. The M2/ is deeply worn and has lost part of the hypocone. The mesial cingulum is broad, and there is a well developed basal pillar in the buccal end of the median transverse valley.
Lower teeth of
The M3/ is moderately worn, with the talon in a position lingual to the midline of the crown. This makes the lingual surface of the tooth almost straight from front to back, whereas the buccal surface curves markedly towards the rear. Most of the paracone is broken off, and there is dentine exposed on the protocone. The talon complex is mesio-distally narrow.
In Paris there is a left mandible fragment containing a worn m/3 (MNHN VAS 16) which has written on it in the handwriting of E. Lartet, the words Villefranche d'A. In the m/3 there is dentine exposed on all the main cusps and accessory cusplets, as well as on the hypoconulid. The enamel surrounding the enamel islands is thick, giving rise to rounded cusps with small indentations where the Fürchen are preserved (Hünermann,
Upper teeth of
MNHN Si 319 is a left maxilla fragment with the canine alveolus, the P1/, P2/ and P3/. There are short diastemata (5 mm) between the canine alveolus and the P1/, and between the latter tooth and the P2/ (specimen Si 319). The canine alveolus reveals that the tooth possessed a short root, and that it had a highly compressed section. The teeth in the maxilla are moderately (P1/-P2/) to heavily worn (P3/). The P1/ has two stout roots, the anterior one smaller than the posterior one. The main cusp lies above the gap between the two roots. The precrista descends from the apex of the main cusp towards the low mesial cusplet which has broken off leaving low buccal and lingual cingular crests. The distal crista bifurcates distally, with one crest slanting disto-buccal, the other distally. There is a low lingual cingulum walling off a shallow disto-lingual basin. The P2/ is a larger version of the P1/ with the main structures enhanced. The disto-buccal crista swells into a low cusplet. The P3/ differs from the preceding teeth by being considerably larger and taller, and in possessing three roots. The crown is heavily worn, but it is possible to make out a well formed mesial cusplet at the end of the mesial crista, from which cingular crests run rootwards buccally and lingually. A remnant of the lingual cingulum remains, but the linguo-distal cusplet has been worn away. In lateral view, it is possible to observe a distinct vertical offset between the cervix of the P1/-P2/ on the one hand, and the P3/ on the other, the distal part of the crown of P2/ touching the root of the P3/ about 3.5 mm beneath the mesial cusplet of the P3/ and about 8 mm beneath its occlusal surface. This specimen is similar in all respects to
MNHN Si 320 is interpreted to be an upper right P1/, but the crown is slightly damaged, such that it might represent a deciduous rather than a permanent tooth.
MNHN Si 321 is a left P1/ similar to that in the maxilla described above.
A right P3/ in the sample (SML 463) is in light wear.
The unworn P4/, MNHN SML 78, has a buccal main cusp which is subdivided at its apex into two points, a large mesial one, and a smaller, lower one immediately behind it. The lingual main cusp (protocone) is lower than the buccal one, and sends a precrista towards the midline of the mesial margin of the tooth but it does not join the mesial cingulum. The paracone sends two crests anteriorly, one directly towards a low cusplet at the buccal end of the mesial cingulum (parastyle), the other from its lingual surface anteriorly towards the midline of the tooth.The posterior cingulum is beaded and swells in the end of the sagittal valley to form a low distal accessory cusplet. The cingulum weakens buccally but is continuous from front to rear. There is no sign of the presence of cusplets in the sagittal valley. Both the buccal and lingual surfaces of the crown are strongly flared, making the tooth considerably broader than long.
The P4/s in the sample (MNHN SML 234, 337) are quite variable in dimensions, but all show the same basic morphology as the germ described above.
MNHN SML 324 is a worn right D4/. It is constructed along the lines of a permanent molar, but has a more trapezoidal occlusal outline, and thinner enamel.
An upper third molar from Simorre (MNHN SM 290) is broad distally due to relatively great development of the structures lingual to the hypocone, and the poor development of the talon complex.
Tetraconodont anterior upper premolars from Simorre, France, housed in the Muséum National d'Histoire Naturelle, Paris. A) SML 320, right P1/,
The first conclusion is that tetraconodont fossils from Villefranche d'Astarac represent two distinct taxa: 1)
The second conclusion is that the second tetracondont species from Villefranche d'Astarac is morphometrically similar to
Tetraconodont upper fourth premolars from Simorre, France, housed in the Muséum National d'Histoire Naturelle, Paris. A) SML 464, right P4/,
The discovery of the long “missing” fossil collections from Simorre and Villefranche d'Astarac studied by Lartet (
Tetraconodont lower incisor, lower canine and lower premolars from Simorre, France, housed in the Muséum National d'Histoire Naturelle, Paris. A) SML 683, right i/2,
Selecting a lectotype from the sample of bunodont suids from Villefranche d'Astarac is not a simple affair. Our examination of the fossils reveals that there are two morphotypes of bunodont suids at Villefranche d'Astarac, both with enlarged posterior premolars and reduced anterior ones. One is comparable in morphology to the material traditionally thought of as representing
MNHN Si 319, left maxilla of
Several authors (Hünermann,
To complicate matters, fossils labelled Le Fousseret, including a palate erroneously said to be from the site, were used by Van der Made (
Tetraconodont upper canine and upper and lower molars from Simorre, France, housed in the Muséum National d'Histoire Naturelle, Paris. A) SML 1144, left m/2,
In order to respect the concept of
Bivariate and univariate plots of the upper (top) and lower cheek teeth (middle and bottom) of
Finally, we stress the fact that the rediscovery of the Lartet (
Access to fossil collections was accorded by many colleagues: Herbert Lutz and Thomas Engel at the Naturhistorisches Museum, Mainz, Oliver Sandrock and Eric Milson at the Hessisches Landesmuseum, Darmstadt. We have been helped by curators in London (Andy Currant, Pip Brewer, Emma Bernard), München (Gertrud Rössner), Stuttgart (Reinhard Ziegler), Vienna (Ursula Göhlich), Graz (Martin Gross), Madrid (Jorge Morales, Patricia Perez), Lyon (Abel Prieur, Emmanuel Robert, Didier Berthet), Toulouse (Francis Duranton), Basel (Loïc Costeur, Martin Schneider), Paris (Christine Argot, Claire Sagne), Universidad Complutense (Manuel Hernandez), Museo San Isidro, Madrid (Mercedes Gamazo), Instituto Geologico y Minero de España (Alfonso Arribas, Guiomar Garrido), Institut Català de Paleontologia Miquel Crusafont, Sabadell (Salvador Moyà-Solà, Dani DeMiguel, Laura Celia) – we thank them all. Carlo Sarti (University of Bologna) kindly sent images of the cast of the mandible from Le Fousseret.