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VOL. 28, N° 2, 2009 Revue De Paléobiologie, Genève (Décembre 2009) 28 (2) : 471-489 ISSN 0253-6730

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VOL. 28, N° 2, 2009 Revue De Paléobiologie, Genève (Décembre 2009) 28 (2) : 471-489 ISSN 0253-6730 1661-5468 VOL. 28, N° 2, 2009 Revue de Paléobiologie, Genève (décembre 2009) 28 (2) : 471-489 ISSN 0253-6730 discussion, evolution and new interpretation of the Tornquistes Lemoine, 1910 (Pachyceratidae, Ammonitina) with the exemple of the Vertebrale Subzone sample (Middle oxfordian) of southeastern France Didier BerT*, 1 Abstract The Cheiron Mountain (Alpes-Maritimes, southeastern France) sample of Tornquistes Lemoine was collected in the Arkelli Biohorizon (Vertebrale Subzone, Plicatilis Biozone). Its study reveals its homogeneity whereas its morphology is between two nominal and classical species of literature : Tornquistes tornquisti (de LorioL) and Tornquistes oxfordiense (tornquist). It appears that the features usually taken into account to establish specific denominations in this genus (whorl section thickness, strength and density of the ornamentation, widening of the umbilicus) are in fact manifestations of the laws of covariation of the characteristics, and the extreme morphologies are interrelated by all intermediaries. There is now no taxonomical reason not to consider all the nominal taxa described in the Plicatilis Biozone as a single paleobiological species : Tornquistes helvetiae (tornquist). On the other hand, the stratigraphic polarity of the position of the primary ribs point of bifurcation (which decreases through time) is a major evolutionary feature in Tornquistes. It now allows defining at least three, maybe four, successive chronospecies : (1) (?) Tornquistes greppini (de LorioL), (2) Tornquistes leckenbyi (ArkeLL), (3) Tornquistes helveticus (JeAnnet) and (4) Tornquistes helvetiae (tornquist). Finally, although Protophites eBrAy has often been regarded as a microconch, it is clearly not the one of Tornquistes. The oldest species of Protophites now recognized is Protophites chapuisi (de LorioL) at the top of the Mariae Biozone (Praecordatum Subzone). Key words Ammonoidea, Tornquistes, Middle Oxfordian (Upper Jurassic), Arkelli Biohorizon, intraspecific variability, evolution, palaeobiology, Southeastern France. résumé discussion, évolution et nouvelle interprétation des Tornquistes Lemoine, 1910 (Pachyceratidae, Ammonitina) avec l’exemple de l’échantillon de la sous-zone à Vertebrale (oxfordien moyen) du sud-est de la France. – L’étude de l’échantillon de Tornquistes Lemoine récolté dans le biohorizon à Arkelli (sous-zone à Vertebrale, biozone à Plicatilis) de la Montagne du Cheiron (Alpes- Maritimes, sud-est de la France) révèle qu’il est homogène mais situé morphologiquement entre deux espèces nominales classiques de la littérature : Tornquistes tornquisti (de LorioL) et Tornquistes oxfordiense (tornquist). Il apparaît que les caractères longtemps pris en compte pour établir des coupures spécifiques dans ce genre (épaisseur de tour, vigueur et densité de l’ornementation, ouverture de l’ombilic) sont en fait la manifestation des lois de covariation des caractères, et les morphologies extrêmes sont reliées entre elles par tous les intermédiaires. Sur le plan taxinomique, il n’existe à présent aucune raison de ne pas considérer tous les taxons nominaux décrits dans la biozone à Plicatilis comme une seule et même espèce paléobiologique à grande variabilité : Tornquistes helvetiae (tornquist). D’autre part, la polarité stratigraphique de la position du point de bifurcation des côtes primaires, dont la hauteur décroît au cours du temps, est un caractère évolutif majeur chez Tornquistes. Il permet à présent de définir au moins trois, peut-être quatre, chronoespèces valides successives : (1) (?) Tornquistes greppini (de LorioL), (2) Tornquistes leckenbyi (ArkeLL), (3) Tornquistes helveticus (JeAnnet) et (4) Tornquistes helvetiae (tornquist). enfin, si Protophites eBrAy a souvent été considéré comme une forme microconque, il n’est clairement pas le dimorphe de Tornquistes. La plus ancienne espèce de Protophites à présent reconnue est Protophites chapuisi (de LorioL), au sommet de la biozone à Mariae (Oxfordien inférieur). Mots-clés Ammonoidea, Tornquistes, Oxfordien moyen (Jurassique supérieur), Biohorizon à Arkelli, Variabilité intraspécifique, evolution, Paléobiologie, sud-est de la France. I. IntroductIon recent works made in the Cheiron Mountain area (Alpes- Maritimes, southeastern France - Fig. 1) (Bert, 2003, This work is a further contribution to the study of the 2004, 2008 ; Bert et al., 2003 ; Bert & Bonnot, 2004). Subtethyan ammonite fauna of the Arkelli Biohorizon The genus Tornquistes Lemoine, 1910 is very scarce in at the top of the Vertebrale Subzone (Plicatilis Biozone, the Arkelli Biohorizon of this locality since it represents Middle Oxfordian). This research is a continuation of only 0.69 % of the total faunal ammonite (see Bert et al., * Corresponding address : Place de l’Eglise, F-04170 La Mure-Argens, France ; [email protected] 1 Université de Bourgogne, Laboratoire Biogéosciences, UMR CNRS 5561, 6 bd Gabriel, F-21000 Dijon, France. 472 D. BerT Fig.1 : Geographical location of the area listed. In blue are the Callovo-Oxfordian layers of the Cheiron Mountain (Alpes-Maritimes, southeastern France). 2003, fig. 3), and less than ten specimens were collected purely descriptive point of view, it is much less in terms in a single layer (see op. cit. for the representation of the of paleobiology (thierry & ChArpy, 1982, p. 645-646). outcrop-section). Under these conditions no real biometric In fact by the GAuse principle of competitive exclusion study could be undertaken. But due to their morphological it appears improbable that so many similar species that homogeneity and their isochrony all the specimens were potentially occupied the same ecological niche have been treated as one sample. The biostratigraphic framework coexisted such a long time without competition (mAyr, used for this work (Lower and Middle Oxfordian) is 1974, p. 48-49). This fact has led these authors to group that proposed by CAriou et al. (1997) and ammended some of the literature’s “species” together as variants of by Bert (2004, p. 11-12, fig. 1) especially when using other taxa. the Arkelli Biohorizon (GłowniAk, 2000) at the top of The goals of the present work are to study the Cheiron the Vertebrale Subzone, and the overlying Antecedens Mountain sample of Tornquistes, to clarify its taxonomic Biohorizon at the base of the Antecedens Subzone. position in relation to the literature nominal species, The genus Tornquistes Lemoine, 1910 was previously and to verify if these taxa have a real paleobiological revised by ChArpy (1976) then by thierry & ChArpy legitimacy from the variability (studied in the literature (1982). These autors have studied a total of over and unpublished datas). For this purpose a review of new 200 specimens divided into nine morphological nominal or currently accepted criteria for the characterization of groups for the single Plicatilis Biozone. Nevertheless, these taxa was performed. Finally, the taxonomic and the significant statistical analysis did not permit them to evolutionary implications are discussed. establish any clear division between most of these taxa. If this taxonomic multiplicity proves satisfactory of a Discussion, evolution and new interpretation of the Tornquistes Lemoine, 1910 473 II. the TornquisTes At the toP oF the Lemoine, 1910 (mArChAnd et al., 2002 ; this work). VertebrAle Subzone oF SoutheAStern The presence of such parabolic nodes is clearly not a FrAnce Stephanocerataceae feature but rather Perisphinctaceae. order Ammonoidea ZitteL, 1884 Suborder Ammonitina Hyatt, 1889 Family Pachyceratidae Buckman, 1918 Superfamily Perisphinctaceae Steinmann, 1890 Genus Tornquistes Lemoine, 1910 The classification of the family Pachyceratidae type species : Macrocephalites helvetiae tornquist, BuCkmAn, 1918 has been problematic for a long time. 1894 in tornquist, 1894, p. 8-10, pl.1, fig. 1a-d. For some authors it should be classified within the The specimen n° J19574 (coll. Basel Natural History Stephanocerataceae neumAyr, 1875 (douviLLé, 1912 ; Museum) from Anvill (Swiss) was featured by ChArpy ArkeLL, 1950, 1952 ; ArkeLL et al., 1957 ; ChArpy, (1976) as a neotype of Tornquistes helvetiae (tornquist, 1976 ; BourseAu, 1977 ; ChArpy & thierry, 1977 ; 1894). But this designation is not valid because this thesis thierry & ChArpy, 1982 ; Besnosov & miChAiLovA, remains unpublished. Moreover, this specimen has been 1983, 1991 ; GiL et al., 1985 ; krishnA & thierry, objected as a neotype by BourseAu et al., 1979 because 1987). So for ArkeLL (1950, and in ArkeLL et al., of too large ornamental differences from the original 1957) the Pachyceratinae are Cardioceratidae hyAtt, type-specimen. The same specimen was included in the 1892 intermediate between the Cadoceratinae hyAtt, morph kobyi de LorioL, 1896 by thierry & ChArpy 1900 and the Cardioceratinae hyAtt. On the other hand (1982, p. 633, pl. 2, fig. 1), therefore it was not proposed ArkeLL proposes the Tulitidae BuCkmAn, 1921 as again as a neotype of Tornquistes helvetiae (tornquist). ancestors of the Pachyceratidae and he classified them But in agreement with ornamental parameters (height of directly between the Sphaeroceratidae neumAyr, 1875 the bifurcation point of the ribs over half of the flanks - see discussion) this specimen is probably not of the and the Macrocephalitidae BuCkmAn, 1922. In contrast, for other authors the Pachyceratidae schould tornquist’s species neither of the de LorioL’s one : it could be in fact older than the Plicatilis Biozone. In be classified within the Perisphinctaceae steinmAnn, addition the Bukowkii
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