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Vol. 30, N° 2, 2011 VOL. 30, N° 2, 2011 Revue de Paléobiologie , Genève (décembre 2011) 30 (2): 619-684 ISSN 0253-6730 The Oppeliidae of the Acanthicum Zone (Upper Kimmeridgian) from Mount Crussol (Ardèche, France): ontogeny, variability and dimorphism of the genera Taramelliceras and Streblites (Ammonoidea) Cyril B AUDOUIN 1* , Patrick B OSELLI 2* & Didier B ERT 3* Abstract The Kimmeridgian outcrops of the Mount Crussol (in Ardèche), already long known for their abundant ammonites, contain many Oppeliidae. The study of the Acanthicum Zone (Upper Kimmeridgian) enables us to conclude that the microconchs of Taramelliceras compsum ( OPPEL ) and Streblites weinlandi ( OPPEL ) correspond to forms previously described by the authors as Glochiceras (Lingulaticeras ) crenosum QUENSTEDT and Creniceras dentatum ( REINECKE 7KHVWXG\RIDVLJQLÀFDQWVDPSOHIURPDSUHFLVHKRUL]RQ leads to treat both T. compsum ( OPPEL ) and 7SVHXGRÁH[XRVXP ( FAVRE ) as two morphological elements from the same paleobiological species. This is the same for S. weinlandi ( OPPEL ) and S. levipictus ( FONTANNES ), which are morphologically very close. Finally, studying the genera’s variability highlights mechanisms underlying it (heterochrony of the development and “laws” of covariation). Keywords Ammonoidea; Oppeliidae; Taramelliceras ; Streblites ; Ochetoceras ; Upper Kimmeridgian (Upper Jurassic); Crussol, Ardèche (France). Résumé Les Oppeliidae de la zone à Acanthicum (Kimméridgien supérieur) de la montagne de Crussol (Ardèche, France): ontogenèse, variabilité et dimorphisme des genres Taramelliceras et Streblites (Ammonoidea).- /HV DIÁHXUHPHQWV GX .LPPpULGJLHQ GH OD montagne de Crussol (Ardèche), bien connus pour leur richesse en ammonites depuis longtemps, contiennent en particulier une DERQGDQWHIDXQHG·2SSHOLLGDH/HXUpWXGHSRXUOD]RQHj$FDQWKLFXP .LPPpULGJLHQVXSpULHXU SHUPHWG·pWDEOLUTXHOHVPLFURFRQTXHV de Taramelliceras compsum ( OPPEL ) et Streblites weinlandi ( OPPEL FRUUHVSRQGHQWjGHVIRUPHVGpFULWHVSUpFpGHPPHQWSDUOHVDXWHXUV comme des représentants respectifs de Glochiceras ( Lingulaticeras ) crenosum QUENSTEDT et Creniceras dentatum ( REINECKE ). L’étude G·XQpFKDQWLOORQLPSRUWDQWSURYHQDQWG·XQKRUL]RQUHVWUHLQWFRQGXLWpJDOHPHQWjFRQVLGpUHU T. compsum ( OPPEL ) et 7SVHXGRÁH[XRVXP (FAVRE FRPPHGHX[H[SUHVVLRQVPRUSKRORJLTXHVGHODPrPHHVSqFHSDOpRELRORJLTXH,OHQHVWGHPrPHSRXU S. weinlandi ( OPPEL ) et S. levipictus ( FONTANNES TXLVRQWGHPRUSKRORJLHVWUqVSURFKHV(QÀQO·pWXGHGHODYDULDELOLWpGHFHVJHQUHVSHUPHWGHPHWWUHHQ pYLGHQFHOHVPpFDQLVPHVTXLODVRXVWHQGHQW KpWpURFKURQLHVGXGpYHORSSHPHQWHW©ORLVªGHFRYDULDWLRQ Mots-clés Ammonoidea ; Oppeliidae ; Taramelliceras ; Streblites ; Ochetoceras .LPPpULGJLHQ VXSpULHXU -XUDVVLTXH VXSpULHXU &UXVVRO Ardèche (France). I. INTRODUCTION monographs, but in a very typological way and without a very accurate stratigraphic calibration because of The historical site of the Mount Crussol (Fig. 1 and 2) their ancient collection (no bed-by-bed tracking). Since has provided many Kimmeridgian ammonites to the then, several works have highlighted the importance of SUHYLRXV DXWKRUV ZLWK D VLJQLÀFDQW SDUW FRQVWLWXWHG WKH LQWUDVSHFLÀF YDULDELOLW\ ZLWKLQ WKH 2SSHOLLGDH IURP by the representatives of the family Oppeliidae various ages ( HÖLDER , 1955; PALFRAMAN , 1966), and DOUVILLÉ , 1890. This family was particularly studied by the existence of a probably sexual dimorphism for that DUMORTIER & F ONTANNES (1876) and then FONTANNES family ( PALFRAMAN , 1966; M AKOWSKI , 1962; Z IEGLER , (1879). Many species of the genera Taramelliceras DEL 1974; Q UEREILHAC , 2009; K EUPP & R IEDEL , 2009). In CAMPANA , 1904, Streblites HYATT , 1900, Creniceras the main case of Taramelliceras , HÖLDER (1955) brought MUNIER -C HALMAS , 1892, Glochiceras HYATT , 1900 together different species in large morphological and and Ochetoceras HAUG , 1855 were described in their stratigraphic groups illustrating the polymorphism of this 1 79, rue Pierre Julien, F-26200 Montélimar, France; [email protected] 2 57 bis, avenue des Patriotes, F-26300 Bourg-de-Péage, France; [email protected] 3 Université de Bourgogne, Laboratoire Biogéosciences, UMR CNRS 5561, 6 bd Gabriel, F-21000 Dijon, France; [email protected]. * Groupe d’étude en Paléobiologie et biostratigraphie des Ammonites (G.P.A), F-04170 La Mure-Argens, France. 620 C. BAUDOUIN , P. BOSELLI & D. B ERT )LJ *HRJUDSKLFVHWWLQJDQGVLPSOLÀHGJHRORJLFDOPDSRIWKH&UXVVRODUHD genus. This applies to the broad group of Taramelliceras Submediterranean realm developed by the French compsum ( OPPEL , 1863) at the Upper Kimmeridgian. Group for the Study of the Jurassic ( HANTZPERGUES In genus Streblites , ZIEGLER UHFRJQL]HG VRPH et al ., 1997), completed and updated with the works of microconchs corresponding to forms assigned to the SCHWEIGERT (1999) and BAIER & S CHWEIGERT (2001). species “ Creniceras dentatum ” ( REINECKE , 1818); he noted that the stratigraphic range of genus Streblites coincides exactly with that of “ C. dentatum ” ( REINECKE ). II. GEOLOGICAL AND BIOSTRATIGRAPHICAL SETTING The recent harvest (C.B. & P.B.) of many Oppeliidae, done with a good stratigraphic tracking in the lower part of The historic area of Mount Crussol is located in Ardèche, the Upper Kimmeridgian in Mount Crussol, now allows RQ WKH ZHVWHUQ ÁDQN RI WKH 5K{QH 9DOOH\ GRZQ VRXWK XVWRFRQVLGHUWKHLUSDOHRQWRORJLFDOUHYLHZIRUDVSHFLÀF RIWKH5K{QH,VqUHFRQÁXHQFHMXVWDFURVV9DOHQFHFLW\ level of the Acanthicum Zone (Upper Kimmeridgian) 'U{PH)UDQFH 7KLVPRXQWULVLQJXSWRPKLJKW WDNLQJ LQWR DFFRXQW WKH LQWUDVSHFLÀF YDULDELOLW\ DQG is a small calcareous massif surrounded by Quaternary modern concept of the species in paleontology. The DOOXYLXP RI WKH JUHDW SODLQ RI 9DOHQFH )LJ 7KLV biostratigraphic framework used in this work (Fig. 3) site has long been known for its fossils abundance, and UHVXPHV WKH .LPPHULGJLDQ VWDQGDUG ]RQDWLRQ RI WKH numerous Jurassic outcrops dating from Toarcian to The Oppeliidae of the Acanthicum Zone (Upper Kimmeridgian) from Mount Crussol 621 Fig. 2: Localisation of the outcrop sections of Mount Crussol (from ATROPS S 7KH0DOOHWTXDUU\LV) 622 C. BAUDOUIN , P. BOSELLI & D. B ERT Fig. 3: Zonation of the Kimmeridgian (from HANTZPERGUES et al ., 1997). Tithonian can be observed. However the massif bulk is However, the sections he published do not exceed the made of limestones and marls from the Oxfordian and top of the Divisum Zone. In our work, the most recent WKH.LPPHULGJLDQVWDJHV7KDQNVWRPDQ\ROGTXDUULHVDV levels of those sections were completed using the ZHOODVQDWXUDORXWFURSVWKHVHTXHQFHRIWKHVHOHYHOVFDQ same numbering (Fig. 4 and 5). According to ATROPS , be observed (Fig. 2). The Jurassic of Mount Crussol has the Acanthicum Zone (and thus the base of Upper EHHQWKHVXEMHFWRIQXPHURXVZRUNVGRQHDORQJWLPHDJR Kimmeridgian) starts from bed No. 175, but beds No. (SAUTIER , 1854; L ORY , 1860; O PPEL , 1865; H UGUENIN , 175 to 178 form a very thick and currently inaccessible 1874; D UMORTIER & F ONTANNES , 1876; F ONTANNES , set that cannot be studied in our work. The Acanthicum 1879; R ICHE & R OMAN , 1921; R OMAN , 1950 - see Zone continues at least to the bed No. 197, but its upper HÖLDER & Z IEGLER , 1959 for a complete biography) OLPLWVWLOOKDVWREHFODULÀHG and more recently by ATROPS (1982) who focused his research on the Lower Kimmeridgian. The Oppeliidae studied in this work are mainly from the The log sections given by ATROPS (1982, p. 275-291, orderly working of beds No. 193 and 195 which form table 46-47) are a valuable and indispensable basis in a thin and easily accessible bundle containing abundant the study of the Kimmeridgian of the Mount Crussol. fauna. These levels are particularly visible around spot The Oppeliidae of the Acanthicum Zone (Upper Kimmeridgian) from Mount Crussol 623 Fig. 5: Outcrop section of the Acanthicum Zone of the Mallet TXDUU\ 1RDQGDWWKHWRSRIWKHJUHDW0DOOHWTXDUU\ )LJ 2, 4-5). Oppeliidae and Aspidoceratidae are predominant in the ammonite fauna of those beds (Fig. 6 and 7): Oppeliidae such as Taramelliceras compsum ( OPPEL , 1863) ([M] & [m]), Streblites weinlandi ( OPPEL , 1863) ([M] & [m]), Ochetoceras canaliferum ( OPPEL , 1863), and Aspidoceratidae: Aspidoceras acanthicum ( OPPEL , 3O ;, ÀJ Orthaspidoceras lallierianum (D’O RBIGNY 3O;,,ÀJ Sutneria cyclodorsata (MOESCH 3O;,,ÀJ 7KHUHDUHDOVR Nebrodites hospes ( NEUMAYR 3O ;, ÀJ Nebrodites gr. agrigentinus ( GEMMELLARO 3O ;,, ÀJ 1), representatives of the genus Discosphinctoides OLORIZ 3O;,ÀJ3O;,,ÀJ DQGUDUHO\ Aulacostephanus phorcus ( FONTANNES , 1876) (Pl. XII, ÀJ Phylloceras praeposterium ( FONTANNES , 1875) 3O;,ÀJ P. aff. VD[RQLFXP NEUMAYR , 1871 (Pl. XI, ÀJ Holcophylloceras polyolcum ( BENECKE , 1866) (Pl. ;,,ÀJ Ptychophylloceras ptychoicum ( QUENSTEDT , 3O;,ÀJ Lytoceras polycyclum NEUMAYR , 3O;,,ÀJ DQGWKHUHDUHDOVRVRPHQDXWLOXV with sinuous septa [ Pseudaganides cf. pseudaganiticus (SCHLOTHEIM @ 3O;,ÀJ $IHZEHOHPQLWHV Fig. 4: Section of the Acanthicum Zone of Mount Crussol. gastropods and bivalves complete the fauna. 624 C. BAUDOUIN , P. BOSELLI & D. B ERT Fig. 6: The ammonites’ genera in bed No. 193 (N=212). III. GOALS AND METHODS name. This approach is particularly convenient if the intermediaries between various possible morphotypes are The choice of a “paleobiological” approach UHFRJQL]HG LQWUDVSHFLÀFSRO\PRUSKLVP 7KXVHYHQLIDW Species concept in paleontology has undergone ÀUVWJODQFHWKHSDOHRELRORJLFDODSSURDFKVHHPVWREHDQ VLJQLÀFDQW FKDQJHV VLQFH WKH ÀUVW SDUW RI WKH WZHQWLHWK apparent loss of signal in the morphological information, century when the “typological species”
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