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Ammonites and Nannofossils from the Campanian of Nalzen (Ariège, France)
AMMONITES AND NANNOFOSSILS FROM THE CAMPANI . OF NALZEN (ARIEGE, FRANCE) W. JAMES KENNEDY Department of Bartb Sciences, Univers@of Oxford, ParksRoad, Oxford, OX13PR, Grande-Bretagno.
MICHEL HANSOTrE Cos, 09000 Foix, France.
MICHEL BILOTrE Laboratoirada adologl~sddimentaira at Paldonto~gie, URA1405, UniversiMPaul Sabaliar,39 alldesJules Guesde,F-31062 ToulousaC$dex.
JACQUELINE B U R N E T r Department of GeologicalSciences, Untvars@CollageLondon, GowarStreet, London WCIE6BT, Grande-Bretagna.
RI~SUM~. La formation campanienne de Saint-Cirac, dans l'unit6 nord-pyr6n6ene du "Bassin" de Nalzen vient de livrer sa premi6re faune d'ammonites (plus de 100 individus), avec Desmophyllites diphylloides (FORBES, 1846), Eupachydiscus cf. isculensis (REDTENBACHER, 1873), Hoplitoplacenticeras (Hoplitoplacenticeras) marroti (CoQUAND, 1859), H. (H.) sp. juv., G~yptoxoceras aquisgranense (SEHL~2rER, 1872a), Baculites sp. et Scaphites (Scaphites) haugi DE GROSSOUVRE, 1894. C'est une laurie de la classique zone ~t marrot/, de la partie inf6rieure du Campanien sup6fieur dans la r6gion type en Aquitaine. Les ammonites sont associ6es /i une nannoflore fiche de plus de 90 esp6ces ; l'association de Broinsonia parca avec Bukryaster hayi et Marthastedtes furcatus indique la sous-zone CC18B du Campanien dans le sch6ma de Sissingh (1977, 1978) et Perch-Nielsen (1985). AMMONITESET NANNOFOSSILESDU CAMPANIENDE NALZEN(ARII~GE,FRANCE)
ABSTRACT The Marnes de Saint-Cirac (Campanian) in the north-pyrenean unit known as the Nalzen "Basin" has yielded an ammonite fauna of more than 100 specimens, with Desmophyllites diphylloides (FORBES, 1846), Eupachydiscus cf. isculensis (REDTENBACHER, 1873), Hoplitoplacenticeras (Hoplitoplacenticeras) marroti (COQUAND, 1859), H. (H.) sp. juv., Glyptoxoceras aquisgranense (SCHL~rER, 1872a), Baculites sp. and Scaphites (Scaphites) haugi DE GROSSOUVRE, 1894. This is a fauna of the classic marroti zone of the lower Upper Campanlnr~ in the type region in Aquitaine. The ammonites are associated with a rich nannoflora of more than 90 species ; the association of Broinsonia parca with Bukryaster hayi and Marthasterites furcatus indicates subzone CC18B of the Campanian in the scheme of Sissingh (1977, 1978), et Perch-Nielsen (1985).
INTRODUCTION
bulloides VOLGER, a. rosetta (CARSEY), G. ventricosa WnrrE, Rosita fomicata (PLUMMER) (Bilotte 1975,
In the north pyrenean unit known as the N',dzen "Basin", the Marnes of Saint-Cirac have been referred to the Campanian sensu lato on the basis of the rudistids (Toucas 1903 ; Rami~re de Fontanier 1933 ; Astre 1954; Bilotte 1985), benthic, and rare planktonic foraminifera present (Fallotia cf. colomi SILVF_SI~, Abrardia cf. mosae (HOFKER), Globotruncana arca (CuSIIMAN), G.
1985). The discovery by one of us (M.H.) of a horizon rich in ammonites, including the most remarkable heteromorphs yet described from the Upper Cretaceous of France, allows the dating of a part of the sequence as lower Upper Campanian. The ammonites are associated with a rich nannoflora, which in turn permits a direct correlation of macro- and nannofossil zones.
Manuscrit accept6 d6finitivement le 22.01.1992
eobios, 199~ 25, fasc. 2 i 9 263-278 1)
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264 GEOLOGICAL SETTING AND P A L A E O G E O G R A P H Y (M.B.)
PALAEOGEOGRAPHIC AND SEDIMENTOLOGICAL CONTEXT
STRUCTURAL CONTEXT
During the later part of the Late Cretaceous, the subpyrenean trough had an Atlantic polarity, bounded to the north by the Aquitaine platform (Ricateau & Villemin 1973). The upper Cretaceous of the Nalzen "Basin" was deposited on the southern side of the trough, as is indicated by the nature and distribution of facies within the Formation de Saint-Cirac (Bilotte 1990; r,g. 2).
To the east of the valley of the Ari6ge, from the meridian of Foix to that of B61esta, the Nalzen "Basin" extends for some 25 kilometres approximately. Situated in the north pyrenean zone, it has tectonic boundaries; the Pech de Foix fault to the north, the frontal thrust of the Saint-Barth616my massif to the south, and the Croquier fault to the east (fig. 1). The Nalzen "Basin" is a complex structure, in which three major units can be recognised (Bilotte 1975, 1985, Bilotte et al. 1988). The northern Saint-Cirac unit is a narrow slice in the form of an anticlinorium, in which the upper part of the Upper Cretaceous is represented by the Formation de SaintCirac.
This unit is between 300 and 500 m thick, although its full extent cannot be determined. The dominant lithology is of blue, highly argillaceous marls, with two associated facies : - fine - and medium-grained sandstones up to 1 m thick, with parallel, sharp upper and lower boundaries ; - rudistid horizons ; in situ in the north east, with biocoenoses at Benaix, Villeneuve d'Olmes, and P6r6ille d'en bas, and resedimented in the south west, with thanatocoenoses at St. Cirac (Catufet, Peyriguel, Pujet ; fig. 2). THE AMMONITE HORIZON (M.H.)
_
5 ~.~.~s,, . /
:)
St.Cirac r V//////////////~c fmt. I
The realignment of the D.117 road between Catufet and Manau exposed about a dozen metres of the Marnes de Saint-Cirac in a blue marl facies, dipping at 20~ to the SSE, as is indicated by rare thin beds of sandstone. The fossil horizon is exposed to a thickness of 1.5 m, at the base of the talus on the south side of the road (Foix 1 : 50,000 topographic sheet, coordinates x = 549, 27 ; y -70, 60, altitude 515 m (Fig. 3).
AGE OF THE AMMONITE FAUNA
(W.J.K.) The ammonite assemblage from the Formation de SaintCirac of the Nalzen Basin comprises : - Desmophyllites diphylloides (FORBES, 1846) (rare) -Eupachydiscus of. isculensis (REDTENBACHER, 1873) (rare) -
Hoplitoplacenticeras
(Hoplitoplacenticeras)
marroti
(COQUAND, 1859) (rare) - Hoplitoplacenticeras
(Hoplitoplacenticeras)
sp. juv.
(rare)
9 4J
ST.
BARTHELEMY
;l~ SAULT
- Glyptoxoceras aquisgranense (SCHLI2IT~, 1872a) (com-
mon) Baculites sp. (rare) - Scaphites (Scaphites) haugi DE GROSSOUVRE, 1894 -
F i g u r e 1 - G e o g r a p h i c a n d geological s i t u a t i o n of the F o r m a t i o n de St. Cirae in the N a l z e n Basin. A : A n t i c l i n o r i u m of St.-Cirac. B : i n t e r b e d d e d m a r l s a n d b i o c o n s t r u e t i o n a l limestone. C : marls, t : N o r t h - P y r e n e a n frontal thrust. 2 : Pech de Foix fault. 3 : M a s s i f de Saint-Barth~16my frontal thrust. 4 : C r o q u i e r fault. 5 : subp y r e n e a n frontal thrust. Situation gaograpbtqua a gdologtqua de la Formation de Salnt-girac darts I* "b~fln" de Na/zen. A : anttcltnortum de St-Ctrac. B : altamancas marnes.calcatres bloconatrutts. 1 : chevauchemant frontal nord-pyrdmdan. 2 : acctdunt radridlonal du Peals de Fotx. 3 : d a ~ a ~ a n t fiontal du Massif de S a i n t - B a b y . 4 :#ill, de Croquter. 5d~vaudaeraent frontal sous.pyrdnd,n.
(rare). The classic division of the Campanian in the type region of Aquitaine (de Grossouvre 1894, 1901 ; Kennedy 1986) is : Upper Campanian : Bostrychoceras potyplocurn zone ; Hoplitoplacenticeras marroti zone.
265
FUTURE
N
NORTH PYRENEAN FAULT
I I LABARRE
SANDSTONE
ST. CIRAC
buitd up
fmt marls
~PkArNE DEui/OtlX
PLATFORM
~
~NALZEN
"basin"
" block)
SUB- PYRENEAN TROUGH VRudistids
Figure 2 - Cross section of the sub-pyrenean basin and its margins during the Campanian at the meridian of the Nalzen basin. Coupe scbdmatklue du bassin sous-pyrdn#en el de ses marges, au Campanien, au rndridten du "basstn" de Nalzen.
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FOIX
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Figure 3 - The fossil locality. A : after the "Carte Geographique de la France" 1:50.000 Foix, sheet (XXI-47). B : General view of the outcrop during road works in the summer of 1989. F : marks the fossiliferous horizon. Situation du gtsementfossilifere. A : d'apr#s la Carte G#ograpbique de la France d 1/50000, feuille de Foix (XXI-47). B : vue d'ensemble des affleurements pendant les travaux d'dlargissement fourier de I'dM 1989. F : po~'tion de l'horizon
fossa~ere. Lower Campanian : Menabites (Delawarella) delawarensis zone ; Placenticeras bidorsatum zone. H. (H.) marroti and S. (S.) haugi are only known from the marroti zone, and the age of the Nalzen assemblage is thus lower Upper Campanian. The nannoflora recovered from the matrix of the ammonites included more than 90 species, and as discussed further below, indicates the Campanian subzone CC18B of Sissingh (1977, 1978) et Perch-Nielsen (1985).
This provides the first direct dating of the lower Upper Campanian marrot/ ammonite zone against the Bannofossil standard.
THE NANNOFLORA (J.B.) The nannoflora recovered in association with the ammonites described above is as follows (for full details of authors of species and combinations see Perch-Nielsen 1985) :
266 Ahmuellerella octoradiata (GORKA, 1957) REINHARDT, 1966 A. regulatis (GORKA, 1957) VERBEEK,1977 Amphizygus brooksii BUKRY,1969 A. papillatus BUKRY,1969 Arkhangelskiella specillata VEKSHINA,1959 Biscutum ellipticum (G6RKA, 1957) GRI3N in GRCrN & ALLEMANN,1975 Braarudosphaera bigelowi (GRAN & BRAARUD, 1935) DEFLANDRE,1947 Broinsonia enormis (SHUMENKO,1%8) MANIVrr, 1971 B. parca (STRADNER,1%3) BUKRu 1969 B. signata (NO~L, 1969) NOL, 1970 Bukryaster hayi (BUKRY, 1969) PRINS & SISSINGH in SISSINGH,1977 Calculites additus (WIND & WISE in WISE & WIND, 1977) PERCH-NIELSEN,1984 C. obscurus (DEH_ANDRE, 1959) PRINS & SISSINGHin 81SSINGH, 1977 C. ovalis (STRADNER, 1963) PRINS & SISSINGH in NISSINGH,1977 Chiastozygus amphipons (BRAMLETYE & MARTINI, 1964) GARTNER, 1968 C. bifadus BUKRY,1969 C. h'tteradus (GORKA, 1957) MANWIT, 1971 Corollithon exiguum STRADNER,1961 C. Kennedyi CRUX, 1981 C. signum STRADNER,1963 Cretarhabdus conicus BRAMLLrlq~ & MARTINI, 1964 Cribrocorona gallica (STRADNER, 1963) PERCHNIELSEN, 1973 Cdbrosphaerella ehrenbergii (ARKHANGELSKY, 1912) DEFLANDREin PIVEI~AU, 1952 Cyclagelosphaera rotaclypeata BUKRY,1969 Cylindralithus biarcus BUKRY, 1969 C. nudus BUKRY, 1969 Discorhabdus ignotus (GORKA, 1957) PERCH-NIELSEN, 1968 Eiffellithus eximius (STOVER, 1966) PERCH-NIELSEN, 1968 E. gorkae REINHARDT,1965 E. parallelus PERCH-NIELSEN,1973 E. turriseiffelii ( D E - - R E , 1954) REINHARDT,1965, BUKRY, 1969 Eprolithusfloralis (STRADNER,1962) STOVER, 1966
E. moratus (STOVER, 1966) Gartnerago obliquum (STRADNER, 1963) REINHARDT, 1970 Gephyrorhabdus coronadventis (REINHARDT, 1966) HILL, 1976 Haqius circumradiatus (STOVER, 1966) ROTH, 1968 Helicolithus anceps (G6RKA, 1957) NO~L, 1970 H. trabeculatus (G6RKA, 1957) VERBEEK,1977 Lithastdnus gdllii STRADNER, 1962 Lithraphidites camiolensis DEFI.ANDRE,1963 Lucianorhabdus arcuatus FORCHHEIMER,1972 L. cayeuxii DEFLANDRE,1959 L. quadrifidus FORCHHEIMER,1972 L. maleformis REINHARDT,1966 Manivitella pemmatoidea (DEFLANDRE in MANIVrr, 1%5) THIERSrEIN,1971 Marthasterites furcatus (DEFLANDRE, 1954) DEFLANDRE, 1959 ; BUKRY,1969 Microrhabdulus decoratus DEFLANDRE,1959 M. undosus PERCH-NIEI~EN,1973 Micula concava (STRADNER in MARTINI & STRADNER, 1960) BUKRY,1969, VERBEEK,1976 M. cubiformis FORCHHEIMER,1972 M. quadrata ? (STRADNER,1961) PERCH-NIELSEN,1984 M. staurophora (GARDET, 1955) STRADNER, 1963 (= M. decussata VEKSmNA,1959) Nannoconus elongatus BRONNIMANN,1955 N. multicadus D ~ R E & D~RE, 1959 Octolithus multiplus (PERCH-NmLSEN, 1973) ROMEIN, 1979 Placozygus fibuliformis (REINHARDT,1964) HOFFMANN, 1970 Prediscosphaem arkhangelskyi (REINHARDT, 1965) PERCH-NmLSEN,1984 P. cretacea (ARKHANGELSKY,1912) GARTNER,1%8 P. grandis PERCH-NIELSEN,1979 P. cf. 1". grandis PEP.Ca-I-NIELSEn,1979 P. microrhabdulina PERCH-NIELSEN,1973 P. ponticula (BUKRY,1959), PERCH-NIELSEN,1984 P. spinosa (BRAMtEI~ & MARTINI, 1964) GARTNER, 1%8 P. stoveri (PERCH-NIELSEn, 1968) SHAr~K & SaXAONER, 1971 Prolatipatella multicatinata GARTNER, 1968 Quadrum ? sp.
PLATE 1 Figs. 1, 18, 13 - Scaphites (Scaphites) haugi DE GROSSOUVRE, 1894. 1, UPS/MH/47 ; 8, UPS/MH/18 ; 13, UPS/MH/19. Figs. 2, 5- Hoplitoplacenticeras (Hoplitoplacenticeras) marroti (COQUAND, 1859). 2, UPS/MH/06 ; 5, UPS/MH/07. Fig. 3 - Desmophyllites diphylloides (FORBES, 1846). UPS/MH/01. Fig. 4 - Hoplitoplacenticeras (Hoplitoplacenticeras) sp. juv. UPS/MH/11. Figs. 6, 7, 11, 12, 14-19 - Glyptoxoceras aquisgranensis (Schliiter, 1872a). 6, UPS/MH/22 ; 7, UPSfMH/20 ; 11, UPS/MH/23 ; 12, UPS/MH/21 ; 14, UPS/MH/24 ; 15, UPS/MH/25 ; 16, UPS/MFU26 ; 17, UPS/MFU27 ; 18, UPS/MH/28 ; 19, UPS/MH/29. All specimens are from the Formation de Saint-Cirac between Catufet and Manau, Ari6ge. Figs. 1, 3, 6, 8, 9, 10, 11, 13-19 are x I ; Figs. 2, 4, 7, 12, 19 are x 4 ; Fig. 5 is x 3.5.
Geobios n ~ 25, fasc. 2
PI. 1 W.J. Kennedy, M. Hansotte, M. Bilotte & +1. Burnett
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268 Reinhardtites anthophorus (DEFLANDRE, 1959) PF_~CHNIELSEN, 1968 R. levis PRINS & SISSINGHin SISSINGH, 1977 Repaguhtm parvidentatum (DEbX~CDRE & FERT in DEFI~NDRE, 1954) FORCHHo.IMER,1972 Retecapsa angustiforata BLACK, 1971 tL crenulata (BRAMLEYrE & MAR'nNL 1964) GRON in GRON & ALLEMANN, 1975 Rhagodiscus acMyostaurion (HILL, 1976) R. angus~s (STRADNFA~1963) REINHARDT, 1971 R. plebeius PERCH-NIELSEN, 1968 R. reniformis PERCH-NIELSI~, 1973 R. splendens (DEFI_ANDRE, 1953) VERBEEK, 1977 Rotelapillus crenulatus (STOVER, 1966) PF~CH-NIELSEN, 1984 Scapholithus fossilis DEFI~\'DRE, 1954 Sollasites horticus (STRADNER et aL in STRADNER & ADAMIKER, 1966) CEPEK & HAY, 1969 Staurolithites mielnicensis (G6RKER & HAY, 1969 Staurolithites rnielnicensis (GORKA, 1957) PERCHNIELSnN,1968 Sensu CRUX in LORD, 1982 Thoracosphaera saxea STRADNER, 1961 Tranolithus gabalus STOVER, 1966 T. minimus (BUKRY, 1969) PERCH-NIElSEN, 1984 T. orionatus (REINHARDT, 1966) RFANHARDT, 1966 (= T. phacelosus STOVER, 1966) T. salillum (NO~L, 1965) CRUX, 1981 Vekshbzella angusta (STOVER, 1966) VF.RBEEK, 1977 Watznaueda barnesae (BLACK in BLACK & BARNES, 1959) PERCH-NIELSEN, 1968 W. biporta BUKRY, 1969 14. rnanivitae BUKRY, 1973 W. ovata BUKRY, 1969 W. quadriradiata BUKRY, 1969 Zeugrhabdotus acanthus R E n ~ , D T , 1965 Z. bipeqbratus (GAgTNE~ 1968) Z. embevgi (NO~L, 1958) PERCH-NIELSEN, 1984 Z. compactus (BUKRY, 1969) (= Rectapontis cornpactus (BUKRY, 1969) VAROL & JAKUBOWSKI,1989 Z. erectus (DEFLANDRE, 1954) REINHARDT, 1965
The nannoflora is relatively well-preserved, and shows Tethyan affinities, the species Bulayaster hayi and Marthastedtes furcatus being present. The cooccurrence of Broinsonia parca, B. hayi and M. furcatus (with Staurolithites rnielnicensis indicates the cosmo-
politan natmofossil zone CC18B (Sissingh 1977, 1978 ; Perch-Nielsen 1985). Some reworked (?) Cenomanian species are present : Corollithon kennedyi and Microstaurus chiastius. In addition, some younger species are present, e.g.P, stoveri and Prolatipatella multica~inata, but index species of Ceratolithoides and Quadrum are absent and Lithastrinus grillii occurs in moderate numbers, all pointing to the present zonal assignation. CONVENTIONS The following abbreviations are used to indicate the repositories of specimens mentioned in the text : BMNH : Natural HistoryMuseum, London. UPS : Universit6 Paul Sabatier, Toulouse. MDA : Mus6e D6partemental de l'Ari6ge, Foix SYSTEMATIC PAl AEONTOLOGY (W.J.K.) Order AMMONOIDEA Zittel, 1884 Suborder AMMONITINA Hyatt, 1889 Superfamily DESMOCERATACEAE Zittel, 1895 Family DESMOCERATIDAE Zittel, 1895 Subfnmily DESMOCERATINAE Zittel, 1895 Genus Desmophyllites SPATH, 1929 [= Schlateda Do. GROSSOUVRO.,1894, p. 126 (non FR_rrsen /n FP,rrSCH & KAFKA, 1887, p. 33) ; Schli~tericeras COLLIGNON, 1938, p. 92 (non HYA'IT, 1903, p. 110)]. Type species - Desmoceras larteti SEUNES, 1891, p. 19, pl. 12(3), fig. 2 ; pl. 13(4), figs. 2, 3, by subsequent designation by Spath, 1921, p. 46, as type species of Schtiiteria, of which Desmophyllites is replacement name. DESMOPHYLLITES 1846) : pl. 1, fig. 3.
DIPHYLLOIDEA
(FORBES,
1846 -Ammonites diphylloides FORBES, p. 105, pl. 8, fig. 8. 1894 Desmoceras pyrenaicum DE GROSSOUVRE, (pars), p. 168, pl. 37, fig. 9 ; non pl. 25, fig. 2.
PLATE 2 Figs. 1-5, 9-15 - Glyptoxoceras aquisgranensis (SCHLOTER, 1872a). 1, UPS/MH/30 ; 2, UPS/MH/31 ; 3, UPS/MH/32 ; 4, UPS/MH/33 ; 5, UPS/MH/34 ; 9, UPS/MH/35 ; 10, UPS/MH/36 ; 11, 14, UPS/MH/37 ; 12, UPS/MH/38 ; 13, UPS/MH/39 ; 15, UPS/Mtt/41. Figs. 6, 7 - Hoplitoplacenticeras (Hoplitoplacenticeras) marroti (COQUAND, 1859). 6, UPS/MH/08 ; 7, UPS/MH/09. Fig. 8 - Baculites sp. UPS/MH/14. All specimens are from the Formation de Saint-Cirac between Catufet and Manau, Ari~ge. Figs 1-3, 5, 11-15 are x 4 ; Fig. 4 is x 2 ; Figs. 6, 7 and 8 are x 1.
Geobios n ~ 25, fasc. 2
PI. 2
WJ. Kennedy, M. Hansotte, M. Bilotte & J. Burner
270 1985 - Desmophyllites ch'phylloides (FORBES, 1846) ; Henderson & McNamara, p. 54, pl. 4, figs. 1-4 (with synonymy). 1989 - Desmophyllites ; Kennedy, text-fig. 17d. 1989 - Desmophyllites diphylloides (FORBES) ; Haggart, p. 193, pl. 8.6, fig. 10 with additional synonymy). Types - Lectotype, by subsequent designation by Matsumoto & Obata 1955 (p. 122) is BMNH C'22682, the original of Forbes 1846, pl. 8, fig. 8 ; paralectotypes are BMNH C22683-5, all from the Upper Maastrichtian Valudavur Formation of Pondlcherry, south India. The lectotype was refigured by Howarth, 1965, pl. 11, fig. 3. Material - Three specimens, UPS/MH/01 ; MDA/MH/ 02-03. a n d d i s c u s s i o n - These specimens are partpyritised, and crushed. The best-preserved specimen is 20 mm in diameter, with a minute, deep umbilicus. The umbilical shoulder is narrowly rounded, the flanks flattened and subparallel, the ventrolateral shoulders broadly rounded, and the venter somewhat flattened. The shell surface is smooth, but for traces of straight, prorsiradiate growth lines. Little of the suture is exposed, but UPS/MH/01 shows a narrow, moderately incised E/L. Description
Discussion - These specimens differ in no significant respects from well-preserved juveniles we have studied from the Upper Santonian of the Corbitres. Henderson & McNamara (1985, p. 54, pl. 4, figs. 1-4) provide a full revision of this species and discuss differences from others referred to the genus.
EUPACHYDISCUS ef. ISCULENSIS (REDTENBACHER, 1873) : pl. 1, figs. 9, 10. compare : 1873 -Ammonites isculensis REDTENBACHER, p. 122, pl. 29, fig. 1. 1894 - Pachydiscus isculensis REDTENI3ACHER ; de Grossouvre, p. 185, pl. 22, fig. 2 ; pl. 26, fig. 1 ; pl. 37, fig. 1. 1986 - Eupachydiscus isculensis REDTENBACHER, 1873 ; Kennedy, p. 52, pk 2, figs. 5, 6 ; pl. 10, figs. 13, 14 ; text-fig. 19 (with synonymy). Holotype - By monotypy, an unregistered specimen in the Obert~sterreichs Landesmuseum, Austria, from the Santonian Gosau Beds between Ischl and Kohlbuch, Austria, the original of Redtenbacher 1873, pl. 29, fig. 1, reiUustrated by Kennedy 1986, text-fig. 19. Material
-
Two specimens, UPS/MH/04 ; MDA/MH/
05. Description - The best-preserved of the specimens is UPS/MH/04 (pl. 1, figs. 9, 10), a wholly septate juvenile 33 mm in diameter. Coiling is very involute, with a depressed, reniform whorl section and an estimated whorl breadth to height ratio of 1.25. Ribs arise either singly or in pairs from umbilical buUae, and are strong, narrow, straight and prorsiradiate on the flanks, strengthening markedly on the ventrolateral shoulders, and crossing the venter in a broad convexity. The suture is deeply and intricately incised, with narrow-stemmed E/L and L/U2. - Although small, these specimens differ in no significant respects from juveniles figured by de Grossouvre (1894, pl. 22, fig. 2 ; pl. 26, fig. 1 ; pl. 37, fig. 1), Summesberger (1979, p. 136, pl. 7, figs. 30, 31 ; pl. 8, fig. 32 ; text-figs. 20-22) and others. Discussion
- Desmophyllites diphylloides has a recorded range of Lower Santouian to Upper Maastrichtian. It is known from southwestern France (Corbitres, Aude, as well as the present records), Tuni.~ia, Angola, Zululand and Pondoland (South Africa), Madagascar, Argentina, California, Arkansas, British Columbia, Alaska, Japan, Australia and south India. Occurrence
Family PACHYDISCIDAE Spath, 1922 Genus Eupachydiscus SPA'If-I, 1922 [= Mesopachydiscus YABE & SHIMIZU, 1926] Type species - Ammonites isculensis RED'I'E.Nq3ACHER, 1873, p. 122, pl. 29, fig. 1, by original'designation.
Occurrence - Eupachydiscus isculensis ranges from the middle of the Santonian to mid-Campanian. It is known from Aquitaine and the Corbitres in France, northern Spain, Austria, Roumania, Zululand (South Africa), and Madagascar. Superfamily HOPLITACEAEA H. DouviUt, 1890 Family PLACENTICERATIDAE Hyatt, 1900 Genus and Subgenus Hoplitoplacenticeras PAULCKE, 1907
PLATE 3 Figs. 1-9 - Glyptoxoceras aquisgranensis (SCHLf..)TER, 1872a). 1, UPS/MH/42 ; 2, UPS/MH/43 ; 3, UPS/MH/44 ; 4, UPS/MH/45 ; 5, UPS,qVIH/46 ; 6, UPS/MH/47 ; 7, UPS/MH/48 ; 8, UPS/MH/49 ; 9, UPS/MH/50. All specimens are from the Formation de Saint-Cirac between Catufet and Manau, Aritge. All figures are x 1.
Geobios n ~ 25, fasc. 2
PI. 3
WJ. Kennedy, M. Hansotte, M. Bilotte & J. Bttrnett
272
[ = Dechenoceras KAYSER, 1924, p. 175]
Material - Five specimens, UPS/MH/06 to 09 ; MDA/ MH/10.
the venter. UPS/MH/07 (pl. 1, fig. 5) is 18 mm in diameter, and has acquired a small inner ventrolateral clavus. Adult ornament is shown by UPS/MH/09 (pl. 2, fig. 7), 57_5 mm in diameter, and UPS/MH/08 (pl. 2, fig. 6), a complete adult with an estimated diameter of 70 mm. Coiling is moderately involute, with a small, shallow umbilicus that comprises 22% of the diameter, and has a low, flattened wall and narrowly rounded umbilical shoulder. There are sixteen crowded umbilical bullae per whorl, perched on the umbilical shoulder, and connected to the umbilical seam by a strong, concave n'b. The bullae give rise to one, sometimes two ribs, while these long ribs may bifurcate around mid flank, and intercalated ribs arise low on the flank to give a total of 40 ribs per whorl. The ribs are strong, narrower than the interspaces and prorsiradiate. They are straight on the inner flank, but flex forwards and are convex at mid-flank, thence flexing back and concave on the outer flank before sweeping forwards to weak, bullate inner ventrolateral tubercles that disappear at the be~nning of the outer whorl, and outer ventrolateral clavi that weaken and disappear on the final sector of body chamber prior to the adult aperture. This is also marked by a weakening of umbilical buUae and ribs, and a proliferation of growth lines and striae. The sutures were not seen.
Description - The earliest growth stages seen are a specimen 4 mm in diameter, MDA/MH/10. This is globose and smooth for the first half whorl, after which blunt umbilical bullae, five per half whorl, appear. These give rise to low, broad, straight prorsiradiate ribs, either singly or in pairs, which terminate in welldeveloped outer ventrolateral clavi. UPS/MH/06 (pl. 1, fig, 2) is 7 mm in diameter, and has similar ornament, showing the clavi alternating in position on either side of
Discussion - The simple ribbing and tuberculation immediately distinguish these specimens from species such as H. (H.) coesfeliense (SCHLt2rER, 1867) (p. 15, pl. 1, figs. 1-5 ; see Kennedy, 1986, p. 73, pl.9, figs. 9, 10 ; text-fig. 27b, ~ f), H. (H.) costulosum (SCHLt:rrER, 1876) (p. 159, pl. 44, figs. 1-4) and H. (hemfoerdiceras) lemfoerdense (SCHLOTER, 1872b) ( = as nomen novum for Ammonites scaphitoides SCHL(3"rER, 1872a, p.63, pl. 19, figs. 1, 2, non COQUAND, 1853, p. 442, pl. 14, figs. 9,
Type species - Hoplites - Placenticeras plasticus PAULCKE, 1907, p. 186 : ICZN Opinion 555, 1959 ; name n ~ 1629. HOPLITOPLACENTICERAS (HOPLITOPLACENT1CERAS) MARROT1 (CoOUAND, 1859) : pl. 1, figs. 2, 5 ; pl. 2, figs. 6, 7. 1859 - Ammonites marroti COQUAND, p:.70. 1984 - Hoplitoplacenticeras vari (SCHLUTER) ; Bilotte, pl. 37, figs. 2, 3. 1986- Hoplitoplacenticeras (Hoplitoplacenticeras) marroti (COOUAND, 1859) ; Kennedy, p. 70, pl. 2, figs. 3, 4; pl. 9, figs. 1-8, 11-12 ; pl. 10, figs. 1-12 ; pl. 12, figs. 1, 2. Type - Holotype, by monotypy, is the specimen from R~erac, Dordogne, mentioned by Coquand (1859, p. 995), refigured by Kennedy 1986 (pl. 9, figs. 5, 6), in the Sorbonne Collection, now in the Universit6 Pierre et Marie Curie, Paris.
PLATE 4 Fig. 1 - Marthasterites furcatus DEFLANDRE. XP, x1250 ; Neg. n ~ UCL-3467-13. Fig. 2 - Lucianorhabdus maleformis REINHARDT. XP, x 1250 ; Neg. n ~ UCL-3467-4. Fig. 3 - Lucianorhabdus quadrifidus FORCHHEIMER. XP, x 460 ; Neg. n ~ UCL-3467-3. Fig. 4 - Calculites ovalis (STRADNER) PRINS & SISSINGH Xp, x 1500 ; Neg. 11" UCL-3467-7. Fig. 5 - Quadrum sp. TL, x 1670 ; Neg. n ~ UCL-3467-23. Figs 6, 7 - Lithastn'nus grillii STRADNER. 6. XP, x 1250 ; Neg. n ~ UCL-3"467-28.7. TL, x 1750 ; Neg. n ~ UCL-3467-31. Fig. 8 - Bukryaster hayi (BUKRY) PRINS & SISSINGH. XP, x 1400 ; Neg. n* UCL-3467-32. Fig. 9 - Broinsonia parca (STRADNER) BUKRY. XP, x 1500 ; Neg. n ~ UCL-3467-16. Fig. 10 - Eiffellithus eximius (STOVER) PERCH-NIELSEN. XP, x 1750 ; Neg. n ~ UCL-3467-11. Fig. 11 - Reinhardtites anthophorus (DEPT_.ANDRE) PERCH-NIELSEN. XP, x 1875 ; Neg. n ~ UCL-3467-15. Fig. 12 - Tranolithus orionatus (REINHARDT) REINHARDT. XP, x 1250 ; Neg. n ~ UCL-3467-5, Fig. 13 - Zeugrahabdotus biperforatus (GARTNER). XP, x 1875 ; Neg. n ~ UCL-3467-14. Fig. 14 - Prediscosphaera grandis PERCIt-NIELSEN. XP, x 1410 ; Neg. n ~ UCL-3467-29. Figs 15, 16 - Watznaueria quadriperforata BUKRu 15. XP, x 1.830 ; Neg. n ~ UCL-3467-18. 16. TL, x 1830 ; Neg. n ~ UCL-3467-19. All negatives cited are preserved in the collections of the Micropalaeontology Unit, Department of Geological Sciences, University College of London. XP = crossed polars ; T : = transmitted light.
Geobios n ~ 25, fast. 2
PI. 4 W.J, Kennedy, M. Hansotte, M. Bilotte & J. Burnett
274 10). H. (H.) vari ( S C H L ~ 1872b) (p. 92 ; nomen novum for Ammonites sttiatocostatus SCHLOTER~1872a~ p. 65, pl. 20, figs. 1-4, non MF.NF_.GHINI,1854, pl. 28) has a rather .~imilar rib style (see Kennedy 1986, text-fig. 25), but here the ribs are less flexuous, and the whole of the shell surface covered in prominent lirae, not seen in the present specimens, which have shell preserved. H. (H.) marrofi (COQUAND, 1859) (see revision in Kennedy 1986, p. 70, pl. 2, figs. 3, 4 ; pl. 9, figs. 1-8, 11-12 ; pl. 10, figs. 1-2 ; pl. 10, figs. 1-i2 ; pl. 12, fags. 1, 2) have 12-14 umbilical bullae per whorl and up to 20 ribs per half whorl in specimens from Aquitaine, with rather blunt ribs in some variants (Kennedy 1986, pl. 9, figs. 7, 8), whereas others (Kennedy 1986, pL 9, figs. 5, 6) are closely comparable to the present specimens, apart from preservation. Occurrence - In Aq#taine, specimens are labelled as being from Assizes pZand p3 of A m a u d (J878), but de Grossouvre (1894, 1901) noted it from p only, which has been our experience. The species occurs at a comparable level at Ressec (Dreuilhe anticline) as well as the present occurrences in Nalzen. It may occur at Loesfeld, Westphalia, Germany, Vijlen near Cotessen on the Belgium-Netherlands border, in Portugal, Israel, Soviet Central Asia, Madagascar and Texas, USA. HOPLITOPLACENTICERAS (HOPLITOPLACENTICERAS) SP. JUV. : pl. 1, fig. 4.
Material - 3 specimens UPS/MH/11 and MDA/MH/ 12-13. - The largest of these specimens is 10 mm in diameter. They differ from H. (H.) marroti, described above, in their much finer, more numerous ribs, with many more ventrolateral elavi per whorl, suggesting the presence of a second Hoplitoplacenticeras spedes. Discussion
Suborder ANCYLOCERATINA Wiedman% 1966 Superfamily T U R R I L I T A C E A E Gill, 1871 Family DIPLOMOCERATIDAE Spath, 1926 Subfamily DIPLOMOCERATINAE Spath, 1926 Genus Gtyptoxoceras SPATH, 1925 [= Neohamites BRUNNSCHWEILER, 1966] Type species - Hamites rugatus FORBES, 1846, p. 116, pl. 11, fig. 6, by orional designation (Spath 1925, p. 30, as Hamites (Anisoceras) rugatus (FORBES) KOSSMAT). GLYPTOXOCERAS AQUISGRANENSE ( S c I - I L ~ 1872a) : pl. 1, figs. 6, 7, 11, 12, 14-19 ; pl. 2, figs. 1-5, 9-15; pl. 3, figs. 1-9 1872a - Toxoceras (?)Aquisgranense SCI-ILOI'ER,p. 102, pl. 31, figs. 6-9.
1887 - Hamites aquisgranensis SCHL~'ER ; Holzapfel, p. 66, pl. 5, rag. 8. 1925 - Hamites aquisgranensis SCI-ILOrER ; Diener, p. 66. Type - Lectotype, here designated, is the original of Schlfiter 1872a, p. 102, pl. 31, figs. 6-9, an unregistered specimen in the Collections of the Geologisches und Paliiontologisches Institut of Bonn University, and from the lower sandy layer with Belemnitella quadrata at the foot of Luseberges, near Aachen, Germany. The spedmen is probably of upper Lower Campanian age, for Schlfiter (1872, pl. 24) illustrated a late form of Scaphites (Scaphites) hippocrepis (DEKAY, 1828) from the same locality. - Almost 100 specimens, UPS/MH/20 to 50 ; MDA/MH/51 to 113. Material
Description - Complete shells are up to 80 mm in diameter, and consist of three distinct ontogenetic stages: an initial straight shaft, followed by a brief helical stage, and then an open plani~piral stage. The protoconch is preserved in UPS/MH/37 (pl. 2, fig. 11 ; counterpart shown in pl. 2, fig. 14). It is overpyritised, with an apparent diameter of 0.4 ram. The succeeding straight growth stage is in the form of a slowly expanding smooth shah up to 20 mm long (191. 1, fig. 12 ; pl. 2, figs. 5, 9, 11-14). Immediately prior to the second growth stage the expansion rate of the shaft increases markedly (pl. 1, fig. 12 ; pl. 2, figs. 5, 12) and be~n.~ to coil, for half a whorl with a very high translation rate (191.2, fig. 12), but thereafter with a low translation rate, to form a helix of at least 5 whorls, the whorls barely in contact, with a circular whorl section. Internal moulds are smooth (pl. 2, fig. 4), but where the shell is preserved, the last three whorls of the spire are ornamented by delicate equal prorsiradiate and feebly flexuous ribs (pl. 2, figs. 2, 10, 13, 15). The spire is succeeded by a brief section of shell on which the translation rate increases markedly to give at most half a whorl of helical coil (pl. 2, figs. 10, 13, 15), succeeded by the third and final growth stage, a planispire of at least two whorls ; the complete ontogeny but for the final section of body chamber is shown in pl. 3, fig. 9. All specimens are crushed. Ornament consists Of sharp, narrow rursiradiate ribs ; the density is initially high, with a rib index of up to 9 (pL 1, fig. 14 ; pl. 3, figs. 1, 5, 9), decreasing to 5 or 6 in some body chamber fragments (pl. 3, fig. 6), although rib density may increase again towards the adult aperture, as shown in the largest, poss~ly adult specimen in the collection (pl. 3, fig. 7). The ribs are generally narrower than the interspaces, weak on the dorsum, but strengthening across the flank, straight on the dorsal flank but flexed forwards in many specimens and feebly concave on the ventral part of the flank. The ribs appear to be at their maximum development over the venter. The suture is typical for the genus, with rectangular little-incised bifid lobes and saddles.
275 This remarkable suite of specimens show for the f'trst time the complete ontogeny of a Glyptoxoceras species. The type material of the type and other species from the Upper Maastrichtian of south India, preserved in the collections of the Natural History Museum in London (Kennedy and Henderson, in press) includes only fragments of loose open helices, more complex examples of which were illustrated, in the case of Glyptoxoceras indicum (FORBES, 1846) (p. 116 (pars), non pl. 11, fig. 4) by Matsumoto (1959, p. 167, pl. 41, figs. 2-6 ; text-fig. 80). These show G. indicum to have a much larger, lower helix than the present species, and an adult shell in the form of an ellipse. The Upper Maastrichtian Glyptoxoceras rugatum (FORBES, 1846) (p. 116, pl. 11, fig. 6) has a compressed whorl section with a rib index of 6-7 in medium-sized specimens from south India. Revision of material from the Miria Formation of western Australia (Henderson, Kennedy & McNamara, in press) show adults of the species to have been up to a metre across, with irregular coiling, long straight or slightly curved sections being linked by short sectors defining an obtuse angle. Discussion
-
Of other European species, Glyptoxoceras vaalsiensis HOLZAPFEL, 1887 (p. 66, pl. 5(2), figs. 6a-c, 7) has very coarse, distant annular ribs that do not weaken on the dorsum according both to the original figures and specimens in the Rheinisch-Westf/ilischen Technischen Hochschule, Aachen, and has highly simplified sutures. The Hamites roemeri GEINrI'z of Holzapfel (1887, pl. 5(2), figs. 4, 5) has much more distant and coarse ribs than the present species.
Ancyloceras retrorsum SCHLOTER~ 1872 (p. 97, pl. 30, figs. 5-8) was revised by Kennedy (1986, p. 106, pl. 16, figs. 1-4, 6, 7 ; pl. 17, figs. 1, 2 ; text-fig. 38), who referred it to Neoglyptoxoceras COLLIGNON, 1969, and reillustrated the type material. This is a much larger species than G. aquisgranense, and appears to have an elliptical rather than circular coil, with juvenile fragments indistinguishable from the present species.
BACULITES SP. : pl. 2, fig. 8. Material - Three specimens, UPS/MH/14 and MDA/ MH 15-16. Discussion - The genus Baculites is represented by fragments only in the present collections. The largest is 31 mm long, with a slowly expanding shell and very distant dorsal nodes on the flank, suggesting it belongs to the widely occurring Baculites capensis WOODS, 1906, group (see revision in Klinger & Kennedy 1977, p.71, text-figs. 2a-f, 3g).
Superfamily SCAPHITACEAE Gill, 1871 Family SCAPHITIDAE Gill, 1871 Subfamily SCAPHITINAE Gill, 1871 Genus and subgenus Scaphites PARK1NSON, 1811 [= Anascaphites HYATE, 1900, p. 572 ; Jahnnites HYATI', 1900, p. 572 ; Holcoscaphites NOWAK, 1911, p.
564] Type species - Scaphites equalis J. SOWERBY, 1813, p. 53, pl. 18, figs. 1-3.
SCAPHITES (SCAPHITES) HAUGI DE GROSSOUVRE, 1894 : pl. 1, f~s. 1, 8, 13 ; text-fig. 4 1894 - Scaphites haugi A. DE GROSSOUVRE sp., p. 244, pl. 31, fig. 5. 1925 - Scaphites haugi DE GROSSOUVRE ; Diener, p. 199. 1928 - Discoscaphites? hau# DE GROSSOUVRE ; Reeside, p. 30. 1970 - Scaphites cf. haugi DE GROSSOLWRE ; Atabekian & Akopyan, p. 39, pl. 2, figs. 3, 4. 1986 - Scaphites (Scaphites) haugi DE GROSSOUWRE ; Kennedy, p. 116, pl. 21, figs. 12-16. Holotype - By monotypy, is the original of de Grossouvre 1894, pl. 31, fig. 5, an unregistered specimen in the Sorbonne Collections, now housed in the Universit6 Pierre et Marie Curie, Paris, ex Arnaud Collection, from
Occurrence - The type material is from the Vaals Formation, and probably of upper Lower Campanian age, as is material described by Holzapfel (1887) ; the present material is thus somewhat younger.
ii / / I
Family BACULITIDAE Gill, 1871 Genus Baculites LAMAP,CK, 1799
[ = Homaloceratites HUPSCH, 1768, p. 110 (non binomen) ; Euhomaloceras SPATII, 1926, p. 80] Type species - Baculites vertebralis LAMARCK, 1801, p. 103, by subsequent designation by Meek (1876, p. 391).
I
Figure 4 - Suture line of Scapbaes ($capbttes) baugi (DE GROSSOOVRE, 1894). UPS/MH/17 (see also pl. 1, fig. 1). Bar scale is 10 mm. Ztgne do s~x~e de Scapbttes (Scapbttes) baugt (D~ oRossovn~,
1894).
276 Assize p3 of Arnaud (1878) at TauiUard (Charente), ref'~ured by Kennedy 1986, pl. 21, figs. 12-14. Material - Three specimens, UPS/MH/17 to 19. Description - All the specimens are crushed. One is an incomplete phragmocone 19 mm in diameter, a second an incomplete microconch with a phragmocone 22 mm in diameter and part of the body chamber shaft preserved. Coiling of the phrao~mocone is very involute, with a tiny umbilicus comprising 12% of the diameter, deep, with a flattened wall and narrowly rounded shoulder. Twelve distant, coarse, straight, prorsiradiate primary n'bs arise at the umbilical seam~ and strengthen across the umbilical wall and shoulder, terminating in strong, conical ventrolateral tubercles. These give rise to pairs of weaker secondary ribs, with a .~ingle short intercalated r ~ between, all ribs crossing the venter without diminutionThe umbilical wall of the shaft of the body chamber is concave, and the wall follows a concave course in profile, such that all of the umbilicus of the phragmocone is exposed. There are four very coarse, distant ribs on the section preserved. These arise at a strong conical to feebly bullate umbilical tubercle, are straight and prorsiradiate on the flanks and terminate in strong ventrolateral clavi. The venter is poorly preserved, but groups of secondary and intercalated ribs are present. The third specimen is a macroconch with a phragmocone an estimated 36 mrn ill diameter, and a total length to the shell of 56 mm~ Phragmocone ornament is as in the microconch, with in addition, a few long secondary ribs. The body chamber is much higherwhorled than that of the microconch, with a convex umbilical wall that is straight when viewed in profile, partially occluding the umbilicus of the phragmocone. There are five conical to feebly bullate umbilical tubercles on the body chamber. The adapical one is weak, the second and third massive, and those on the final hook much weakened. They give rise to one or two low, broad primary ribs that terminate in coarse ventrolateral clavi that are restricted to the shaft and initial sector of the hook. The clavi give rise to groups of two or three coarse secondary ribs that loop to the clavi on the opposite flank, and there are. some intercalated fibs. Umbilical tubercles are very weak, and ventrolateral tubercles absent from the final sector of shell prior to the adult aperture, which is ornamented by narrow ribs on outermost flank, ventrolateral shoulders and venter. The suture is only partially exposed (text-fig. 4). - Kennedy (1986, p. 116) redescribed the type specimen of this species, and discussed differences from the most closely allied species. The older S. (S.) hippocrepis (DEKAY, 1828) is generally smaller, the spire has a rounded whorl section with weak or no tubercles, while the body chamber is weakly tuberculate, and may have a lateral tubercle in Discussion
some specimens (see Cobban, 1969, for a full review of the hippocrepis group). The younger S. (S.) binodosus (ROEMER, 1841) (p. 90, pl. 13, fig. 16 ; of which S. (S.) inflatus ROEMER, 1841 (p. 90, pl. 14, fig. 3) is the macroconch) is discussed by Kennedy (1986, p. 116, text-figs. 39a-e). This is a much larger species than S. (S.) haugi, with a quite different shell shape, the microconchs with numerous umbilical tubercles on the body chamber, with broad, flattened triangular ribs; macroconchs have only feeble umbilical bullae on the body chamber, with numerous weak flank ribs and small ventrolateral clavi. Occurrence - Apart from the type occurrence, there is a second, topotype specimen, and a third from division Campanian 3 of Plate 1 (1977) near Prrigueux (Dordogue). Atabekian & Akopian (1970) recorded Scaphites el. haug/from the Campanian of Soviet Armenia. Acknowledgements - Burnett acknowledges the financial support of the Natural Environment Research Council (UK) under the terms of Grant GR3/6767, and technical support of the staff of the Micropalaeontology Unit of the Department of Geological Sciences, University College, London. Kennedy acknowledges the financial support of the Natural Environment Research Council (UK) under the terms of Grant GR3/6501a, and the technical assistance of the staff of the Department of Earth Sciences, Oxford and Geological Collections, University Museum, Oxford.
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278 RAM~RE De FONTANIER E. 1933 - Les terrains cr6tac6s entre Foix et B61esta. t~tude stratigraphique et tectonique. Bull. Soc. Hist. nat. Toulouse, 64 : 369-468. REDTENBACHER A. 1873 - Die Cephalopodenfauna der Gosauschichten in den nordSstlichen Alpen. Abh. K.-K. geol. Reichsanst., 5 : 91-140. REESIDE J.B. Jr. 1928 - The scaphites, an Upper Cretaceous ammonite group. Prof. Pap. U..S. geol. Surv., I$0-B : 21-36. RICATF_AU R. & VILLEMIN J. 1973 - Evolution au Cr6tac6 sup6rieur de la pente s6parant le domaine de plate-forme du siUon sous-pyr6n6en en Aquitaine m6ridionale. Bull. Soc. gdol. Fr., (7), 15 : 30-39. ROEMER A. 1840-1841 - Die Versteinerungen des norddeutschen Kreidegebirges. (1-48, pls. 1-7, 1840 ; 49-145, pls. 8-16, 1841). Hzhn'schen Hofbuchhandlung, Hznnover : 145 p. SCHLU'TER C. 1867 - Beitrag zur Kenntniss der fiingsten Ammoneen Norddeutsehlands. A. Henry, Bonn :36 p. SCHLWIER C. 1871-1876 - Cephalopoden der oberen deutschen Kreide. Palaeontographica, 21 : 1-24, pls. 1-8 (1871) ; 21 : 25-120, pls. 9-35 (1872a) ; 24 : 1-144
(121-264) + x, pts. 36-55 (1876). SCHLI]TER C. 1827b Verb. naturh. Ver. preuss. Rheine, 29, 1 : 91-92. SEUNES J. 1891 - Contribution a l'6tude des C6phalopodes du Cr6tac6 Sup6rieur de France. I. Ammonites du Calcaire ~ Baculites du Cotentin (Suite). 1I. Ammonites du Campanien de la r6gion souspyr6n6enne. D6partment de Landes. Mdm. Soc. gdol. Fr., S~r. Pal~ont., 2, m6m. 2 : 8-22. SISSINGH W. 1977 Biostrafigraphy of Cretaceous calcareous nznnoplankton. Geol. Mijnbouw, 56 : 37-65. -
SISSINGH W. 1978 - Microfossil stratigraphy and stagestratotypes of the Cretaceous. GeoL Mi]nbouw, 57 : 433-440. SOWERBY J. 1813 - The Mineralogy Conchology of Great Britain. The Author, London, 1 : 10-44. SPATH L.F. 1921 - On U p p e r Cretaceous Ammonoidea from Pondoland.Ann. Durban Mus., 3 : 39-56. SPATH L.F. 1922- On the Senonian ammonite fauna of Pondoland. Trans. R. Soc. S. Aft'., 10 : 113-147. SPATH L.F. 1925 - On Senonian Ammonoidea from Jamaica. Geol. Mag., 62 : 28-32. SPATH L.F. 1926 - On new ammonites from the English Chalk. Geol. Mag., 63 : 77-83, table. SPATH L.F. 1929 - Corrections of cephalopod nomenclature. Naturalist, 871 : 269-271. SUMMESBERGER H . 1979 - Eine obersantone Ammonitenfauna aus dem Beeken von Gosau (OberSsterreich).Ann. Naturhistor. Mus. Wien, 82:109-176. TOUCAS A. 1903 - Etudes sur la classification et l'6volution des Hippurites. Mdm. Soc. gdol. Ft., ser. Paldont., 30:128 p. WmDMANN J. 1966 - Stammesgeschichte nnd system den posttriadischen ammonoideen ; ein iiberblick. N. .lb. geol. Palaont. Abh., 125 : 49-79 ; 127 : 13-81. WOODS H. 1906 - The Cretaceous fauna of Pondoland. Ann. S. Af. Mus., 4 : 275-350. YABE H . & SI-nMIZU S. 1926 - A study of the genus "Parapachydiscus" HYATI'. Proc. Imp. Acad., 2 : 171-173. ZrrrEL K.A. YON 1884 - Handbuch der Palaeontologie. 1, Abt. 2 ; Lief 3, Cephalopoda. R. Oldenbourg, Munich & Leipzig : 329-522. ZrrrEL K.A. YON 1895 - Gmndzage der Palaeontologie (Palaeozoologie). R. Oldenbourg, Munich & Leipzig: vii + 972 p.
MICHEL HANSOTrE Cos, 09000 Foix, France.
MICHEL BILOTrE Laboratoirada adologl~sddimentaira at Paldonto~gie, URA1405, UniversiMPaul Sabaliar,39 alldesJules Guesde,F-31062 ToulousaC$dex.
JACQUELINE B U R N E T r Department of GeologicalSciences, Untvars@CollageLondon, GowarStreet, London WCIE6BT, Grande-Bretagna.
RI~SUM~. La formation campanienne de Saint-Cirac, dans l'unit6 nord-pyr6n6ene du "Bassin" de Nalzen vient de livrer sa premi6re faune d'ammonites (plus de 100 individus), avec Desmophyllites diphylloides (FORBES, 1846), Eupachydiscus cf. isculensis (REDTENBACHER, 1873), Hoplitoplacenticeras (Hoplitoplacenticeras) marroti (CoQUAND, 1859), H. (H.) sp. juv., G~yptoxoceras aquisgranense (SEHL~2rER, 1872a), Baculites sp. et Scaphites (Scaphites) haugi DE GROSSOUVRE, 1894. C'est une laurie de la classique zone ~t marrot/, de la partie inf6rieure du Campanien sup6fieur dans la r6gion type en Aquitaine. Les ammonites sont associ6es /i une nannoflore fiche de plus de 90 esp6ces ; l'association de Broinsonia parca avec Bukryaster hayi et Marthastedtes furcatus indique la sous-zone CC18B du Campanien dans le sch6ma de Sissingh (1977, 1978) et Perch-Nielsen (1985). AMMONITESET NANNOFOSSILESDU CAMPANIENDE NALZEN(ARII~GE,FRANCE)
ABSTRACT The Marnes de Saint-Cirac (Campanian) in the north-pyrenean unit known as the Nalzen "Basin" has yielded an ammonite fauna of more than 100 specimens, with Desmophyllites diphylloides (FORBES, 1846), Eupachydiscus cf. isculensis (REDTENBACHER, 1873), Hoplitoplacenticeras (Hoplitoplacenticeras) marroti (COQUAND, 1859), H. (H.) sp. juv., Glyptoxoceras aquisgranense (SCHL~rER, 1872a), Baculites sp. and Scaphites (Scaphites) haugi DE GROSSOUVRE, 1894. This is a fauna of the classic marroti zone of the lower Upper Campanlnr~ in the type region in Aquitaine. The ammonites are associated with a rich nannoflora of more than 90 species ; the association of Broinsonia parca with Bukryaster hayi and Marthasterites furcatus indicates subzone CC18B of the Campanian in the scheme of Sissingh (1977, 1978), et Perch-Nielsen (1985).
INTRODUCTION
bulloides VOLGER, a. rosetta (CARSEY), G. ventricosa WnrrE, Rosita fomicata (PLUMMER) (Bilotte 1975,
In the north pyrenean unit known as the N',dzen "Basin", the Marnes of Saint-Cirac have been referred to the Campanian sensu lato on the basis of the rudistids (Toucas 1903 ; Rami~re de Fontanier 1933 ; Astre 1954; Bilotte 1985), benthic, and rare planktonic foraminifera present (Fallotia cf. colomi SILVF_SI~, Abrardia cf. mosae (HOFKER), Globotruncana arca (CuSIIMAN), G.
1985). The discovery by one of us (M.H.) of a horizon rich in ammonites, including the most remarkable heteromorphs yet described from the Upper Cretaceous of France, allows the dating of a part of the sequence as lower Upper Campanian. The ammonites are associated with a rich nannoflora, which in turn permits a direct correlation of macro- and nannofossil zones.
Manuscrit accept6 d6finitivement le 22.01.1992
eobios, 199~ 25, fasc. 2 i 9 263-278 1)
I!
264 GEOLOGICAL SETTING AND P A L A E O G E O G R A P H Y (M.B.)
PALAEOGEOGRAPHIC AND SEDIMENTOLOGICAL CONTEXT
STRUCTURAL CONTEXT
During the later part of the Late Cretaceous, the subpyrenean trough had an Atlantic polarity, bounded to the north by the Aquitaine platform (Ricateau & Villemin 1973). The upper Cretaceous of the Nalzen "Basin" was deposited on the southern side of the trough, as is indicated by the nature and distribution of facies within the Formation de Saint-Cirac (Bilotte 1990; r,g. 2).
To the east of the valley of the Ari6ge, from the meridian of Foix to that of B61esta, the Nalzen "Basin" extends for some 25 kilometres approximately. Situated in the north pyrenean zone, it has tectonic boundaries; the Pech de Foix fault to the north, the frontal thrust of the Saint-Barth616my massif to the south, and the Croquier fault to the east (fig. 1). The Nalzen "Basin" is a complex structure, in which three major units can be recognised (Bilotte 1975, 1985, Bilotte et al. 1988). The northern Saint-Cirac unit is a narrow slice in the form of an anticlinorium, in which the upper part of the Upper Cretaceous is represented by the Formation de SaintCirac.
This unit is between 300 and 500 m thick, although its full extent cannot be determined. The dominant lithology is of blue, highly argillaceous marls, with two associated facies : - fine - and medium-grained sandstones up to 1 m thick, with parallel, sharp upper and lower boundaries ; - rudistid horizons ; in situ in the north east, with biocoenoses at Benaix, Villeneuve d'Olmes, and P6r6ille d'en bas, and resedimented in the south west, with thanatocoenoses at St. Cirac (Catufet, Peyriguel, Pujet ; fig. 2). THE AMMONITE HORIZON (M.H.)
_
5 ~.~.~s,, . /
:)
St.Cirac r V//////////////~c fmt. I
The realignment of the D.117 road between Catufet and Manau exposed about a dozen metres of the Marnes de Saint-Cirac in a blue marl facies, dipping at 20~ to the SSE, as is indicated by rare thin beds of sandstone. The fossil horizon is exposed to a thickness of 1.5 m, at the base of the talus on the south side of the road (Foix 1 : 50,000 topographic sheet, coordinates x = 549, 27 ; y -70, 60, altitude 515 m (Fig. 3).
AGE OF THE AMMONITE FAUNA
(W.J.K.) The ammonite assemblage from the Formation de SaintCirac of the Nalzen Basin comprises : - Desmophyllites diphylloides (FORBES, 1846) (rare) -Eupachydiscus of. isculensis (REDTENBACHER, 1873) (rare) -
Hoplitoplacenticeras
(Hoplitoplacenticeras)
marroti
(COQUAND, 1859) (rare) - Hoplitoplacenticeras
(Hoplitoplacenticeras)
sp. juv.
(rare)
9 4J
ST.
BARTHELEMY
;l~ SAULT
- Glyptoxoceras aquisgranense (SCHLI2IT~, 1872a) (com-
mon) Baculites sp. (rare) - Scaphites (Scaphites) haugi DE GROSSOUVRE, 1894 -
F i g u r e 1 - G e o g r a p h i c a n d geological s i t u a t i o n of the F o r m a t i o n de St. Cirae in the N a l z e n Basin. A : A n t i c l i n o r i u m of St.-Cirac. B : i n t e r b e d d e d m a r l s a n d b i o c o n s t r u e t i o n a l limestone. C : marls, t : N o r t h - P y r e n e a n frontal thrust. 2 : Pech de Foix fault. 3 : M a s s i f de Saint-Barth~16my frontal thrust. 4 : C r o q u i e r fault. 5 : subp y r e n e a n frontal thrust. Situation gaograpbtqua a gdologtqua de la Formation de Salnt-girac darts I* "b~fln" de Na/zen. A : anttcltnortum de St-Ctrac. B : altamancas marnes.calcatres bloconatrutts. 1 : chevauchemant frontal nord-pyrdmdan. 2 : acctdunt radridlonal du Peals de Fotx. 3 : d a ~ a ~ a n t fiontal du Massif de S a i n t - B a b y . 4 :#ill, de Croquter. 5d~vaudaeraent frontal sous.pyrdnd,n.
(rare). The classic division of the Campanian in the type region of Aquitaine (de Grossouvre 1894, 1901 ; Kennedy 1986) is : Upper Campanian : Bostrychoceras potyplocurn zone ; Hoplitoplacenticeras marroti zone.
265
FUTURE
N
NORTH PYRENEAN FAULT
I I LABARRE
SANDSTONE
ST. CIRAC
buitd up
fmt marls
~PkArNE DEui/OtlX
PLATFORM
~
~NALZEN
"basin"
" block)
SUB- PYRENEAN TROUGH VRudistids
Figure 2 - Cross section of the sub-pyrenean basin and its margins during the Campanian at the meridian of the Nalzen basin. Coupe scbdmatklue du bassin sous-pyrdn#en el de ses marges, au Campanien, au rndridten du "basstn" de Nalzen.
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.
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Figure 3 - The fossil locality. A : after the "Carte Geographique de la France" 1:50.000 Foix, sheet (XXI-47). B : General view of the outcrop during road works in the summer of 1989. F : marks the fossiliferous horizon. Situation du gtsementfossilifere. A : d'apr#s la Carte G#ograpbique de la France d 1/50000, feuille de Foix (XXI-47). B : vue d'ensemble des affleurements pendant les travaux d'dlargissement fourier de I'dM 1989. F : po~'tion de l'horizon
fossa~ere. Lower Campanian : Menabites (Delawarella) delawarensis zone ; Placenticeras bidorsatum zone. H. (H.) marroti and S. (S.) haugi are only known from the marroti zone, and the age of the Nalzen assemblage is thus lower Upper Campanian. The nannoflora recovered from the matrix of the ammonites included more than 90 species, and as discussed further below, indicates the Campanian subzone CC18B of Sissingh (1977, 1978) et Perch-Nielsen (1985).
This provides the first direct dating of the lower Upper Campanian marrot/ ammonite zone against the Bannofossil standard.
THE NANNOFLORA (J.B.) The nannoflora recovered in association with the ammonites described above is as follows (for full details of authors of species and combinations see Perch-Nielsen 1985) :
266 Ahmuellerella octoradiata (GORKA, 1957) REINHARDT, 1966 A. regulatis (GORKA, 1957) VERBEEK,1977 Amphizygus brooksii BUKRY,1969 A. papillatus BUKRY,1969 Arkhangelskiella specillata VEKSHINA,1959 Biscutum ellipticum (G6RKA, 1957) GRI3N in GRCrN & ALLEMANN,1975 Braarudosphaera bigelowi (GRAN & BRAARUD, 1935) DEFLANDRE,1947 Broinsonia enormis (SHUMENKO,1%8) MANIVrr, 1971 B. parca (STRADNER,1%3) BUKRu 1969 B. signata (NO~L, 1969) NOL, 1970 Bukryaster hayi (BUKRY, 1969) PRINS & SISSINGH in SISSINGH,1977 Calculites additus (WIND & WISE in WISE & WIND, 1977) PERCH-NIELSEN,1984 C. obscurus (DEH_ANDRE, 1959) PRINS & SISSINGHin 81SSINGH, 1977 C. ovalis (STRADNER, 1963) PRINS & SISSINGH in NISSINGH,1977 Chiastozygus amphipons (BRAMLETYE & MARTINI, 1964) GARTNER, 1968 C. bifadus BUKRY,1969 C. h'tteradus (GORKA, 1957) MANWIT, 1971 Corollithon exiguum STRADNER,1961 C. Kennedyi CRUX, 1981 C. signum STRADNER,1963 Cretarhabdus conicus BRAMLLrlq~ & MARTINI, 1964 Cribrocorona gallica (STRADNER, 1963) PERCHNIELSEN, 1973 Cdbrosphaerella ehrenbergii (ARKHANGELSKY, 1912) DEFLANDREin PIVEI~AU, 1952 Cyclagelosphaera rotaclypeata BUKRY,1969 Cylindralithus biarcus BUKRY, 1969 C. nudus BUKRY, 1969 Discorhabdus ignotus (GORKA, 1957) PERCH-NIELSEN, 1968 Eiffellithus eximius (STOVER, 1966) PERCH-NIELSEN, 1968 E. gorkae REINHARDT,1965 E. parallelus PERCH-NIELSEN,1973 E. turriseiffelii ( D E - - R E , 1954) REINHARDT,1965, BUKRY, 1969 Eprolithusfloralis (STRADNER,1962) STOVER, 1966
E. moratus (STOVER, 1966) Gartnerago obliquum (STRADNER, 1963) REINHARDT, 1970 Gephyrorhabdus coronadventis (REINHARDT, 1966) HILL, 1976 Haqius circumradiatus (STOVER, 1966) ROTH, 1968 Helicolithus anceps (G6RKA, 1957) NO~L, 1970 H. trabeculatus (G6RKA, 1957) VERBEEK,1977 Lithastdnus gdllii STRADNER, 1962 Lithraphidites camiolensis DEFI.ANDRE,1963 Lucianorhabdus arcuatus FORCHHEIMER,1972 L. cayeuxii DEFLANDRE,1959 L. quadrifidus FORCHHEIMER,1972 L. maleformis REINHARDT,1966 Manivitella pemmatoidea (DEFLANDRE in MANIVrr, 1%5) THIERSrEIN,1971 Marthasterites furcatus (DEFLANDRE, 1954) DEFLANDRE, 1959 ; BUKRY,1969 Microrhabdulus decoratus DEFLANDRE,1959 M. undosus PERCH-NIEI~EN,1973 Micula concava (STRADNER in MARTINI & STRADNER, 1960) BUKRY,1969, VERBEEK,1976 M. cubiformis FORCHHEIMER,1972 M. quadrata ? (STRADNER,1961) PERCH-NIELSEN,1984 M. staurophora (GARDET, 1955) STRADNER, 1963 (= M. decussata VEKSmNA,1959) Nannoconus elongatus BRONNIMANN,1955 N. multicadus D ~ R E & D~RE, 1959 Octolithus multiplus (PERCH-NmLSEN, 1973) ROMEIN, 1979 Placozygus fibuliformis (REINHARDT,1964) HOFFMANN, 1970 Prediscosphaem arkhangelskyi (REINHARDT, 1965) PERCH-NmLSEN,1984 P. cretacea (ARKHANGELSKY,1912) GARTNER,1%8 P. grandis PERCH-NIELSEN,1979 P. cf. 1". grandis PEP.Ca-I-NIELSEn,1979 P. microrhabdulina PERCH-NIELSEN,1973 P. ponticula (BUKRY,1959), PERCH-NIELSEN,1984 P. spinosa (BRAMtEI~ & MARTINI, 1964) GARTNER, 1%8 P. stoveri (PERCH-NIELSEn, 1968) SHAr~K & SaXAONER, 1971 Prolatipatella multicatinata GARTNER, 1968 Quadrum ? sp.
PLATE 1 Figs. 1, 18, 13 - Scaphites (Scaphites) haugi DE GROSSOUVRE, 1894. 1, UPS/MH/47 ; 8, UPS/MH/18 ; 13, UPS/MH/19. Figs. 2, 5- Hoplitoplacenticeras (Hoplitoplacenticeras) marroti (COQUAND, 1859). 2, UPS/MH/06 ; 5, UPS/MH/07. Fig. 3 - Desmophyllites diphylloides (FORBES, 1846). UPS/MH/01. Fig. 4 - Hoplitoplacenticeras (Hoplitoplacenticeras) sp. juv. UPS/MH/11. Figs. 6, 7, 11, 12, 14-19 - Glyptoxoceras aquisgranensis (Schliiter, 1872a). 6, UPS/MH/22 ; 7, UPSfMH/20 ; 11, UPS/MH/23 ; 12, UPS/MH/21 ; 14, UPS/MH/24 ; 15, UPS/MH/25 ; 16, UPS/MFU26 ; 17, UPS/MFU27 ; 18, UPS/MH/28 ; 19, UPS/MH/29. All specimens are from the Formation de Saint-Cirac between Catufet and Manau, Ari6ge. Figs. 1, 3, 6, 8, 9, 10, 11, 13-19 are x I ; Figs. 2, 4, 7, 12, 19 are x 4 ; Fig. 5 is x 3.5.
Geobios n ~ 25, fasc. 2
PI. 1 W.J. Kennedy, M. Hansotte, M. Bilotte & +1. Burnett
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268 Reinhardtites anthophorus (DEFLANDRE, 1959) PF_~CHNIELSEN, 1968 R. levis PRINS & SISSINGHin SISSINGH, 1977 Repaguhtm parvidentatum (DEbX~CDRE & FERT in DEFI~NDRE, 1954) FORCHHo.IMER,1972 Retecapsa angustiforata BLACK, 1971 tL crenulata (BRAMLEYrE & MAR'nNL 1964) GRON in GRON & ALLEMANN, 1975 Rhagodiscus acMyostaurion (HILL, 1976) R. angus~s (STRADNFA~1963) REINHARDT, 1971 R. plebeius PERCH-NIELSEN, 1968 R. reniformis PERCH-NIELSI~, 1973 R. splendens (DEFI_ANDRE, 1953) VERBEEK, 1977 Rotelapillus crenulatus (STOVER, 1966) PF~CH-NIELSEN, 1984 Scapholithus fossilis DEFI~\'DRE, 1954 Sollasites horticus (STRADNER et aL in STRADNER & ADAMIKER, 1966) CEPEK & HAY, 1969 Staurolithites mielnicensis (G6RKER & HAY, 1969 Staurolithites rnielnicensis (GORKA, 1957) PERCHNIELSnN,1968 Sensu CRUX in LORD, 1982 Thoracosphaera saxea STRADNER, 1961 Tranolithus gabalus STOVER, 1966 T. minimus (BUKRY, 1969) PERCH-NIElSEN, 1984 T. orionatus (REINHARDT, 1966) RFANHARDT, 1966 (= T. phacelosus STOVER, 1966) T. salillum (NO~L, 1965) CRUX, 1981 Vekshbzella angusta (STOVER, 1966) VF.RBEEK, 1977 Watznaueda barnesae (BLACK in BLACK & BARNES, 1959) PERCH-NIELSEN, 1968 W. biporta BUKRY, 1969 14. rnanivitae BUKRY, 1973 W. ovata BUKRY, 1969 W. quadriradiata BUKRY, 1969 Zeugrhabdotus acanthus R E n ~ , D T , 1965 Z. bipeqbratus (GAgTNE~ 1968) Z. embevgi (NO~L, 1958) PERCH-NIELSEN, 1984 Z. compactus (BUKRY, 1969) (= Rectapontis cornpactus (BUKRY, 1969) VAROL & JAKUBOWSKI,1989 Z. erectus (DEFLANDRE, 1954) REINHARDT, 1965
The nannoflora is relatively well-preserved, and shows Tethyan affinities, the species Bulayaster hayi and Marthastedtes furcatus being present. The cooccurrence of Broinsonia parca, B. hayi and M. furcatus (with Staurolithites rnielnicensis indicates the cosmo-
politan natmofossil zone CC18B (Sissingh 1977, 1978 ; Perch-Nielsen 1985). Some reworked (?) Cenomanian species are present : Corollithon kennedyi and Microstaurus chiastius. In addition, some younger species are present, e.g.P, stoveri and Prolatipatella multica~inata, but index species of Ceratolithoides and Quadrum are absent and Lithastrinus grillii occurs in moderate numbers, all pointing to the present zonal assignation. CONVENTIONS The following abbreviations are used to indicate the repositories of specimens mentioned in the text : BMNH : Natural HistoryMuseum, London. UPS : Universit6 Paul Sabatier, Toulouse. MDA : Mus6e D6partemental de l'Ari6ge, Foix SYSTEMATIC PAl AEONTOLOGY (W.J.K.) Order AMMONOIDEA Zittel, 1884 Suborder AMMONITINA Hyatt, 1889 Superfamily DESMOCERATACEAE Zittel, 1895 Family DESMOCERATIDAE Zittel, 1895 Subfnmily DESMOCERATINAE Zittel, 1895 Genus Desmophyllites SPATH, 1929 [= Schlateda Do. GROSSOUVRO.,1894, p. 126 (non FR_rrsen /n FP,rrSCH & KAFKA, 1887, p. 33) ; Schli~tericeras COLLIGNON, 1938, p. 92 (non HYA'IT, 1903, p. 110)]. Type species - Desmoceras larteti SEUNES, 1891, p. 19, pl. 12(3), fig. 2 ; pl. 13(4), figs. 2, 3, by subsequent designation by Spath, 1921, p. 46, as type species of Schtiiteria, of which Desmophyllites is replacement name. DESMOPHYLLITES 1846) : pl. 1, fig. 3.
DIPHYLLOIDEA
(FORBES,
1846 -Ammonites diphylloides FORBES, p. 105, pl. 8, fig. 8. 1894 Desmoceras pyrenaicum DE GROSSOUVRE, (pars), p. 168, pl. 37, fig. 9 ; non pl. 25, fig. 2.
PLATE 2 Figs. 1-5, 9-15 - Glyptoxoceras aquisgranensis (SCHLOTER, 1872a). 1, UPS/MH/30 ; 2, UPS/MH/31 ; 3, UPS/MH/32 ; 4, UPS/MH/33 ; 5, UPS/MH/34 ; 9, UPS/MH/35 ; 10, UPS/MH/36 ; 11, 14, UPS/MH/37 ; 12, UPS/MH/38 ; 13, UPS/MH/39 ; 15, UPS/Mtt/41. Figs. 6, 7 - Hoplitoplacenticeras (Hoplitoplacenticeras) marroti (COQUAND, 1859). 6, UPS/MH/08 ; 7, UPS/MH/09. Fig. 8 - Baculites sp. UPS/MH/14. All specimens are from the Formation de Saint-Cirac between Catufet and Manau, Ari~ge. Figs 1-3, 5, 11-15 are x 4 ; Fig. 4 is x 2 ; Figs. 6, 7 and 8 are x 1.
Geobios n ~ 25, fasc. 2
PI. 2
WJ. Kennedy, M. Hansotte, M. Bilotte & J. Burner
270 1985 - Desmophyllites ch'phylloides (FORBES, 1846) ; Henderson & McNamara, p. 54, pl. 4, figs. 1-4 (with synonymy). 1989 - Desmophyllites ; Kennedy, text-fig. 17d. 1989 - Desmophyllites diphylloides (FORBES) ; Haggart, p. 193, pl. 8.6, fig. 10 with additional synonymy). Types - Lectotype, by subsequent designation by Matsumoto & Obata 1955 (p. 122) is BMNH C'22682, the original of Forbes 1846, pl. 8, fig. 8 ; paralectotypes are BMNH C22683-5, all from the Upper Maastrichtian Valudavur Formation of Pondlcherry, south India. The lectotype was refigured by Howarth, 1965, pl. 11, fig. 3. Material - Three specimens, UPS/MH/01 ; MDA/MH/ 02-03. a n d d i s c u s s i o n - These specimens are partpyritised, and crushed. The best-preserved specimen is 20 mm in diameter, with a minute, deep umbilicus. The umbilical shoulder is narrowly rounded, the flanks flattened and subparallel, the ventrolateral shoulders broadly rounded, and the venter somewhat flattened. The shell surface is smooth, but for traces of straight, prorsiradiate growth lines. Little of the suture is exposed, but UPS/MH/01 shows a narrow, moderately incised E/L. Description
Discussion - These specimens differ in no significant respects from well-preserved juveniles we have studied from the Upper Santonian of the Corbitres. Henderson & McNamara (1985, p. 54, pl. 4, figs. 1-4) provide a full revision of this species and discuss differences from others referred to the genus.
EUPACHYDISCUS ef. ISCULENSIS (REDTENBACHER, 1873) : pl. 1, figs. 9, 10. compare : 1873 -Ammonites isculensis REDTENBACHER, p. 122, pl. 29, fig. 1. 1894 - Pachydiscus isculensis REDTENI3ACHER ; de Grossouvre, p. 185, pl. 22, fig. 2 ; pl. 26, fig. 1 ; pl. 37, fig. 1. 1986 - Eupachydiscus isculensis REDTENBACHER, 1873 ; Kennedy, p. 52, pk 2, figs. 5, 6 ; pl. 10, figs. 13, 14 ; text-fig. 19 (with synonymy). Holotype - By monotypy, an unregistered specimen in the Obert~sterreichs Landesmuseum, Austria, from the Santonian Gosau Beds between Ischl and Kohlbuch, Austria, the original of Redtenbacher 1873, pl. 29, fig. 1, reiUustrated by Kennedy 1986, text-fig. 19. Material
-
Two specimens, UPS/MH/04 ; MDA/MH/
05. Description - The best-preserved of the specimens is UPS/MH/04 (pl. 1, figs. 9, 10), a wholly septate juvenile 33 mm in diameter. Coiling is very involute, with a depressed, reniform whorl section and an estimated whorl breadth to height ratio of 1.25. Ribs arise either singly or in pairs from umbilical buUae, and are strong, narrow, straight and prorsiradiate on the flanks, strengthening markedly on the ventrolateral shoulders, and crossing the venter in a broad convexity. The suture is deeply and intricately incised, with narrow-stemmed E/L and L/U2. - Although small, these specimens differ in no significant respects from juveniles figured by de Grossouvre (1894, pl. 22, fig. 2 ; pl. 26, fig. 1 ; pl. 37, fig. 1), Summesberger (1979, p. 136, pl. 7, figs. 30, 31 ; pl. 8, fig. 32 ; text-figs. 20-22) and others. Discussion
- Desmophyllites diphylloides has a recorded range of Lower Santouian to Upper Maastrichtian. It is known from southwestern France (Corbitres, Aude, as well as the present records), Tuni.~ia, Angola, Zululand and Pondoland (South Africa), Madagascar, Argentina, California, Arkansas, British Columbia, Alaska, Japan, Australia and south India. Occurrence
Family PACHYDISCIDAE Spath, 1922 Genus Eupachydiscus SPA'If-I, 1922 [= Mesopachydiscus YABE & SHIMIZU, 1926] Type species - Ammonites isculensis RED'I'E.Nq3ACHER, 1873, p. 122, pl. 29, fig. 1, by original'designation.
Occurrence - Eupachydiscus isculensis ranges from the middle of the Santonian to mid-Campanian. It is known from Aquitaine and the Corbitres in France, northern Spain, Austria, Roumania, Zululand (South Africa), and Madagascar. Superfamily HOPLITACEAEA H. DouviUt, 1890 Family PLACENTICERATIDAE Hyatt, 1900 Genus and Subgenus Hoplitoplacenticeras PAULCKE, 1907
PLATE 3 Figs. 1-9 - Glyptoxoceras aquisgranensis (SCHLf..)TER, 1872a). 1, UPS/MH/42 ; 2, UPS/MH/43 ; 3, UPS/MH/44 ; 4, UPS/MH/45 ; 5, UPS,qVIH/46 ; 6, UPS/MH/47 ; 7, UPS/MH/48 ; 8, UPS/MH/49 ; 9, UPS/MH/50. All specimens are from the Formation de Saint-Cirac between Catufet and Manau, Aritge. All figures are x 1.
Geobios n ~ 25, fasc. 2
PI. 3
WJ. Kennedy, M. Hansotte, M. Bilotte & J. Bttrnett
272
[ = Dechenoceras KAYSER, 1924, p. 175]
Material - Five specimens, UPS/MH/06 to 09 ; MDA/ MH/10.
the venter. UPS/MH/07 (pl. 1, fig. 5) is 18 mm in diameter, and has acquired a small inner ventrolateral clavus. Adult ornament is shown by UPS/MH/09 (pl. 2, fig. 7), 57_5 mm in diameter, and UPS/MH/08 (pl. 2, fig. 6), a complete adult with an estimated diameter of 70 mm. Coiling is moderately involute, with a small, shallow umbilicus that comprises 22% of the diameter, and has a low, flattened wall and narrowly rounded umbilical shoulder. There are sixteen crowded umbilical bullae per whorl, perched on the umbilical shoulder, and connected to the umbilical seam by a strong, concave n'b. The bullae give rise to one, sometimes two ribs, while these long ribs may bifurcate around mid flank, and intercalated ribs arise low on the flank to give a total of 40 ribs per whorl. The ribs are strong, narrower than the interspaces and prorsiradiate. They are straight on the inner flank, but flex forwards and are convex at mid-flank, thence flexing back and concave on the outer flank before sweeping forwards to weak, bullate inner ventrolateral tubercles that disappear at the be~nning of the outer whorl, and outer ventrolateral clavi that weaken and disappear on the final sector of body chamber prior to the adult aperture. This is also marked by a weakening of umbilical buUae and ribs, and a proliferation of growth lines and striae. The sutures were not seen.
Description - The earliest growth stages seen are a specimen 4 mm in diameter, MDA/MH/10. This is globose and smooth for the first half whorl, after which blunt umbilical bullae, five per half whorl, appear. These give rise to low, broad, straight prorsiradiate ribs, either singly or in pairs, which terminate in welldeveloped outer ventrolateral clavi. UPS/MH/06 (pl. 1, fig, 2) is 7 mm in diameter, and has similar ornament, showing the clavi alternating in position on either side of
Discussion - The simple ribbing and tuberculation immediately distinguish these specimens from species such as H. (H.) coesfeliense (SCHLt2rER, 1867) (p. 15, pl. 1, figs. 1-5 ; see Kennedy, 1986, p. 73, pl.9, figs. 9, 10 ; text-fig. 27b, ~ f), H. (H.) costulosum (SCHLt:rrER, 1876) (p. 159, pl. 44, figs. 1-4) and H. (hemfoerdiceras) lemfoerdense (SCHLOTER, 1872b) ( = as nomen novum for Ammonites scaphitoides SCHL(3"rER, 1872a, p.63, pl. 19, figs. 1, 2, non COQUAND, 1853, p. 442, pl. 14, figs. 9,
Type species - Hoplites - Placenticeras plasticus PAULCKE, 1907, p. 186 : ICZN Opinion 555, 1959 ; name n ~ 1629. HOPLITOPLACENTICERAS (HOPLITOPLACENT1CERAS) MARROT1 (CoOUAND, 1859) : pl. 1, figs. 2, 5 ; pl. 2, figs. 6, 7. 1859 - Ammonites marroti COQUAND, p:.70. 1984 - Hoplitoplacenticeras vari (SCHLUTER) ; Bilotte, pl. 37, figs. 2, 3. 1986- Hoplitoplacenticeras (Hoplitoplacenticeras) marroti (COOUAND, 1859) ; Kennedy, p. 70, pl. 2, figs. 3, 4; pl. 9, figs. 1-8, 11-12 ; pl. 10, figs. 1-12 ; pl. 12, figs. 1, 2. Type - Holotype, by monotypy, is the specimen from R~erac, Dordogne, mentioned by Coquand (1859, p. 995), refigured by Kennedy 1986 (pl. 9, figs. 5, 6), in the Sorbonne Collection, now in the Universit6 Pierre et Marie Curie, Paris.
PLATE 4 Fig. 1 - Marthasterites furcatus DEFLANDRE. XP, x1250 ; Neg. n ~ UCL-3467-13. Fig. 2 - Lucianorhabdus maleformis REINHARDT. XP, x 1250 ; Neg. n ~ UCL-3467-4. Fig. 3 - Lucianorhabdus quadrifidus FORCHHEIMER. XP, x 460 ; Neg. n ~ UCL-3467-3. Fig. 4 - Calculites ovalis (STRADNER) PRINS & SISSINGH Xp, x 1500 ; Neg. 11" UCL-3467-7. Fig. 5 - Quadrum sp. TL, x 1670 ; Neg. n ~ UCL-3467-23. Figs 6, 7 - Lithastn'nus grillii STRADNER. 6. XP, x 1250 ; Neg. n ~ UCL-3"467-28.7. TL, x 1750 ; Neg. n ~ UCL-3467-31. Fig. 8 - Bukryaster hayi (BUKRY) PRINS & SISSINGH. XP, x 1400 ; Neg. n* UCL-3467-32. Fig. 9 - Broinsonia parca (STRADNER) BUKRY. XP, x 1500 ; Neg. n ~ UCL-3467-16. Fig. 10 - Eiffellithus eximius (STOVER) PERCH-NIELSEN. XP, x 1750 ; Neg. n ~ UCL-3467-11. Fig. 11 - Reinhardtites anthophorus (DEPT_.ANDRE) PERCH-NIELSEN. XP, x 1875 ; Neg. n ~ UCL-3467-15. Fig. 12 - Tranolithus orionatus (REINHARDT) REINHARDT. XP, x 1250 ; Neg. n ~ UCL-3467-5, Fig. 13 - Zeugrahabdotus biperforatus (GARTNER). XP, x 1875 ; Neg. n ~ UCL-3467-14. Fig. 14 - Prediscosphaera grandis PERCIt-NIELSEN. XP, x 1410 ; Neg. n ~ UCL-3467-29. Figs 15, 16 - Watznaueria quadriperforata BUKRu 15. XP, x 1.830 ; Neg. n ~ UCL-3467-18. 16. TL, x 1830 ; Neg. n ~ UCL-3467-19. All negatives cited are preserved in the collections of the Micropalaeontology Unit, Department of Geological Sciences, University College of London. XP = crossed polars ; T : = transmitted light.
Geobios n ~ 25, fast. 2
PI. 4 W.J, Kennedy, M. Hansotte, M. Bilotte & J. Burnett
274 10). H. (H.) vari ( S C H L ~ 1872b) (p. 92 ; nomen novum for Ammonites sttiatocostatus SCHLOTER~1872a~ p. 65, pl. 20, figs. 1-4, non MF.NF_.GHINI,1854, pl. 28) has a rather .~imilar rib style (see Kennedy 1986, text-fig. 25), but here the ribs are less flexuous, and the whole of the shell surface covered in prominent lirae, not seen in the present specimens, which have shell preserved. H. (H.) marrofi (COQUAND, 1859) (see revision in Kennedy 1986, p. 70, pl. 2, figs. 3, 4 ; pl. 9, figs. 1-8, 11-12 ; pl. 10, figs. 1-2 ; pl. 10, figs. 1-i2 ; pl. 12, fags. 1, 2) have 12-14 umbilical bullae per whorl and up to 20 ribs per half whorl in specimens from Aquitaine, with rather blunt ribs in some variants (Kennedy 1986, pl. 9, figs. 7, 8), whereas others (Kennedy 1986, pL 9, figs. 5, 6) are closely comparable to the present specimens, apart from preservation. Occurrence - In Aq#taine, specimens are labelled as being from Assizes pZand p3 of A m a u d (J878), but de Grossouvre (1894, 1901) noted it from p only, which has been our experience. The species occurs at a comparable level at Ressec (Dreuilhe anticline) as well as the present occurrences in Nalzen. It may occur at Loesfeld, Westphalia, Germany, Vijlen near Cotessen on the Belgium-Netherlands border, in Portugal, Israel, Soviet Central Asia, Madagascar and Texas, USA. HOPLITOPLACENTICERAS (HOPLITOPLACENTICERAS) SP. JUV. : pl. 1, fig. 4.
Material - 3 specimens UPS/MH/11 and MDA/MH/ 12-13. - The largest of these specimens is 10 mm in diameter. They differ from H. (H.) marroti, described above, in their much finer, more numerous ribs, with many more ventrolateral elavi per whorl, suggesting the presence of a second Hoplitoplacenticeras spedes. Discussion
Suborder ANCYLOCERATINA Wiedman% 1966 Superfamily T U R R I L I T A C E A E Gill, 1871 Family DIPLOMOCERATIDAE Spath, 1926 Subfamily DIPLOMOCERATINAE Spath, 1926 Genus Gtyptoxoceras SPATH, 1925 [= Neohamites BRUNNSCHWEILER, 1966] Type species - Hamites rugatus FORBES, 1846, p. 116, pl. 11, fig. 6, by orional designation (Spath 1925, p. 30, as Hamites (Anisoceras) rugatus (FORBES) KOSSMAT). GLYPTOXOCERAS AQUISGRANENSE ( S c I - I L ~ 1872a) : pl. 1, figs. 6, 7, 11, 12, 14-19 ; pl. 2, figs. 1-5, 9-15; pl. 3, figs. 1-9 1872a - Toxoceras (?)Aquisgranense SCI-ILOI'ER,p. 102, pl. 31, figs. 6-9.
1887 - Hamites aquisgranensis SCHL~'ER ; Holzapfel, p. 66, pl. 5, rag. 8. 1925 - Hamites aquisgranensis SCI-ILOrER ; Diener, p. 66. Type - Lectotype, here designated, is the original of Schlfiter 1872a, p. 102, pl. 31, figs. 6-9, an unregistered specimen in the Collections of the Geologisches und Paliiontologisches Institut of Bonn University, and from the lower sandy layer with Belemnitella quadrata at the foot of Luseberges, near Aachen, Germany. The spedmen is probably of upper Lower Campanian age, for Schlfiter (1872, pl. 24) illustrated a late form of Scaphites (Scaphites) hippocrepis (DEKAY, 1828) from the same locality. - Almost 100 specimens, UPS/MH/20 to 50 ; MDA/MH/51 to 113. Material
Description - Complete shells are up to 80 mm in diameter, and consist of three distinct ontogenetic stages: an initial straight shaft, followed by a brief helical stage, and then an open plani~piral stage. The protoconch is preserved in UPS/MH/37 (pl. 2, fig. 11 ; counterpart shown in pl. 2, fig. 14). It is overpyritised, with an apparent diameter of 0.4 ram. The succeeding straight growth stage is in the form of a slowly expanding smooth shah up to 20 mm long (191. 1, fig. 12 ; pl. 2, figs. 5, 9, 11-14). Immediately prior to the second growth stage the expansion rate of the shaft increases markedly (pl. 1, fig. 12 ; pl. 2, figs. 5, 12) and be~n.~ to coil, for half a whorl with a very high translation rate (191.2, fig. 12), but thereafter with a low translation rate, to form a helix of at least 5 whorls, the whorls barely in contact, with a circular whorl section. Internal moulds are smooth (pl. 2, fig. 4), but where the shell is preserved, the last three whorls of the spire are ornamented by delicate equal prorsiradiate and feebly flexuous ribs (pl. 2, figs. 2, 10, 13, 15). The spire is succeeded by a brief section of shell on which the translation rate increases markedly to give at most half a whorl of helical coil (pl. 2, figs. 10, 13, 15), succeeded by the third and final growth stage, a planispire of at least two whorls ; the complete ontogeny but for the final section of body chamber is shown in pl. 3, fig. 9. All specimens are crushed. Ornament consists Of sharp, narrow rursiradiate ribs ; the density is initially high, with a rib index of up to 9 (pL 1, fig. 14 ; pl. 3, figs. 1, 5, 9), decreasing to 5 or 6 in some body chamber fragments (pl. 3, fig. 6), although rib density may increase again towards the adult aperture, as shown in the largest, poss~ly adult specimen in the collection (pl. 3, fig. 7). The ribs are generally narrower than the interspaces, weak on the dorsum, but strengthening across the flank, straight on the dorsal flank but flexed forwards in many specimens and feebly concave on the ventral part of the flank. The ribs appear to be at their maximum development over the venter. The suture is typical for the genus, with rectangular little-incised bifid lobes and saddles.
275 This remarkable suite of specimens show for the f'trst time the complete ontogeny of a Glyptoxoceras species. The type material of the type and other species from the Upper Maastrichtian of south India, preserved in the collections of the Natural History Museum in London (Kennedy and Henderson, in press) includes only fragments of loose open helices, more complex examples of which were illustrated, in the case of Glyptoxoceras indicum (FORBES, 1846) (p. 116 (pars), non pl. 11, fig. 4) by Matsumoto (1959, p. 167, pl. 41, figs. 2-6 ; text-fig. 80). These show G. indicum to have a much larger, lower helix than the present species, and an adult shell in the form of an ellipse. The Upper Maastrichtian Glyptoxoceras rugatum (FORBES, 1846) (p. 116, pl. 11, fig. 6) has a compressed whorl section with a rib index of 6-7 in medium-sized specimens from south India. Revision of material from the Miria Formation of western Australia (Henderson, Kennedy & McNamara, in press) show adults of the species to have been up to a metre across, with irregular coiling, long straight or slightly curved sections being linked by short sectors defining an obtuse angle. Discussion
-
Of other European species, Glyptoxoceras vaalsiensis HOLZAPFEL, 1887 (p. 66, pl. 5(2), figs. 6a-c, 7) has very coarse, distant annular ribs that do not weaken on the dorsum according both to the original figures and specimens in the Rheinisch-Westf/ilischen Technischen Hochschule, Aachen, and has highly simplified sutures. The Hamites roemeri GEINrI'z of Holzapfel (1887, pl. 5(2), figs. 4, 5) has much more distant and coarse ribs than the present species.
Ancyloceras retrorsum SCHLOTER~ 1872 (p. 97, pl. 30, figs. 5-8) was revised by Kennedy (1986, p. 106, pl. 16, figs. 1-4, 6, 7 ; pl. 17, figs. 1, 2 ; text-fig. 38), who referred it to Neoglyptoxoceras COLLIGNON, 1969, and reillustrated the type material. This is a much larger species than G. aquisgranense, and appears to have an elliptical rather than circular coil, with juvenile fragments indistinguishable from the present species.
BACULITES SP. : pl. 2, fig. 8. Material - Three specimens, UPS/MH/14 and MDA/ MH 15-16. Discussion - The genus Baculites is represented by fragments only in the present collections. The largest is 31 mm long, with a slowly expanding shell and very distant dorsal nodes on the flank, suggesting it belongs to the widely occurring Baculites capensis WOODS, 1906, group (see revision in Klinger & Kennedy 1977, p.71, text-figs. 2a-f, 3g).
Superfamily SCAPHITACEAE Gill, 1871 Family SCAPHITIDAE Gill, 1871 Subfamily SCAPHITINAE Gill, 1871 Genus and subgenus Scaphites PARK1NSON, 1811 [= Anascaphites HYATE, 1900, p. 572 ; Jahnnites HYATI', 1900, p. 572 ; Holcoscaphites NOWAK, 1911, p.
564] Type species - Scaphites equalis J. SOWERBY, 1813, p. 53, pl. 18, figs. 1-3.
SCAPHITES (SCAPHITES) HAUGI DE GROSSOUVRE, 1894 : pl. 1, f~s. 1, 8, 13 ; text-fig. 4 1894 - Scaphites haugi A. DE GROSSOUVRE sp., p. 244, pl. 31, fig. 5. 1925 - Scaphites haugi DE GROSSOUVRE ; Diener, p. 199. 1928 - Discoscaphites? hau# DE GROSSOUVRE ; Reeside, p. 30. 1970 - Scaphites cf. haugi DE GROSSOLWRE ; Atabekian & Akopyan, p. 39, pl. 2, figs. 3, 4. 1986 - Scaphites (Scaphites) haugi DE GROSSOUWRE ; Kennedy, p. 116, pl. 21, figs. 12-16. Holotype - By monotypy, is the original of de Grossouvre 1894, pl. 31, fig. 5, an unregistered specimen in the Sorbonne Collections, now housed in the Universit6 Pierre et Marie Curie, Paris, ex Arnaud Collection, from
Occurrence - The type material is from the Vaals Formation, and probably of upper Lower Campanian age, as is material described by Holzapfel (1887) ; the present material is thus somewhat younger.
ii / / I
Family BACULITIDAE Gill, 1871 Genus Baculites LAMAP,CK, 1799
[ = Homaloceratites HUPSCH, 1768, p. 110 (non binomen) ; Euhomaloceras SPATII, 1926, p. 80] Type species - Baculites vertebralis LAMARCK, 1801, p. 103, by subsequent designation by Meek (1876, p. 391).
I
Figure 4 - Suture line of Scapbaes ($capbttes) baugi (DE GROSSOOVRE, 1894). UPS/MH/17 (see also pl. 1, fig. 1). Bar scale is 10 mm. Ztgne do s~x~e de Scapbttes (Scapbttes) baugt (D~ oRossovn~,
1894).
276 Assize p3 of Arnaud (1878) at TauiUard (Charente), ref'~ured by Kennedy 1986, pl. 21, figs. 12-14. Material - Three specimens, UPS/MH/17 to 19. Description - All the specimens are crushed. One is an incomplete phragmocone 19 mm in diameter, a second an incomplete microconch with a phragmocone 22 mm in diameter and part of the body chamber shaft preserved. Coiling of the phrao~mocone is very involute, with a tiny umbilicus comprising 12% of the diameter, deep, with a flattened wall and narrowly rounded shoulder. Twelve distant, coarse, straight, prorsiradiate primary n'bs arise at the umbilical seam~ and strengthen across the umbilical wall and shoulder, terminating in strong, conical ventrolateral tubercles. These give rise to pairs of weaker secondary ribs, with a .~ingle short intercalated r ~ between, all ribs crossing the venter without diminutionThe umbilical wall of the shaft of the body chamber is concave, and the wall follows a concave course in profile, such that all of the umbilicus of the phragmocone is exposed. There are four very coarse, distant ribs on the section preserved. These arise at a strong conical to feebly bullate umbilical tubercle, are straight and prorsiradiate on the flanks and terminate in strong ventrolateral clavi. The venter is poorly preserved, but groups of secondary and intercalated ribs are present. The third specimen is a macroconch with a phragmocone an estimated 36 mrn ill diameter, and a total length to the shell of 56 mm~ Phragmocone ornament is as in the microconch, with in addition, a few long secondary ribs. The body chamber is much higherwhorled than that of the microconch, with a convex umbilical wall that is straight when viewed in profile, partially occluding the umbilicus of the phragmocone. There are five conical to feebly bullate umbilical tubercles on the body chamber. The adapical one is weak, the second and third massive, and those on the final hook much weakened. They give rise to one or two low, broad primary ribs that terminate in coarse ventrolateral clavi that are restricted to the shaft and initial sector of the hook. The clavi give rise to groups of two or three coarse secondary ribs that loop to the clavi on the opposite flank, and there are. some intercalated fibs. Umbilical tubercles are very weak, and ventrolateral tubercles absent from the final sector of shell prior to the adult aperture, which is ornamented by narrow ribs on outermost flank, ventrolateral shoulders and venter. The suture is only partially exposed (text-fig. 4). - Kennedy (1986, p. 116) redescribed the type specimen of this species, and discussed differences from the most closely allied species. The older S. (S.) hippocrepis (DEKAY, 1828) is generally smaller, the spire has a rounded whorl section with weak or no tubercles, while the body chamber is weakly tuberculate, and may have a lateral tubercle in Discussion
some specimens (see Cobban, 1969, for a full review of the hippocrepis group). The younger S. (S.) binodosus (ROEMER, 1841) (p. 90, pl. 13, fig. 16 ; of which S. (S.) inflatus ROEMER, 1841 (p. 90, pl. 14, fig. 3) is the macroconch) is discussed by Kennedy (1986, p. 116, text-figs. 39a-e). This is a much larger species than S. (S.) haugi, with a quite different shell shape, the microconchs with numerous umbilical tubercles on the body chamber, with broad, flattened triangular ribs; macroconchs have only feeble umbilical bullae on the body chamber, with numerous weak flank ribs and small ventrolateral clavi. Occurrence - Apart from the type occurrence, there is a second, topotype specimen, and a third from division Campanian 3 of Plate 1 (1977) near Prrigueux (Dordogue). Atabekian & Akopian (1970) recorded Scaphites el. haug/from the Campanian of Soviet Armenia. Acknowledgements - Burnett acknowledges the financial support of the Natural Environment Research Council (UK) under the terms of Grant GR3/6767, and technical support of the staff of the Micropalaeontology Unit of the Department of Geological Sciences, University College, London. Kennedy acknowledges the financial support of the Natural Environment Research Council (UK) under the terms of Grant GR3/6501a, and the technical assistance of the staff of the Department of Earth Sciences, Oxford and Geological Collections, University Museum, Oxford.
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