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Acta Geologica Polonica, Vol. 54 (2004), No.3, pp. 369-380

Middle Cenomanian ammonites from the type section of the Sanandinovo Formation of northern Bulgaria

WILLIAM J. KENNEDyl & NIKOLAI JOLKICEV2

10xford University Museum of Natural History, Parks Road, Oxford OXI 3PW, United Kingdom E-mail: [email protected] 2Department of Geology & Geography, University of Sofia, Russki Boulevard 15, Sofia 1000, Bulgaria

ABSTRACT:

KENNEDY, W:J. & JOLKICEV, N. 2004. Middle Cenomanian ammonites from the type section of the Sanandinovo Formation of northern Bulgaria. Acta Geologica Polonica, 54 (3), 369-380. Warszawa.

The type section of the Sanandinovo Formation between the villages of Sanandinovo and Novachene in northern Bulgaria has yielded numerous ammonites from packages 7-9 of the sequence recognised by previous authors. The assemblage is: Tetragonites subtimotheanus WIEDMANN, 1962, Puzosia (Puzosia) maY0l7ana (D'ORBIGNY, 1841), Acanthoceras jukesbrownei (SPATH, 1926), Protacanthoceras tubercula tum tuberculatum THOMEL, 1972, Calycoceras (Newboldiceras) asiaticum asiaticum (JIMBO, 1894), C. (N.) asiaticum spinosum (KOSSMAT, 1897), C. (N.) planecosta tum (KOSSMAT, 1897), Calycoceras (Proeucalycoceras) picteti WRIGHT & KENNEDY, 1990, and Scaphites equalis J. SOWERBY, 1813. The assemblage indicates the upper Middle Cenomanian Acanthoceras jukesbrownei Zone of Western Europe. We have seen no evidence for either older or younger Cenomanian ammonite zones from the sequence.

Key words: Ammonites, Cenomanian, Bulgaria.

INTRODUCTION ------erosional contact ------ Packet lO-gray, indistinctly bedded, clayey marls with The Sanandinovo Formation is a 78 m thick extremely rare interbeds of fine-bedded (10-15 cm) sequence of marls with rarer interbeds of marly lime clayey limestones - 36.00 m stone that has a type section between the villages of Packet 9-gray, indistinctly bedded, clayey marls with Sanandinovo and Novachene in northern Bulgaria extremely rare interbeds of fine-bedded (10-20 cm) (JOLlGCEV, 1987, 1989; JOLlGCEV & al. 1988; Text-fig. 1). microgranular clayey limestones - 15.00 m The formation overlies glauconitic sandstones assigned Packet 8-marls, slightly aleuritic, yellowish, indistinctly to the Upper Albian Stoliczkaia dispar Zone, and is in bedded with frequent limonite concretions - 5.00 m turn overlain by glauconitic sandstones of the Packet 7-marls (opposite the bridge on the Osam River), Belobardo Formation, the base of which is assigned to gray, massive, commonly including muscovite flakes - the Upper Turonian on the basis of the occurrence of 5.00m Subprionocyclus hitchinensis (BILLINGHURST, 1927) and Packet 6-marls, gray, indistinctly bedded, locally slightly other fossils. It is divided into six sediment packets aleuritic - 10.00 m (Text-fig. 2), following JOLlGCEV & al. 1988, as follows: Packet 5 -marls, grayish-whitish, indistinctly bedded, clayey, at the base slightly arenitic, with glauconite - 7.00 m - glauconitic sandstones (Be1obardo Formation-Upper ------gradual transition ------ Turonian - Lower Campanian) Packet 4-(Dispar Zone) - glauconitic sandstones 370 w.J. KENNEDY & N. JOLKICEV

The microfaunas of the Formation were listed by Ammonite evidence for a Lower Cenomanian date JOLKICEV & al. (1988), who also provided a preliminary for the lower part of the Sanandinovo Formation is lack listing of the ammonite faunas. On the basis of these ing in the material studied here. NIKOLOV & NACHEV records, and those of NIKOLOV & NACHEV (1962) in cor (1962) recorded Mantelliceras mantelli (J. SOWERBY, 1814) relative sediments in the Pleven district, they recognised a and Mantelliceras martimpreyi (COQUAND, 1862) from the Lower Cenomanian Mantelliceras mantelli Zone, to which correlative of package 5 in the Pleven district. We have they assigned packets 5 and 6, a Middle Cenomanian been unable to examine these specimens; their M mantel Acanthoceras rhotomagense Zone, to which they assigned li (1962, pI. 1, figs 1, 2) is generically indeterminate from packets 7, 8, and part of 9, and an Upper Cenomanian the figures. Their M martimpreyi (1962, pI. 1, fig. 3) does Calycoceras naviculare Zone, to which they assigned the not belong to that species, which is a junior synonym of upper part of packet 9, and packet 10. Mantelliceras saxbii (SHARPE, 1857), and is again generi The currently widely recognized ammonite zonation cally indeterminate from the figure. of the Cenomanian stage in Western Europe is as follows: At the type section, ammonites are present from the basal part of package 7 to the upper part of package 9. The Substage Zone Subzone assemblage is: Tetragonites subtimotheanus (WIEDMANN, 1962) Puzosia (Puzosia) mayoriana (D'ORBIGNY, 1841), u { Neocardioceras juddii Acanthoceras jukesbrownei (SPATH, 1926), Protacanthoceras pper . . . . Metozcoceras geslznzanum tuberculatum tuberculatum TROMEL, 1972, Calycoceras Cenomaman Calycoceras gue/'angeli (Newboldiceras) asiaticum asiaticum (JIMBO, 1894), C. (N.) asiaticum spinosum (KOSSMAT, 1897), C. (N.) planecosta M'ddl {ACanthoceras jukesbrownei .. tum (KOSSMAT, 1897), Calycoceras (Proeucalycoceras) I e { Tumbtes acutus . Acanthoceras rhotomagense picteti WRIGHT & KENNEDY, 1990, and Scaphites equalis J. CenomanIan .. .. Tunilites costatllS Cunnzngtomceras cunnzngtom SOWERBY, 1813. The range and occurrence of all of these species is documented in detail by WRIGHT & KENNEDY Lower {Mantelliceras dixoni (1984, 1987, 1990, 1995, 1996), and the assemblage can be Mantelliceras saxbii confidently assigned to the upper Middle Cenomanian Cenomanian Mantelliceras mantelli { Neost. carcitanense Acanthoceras jukesbrownei Zone. We have seen no evi-

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Fig.!. Lccality map showing the position of the type section of the Sanandinovo Formation, which is between the villages of Sanandinovo and Novachene, in northern Bulgaria 21 qq t--' ALBIAN CIJJ. m CENOMANIAN -;, STAGE ;:; (part) po'" co :IJ1JJ ~. 0» 8 Z:IJ SANADINOVO -0 FORMATION ""[- ~o '" '""Tl " ...... Os:: SEDIMENT g. ..j:::.. (J1 0) -....I (X) co o· 0 PACKET "0 :z '"'& ,,":.:-.':', "(/) po :~·:··4·~· ~ ~ ~ II" ~ I"- ~ 1"-1- r I~ "--~ ~-I" po "f- '" " " " " " "'"0 "&. '" '" ~ c···tt ~ ~ ~ I ~II"- I~ I~ I" I I I sr "0 ~ '" ~ " I" G~R ~ 0 .::.: .. :.• >:.: .. : " " "" " " " " " 6 & 61 1"-1- r ,,- u § : ;. I" I" I- " "f.-" "f- ",. ~4 ~ " po ...... '"~ " po g. " " " " " " " " " " " " " c¥" ...... c.v THICKNESS ~ "!", -....I (J1 (J1 o '" 0 (J1 0) IN METRES ""(S."'" o..q'" U). >-c ~ 6". ~ cr" " ~ ~ a. Tetragonites subtimotheanus o· 0 ~ " cr"" I ~ 2., " S ? • Puzosia mayoriana ~ 8 " 0 " ~ 8. "& I '" ~ :5." Acanthoceras jukes brownei ;;J 0.. ~ o ~ I ~ g ""~ tll 5' 2., • • Protacanthoceras tubercula tum tuberculatum c:: 0.. " po s· '" I I I & " "'"&. - Calycoceras (N) asiaticum asiaticum "(t ~ ;:s. "0 "0 I po c. (N) asiaticum spinosum "0.. Z S] '"(") • I. c. (N) planecostatum "'" ,"" s· • Calycoceras (Proeucalycoceras) picteti 0 "8- I ~ - Scaphites equalis "tll & r:; -...J p" I '"'"-' 372 w.J. KENNEDY & N. JOLKILEV dence for either older or younger Cenomanian ammonite Wb = -(-); WI) = 36.7(45.5%); U = 19.3(23.9). The spec zones. imen retains a 2400 sector of body chamber, and shows eccentric coiling over the last half of the outer whorl, sug gesting it may be adult. The umbilicus is deep, with a flat SYSTEMATIC PALAEONTOLOGY tened, subvertical wall, and narrowly rounded umbilical shoulder. The flanks are flattened and subparallel, the All dimensions are given in millimetres as follows: D ventrolateral shoulders broadly rounded, and the venter = diameter; Wb = whorl breadth; Wh = whorl height; U very feebly convex. The compressed whorl section (PI. 1, = umbilicus. Figures given in parentheses are dimensions Figs 1, 3) is a result of the post-mortem crushing. There expressed as a percentage of diameter. Repositories of are an estimated four constrictions on the half whorl that specimens are indicated as follows: GPSU: Department comprises the adapertural end of the phragmocone and of Geology and Geography, University of Sofia; BMNH: the adapical cnd of the body chamber. The constrictions The Natural History Museum, London; OUM: Oxford are narrow, faint, markedly prorsiradiate, very feebly con University Museum of Natural History. cave on the inner to middle flank, broadly convex on the outermost flank and ventrolateral shoulder, sweeping backwards, and crossing the venter in a relatively deep Order Ammonoidea ZITTEL, 1884 concavity. The constrictions are accompanied by growth Suborder Lytoceratina HYATT, 1889 lines and striae, and ill-defined adapical collars, most Superfamily Tetragonitaceae HYATT, 1900 obvious on the ventrolateral shoulders. There is a faint, Family Tetragonitidae HYATT, 1900 low, rounded mid-ventral ridge. Genus Tetragonites KOSSMAT, 1895 DISCUSSION: See WIEDMANN (1973) and THOMEL TYPE SPECIES: Ammonites timotheanus PICfET, 1847, (1987). p. 295, pI. 2, fig. 6; pI. 3, figs 1, 2, by original designation. OCCURRENCE: The total stratigraphic range of T. sub timotheanus and its subspecies is upper Lower Albian to Tetragonites subtimotheanus WIEDMANN, 1962 Upper Cenomanian. The geographic range extends from (PI. 1, Figs 1-3; PI. 5, Fig. 5) SE France to southern England, Bulgaria, Crimea, Zululand (South Africa), Madagascar, south India, 1973. Tetragonites subtimotheanus macleami WIEDMANN, p. 595, Alaska, Oregon and British Columbia. pI. 4, fig. 1; pI. 5, figs 1-3; text-fig. 3. 1984. Tetragonites subtimotheanus subtimotheanus WIEDMANN, 1962; WRIGlrr & KENNEDY, p. 40, pI. 1, figs 3, 4; text-fig. 19 Order Ammonitina HYATT, 1889 (with synonymy). Superfamily Desmocerataceae ZITTEL, 1895 1987. Tetragonites subtimotheanus maclearni WIEDMANN; Family Desmoceratidae ZITTEL, 1895 TROMEL, p. 16, pI. 2, figs 5, 6; pI. 3, fig. 2; pI. 4, figs 1-3, 8- Subfamily Puzosiinae SPATH, 1922 10; pI. 5, figs 1, 2, 6-8 (with synonymy). Genus Puzosia BAYLE, 1878 1988. MesogaUlilyceras ieptonema (SHARPE); JOLI(]CEV & ai., p. 27. Subgenus Puzosia BAYLE, 1878 1988. Mesogaud/yceras cf.ieptonema (SHARPE); JOLI(]CEV & ai., p.27. TYPE SPECIES: Ammonites planulatus 1. DE C. 1994. Tetragonites subtimotheanus WIEDMANN, 1962a; KENNEDY, SOWERBY (1827, p. 134, pI. 570, fig. 5) (non SCHLOTHEIM, p. 217, pI. 2, fig. 15; pI. 4, figs 5,6,9-14,18-22. 1820, p. 59 = Ammonites mayorianus D'ORBIGNY, 1841, p. 267, pI. 79, figs 1-3). WRIGHT & KENNEDY (1984, p. 54) TYPE: The holotype, by original designation, is the origi clarify the name of the type species. nal of KOSSMAT (1895, pI. 17(3), fig. 13), from the lower part of the Utatur Group of south India, and is probably of Cenomanian age. Puzosia (Puzosia) mayoriana (D'ORBIGNY, 1841) (PI. 1, Figs 4-6) MATERIAL: GPSU 2603, 2612. 1827. Ammonites planuiatus J. DE C. SOWERBY, p. 597, pI. 570, fig. DESCRIPTION: 2612 (PI. 5, Fig. 5) is a very laterally 5 (non SCHLOTHEIM, 1820, p. 59). crushed individual 51 mm in diameter, with U = 26% of 1841. Ammonites mayorianus D'ORBlGNY, p. 267, pI. 79, figs 1-3. diameter. 2603 (PI. 1, Figs 1-3) is less crushed, 50.7 mm in 1984.Puzosia (Puzosia) mayoriana (D'ORBIGNY, 1841); WRIGHf diameter, with the following dimensions: D = 50.7(100); & KENNEDY, p. 55, pI. 3, figs 1,2,4,6,9-12; pI. 4, figs 1,2, MIDDLE CENOMANIAN AMMONITES FROM BULGARIA 373

5-7; text-figs la, b; 2c, h, m; 3n-r; 4a-e (with full synonymy). DISCUSSION: See WRIGHT & KENNEDY (1984, p. 87), 1985. Puzosia (puzosia) cf. sUbpianuiata (SCHLUTER, 1871); and COOPER & KENNEDY (1987, p. 106). IMMEL & SEYED-EMAMI, p. 91. 1987. Puzosia (Puzosia) mayoriana (D'ORBIGNY, 1841); COOPER OCCURRENCE: Upper Albian to Upper Cenomanian, & KENNEDY, p. 106, figs 1-7, 9, 10. Europe, Africa, South India, Japan. 1987. Puzosia (Puzosia) mayO/iana (D'ORBIGNY, 1841); WRIGHT & KENNEDY, p. 142, pI. 47, fig. 4. 1991. Puzosia (Puzosia) mayoriana (D'ORBIGNY, 1841); Superfamily Acanthocerataceae DE GROSSOUVRE, 1894 DELAMEITE & KENNEDY, p. 442, fig. 8.26. Family Acanthoceratidae DE GROSSOUVRE, 1894 1994. Puzosz'a (Puzosia) pianulata (D'ORBIGNY, 1841); KENNEDY, Subfamily Acanthoceratinae DE GROSS OUVRE, 1894 p. 220, pI. 2, figs 10-12, 18; pI. 5, figs 1-9. Genus Acanthoceras NEUMAYR, 1875 1996. Puzosia (Puzosia) mayoriana (D'ORBIGNY, 1841); KENNEDY in GALE & ai., p. 552, figs lOf; 11k, I; 14h-n. TYPE SPECIES: By the subsequent designation of DE 1996. Puzosia (Puzosia) mayoriana (D'ORBIGNY, 1841); GROSSOUVRE (1894, p. 27) Ammonites rhotomagensis KENNEDY, BlLOITE & HANSOITE, p. 312, PI. 39, Figs 4, 5 BRONGNIART (1822, p. 83, 391, pI. 6 fig. 2). (with additional synonymy). 1998. Puzosz'a (Puzosia) mayO/iana (D'ORBIGNY, 1841); KAPLAN & ai., p. 71, pI. 1, figs 12, 13; pI. 9, figs 1, 11, 12; pI. 10, figs Acanthoceras jukesbrownei (SPATH, 1926) 3-5. (PI. 2, Figs 9-12; PI. 3, Fig. 12) 1998. Puzosia mayoriana (D'ORBIGNY, 1841); LEHMANN, p. 15, pI. 1, figs 3, 4, 6; pI. 2, fig. 7. 1926. Protacanthoceras jukesbrownei SPATH, p. 82. 1984. Acanthoceras jukesbrownei (SPATH); KAPLAN & ai., pI. 6, LECTOTYPE: By the subsequent designation of fig. 2. WRIGHT & WRIGHT, 1951, p. 35): BMNH C9381, from 1986.Acanthoceras jukesbrownei (SPATH); KENNEDY, pI. 7, figs 3, 4. the Lower Chalk of Hamsey near Lewes, Sussex. This 1987b.Acanthoceras jukesbrownei (SPATH, 1926a); WRIGHT & specimen was designated lectotype of planulatus of J. DE KENNEDY, p. 191, pI. 49, figs 2-4; pI. 50, figs 1-5; pI. 51, figs C. SOWERBY for which D'ORBIGNY proposed mayorianus 1-7; text-figs 55-62; 64c, 65e, f, h-j, I, m; 66e; 67h-l (with as replacement name. full synonymy). 1988. Acanthoceras jukesbrownei (SPATH, 1926a); LEHMANN, p. MATERIAL: GPSU 3955. 143, figs 12-15. 1988.Acanthoceras rhotomagense (BRONGNIART); JOLKICEV & DESCRIPTION: GPSU 3955 is a well-preselVed, lateral al.,p.27. ly crushed composite mould. The dimensions are, D = 1992. Acanthocerasjukesbrownei (SPATH); THOMEL, pI. 34, figs 1, 107.3(100); Wb = 24.6(22.9); Wh = 41.1(38.3); U = 2; pI. 35, pI. 37, figs 3, 4. 31.1(29.0). Coiling is moderately involute, with 77% of the 1994. Acanthoceras jukesbrownei (SPATH); KENNEDY, p. 225, pI. previous whorl covered. The umbilicus is shallow, with a 9, figs 11, 12. low, flattened, outward-inclined umbilical wall. The 1996. Acanthoceras jukesbrownei (SPATH, 1926a); KENNEDY & umbilical shoulder is very narrowly rounded. The whorl ai., p. 313, pI. 40, fig. 5. breadth to height ratio (modified by compaction) is 0.6, ?1997.Acanthoceras cf. jukesbrownei (SPATH); WILMSEN, pI. 19, with very feebly convex, flattened, subparallel flanks, and fig. 4. narrowly arched, rounded venter. There are 7 widely sep 1997. Acanthocerasjukesbrownei (SPAIH); WILMSEN, pI. 38, fig. 1. arated constrictions on the outer whorl. They are straight 1998. Acanthocerasjukesbrownei (SPATH, 1926a); KAPLAN & ai., and prorsiradiate on the inner flank, convex on the mid p. 144, pI. 48, figs 3, 4; pI. 49, fig. 1; pI. 50, figs 1, 2; pI. 62, flank region, flexing back and concave on the outer flank, fig. 3. and sweeping forwards to cross the venter in a marked 1998. Acanthoceras jukesbrownei (SPATH, 1926a); LEHMANN, p. convexity. The edges of the constrictions are strengthened 25, pI. 4, fig. 2. into narrow collar ribs, the adapical more prominent than the adaperturaI. The constrictions are weak on the flanks MATERIAL: GPSU 3942, 2651 but strengthen and become very prominent (notably the adapical one) across the venter (PI. 1, Fig. 6). There are 9- TYPE: Holotype, by monotypy, is BMNH 50162, the orig 13 ribs between the collared constrictions. They parallel inal of SHARPE (1855, pl. 17, fig. 2) [reillustrated by the constrictions, are very weak on the inner flank, but WRIGHT & KENNEDY (1987, pI. 50, fig. 3)] from the base strengthen across the outer flanks, and are at their most ment bed of the Lower Chalk at Man O'War Cave, prominent on the ventrolateral shoulders and venter. Dorset, England. 374 w.J. KENNEDY & N. JOLKJCEV

DESCRIPTION: 3942 is a nucleus, deformed into an 1972. Protacanthoceras tl.lbercl.llatum THOMEL, p. 101, pI. 32, figs ellipse with a maximum preserved diameter of 32 mm (PI. 9-12. 2, Figs 9-12). The coiling appears to have been moderate 1987. Protacanthoceras tubercl.llatum THOMEL, 1972; WRlGIIT & ly evolute, the whorl section depressed trapezoidal in inter KENNEDY, p. 206, pI. 46, fig. 5; pI. 54, figs 1-4; text-fig. 82A, costal section, and depressed polygonal in costal section. C, H; text-fig. 84A-C (with full synonymy). There are an estimated 21 ribs on the outer whorl. Primary 1988. Protacanthoceras compressl.lm (JUKES-BROWNE); JOLKICEV ribs arise at the umbilical seam, and strengthen into coarse & aI., p. 26, 27. bullae, perched on the umbilical shoulder. These give rise to coarse, straight, prorsiradiate ribs that link to strong, TYPE: The holotype, by monotypy, is the original of subspinose irmer ventrolateral tubercles. A coarse rib links TROMEL (1972, pI. 32, figs 9-12), no. 17525 in the TROMEL to weaker outer ventrolateral clavi, and passes straight Collection, housed in the Collections of the Faculte des across the venter, bearing a sub equal siphonal clavus. Sciences, Nice, from the lower Upper Cenomanian of Primaries are separated by single intercalated ribs that Peille, Alpes-Maritimes, France. arise both low and high on the flanks and have no, or only a weak irmer ventrolateral tubercle. These ribs strengthen MATERIAL: GPSU 2579, 2645. to match the primaries, and bear equal outer ventrolateral and siphonal clavi. GPSU 2651 (PI. 3, Fig. 12) comprises a DESCRIPTION: GPSU 2579 (PI. 2, Figs 5-8) is a well 2400 sector of juvenile body chamber with an estimated preserved, laterally crushed composite mould with the original diameter of 124 mm. Coarse primary ribs arise at following dimensions at D = 28.0 (100); Wb: -(-); Wh: well-developed umbilical bullae, and are straight and pror 12.2(43.6); U: 7.4(26.4); the original diameter was over siradiate on the flanks, linking to coarse conical-clavate 31 mm. Coiling is moderately involute, the umbilicus of irmer ventrolateral tubercles. These ribs are separated by moderate depth, with a flattened subvertical wall and single intercalated ribs that arise low on the flank and link narrowly rounded umbilical shoulder. The whorl section to similar irmer ventrolateral tubercles. A low, broad rib is compressed, as a result of post-mortem crushing, with links these tubercles to long outer ventrolateral clavi, and flattened subparallel flanks. Primary ribs arise at the a very broad rib that is wider than the interspaces links to umbilical seam, and strengthen across the umbilical a weak, effacing siphonal clavus (PI. 3, Fig. 12). wall, developing into weak to strong bullae, perched on the umbilical shoulder, six in all on the outer half whorl. DISCUSSION: As noted by WRIGHT & KENNEDY (1987, They give rise to single straight, prorsiradiate, coarse, p. 192), juveniles ofA. jukesbrownei differ from those ofA. feebly convex ribs, with single ribs intercalated between, rhotomagense in the presence of one or several secondary in some cases tenuously linked to the umbilical bulla. ribs between primaries with weaker or no inner ventrolat All ribs bear strong conical to feebly clavate inner ven eral tubercles, rather than regularly alternate long and trolateral tubercles. A broad strong rib connects to short ribs with equal inner ventrolaterals. In middle strong outer ventrolateral clavi, linked across the venter growth short ribs persist to 120 mm at least in early forms by a strong transverse rib, bearing a subequal siphonal ofjukesbrownei, and persist to maturity in late forms. clavus. It is interpreted as an incomplete macroconch. GPSU 2645 (PI. 2, Figs 1, 2) is an incomplete individual OCCURRENCE: Upper Middle Cenomanian, index of preserved to a diameter of 28 mm, associated with a the A. jukesbrownei Zone. Recorded from southern juvenile Calycoceras (Newboldiceras) sp. Ornament is England, the Munster Basin in Westphalia, Germany, coarser, with more widely separated ribs than in GPSU Boulonnais, Haute Normandie, Sarthe, and Provence in 2579, while at the largest preserved diameter, the inner France, Poland, Bulgaria (the present records), and Kopet ventrolateral tubercles are lost and the last two ribs Dag, Turkmenistan. crowd, suggesting the specimen to be a near-complete microconch.

Genus Protacanthoceras SPATH, 1923 DISCUSSION: The representatives of the genus Protacanthoceras were comprehensively reviewed by TYPE SPECIES: Ammonites bunburianus SHARPE (1853, WRIGHT & KENNEDY (1980, 1987), who discuss differ p. 25, pI. 9, fig. 3) by original designation by SPATH (1923, ences between P. tuberculatum mite and other sub p.144). species.

OCCURRENCE: Middle and lower Upper Cenomanian, Protacanthoceras tuberculatum tuberculatum TROMEL, 1972 southern England, France, and Bulgaria (the present (PI. 2, Figs 1, 2, 5-8) records). MIDDLE CENOMANIAN AMMONITES FROM BULGARIA 375

Genus Calycoceras HYATT, 1900, p. 589 DESCRIPTION: GPSU 2649 (PI. 5, Figs 1-4) is a well (ICZN Generic Name No. 1352) preserved juvenile, with a maximum preserved diameter of 48.3 mm. Coiling is moderately evolute; the umbilicus TYPE SPECIES: By designation under the Plenary comprises 26% approximately of the diameter. The whorl Powers (ICZN Opinion No. 557) Ammonites navicularis section has been modified by post-mortem compaction MANTELL (1822, p. 198, pI. 22, fig. 5) (ICZN Specific but appears to have been trapezoidal in intercostal sec name). tion, and polygonal in costal section. There are an estimated 40 or more ribs on the outer whorl. On the adapical half whorl, primary ribs arise at Subgenus Calycoceras (Newboldiceras) THOMEL, 1972 the umbilical seam, and strengthen into prominent sharp bullae, perched on the umbilical shoulder. Straight, TYPE SPECIES: By original designation by THOMEL prorsiradiate ribs strengthen across the flanks and ven (1972, p. 105) Acanthoceras newboldi KOSSMAT, 1897, p. ter, and bear strong subspinose inner ventrolateral 5(112), which is a junior synonym of Acanthoceras rho tubercles, and smaller outer ventrolateral and siphonal tomagense var. asiaticum JIMBO, 1894, p. 177, pI. 20, fig. 1 clavi. The primary ribs are separated by one or two long (WRIGHT & KENNEDY, 1990, p. 239). or short intercalated ribs with no or weak inner ventro lateral tubercles, plus outer ventrolateral and siphonal clavi that are equal in strength to those on the primary Calycoceras (Newboldiceras) asiaticum asiaticum (JIMBO, ribs. On the adapertural half of the outer whorl, bullate 1894) primary ribs are separated by up to 3 long or short inter (PI. 3, Fig. 1; PI. 4, Figs 6, 7; PI. 5, Figs 1-4) calated ribs. The inner ventrolateral clavi decline and are lost, the outer ventrolateral clavi persist, while the 1865.Ammonites Rotomagensis DEFRANCE; STOLICZKA, p. 66 siphonal clavi decline. These changes suggest the speci (pars), including typicus (p. 68) (pars) and var. subcompres men may be a small micro conch. sus (p. 68), pI. 34, figs 3, 4; pl. 35, fig. 1; pl. 36, fig. 1; pI. 37, GPSU 2662 (PI. 4, Figs 6, 7) is a slightly deformed figs 1,2. individual 147 mm in diameter. The penultimate whorl 1894.Acanthoceras rhotomagense var. asiatica lIMBO, p. 177, pl. bears crowded bullate primary ribs with one, rarely two 20, fig. 1. long or short intercalated ribs between. There are an esti 1897. Acanthoceras Newboldi n.sp. (Typische form) KOSSIVIAT, p. 5 mated 42-44 ribs on the outer whorl. Primary ribs arise at (112), pl. 1 (12), figs 2,3; pi. 3 (14), fig. 2. the umbilical seam, and strengthen into prominent bullae 1972. Newboldiceras (Newboldiceras) newboldi (KOSSMAT); perched on the umbilical shoulder, which give rise to rel TROMEL, p. 106 (pars), pl. 40, figs 1-3; non pI. 34, figs 1-3; atively coarse, straight, prorsiradiate ribs, usually singly, pI. 38, figs 1, 2; pI. 39, figs 1-3; pl. 41; ?pl. 42, figs 3, 4. occasionally in pairs, with one or two long or short inter 1988. Mantelliceras cf. mantelli (SOWERBY); lOLIGCEV & al., p. 27. calated ribs between. At the adapical end of the outer 1988. Calycoceras (Cal.) multicostatum (COLLIGNON); lOLIGCEV whorl there are blunt inner ventrolateral tubercles and & al., p. 27. well developed small outer ventrolateral and siphonal 1988. Calycoceras cf. paucinodatum (CRICK); lOLIGCEV & aI., p. clavi, borne on a coarse transverse rib. As size increases, 27. the ribbing coarsens, inner and outer ventrolateral tuber 1990. Calycoceras (Newboldiceras) asiaticum asiaticum (lIMBO, cles persist, but the siphonal clavi are lost, leaving a broad, 1894); WRIGHf & KENNEDY, p. 239, pl. 58, fig. 1; pl. 64, figs feebly convex venter. 1,2; pl. 65, figs 1-3,5,7; pl. 72, fig. 3; text-figs 87a-c; 88f; 97; 98 (with full synonymy). DISCUSSION: Calycoceras (Newboldiceras) astatlcum 1994. Calycoceras (Newboldiceras) asiaticum asiaticum (lIMBO, asiaticum and C. (N.) asiaticum spinosum (KOSSMAT, 1894); KENNEDY, p. 227, pl. 7, figs 7,8; pi. 8, figs 1,2. 1897) are closely allied, and linked by occasional passage 1997. Calycoceras (Newboldiceras) asiaticum asiaticum (lIMBO); forms. They differ in the much more robust tuberculation WILMSEN, pl. 16, fig. 1. of the latter, well-seen in the present collection (compare PI. 4, Figs 1-5 and PI. 4, Figs 6, 7). TYPE: The holotype by monotypy is the original of JIMBO (1894, pI. 20, fig. 1), no. 1-105 in the Collections of the OCCURRENCE: Middle and upper Middle Geological Institute, Tokyo University, from the Middle Cenomanian, rarer in the lower Upper Cenomanian. Cenomanian Trigonia Sandstone of the lkushumbets, Southern England, northern and southern France, Spain, Hokkaido, Japan. Romania, Bulgaria, Tunisia, Zululand (South Africa), Madagascar, south India, and Japan, with possible MATERIAL: GPSU 2630, 2649, 2662, 3950. records from Poland, Israel, and China. 376 WJ. KENNEDY & N. JOLKJ(EV

Calycoceras (Newboldiceras) asiaticum spinosum mm, giving the characteristic ventral profile of adults of (KoSSMAT,1897) the species, with persistent inner and outer ventrolateral (PI. 3, Figs 7,10,11; PI. 4, Figs 1-5; PI. 5, Figs 6-11) clavi, and an evenly arched venter between (KOSSMAT, 1897, PI. 2 (14), Fig. 2b). 1897. Acanthoceras Newboldi var. spinosa KOSSMAT, p. 7 (114), pI. This, and other small specimens appear to be micro 2 (13), figs 2, 3; pI. 3 (14), fig. 1. conchs. In other specimens (GPSU 2572, 2574), the 1972. Newboldiceras (Newboldiceras) spinosum (KOSSMAT); siphonal tubercles persist to diameters of up to 75 mm; THOMEL, p. 110, pI. 42, figs 1, 2; pI. 46, figs 1-3. these may be incomplete macroconchs. 1988.Acanthoceras rhotomagense (BRONGNIART); JOLKICEV & al., p. 27. DISCUSSION: Crowded ribbing, strong, persistent 1988. Calycoceras (Cai.) boulei (COLLIGNON); JOLIaCEV & ai., p. 27. umbilical, inner and outer ventrolateral tubercles, and the 1988. Ellcalycoceras sp.; JOLKICEV & al., p. 27. distinctive arched venter without siphonal clavi make even 1988. Mantelliceras (Promantelliceras) picteti cf. densecostata fragments of this species readily identifiable (PI. 4, Figs 1- THOMEL; JOLKICEV & ai., p. 27. 5). See WRIGHT & KENNEDY (1987, p. 249) for further 1990. Calycoceras (Newboldiceras) asiaticum spinosum (KOSSMAT, observations on the subspecies. 1897); WRIGlIT & KENNEDY, p. 249, pI. 64, fig. 3; pI. 65, figs 4,6; pI. 66, figs 3, 4; pI. 69, fig. 1; pI. 70, fig. 3; text-figs 87a OCCURRENCE: Upper Middle and lower Upper e; 88d, 1; 99; 100; 102; 107k (with full synonymy). Cenomanian, southern England, France, Spain, Romania, 1994. Calycoeeras (Newboldieeras) asiaticum spinosum (KOSSMAT, Bulgaria (the present record), Tunisia, Zululand (South 1897); KENNEDY, p. 228, pI. 9, figs 1-3, 7-9; pI. 10, figs 12, 13. Africa), Madagascar, South India, Tibet(?), Japan and 1997. Calycoceras (Newboldiceras) asiaticumspinosum (KOSSlvIAT); California. WILMSEN, pI. 12, fig. 1; pI. 20, fig. 1.

TYPES: Lectotype, designated by WRIGHT & KENNEDY Calycoceras (Newboldiceras) planecostatum (KOSSMAT, (1990) is the original of KOSSMAT (1897, pI. 2 (13), fig. 2), 1897) from the Utatur Group of Odium, South India. (PI. 3, Figs 2-6, 8, 9) Paralectotypes are the originals of KOSSMAT'S [pI. 2 (13), fig. 3, and pI. 3 (14), fig. 1]. The paralectotype in 1897.Acanthoeeras Newboldi var. planecostata KOSSMAT, p. STOLICZKA (1865, pI. 35, fig. 2) may not belong to this sub 9(116), pI. 2(13), fig. 1. species. 1988. Acanthoceras (Ae.) rhotomagense var. subflexuosum (SPATH); JOLKICEV & al., p. 27. MATERIAL: 11 specimens, GPSU 2572, 2574, 2576, 1990. Calycoeeras (Newboldiceras) planecostatum (KOSSMAT, 2577,2625,2632,2643,2652,3943,3953,6022. 1897); WRIGHT & KENNEDY, p. 252, pI. 61, figs 2, 3; pI. 67, figs 1-4; text-figs. 101e-e (with full synonymy). DESCRIPTION: All specimens are crushed to varying 1994. Calycoceras (Newboldiceras) planecostatum (KOSSMAT, degrees. Coiling is moderately evolute, with an umbilicus 1897); KENNEDY, p. 228, pI. io, figs 2, 3. that ranges from 28% at a diameter of 73 mm in GPSU 1994. Calycoceras (Newboldiceras) planecostatum (KOSSMAT, 2574 (PI. 5, Figs 6-8) to 31 % in GPSU 2572 (PI. 5, Figs 9- 1897); KENNEDY & JUIGNET, p. 50, text-figs Ie; 18a-e; 19a-e. 11). The umbilicus of moderate depth, with a flattened 1996. Calyeoceras (Newboldiceras) planecostatum (KOSSMAT, wall and narrowly rounded umbilical shoulder. The whorl 1897); KENNEDY & ai., p. 314, pI. 40, fig. 3. section is trapezoidal in intercostal section, and polygonal 1998. Calycoceras (Newboldiceras) planecostatum (KOSSMAT, in costal section. Strong primary ribs arise at the umbilical 1897); KAPLAN & ai., p. 158, pI. 26, figs 9-11. seam, and develop into strong, subspinose to spinose bul lae, perched on the umbilical shoulder. These give rise to TYPE: Lectotype, by the subsequent designation of strong prorsiradiate primary ribs, either singly, or in pairs, WRIGHT & KENNEDY (1990, p. 252), is no. 14842 in the the second rib varying from strongly to weakly attached to Collections of the Geological Survey of India, the original the bulla. One or two long or short rihs intercalate of KOSSMAT [1897, pI. 2(13), fig. 1] (WRIGHT & KENNEDY between the bullate ribs. All ribs bear well-developed 1990, text-fig. 100c-e), from the Utatur Group of Odium, inner and outer ventrolateral clavi, the latter linked across south India. the venter by strong rounded ribs. There are an estimated 46 ribs per whorl in GPSU 2574, and 48 in GPSU 2572. MATERIAL: GPSU 296, 2578, 2623, 2632, 2646, 3952. Siphonal clave are present at the smallest diameter seen (GPSU 6022; PI. 3, Fig. 7). In GPSU 2643 (PI. 4, Figs 4, DESCRIPTION: Fragmentary juveniles (PI. 3, Figs 2-6, 5), they have already disappeared by a whorl height of 18 8) have whorl heights of up to an estimated 25 mm. The MIDDLE CENOMANIAN AMMONITES FROM BULGARIA 377 coiling appears to have been moderately evolute, with ?1846.Ammonites cenomanensis D'ARcHIAc, pp. 62,78 (nomen convergent, feebly convex flanks, narrowly rounded ven nudum). trolateral shoulders, and a broad, feebly convex venter. 1863. Ammonites cenomanensis D'ARcHIAc; PICIET, p. 28 iJlars], Prorsiradiate primary ribs arise either singly or in pairs pi. 3, fig. 2; inner whorls of fig. 1, pi. 4. from narrow umbilical bullae, with one or two intercalat 1969. Calycoceras aff. choffati (KOSSMAT); KENNEDY, pi. 18, fig. 4. ed ribs between, both short and long, and in some cases 1971. Calycoceras aff. newboldi ankomakaensis COLLIGNON; extending to the umbilical shoulder as mere striae (PI. 3, KENNEDY, p. 76, pi. 42, fig. 4. Fig. 2). Some juveniles have weak inner ventrolateral 1971. Calycoceras aff. choffati (KOSSMAT); KENNEDY, p. 75iJ1ars], tubercles (PI. 3, Fig. 2), but they are absent in others (PI. pi. 38, fig. 5; pi. 39, fig. 1 only. 3, Figs 3, 4, 8). There are small but well-developed outer 1972. Acanthoceras (Acanthoceras) aff. cenomanense (D'ARcHIAc); ventrolateral clavi on all ribs; and these are linked across TROMEL, p. 141, pi. 68, figs 1, 2. the broad venter by strong transverse to feebly convex 1972.Acanthoceras (Acanthoceras) aff. cenomanense densecosta ribs. tum TROMEL, p. 143, Pi. 44, figs 3-5 (proposed as a variety: GPSU 2623 (PI. 3, Fig. 9) is part of the adult phrag not available under the Rules). mocone of a macroconch that is referred to the species. ?1972. Calycoceras (Gentoniceras) subwiestii (SPATH); TROMEL, p. The inner flanks are broadly convex, the outer flanks flat 76 iJlars], pi. 25, figs 1, 2. tened and convergent, the ventrolateral shoulders broad 1983. Calycoceras cmomanense (D'ARcHIAC, 1846); MOREAU & ly rounded, the venter broad, and feebly convex. Five well al., p. 324, Fig. 6. developed umbilical bullae are present on the 120° frag 1990. Calycoceras (Proeucalycoceras) picteti WRIGHT & KENNEDY, ment. They give rise to single ribs, or a second rib may be p. 264, pi. 54, fig. 2; pi. 64, fig. 4; i. 68, figs 1, 2; pi. 74, fig. 3; weakly attached to a bulla, with one or two long interca pi. 75, figs 2-6; text-figs 110d, g; 113; 114; 116a-d; 117; 119a lated ribs between the bullate ones. They are straight and c. prorsiradiate on the inner flank, strengthening, broaden 1994. Calycoceras (Proeucalycoceras)picteti JUIGNET & KENNEDY, ing and flexing back and feebly convex at mid-flank, and p. 471, figs 1a; 2a, b; 3a-d; 4a-d; 5a-d; 13a. feebly concave on the outer flank, where they link to a 1998. Calycoceras (Proeucalycoceras) picteti WRIGHT & KENNEDY, very effaced inner ventrolateral tubercle. A broad, flat 1990; KAPLAN & aI., p. 164, pi. 51, figs 1-8; pi. 52, figs 1-6. tened rib links to blunt outer ventrolateral clavi, and 1998. Calycoceras picteti WRIGHT & KENNEDY, 1990; LEHMANN, broad, ribbon-like ribs extend across the venter, separat p. 26, pi. 3, fig. 6. ed by narrower interspaces. TYPES: The holotype is OUM K22586 from the upper DISCUSSION: Early loss of all but outer ventrolateral Middle Cenomanian fauna of the Chalk Basement at tubercles, crowded flattened ribs that are wider than the Storridge Hill, Chardstock, Devon; there are numerous interspaces on the venter of the larger fragment, are char paratypes, listed in WRIGHT & KENNEDY (1990, p. 264). acters that compare well with those of the type material and numerous topotype specimens we have studied. MATERIAL: GPSU 3944.

OCCURRENCE: Upper Middle and lower Upper DESCRIPTION: GPSU 3944 (PI. 2, Figs 3, 4) is a later Cenomanian, south India, Madagascar, Zululand (South ally crushed composite mould with a maximum preserved Africa), James Ross Island, Antarctica, Iran, Bulgaria (the diameter of 63.6 mm. Coiling is moderately evolute, the present records), France, Germany and southern whorl section compressed, as a result of post-mortem England. crushing. The inner flanks are feebly convex, the outer flanks convergent, the ventrolateral shoulders narrowly rounded, the venter broad, and flattened. Subgenus Proeucalycoceras THOMEL, 1972 Crowded primary ribs arise from small, elongate bul lae either singly or in pairs, and additional ribs intercalate TYPE SPECIES: By original designation Calycoceras below mid-flank. The ribs are straight and prorsiradiate, (Eucalycoceras) besairiei COLLIGNON, 1937 (p. 37(13), pI. broadening progressively across the flanks and terminating 3, figs 1-4; pI. 8, fig. 5), from the Cenomanian of in narrow outer ventrolateral clavi. These are linked across Madagascar. the venter by low, broad flattened ribs, separated by nar rower interspaces. All bear feeble siphonal clavi.

Calycoceras (?roeucalycoceras) picteti WRIGHT & DISCUSSION: Coiling, flattened flanks, lack of an inner KENNEDY, 1990 ventrolateral tubercle, and the flat venter with broad, flat (PI. 2, Figs 3, 4) tened ribbon-like ribs bearing feeble siphonal clavi sepa- 378 WJ. KENNEDY & N. JOLKICEV

rate this specimen from all others in the collection. In long. Coiling of the spire is very involute, with a tiny, deep these respects it finds a match in juvenile Calycoceras umbilicus. Coarse crowded primary ribs arise at the (Proeucalycoceras) picteti from the upper Middle umbilical seam and are straight and prorsiradiate on the Cenomanian and lower Upper Cenomanian of southern inner to mid flank region. They increase by bi- and trifur England and Sarthe, France, as figured by WRIGHT & cation on the outer flank, where additional ribs coarsen KENNEDY (1987, pI. 75, figs 2-6; text-fig. 119A-C), such dif markedly on the adapical end of the body chamber and ferences as are apparent being due to the dissimilar are linked across the venter by groups of two or three ribs, preservation of the material. with additional ribs intercalated between. This coarsening continues on the remainder of the shaft and adapical part OCCURRENCE: Upper Middle and lower Upper of the hook, with coarse distant flank ribs that strengthen Cenomanian. C. picteti is common in the upper Middle into coarse to weak ventrolateral bullae. These are linked Cenomanian in Sarthe, with numerous specimens from over the venter by groups of up to four ribs, with addi the Sables du Perche, as well as the Craie de Rouen. It tional ribs intercalated between (PI. 6, Figs 9-16). The bul characterizes a similar stratigraphic level elsewhere as in lae weaken on the adapertural part of the hook, and there Charente-Maritime, Haute Normandie and Provence in is a short, completely smooth section of shell immediate France, the Munster Basin, Westphalia, Germany, ly preceding the adult aperture. This is marked by a strong Bulgaria (the present records), Dorset, Somerset and constriction, markedly prorsiradiate on the flanks, and Devon in southern England. It ranges into the lower crossing the venter in a broad convexity (PI. 6, Figs 8, 9, Upper Cenomanian in Dorset and Devon. 11,14,16). One specimen referred to the species (GPSU 2607: PI. 6, Fig. 5) has strong close-spaced flank ribs, but fails to develop ventrolateral tubercles. Suborder Ancyloceratina WIEDMANN, 1966 Specimens regarded as micro conchs have relatively Superfamily Scaphitaceae GILL, 1871 slender body chambers, the umbilical wall of the body Family Scaphitidae GILL, 1871 chamber concave, and barely concealing the umbilicus of Subfamily Scaphitinae GILL, 1871 the spire (PI. 6, Figs 3, 4,9, 10, 14, 15). None of the mate Genus Scaphites PARKINSON, 1811 rial can be assigned to the macro conch of the species with confidence, because of defective preservation. TYPE SPECIES: Scaphites equalis J. SOWERBY, 1813, p. 53, pI. 18, figs 1-3, by original designation by MEEK, 1876, DISCUSSION: Scaphites equalis is readily separated from p.413. S. obliquus J. SOWERBY, 1813 (see revision in WRIGHT & KENNEDY (1996, p. 390, pl.114, figs 7-20; pI. 115, figs 1-17; pI. 116, fig. 6; text-figs 150F, K). The shell of S. obliquus is Scaphites equalis J. SOWERBY, 1813 shorter, the ribs on the shaft are finer, denser, and (PI. 6, Figs 1-16) markedly prorsiradiate; the shaft is constricted adorally of the mid-shaft lateral swelling; there are no, or only incipi 1813. Scaphites equalis J. SOWERBY, 1813, p. 53, pI. 18, figs 1-3. ent ventrolateral tubercles. Scaphites (Scaphites) bituber 1988. Scaphites equalis J. SOWERBY; JouacEv & al., p. 27. culatus SANTAMARIA-ZABALA, 1992 (p. 247, pI. 2, fig. 9) is 1996. Scaphites equalis J. SOWERBY, 1813; WRIGHr & KENNEDY, no more than an Upper Cenomanian intraspecific variant p. 394, pI. 116, figs 1-5,7-11; pI. 117, figs 1-11; pI. 118, figs of S. equalis. 1-13; text-fig. 129B, C (with full synonymy). 1996. Scaphites (Scaphites) bituberculatus SANTAMARIA-ZABALA, OCCURRENCE: Middle and Upper Cenomanian. 1992; WILMSEN & WIESE, p. 95, pI. 1, fig. 1. Southern England, France, Germany, Poland, the Czech 1998. Scaphites equalis J. SOWERBY, 1813; KAPLAN & al., p. 224. Republic, Ukraine, Bulgaria (the present records), the 1998. Scaphites equalis J. SOWERBY, 1813; LEHMANN, p. 41. Caucasus, Turkmenia, Iran, Algeria, Tunisia, and south India. TYPE: The holotype, by the subsequent designation of WRJGHT & KENNEDY (1996, pI. 395), is BMNH 43986, the original of J. SOWERBY (1813, pI. 18, figs 1-3). Acknowledgements:

MATERIAL: 14 specimens, GPSU 2580, 2604-9, 2614, The technical support of the staff of the Department of Earth 2615,2634,2636,2637,2640,2650. Sciences, Oxford, and the Geological Collections, Oxford University Museum of Natural History, is gratefully acknowl DESCRIPTION: All specimens are crushed to varying edged. Francis AMEDRO and Herbert SUMMESBERGER, journal degrees. Complete specimens vary from 22.3 to 27 mm referees, are thanked for their thorough reviews. MIDDLE CENOMANIAN AMMONITES FROM BULGARIA 379

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Manuscript submitted: 10th October 2003 Revised version accepted: 20th April 2004 PLATES 1-6 ACTA GEOLOGICA POLONICA, VOL. 54 WJ. KENNEDY & N. JOLKICEV, PL. 1

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