Annales Societatis Geologorum Poloniae (2013), vol. 83: 329–342.

EARLY AMMONITES FROM THE ŠTRAMBERK LIMESTONE IN THE KOTOUÈ QUARRY (OUTER WESTERN CARPATHIANS, CZECH REPUB LIC)

Zdenìk VAŠÍÈEK1 & Petr SKUPIEN2

1 Insti tute of Geonics, Academy of Sciences of the Czech Repub lic, Studentská 1768, 708 00 Ostrava-Poruba, Czech Re pub lic; e-mail: [email protected] 2 In sti tute of Geo log i cal En gi neer ing, VŠB-Tech ni cal Uni ver sity, 17. listopadu 15, 708 33 Ostrava-Poruba, Czech Repub lic; e-mail [email protected]

Vašíèek, Z. & Skupien, P., 2013. Early Berriasian ammonites from the Štramberk Limestone in the Kotouè Quarry (Outer West ern Carpathians, Czech Re pub lic). Annales Societatis Geologorum Poloniae, 83: 329–342.

Ab stract: Over 100 ammonites were col lected from a block of bed ded shal low-wa ter Štramberk Lime stone at a new lo ca tion at the Kotouè Quarry, near Štramberk (Silesian Unit, Czech Re public). Berriaselline neocomitids, in clud ing jacobi, B. oppeli, Tirnovella allobrogensis, T. cf. allobrogensis, Delphinella consanguinea, Pseudosubplanites cf. grandis and Malbosiceras cf. asper, are the most abundant spe cies in this ammonite as so ci a tion. The olcostephanitid, Spiticeras blancheti, is rare. The index spe cies for the Early Berriasian, B. jacobi, oc curs in large num bers. On the ba sis of the pres ent study, the up per bound ary of the stratigraphic range of the Štramberk Limestone in the type area is dated as early Berriasian (ammonite Berriasella jacobi Zone). Ammonites with ages youn ger than early Berriasian have never been found in the Štramberk area. The depo si tion of the Štramberk Lime stone in the Štramberk area came to an end dur ing that pe riod.

Key words: Ammonites, Berriasian, Štramberk area, Outer Western Carpathians.

Manu script re ceived 7 November 2013, ac cepted 27 December 2013

IN TRO DUC TION The fos sil-rich Štramberk Lime stone (pos si bly pri mar ily Štramberk Limestone forming Cas tle Hill was accessed at to early Berriasian in age) in the Outer West ern the Cas tle Hill Quarry at a strati graphic level, which cor re - Carpathians of Moravia (Czech Re public) has at tracted the sponded to the upper part of the lower Tithonian (Houša, atten tion of palaeontologists since the first half of the nine- 1975, 1990). The ammonite species from this quarry exhib - teenth century. The first author to mention limestones in this its a uni form lower Tithonian charac ter, docu menting this area was Oyenhausen (1822). The term Štramberk Limestone strati graphic in ter val. Ma te rial in the Hohenegger col lec tion for the light-grey to whit ish-grey limestones was intro duced came from other sites, as well. A small por tion of this col - by Hohenegger (1849). From approx i mately 1910 onwards, lection was re cov ered from the Mu nic ipal Quarry (Obecní the quarry at Kotouè Hill has been the main source of mate - lom, Gemeinde Steinbruch) at Štramberk, which was still rial for palaeontological studies. A review of geolog i cal and rather small in size at the time (opened in 1820). Part of Ho- palaeontological papers, address ing the Štramberk Lime- henegger’s mate rial was also ob tained from sites, compris - stone and as so ci ated deeper-wa ter Lower Cre ta ceous de pos - ing the so-called “exotic boulders” of Štramberk Limestone, its from Štramberk and the surround ing area, was presented and taken from differ ent local i ties in the Podbeskydská pa- by Vašíèek and Skupien (2004, 2005). horkatina Highland. These sites contain ing “exotic boul- On the basis of the Štramberk Limestone, the Tithonian ders” and the Munic i pal Quarry at Štramberk yielded spe - Stage was defined (see Oppel, 1865; von Zittel, 1870) as the cies that led to doubts among later authors over the unifor - up permost unit of the Juras sic in the Tethyan Realm. Both mity of the Štramberk Limestone ammonite fauna. of these classic works were based on mate rial from the Ho- The younger Remeš collec tion from the Štramberk Li- henegger collec tion, which was col lected almost exclusi- mestone shows a somewhat differ ent faunal spectrum, com- vely from the type area in the Cas tle Hill (Zámecký vrch) pared to the older col lec tions. This differ ence is ex plained Quarry (Schlossberg Steinbruch), where mining started in by the fact that the Remeš collec tion was ob tained later, be - 1780. There fore, this quarry was se lected as the type area of tween 1870 (when it was ini ti ated by Remeš’s father) and the Štramberk Limestone (Houša, 1968). The body of the about 1930, and thus contains samples from quarries that 330 Z. VAŠÍÈEK & P. SKUPIEN

Fig. 1. Tectonic map of the Outer Western Carpathian area in the Czech Re pub lic were ac tive at that time, as well as pur chases from quarry work ers. The Castle Hill Quarry had been almost com- pletely abandoned in the end of 19th century, and the main sources of Štramberk Limestone were the quarry on Kotouè Hill and the Munic i pal Quarry (Remeš, 1899, 1904). How- ever, the newly opened quarry on Kotouè Hill was the main Fig. 2. Lo ca tion map. A. Top o graphic sit u a tion of bod ies of source of fos sils. Here, the Štramberk Limestone was ex ten - Štramberk Lime stone in the vicin ity of Štramberk. B. Loca tion of sively exposed and extracted during the 1900s, throughout the sec tion (marked with an aster isk) on the tran sition be tween nearly its en tire stratigraphic range be tween the early Titho- lev els 4 (IV) and 3 (III) at Kotouè Quarry nian and early Berriasian. An early Tithonian age is docu - mented by an ammonite assem blage (e.g., Blaschke, 1911) from the former Gutmann Quarry, opened in the oldest, stone. It thus al lows re defi ni tion of the up per stratigraphic southwesterly portion of the Homole body of the Kotouè limit of this clas sic unit in its type area as lower Berriasian, Hill complex in 1881. even though most of other (old) collec tions indi cate a Titho- During a visit to the Uni versity of Tübingen, within the nian age. framework of the Humboldt Fellow ships in Febru ary 1992, one of us (Z. V.) vis ited the Bayerische Staatssammlung für Paläontologie und historische Geologie (Univer sity of Mu - GEO LOG I CAL SET TING nich). A clas sic col lection of ammonites from the Štramberk Limestone, monographed by von Zittel (1868), is housed The Štramberk Limestone in its clas sic form oc curs at there. At the time, the German col leagues, H. Immel and G. several quarries (the Kotouè, Munic i pal, Horní skalka and Schairer, had plas ter casts made of ten orig inal spec imens Cas tle Hill quarries) in the imme di ate vi cin ity of the town of from von Zittel’s col lec tion, rep re sent ing rather small shells. Štramberk (Figs 1, 2) in the form of larger and smaller car- Some of these casts have now been of assis tance in our iden- bonate blocks, smaller blocks, breccias and conglom er ates. ti fi ca tion. The Štramberk Limestone also occurs also within the Creta - New discov er ies of ammonite fauna in a block of Štram- ceous flysch depos its of the Silesian Unit in the Outer West- berk Limestone with conspic uous bedding, from the Kotouè ern Carpathians. Quarry near Štramberk, form the sub ject of the present pa per. The ge ology of the Štramberk area and the nature of the This study provides a system atic as sess ment of the ammoni- megablocks are the subject of contro versy. Houša (1990) tes and their stratigraphic inter preta tion. The ammonite fauna interpreted the carbon ate blocks as tectonic klippen, sepa - is in dic a tive of the lower Berriasian Beriasella jacobi Zone rated from the carbon ate plat form during the course of Sile- and is the youngest, so far reported from the Štramberk Lime- sian Nappe overthrust. In his inter preta tion, the asso ci ated BERRIASIAN AMMONITES FROM THE ŠTRAMBERK LIME STONE 331

deeper-wa ter Early Cre ta ceous de pos its rep re sent ma te rial which filled fissures or cavi ties of differ ent or igin or cover orig i nal sur faces of lime stone bod ies. According to Eliáš (see Eliáš and Stráník, 1963; Eliáš, 1970; Eliáš and Eliášová, 1986), the limestones are embed - ded in base-of-slope conglom er ates and slump bodies within the Cre taceous part of the Tìšín–Hradištì Formation (now Hradištì Forma tion – see Eliáš et al., 2003), consti tut ing an ex treme de vel op ment of the Chlebovice Con glom er ate. This ac cu mu la tion de vel oped be tween the Tithonian and the Turonian (see also Vašíèek and Skupien, 2004; Svobodová et al., 2011). Ac cord ing to Picha et al. (2006), the Štramberk carbon - ate platform, rimmed by coral reefs, was the source of clas- tics and large fragments. Gravi ta tional slides and turbidite currents transported both small and large blocks and frag- ments from the rim (edge) of the platform, as far out as the foot of the adja cent basin. However, during the course of later nappe tec tonic transport during the Neogene, large tec- tonic pieces of the car bon ate platform were sep a rated from the softer, less com petent rocks, sit u ated on the slopes of the platform. The result is a melange, in which larger blocks from the car bonate platform show the char acter istics of kli- ppen. Smaller blocks and debris corre spond to clastic sed i- ments at the foot of the platform. These devel oped in the Early Cre taceous and the early part of Late Cre taceous, in par tic u lar. The Štramberk Limestone is whitish-grey in colour and was depos ited in differ ent settings across the carbon ate plat- Fig. 3. Photo graph of out crop showing the num ber of lay ers form and reef complex, includ ing reefal buildups. Biogenic and sparitic limestones (e.g., biosparites, intrasparites and intrabiosparites) are attrib uted to the core of the reef. The limestone found in some inter vals is very coarse to grav elly, The in ter vals be tween grav ity flows are rep re sented by as it is formed of whole rudist shells (Heterodiceras) and hemipelagic sed iments. The grad ual lateral and ver tical minute coral colo nies. In contrast, the limestone in some transition of the block accu mu la tions into the ambi ent sedi - other inter vals is very fine and micritic. How ever, the com- ment con tra dicts the clas sic the ory re garding the tec tonic mon est type is finely biodetrital limestone. klippen in the Silesian Unit. Tra di tionally, the Štramberk Limestone was be lieved to be Tithonian in age, which may be the correct age for the main stage of reef devel op ment. However, the calpionellids DESCRIP TION OF AMMONITE-BEARING and ammonites pres ent in di cate that the Štramberk Limesto- LIMESTONES ne orig i nated dur ing the lat est Kimmeridgian–ear li est Berria- sian (e.g., Houša, 1990; Eliáš and Vašíèek, 1995; Houša and In the upper most portion of the exit road, running from Vašíèek, 2005). The known calpionellid zonation (Houša and level 4 to level 3 at Kotouè Quarry (approximately10 m be- Vašíèek, 2005) is in dic a tive of the lat est early Tithonian, the fore end; GPS coodinates: 49°35'1.329"N, 18°6'52.689"E; en tire late Tithonian and the ear li est Berriasian in the lime- Fig. 2), there is a block of Štramberk Limestone with con- stone bodies exposed in the Štramberk area. spic u ous bedding (ori enta tion of the strata 280/30; Fig. 3), The Štramberk Lime stone rep re sents sed i ments formed which is found only rarely at other sites in the quarry. Nine on a carbon ate platform during the Late Juras sic and earli est layers (in the section marked 0 to 8) have been identi fied. Creta ceous along the north ern Tethyan margin. This carbo- Their thickness ranges from 20 to 90 cm (see Figs 3, 4). nate platform was affected by block tecton ics during the lat- These lay ers are disrupted by sys tems of fissures, per pen - est Ju ras sic–ear li est Cre ta ceous (e.g., Eliáš and Eliášová, dic u lar to the bed ding. The fis sures are filled with cal cite 1986). The block accu mu la tions form part of the conti nen - sparitic ce ment. Oc ca sion ally, the char ac ter is tics of pres - tal-rise facies of the Baška devel op ment, depos ited in the sure dis so lu tion (sty lo lites) ap pear in these lay ers, which flysch trough of the (hypo thet i cal) Baška Cordil lera. This emphasise the bound aries be tween the lay ers identi fied. up per most Ju ras sic to Up per Cre ta ceous sed i men tary suc- Descrip tion of the layers identi fied from the bottom up cession includes slumps, slides, olistholites and occa sional (the thickness of layers given is the maxi mum value): turbidites, which were fed from the Upper Juras sic to the Layer 0 has a maxi mum thickness of 25 cm. The low - Coniacian carbon ate plat form on the Baška Cordil lera and est-ly ing layer was exposed by exca va tion at the bottom be- its slopes, in cluding the Tithonian–Berriasian reef com plex. low the quarry face. The limestone is de tri tal throughout the 332 Z. VAŠÍÈEK & P. SKUPIEN

Fig. 4. Lithological col umn and range chart of ammonites of the sec tion marked in Fig. 2 BERRIASIAN AMMONITES FROM THE ŠTRAMBERK LIME STONE 333

en tire thick ness, and more fine-grained lay ers al ter nate with crofossils, and does not change in charac ter throughout its coarse-grained ones. In the mid dle part, recrystallisation ap - thick ness. The up per most layer out crop ping is de lim ited at pears. Only a sin gle in de ter minate ammonite frag ment was the top by a marked fault and the bottom of level 3. found. Layer 1 (thickness of 32 cm) is composed of four beds: (1a) 8 cm – finely de tri tal limestone with small cor als; (1b) MA TE RIAL AND AMMONITE 17 cm – mainly coarsely detri tal limestone, contain ing cor - PRES ER VA TION als, brachio pods and sporadic ammonites. In the lower part, recrystallisation is ap par ent; (1c) 5 cm – coarsely de trital, The newly collected mate rial, more than 100 ammoni- recrystallised and in its up per most part (2 cm) finely de tri tal tes, consists partic u larly of fragmentarily preserved, ribbed and even muddy. speci mens of small size. Most frequently, one-third of the Layer 2 (thickness of 37.5 cm) is com posed of three ul timate whorls is well pre served, whereas the re main ing beds: (2a) 18 cm – finely detri tal, contain ing bi valve debris; parts of the spec imens are corroded to vary ing degrees, es- (2b) 6 cm – mainly coarsely de tri tal limestone, con tain ing pecially on the venter. In addi tion, the outer surfaces are corals, bivalves, brachio pods and ammonites, and recrystal- usually matrix covered. The con tin u ation of a more mature lisation is appar ent; (2c) 13.5 cm – finely detri tal and muddy part of the spec imens is in some cases pre served only as its in places, contain ing fragments of corals, bivalves and am- outer outline. The preserved portion usually retains the monites. recrystallised calcitic origi nal shell. The thickness of the Layer 3 (thickness of 59 cm) is composed of four beds: original shell is occa sion ally encrusted by micro bial peri- (3a) 6 cm – finely detri tal, contain ing bivalves, ammonites phyton or by mineral crusts. In some cases, the origi nal shell and small cor als; (3b) 18 cm – rather coarsely de tri tal, with came loose from its inter nal filling dur ing ex trac tion; thus corals, algae, sponges, serpulids, brachio pods and ammoni- rather short portions are im perfectly preserved as in ner tes. Sub stan tial recrystallisation is ap par ent; (3c) 19 cm – moulds. In these, the relicts of suture lines are poorly pre - coarsely de tri tal lime stone, con tain ing ammonites, bi valves served to a small ex tent. In some cases, shells or parts of and brachio pods, follows; (3d) 16 cm – finely detri tal and their whorls were not filled in. In the exposed cavi ties of muddy in places with frag ments of small corals and, sporad - such whorls, tiny cal cite crystals are usually vis ible. The ically, ammonites. Throughout the entire thickness, a total parts of whorls that are not filled are usually broken, as seen of 70 ammonites were found, all largely fragmen tary. both on the surfaces and in cross-sec tions of some shells. Layer 4 (thickness of 18.5 cm) is com posed of three Over all, the major ity of shells ex hibit well pre served orig i- beds: (4a) 6 cm – finely detri tal with bivalves and small cor- nal non-deformed cross-sections or whorl shapes, compa ra - als; (4b) 9 cm – coarsely detri tal limestone, contain ing corals, ble to those from the clas sic French lo cali ties (Toucas, bi valves, gas tro pods, brachi o pods and ammonites (ap prox i - 1890; Mazenot, 1939) and pyritized shells from Tuni sia mately 20 spec i mens). Sub stan tial lime stone recrystallisa- (Arnould-Saget, 1953). Most of the shells stud ied ex pose tion is appar ent; (4c) 3.5 cm – finely detri tal and even muddy only one side. Thus, the whorl width can not be measured di - in places, with frag ments of small corals, brachio pods, bi- rectly, but it can, in some cases, be es timated rel atively ac- valves, sponges and, sporad i cally, ammonoid aptychi. curately from their half-width. The um bili cus and thus, the Layer 5 (thickness of 38 cm) is com posed of three beds: youn ger whorls are usu ally filled with sed i ment, simi lar to (5a) 8 cm – finely detri tal, contain ing bivalves, brachio pods the interspaces be tween the ribs. Prep ara tion of the shells, and small cor als; (5b) 21 cm – mainly coarsely de tri tal with us ing a me chan i cal prep a ra tion nee dle, was usu ally only corals, brachio pods and ammonites. Recrystallisation of suc cess ful in part. limestone and fos sils is appar ent; (5c) 9 cm – finely detri tal Only approx i mately one-third of the shells collected and muddy in places, showing frag ments of small corals and met the nec es sary re quire ments for iden ti fi ca tion as spe cies, echinoid spines. i.e., provid ing at least one measur able diam e ter of the shell Layer 6 (thickness of 57 cm) is com posed of three beds: with a measur able whorl height, a measur able um bili cus (6a) 25 cm – rather coarsely detri tal, contain ing bivalves, width and a quanti fi able rib density (though usually per serpulids, aptychi and algae. Upwards, the layer changes to half-whorl in the present study), both on the um bil ical area finely de tri tal limestone with small cor als; (6b) 17 cm – de - and venter. tri tal, dis play ing de bris of bi valve shells in par tic u lar, al - In collec tion of the au thors, a spe cial cate gory was rep - though entire shells of brachio pods occur in places; (6c) 15 re sented by three im per fectly pre served ju ve nile shells cm finely detri tal to muddy, and a sin gle ammonite was (with diam e ters of 12 to 13 mm) that differ from all of the found in this part. Cup coral colo nies appear in places. others, owing to the presence of evolute coil ing and a style Layer 7 (thickness of 20 cm) is com posed of three beds: of rib bing with pe ri odic um bil i cal and lat eral tu ber cles. (7a) 6 cm – rather coarsely detri tal, contain ing distin guish - These are fea tures, in dic a tive of the ge nus Spiticeras Uhlig, able corals and hy drozo ans. The growth po sition is ev ident 1903, but owing to the poor state of preser va tion and small in some coral colo nies; (7b) 9 cm – finely detri tal, harbour - size, these char ac ter is tics can not be used for iden ti fi ca tion ing distin guish able corals, aptychi, ammonites and brachio - above the ge neric level. The sin gle spec imen of S. blancheti pods; (7c) 5 cm – finely detri tal to muddy, showing a con - rep re sents an ex cep tion. sid er able num ber of aptychi. The ammonite col lec tion of the present study is de pos- Layer 8 (vari able thickness of up to 90 cm) the lime- ited at the Nový Jièín Regional Museum (cata logue num- stone of this layer is finely organodetrital, lacking any ma- bers PL 4145–4163). 334 Z. VAŠÍÈEK & P. SKUPIEN

TAX ON OMY De scrip tion: The shell is small and evolute, with whorls that are a lit tle wider than high. The great est whorl width oc curs in the um - For the taxo nomic descrip tions of the ammonites from bili cal area. Initially, the rela tively flat flanks of the ulti mate whorl the Kotouè Quarry, the authors have fol lowed the system - are in clined to the venter, into which they pass grad u ally. The venter is quite wide and rounded. atic clas si fi ca tion of Wright et al. (1996) and Klein (2005). On the ul ti mate whorl, a wide and quite deep con stric tion that is In accor dance with the 1999 Code of Zoolog i cal Nomen cla - nearly straight and inclined to the aper ture can be seen (not clearly ture (p. 117), the suffix -oidea is used for superfamilies. The visi ble in the pho to graph). Adapertural of this con stric tion, near classi fi ca tion scheme proposed by Arkadev and Bogdanova the um bi li cus, only four simple, rather strong ribs are pre served. (2012), in which Crimean berriasellines were assigned to From each of these ribs, 3–4 weaker ribs split on the flanks at a the superfamily Olcostephanoidea Pavlov, 1892 (or Olco- short distance. Owing to the im per fect pres er va tion of the um bil i - stephanaceae), fol low ing the rec om men da tions by Kvanta- cal area, the basal part appears to bear weak umbil i cal and stronger liani (1999), on the basis of the devel op ment of the sus- lat eral tu ber cles, although this can not be clearly distin guished. pensive lobe U, is not accepted here. Follow ing the basic The ribs are prorsiradiate and cross the venter with out in ter rup - classification of berriasellines, the au thors ac cept the super- tion, show ing a hint of a chev ron. Mea sure ments: The shell had an es timated maxi mum diam e ter of fam ily . ap proxi mately 27 mm. Due to the state of preser va tion, the umbi li - To keep this contri bu tion as brief as possi ble, the syn- cus can not be mea sured. D = 24.3 mm, H = 7.6 (0.31) and B = c. onymies presented include only the most re cent papers, as 10. 2 (0.42); B/H = 1.34, H/B = 0.745. well as those that are of regional inter est or of im portance to Re marks: The size pa rame ters are equiva lent to mea surements the discus sion. For the size param e ters of ammonite shells, noted by Arnould-Saget (1953). Evolute shells with whorls with a the fol low ing ab bre vi a tions are used: D – shell di am e ter width greater than height and umbil i cal and lat eral tu ber cles on ju - (Dmax – maxi mum preserved diam eter), H – whorl height, venile whorls are charac teris tic of the ge nus Spiticeras. With re- U – umbi li cus width and B – whorl width (where measur - gard to the data fur nished by Djanélidzé (1922, p. 98), the pres ent able). Ra tios of the pa rame ters measured to shell di am e ter au thors be lieve, in con trast to pre vi ous au thors, that, from a no- mencla tor ial point of view, the spec i mens described can not be (H/D, U/D, B/D) or the ratio of whorl width to whorl height clas si fied as a sub spe cies (S. pseudogroteanum blancheti), but in - (B/H) are in di cated in brack ets. With re gard to the fact that stead rep resent a distinct spe cies, S. blancheti. some authors (e.g., Kvantaliani et al., 1999) reported the ra- Dis tri bu tion: From the point of view of modern stra tig ra phy, the tio of whorl height to whorl width (H/B), rather than the po si tion of S. blancheti has not been con sidered in de tail for quite B/H ra tio, the pres ent authors give this as well. Rib density some time. A spec i men il lustrated by von Zittel (1868, pl. 16, fig. 3), near the umbi licus (UR) and ventrolaterally (VR) are given as well as all related spec imens, come from Koòákov near Èeský per half-whorl. Tìšín and, thus, are not from Štramberk. Ac cord ing to lit er a ture data (see syn onymy), the spe cies occurs in France and Tuni sia, as well as in the Silesian Unit in the Czech Re public. With ref er ence to the Class von Zittel, 1884 asso ci ated ammonites from Kotouè Quarry de scribed be low, the Subclass von Zittel, 1884 present species can be assumed to be charac ter is tic of the lower Superfamily PERISPHINCTOIDEA Steinmann, 1890 Berriasian (ammonite Berriasella jacobi Zone). Family OLCOSTEPHANITIDAE Haug, 1910 Oc cur rence: Layer 3, along the exit road from level 4 to level 3, at Subfamily SPITICERATINAE Spath, 1924 Kotouè Quarry.

Genus Spiticeras Uhlig, 1903 Family Salfeld, 1921 Type spe cies: Ammonites spitiensis Blanford, 1864 Subfamily Spath, 1922 (Berriasian, In dia), by sub se quent des ig na tion of Ro man (1938, p. 398) Ge nus Berriasella Uhlig, 1905 Type spe cies: Ammonites privasensis Pictet, 1867 Spiticeras blancheti Djanélidzé, 1922 (Berriasian, south east France), by sub se quent des ig na tion Fig. 5A, B of Roman (1938, p. 324) partim 1868 Ammonites Groteanus Oppel – von Zittel, p. 90, pl. The ge nus Berriasella, simi lar to related genera, has been de- 16, figs 3a–c, 4. [non fig. 1a–c = Spiticeras celsum scribed in con sistently in a wide and vari able man ner since stud ies Oppel; non fig. 2a, b = Spiticeras zitteli Djané- by Mazenot (1939). In subse quent de cades, this situ a tion re mained lidzé]. unchanged. Tavera-Benitez (1985) sub di vided this ge nus into four 1922 Spiticeras psudogroteanum var. Blancheti n. sp. – sub gen era, while Wright et al. (1996) fa voured two sub gen era and Djanélidzé, p. 96, pl. 3, fig. 2a–c; text–fig. 21. Klein (2005), in ac cor dance with Tavera-Benitez, four. Amended 1953 Spiticeras (Spiticeras) pseudogroteanum Djané- char ac ter is tics of the ge nus Berriasella have recently been pro- lidzé var. Blancheti Djanélidzé – Arnould-Saget, p. vided by Arkadev and Bogdanova (2004, 2012). In prin ci ple, the 99, pl. 8, figs 9a–c, 10a–c. con cept of sub gen era is aban doned here. 2005 Spiticeras pseudogroteanum blancheti Djanélidzé In the di ag no sis of Berriasella, it is stated that the genus in- – Klein, p. 58. cludes discoidal shells with faintly arched flanks and a rounded Ma te rial: A single frag ment (PL 4145) of a com posite mould with venter. The umbi li cus is wide and stepped. Rib bing is thin, con- a fairly well-pre served third of the ulti mate whorl and with the sisting of simple and bi fur cated ribs. On the venter, the ribs are in - con tin u a tion of the in ner most part of the ul ti mate whorl, only ob - ter rupted by a fur row that is dis tinct on ju ve nile whorls and less servable in the shell outline. Only a small part of the venter is re - clear or ab sent on the ul ti mate whorl. tained. The en tire um bili cal area is poorly preserved. This is the ge neric con cept of Berriasella employed here. BERRIASIAN AMMONITES FROM THE ŠTRAMBERK LIME STONE 335

Berriasella jacobi Mazenot, 1939 Berriasella jacobi dif fers from other berriasellines by hav ing a Fig. 5C, D, G–I nar rower um bi li cus. Ac cord ing to Arkadev and Bogdanova (2012), this spe cies should also dif fer, in dis play ing a higher de - * 1939 Berriasella Jacobi n. sp. – Mazenot, p. 54, pl. 4, figs gree of whorl con vex ity. 1a–b, 2a–b, 3a–b, 4. [non fig. 5 = Lemencia subjacobi Dis tri bu tion: Berriasella jacobi is an in dex spe cies for the base of Donze and Enay]. the Berriasian. This spe cies occurs over a consid er able area cov er- 2005 Berriasella (Berriasella) jacobi Mazenot – Klein, pp. ing almost the entire Tethys, from the Hima la yas through all of 170–171 [cum syn.]. Med i ter ra nean Eu rope to north west Af rica. How ever, many spec i - 2012 Berriasella jacobi Mazenot – Arkadev and Bogda- mens illus trated in the liter a ture are preserved only fragmentarily, nova, p. 144, pl. 4, figs 4–6 [cum syn.]. usually as quite de formed shells, and thus, their iden ti fi ca tion is Ma te rial: More than 20 fairly well-pre served shells and about 30 some times contro ver sial. In the Carpathian area, speci mens de- of frag ments. The best-preserved spec i mens include PL 4146, with scribed by Wierzbowski and Remane (1992) be long to this cat e - a sedi ment-filled origi nal shell. Its ulti mate whorl, which is sligh- gory. tly de formed, is well pre served. Near the ap er ture, a short por tion Oc cur rence: The first abun dant finds of this spe cies from the of the venter is ex posed. Fur ther more, this group of spec i mens in- Štramberk Limestone came from a small classic region near the cludes a ju ve nile one, PL 4149. The largest spec i men, PL 4151, town of Štramberk. Shells of this spe cies were found in lay ers 1 to has a diam e ter of approx i mately 45 mm. Other speci mens are PL 5, i.e., es sen tially through out the en tire sec tion stud ied. 4150–4152, PL 4147 and PL 4148. De scrip tion: The shells are semi-invo lute, with rather narrow whorls of median height and a quite nar row umbi li cus. The whorls Berriasella oppeli (Kilian, 1889) are slightly vaulted. The greatest whorl width oc curs in the lower Fig. 5E, F, J quar ter of the whorl. From there, the flanks grad ually and con tin u- ously in cline to the venter. The flanks are curved over the low um - partim 1868 Ammonites Calisto d’Orb. – von Zittel, p. 100, pl. bili cal wall and pass into the umbi li cus. The flanks pass to the 20, figs 1a–c [lectotype], 2, 3a–c, 4. [non fig. 5a, b]. rel a tively flat, nar row venter. * 1889 Perisphinctes Oppeli n. sp. – Kilian, p. 662. The rib bing is somewhat vari able. On ju ve nile whorls, ribs in partim 1939 Berriasella Oppeli (Kilian) – Mazenot, p. 49, pl. 3, the lower half of the whorl ap pear to be sparser, whereas in the figs 1a–c, 2a–c, 3a, b, 6a, b, 8a–c. [non figs. 4a, b, subse quent small por tion, they show thick en ing. Af ter this re gion, 5a, b]. the ribs again ap pear to be sparsely ar ranged. The ini tially quite 2005 Berriasella (Berriasella) oppeli (Kilian) – Klein, p. thin ribs grad ually strengthen in ac cord with shell growth, and the 173 [cum syn.]. interspaces between the ribs increase. Overall, the ribs are slightly 2012 Berriasella oppeli (Kilian) – Arkadev and Bogda- S shaped; they be gin along the line of coil ing as simple ribs and nova, p. 147, pl. 5, fig. 3. along the line of coiling, they are curved adaperturally within a Ma te rial: A fa vour ably pre served com posite mould with a near- short sec tion. At approx i mately mid-whorl height, almost all ribs complete ulti mate whorl and half of the pre ced ing whorl, stored in bi fur cate nar rowly. On the vent ers of all ju ve niles and some ap par - the col lec tions of the Silesian Museum at Opava (spec i men 17/80- ently adult spec i mens, ribs are in ter rupted. In some shells with a 1980-3181) and a frag ment (less than half a whorl) with a recrys- rela tively large diam e ter, the ven tral fur row is missing, whereas in tallised origi nal shell (PL 4153). oth ers, the ribs are weak ened on the venter. Descrip tion: The shells are semi-evolute. The ulti mate whorl is Measure ments (in mm) and ratios are in Table 1. not high, its height be ing somewhat greater than its width. The Overall, it ap pears that juve nile shells exhibit some what higher whorls are slightly vaulted and in the up per three-quar ters of the whorls than more adult shells. shell height, they incline towards the venter. The greatest width Re marks: The present speci mens are microconchs. These shells oc curs at ap prox i mately mid-height. The venter, which is not are somewhat variable in terms of the param e ters measured, which wide, is mod er ately arched and slightly flat tened in the siphonal is most likely linked to their natu ral variabil ity, includ ing that area. The umbil i cal wall is low and slightly rounded. growth and shell diagenesis. The um bil i cal width is close to val ues The ribs be gin as simple ribs at approx i mately mid-height of the found in the type ma te rial (U/D = 0.27–0.31) as re ported by umbil i cal wall, where they are curved con cavely. The spaces be- Mazenot (1939) and in mate rial de scribed by Arkadev and Bogda- tween in di vid ual ribs are rather large in the lower part of the nova (2012) from Cri mea, but not to those in di cated by Tavera- flanks. At two-thirds of the whorl height, the ribs bi fur cate quite Benitez (1985), who pre sented val ues cor re sponding to a quite nar rowly. The an te rior bi fur cated one runs in the di rec tion of the wide umbi li cus. In con trast, rib den sity per half whorl re mains in simple ribs, whereas the pos te rior is de flected in the op po site di - prin ci ple con stant with shell growth. rec tion. Apart from the di rec tion of the pos te rior ribs, all of the ribs

Ta ble 1 Measure ments (in mm) and ratios of Berriasella jacobi Mazenot, 1939

Spec. No. D H U B B/H H/B UR VR PL 4149 approx. 23.5 10.6 (0.45) 6.7 (0.29) c. 6.6 (0.28) 0.62 1.60 22 PL 4148 max. 31.9 12.6 (0.395) 10.1 (0.32) c. 10.0 (0.31) 0.79 1.26 PL 4148 29.8 11.9 (0.40) 9.2 (0.31) 21 41 PL 4147 max. 37.6 16.2 (0.43) 10.6 (0.28) 22 PL 4147 31.1 13.9 (0.45) 8.4 (0.27) c. 10.3 (0.33) 0.73 1.35 PL 4146 37.8 15.5 (0.41) 10.2 (0.27) c. 11.0 (0.29) 0.71 1.41 22 43 336 Z. VAŠÍÈEK & P. SKUPIEN

in general are rela tively straight and slightly prorsiradiate. On the rupt ing the ribs. Near ing the max i mum diam e ter, this furrow grad - pen ul ti mate whorl, the points of rib bi fur ca tion are clear. In the u ally dis ap pears. pre vail ing por tion of the ul ti mate whorl of the nearly complete Measure ments (in mm) and ratios are in Table 2. The incom - spec i men, in which more than a half of the ulti mate whorl be longs plete smaller shell (spec. PL 4153) has a di ame ter of ap prox i - to the body cham ber, a nar row siphonal fur row is distinct, in ter - mately 40 mm. BERRIASIAN AMMONITES FROM THE ŠTRAMBERK LIME STONE 337

Fig. 5. The Berriasian amonites from the Štramberk Lime stone. Prior to pho tog ra phy, all speci mens were coated with ammo nium chlo - ride. Pho to graphs: K. Mezihoráková, Ostrava. Scale bars equal 10 mm. A, B. Spiticeras blancheti Djanélidzé, 1922; A – lat eral view. On the left, abapertural to the part with ribs, there is a distinct con stric tion; how ever, this is not prop erly vis i ble in the pho to graph; B – ven tral view. PL 4145, layer 3. C. Berriasella jacobi (Mazenot, 1939), PL 4148, layer 5. D, I. Berriasella jacobi (Mazenot, 1939); D – lat eral view showing bi fur cate rib bing, I – ven tral view near ap er ture with siphonal fur row, PL 4146, layer 3. E, F. Berriasella oppeli (Kilian, 1889); E – lat eral view, F – ven tral view. Spec i men 17/80-1980-3181, Silesian Mu seum at Opava; al leg edly from the old 3rd level at Kotouè Quarry, quar ried in the 1960s. G, H. Berriasella jacobi (Mazenot, 1939); G – lateral view, H – ventral view at the start of ulti mate whorl pre served as in ter nal mould with dis tinct siphonal fur row, PL 4147, layer 4. J. Berriasella oppeli (Kilian, 1889), PL 4153, layer 2. K, L. Tirnovella allobrogensis (Mazenot, 1939); K – lat eral view – note dense rib bing, L – ven tral view, PL 4154, layer. 3. M. Tirnovella allobrogensis (Mazenot, 1939), with shell retained, PL 4155, layer 3; N. Tirnovella cf. allobrogensis (Mazenot, 1939), in ter nal mould, PL 4156, layer 3. O, P. Delphinella consanguinea (Retowski, 1893); O – lat eral view with the polygyrate rib bing, P – ven tral view with siphonal fur row. PL 4157, layer 3. Q. Pseudosubplanites cf. grandis (Mazenot, 1939), PL 4158, layer 6. R. Malbosiceras cf. asper (Mazenot, 1939), PL 4159, layer 3

Re marks: The size param e ters of the nearly complete shell from ther more, a frag ment of a ju ve nile shell (PL 4154) is avail able. the Silesian Museum are close to those of the lectotype of B. De scrip tion: The shells are semi-invo lute to invo lute, with high oppeli (in von Zittel, 1868, pl. 20, fig. 1a–c). A char ac ter is tic fea - and nar row whorls and a nar row umbi li cus. The um bil i cal wall is ture, in contrast to other berriasellids, is the some what sparse rib- low and obliquely in clined. The flanks are slightly arched. Their bing and a long ap par ent siphonal fur row. maxi mum width oc curs near the um bil i cal shoul der. The lower Dis tri bu tion: The lower Berriasian (Berriasella jacobi Zone) in part of the whorl is quite flat, and the flanks only be gin to in cline Tu ni sia, Spain, France, Ro mania and at Koòákov in the Silesian to the venter somewhat more markedly at mid-whorl height. The Unit. In Bul garia and Cri mea, B. oppeli occurs in the Berriasella venter is not pre served. jacobi Zone as well as in the overly ing Occitanica Zone. The dense, thin rib bing con sists of ribs that are approx i mately Oc cur rence: The speci men from the Silesian Museum, la belled as equally strong along their entire length. On the umbil i cal wall, the Berriasella carpathica (Zittel), co mes from the old, long-exploi- ribs are curved slightly con cavely in re la tion to the aper ture. On ted level 3 at Kotouè Quarry. The in complete spec i men of the the flanks, the ribs are rather straight initially; then, at approx i - pres ent study was found in Layer 2, along the exit road be tween mately mid-whorl height, they are very slightly S-shaped, and in level 4 and level 3. the up per two-thirds, they are mark edly prorsiradiate. A por tion of the ribs bi fur cate near the um bi li cus. In the lower third to half of the Ge nus Tirnovella Nikolov, 1966 whorl height, a greater por tion of the ribs bifur cate nar rowly. Type spe cies: Berriasella alpillensis Mazenot, 1939 On the op posite, im per fectly pre served side of the larger spec i - men, the orig i nal shell is not preserved. In the in ner mould, a weak - (Berriasian, south east France), by orig i nal des ig na tion of en ing and grad ual dis ap pear ance of the rib bing is clearly visi ble. Mazenot (1939, p. 73) Mea sure ments: PL 4155 has a max i mum di am e ter of ap prox i - mately 44 mm. At this diam e ter, H = c. 21.0 (0.48), U = c. 10.4 Tirnovella allobrogensis (Mazenot, 1939) (0.24), B = c. 10.0 (0.23), B/H = c. 0.48 and H/B = c. 2.1. How- Fig. 5K–M ever, these param e ters are ap proxi mate at best. In one quarter of the ulti mate whorl, at a shell diam e ter of 44 mm, there are 13 ribs * 1939 allobrogensis n. sp. – Mazenot, p. 210, pl. near the um bi li cus and 28–29 ribs ventrolaterally, i.e., ap prox i - 33, fig. 4a, b. mately 26 ribs near the um bi li cus and ap prox i mately 56 ribs on the 1965 Neocomites suprajurensis Mazenot – Houša, p. 532, cir cum fer ence per half whorl. fig. VIII–282. The ju ve nile shell, PL 4154, has an es ti mated max i mum diam e - 2005 Tirnovella allobrogensis (Mazenot) – Klein, p. 254 ter of ap prox i mately 25 mm. At another point on the shell close to [cum syn.]. the max i mum diam e ter, H = 12.8 mm, B = 6.8 mm, B/H = 0.53 and 2012 Tirnovella allobrogensis (Mazenot) – Arkadev and H/B = 1.88. Bogdanova, p. 159, pl. 7, figs 7–9 [cum syn.]. Re marks: Tirnovella allobrogensis is mor pho log i cally close to T. Ma te rial: Four incom plete spec i mens, the best-pre served be ing suprajurensis (Mazenot, 1939). The basic differ ence between PL 4155. Its incom plete ulti mate half whorl, with origi nal recrys- them appears to be shell size, as T. allobrogensis most likely rep re- tallised shell retained, exhib its an imper fectly pre served venter sents a microconch and T. suprajurensis a macroconch. Re gard ing and a small part of the pre ced ing whorl. Shell diam e ter can only be shell diam e ter, the limit between the two species is approx i mately mea sured ap prox i mately. The last quar ter is body cham ber. Fur - 65 mm (Mazenot, 1939, p. 210).

Ta ble 2 Measure ments (in mm) and ratios of Berriasella oppeli (Kilian, 1889)

Spec. No. D H U B B/H H/B UR VR Sil. Mus. 3181 48.4 17.8 (0.37) 18.6 (0.38) c. 15.6 (0.32) c. 0.88 1.14 19 37 45.0 16.5 (0.37) 17.2 (0.38) c. 15.5 (0.34) 15.0 13.5 0.90 1.11 338 Z. VAŠÍÈEK & P. SKUPIEN

The shells are thinly and densely ribbed. Data on den sity and fea ture is seen on one of the pre served surfaces of PL 4155 (de - number of ribs, re ported by Arkadev and Bogdanova (2012, p. scribed above as T. allobrogensis). However, the reverse side, 159: 35 ribs near the umbi li cus and 80 ribs on the cir cumfer ence with the pre served orig i nal shell, does not ex hibit any rib weak en - per whorl at a shell di ame ter of 31.5 mm), are evi dent for a ing. Weak en ing of the ribs in Pseudoneocomites retowskyi from half-whorl, rather than an en tire whorl (see, e.g., spec i men in their Crimea occurs initially only at a shell di ame ter of approx i mately pl. 7, fig. 9). In ad di tion, T. occitanica (Pictet, 1867) and T. 60 mm. The iden ti fi ca tion of PL 4156 as P. retowskyi is there fore subalpina (Mazenot, 1939), in which the rib bing is stron ger and un likely. ribs bear weak umbil i cal tuber cles, appear to be rela tively closely Oc cur rence: Layer 3 along the exit road from level 3 to level 4 at re lated to T. allobrogensis. Kotouè Quarry (to gether with T. allobrogensis). The well-preserved speci men from the Štramberk Limestone il- lustrated by Houša (1965), re ferred to as Neocomites supraju- rensis, ev i dently rep re sents one of the larg est and most per fectly Ge nus Delphinella Le Hégarat, 1971 pre served microconchs of T. allobrogensis. Type spe cies: Hop lites delphinensis Kilian, 1889 Dis tri bu tion: This spe cies is, ac cord ing to Arkadev and Bogda- (Berriasian, south east France), by orig i nal des ig na tion of nova (2012), known from the lower Berriasian Berriasella jacobi Nikolov (1966, p. 639) Zone, spe cif i cally from its higher part, i.e., the Grandis Subzone from the Cri mean Mountain ous, as well as from the Berriasella Delphinella consanguinea (Retowski, 1893) jacobi Zone in the Cau ca sus, Bul garia, south east France, Spain and North Af rica. Fig. 5O, P Oc cur rence: Layer 3, exit road from level 3 to level 4, Kotouè ?partim * 1893 Hop lites consanguineus n. sp. – Retowski, p. Quarry. 268, pl. 12, figs 1a, b. [?non fig. 2]. 1939 Berriasella consanguinea (Retowski) – Mazenot, p. 79, pl. 7, fig. 4a–c. Tirnovella cf. allobrogensis (Mazenot, 1939) 2005 Delphinella consanguinea (Retowski) – Klein, p. Fig. 5N 184 [cum syn.]. Ma te rial: A single in complete and rather poorly pre served in ter - Ma te rial: A sin gle in ter nal mould, rep re sent ing the ul ti mate half- nal mould, show ing in distinct relicts of su tures (PL 4156). whorl of vari able pres er va tion (PL 4157). De scrip tion: The spec i men is invo lute, with high, nar row whorls De scrip tion: The spec i men is semi-evolute, with nar row whorls and a nar row umbi li cus. The whorl flanks are quite flat and grad u - of medium height. The umbi li cus is quite wide. The ul ti mate whorl ally in clined to the venter, which is quite nar row, only slightly is slightly arched. The very low umbil i cal wall is inclined obli- arched and sepa rated from the flanks rela tively clearly. quely to wards the line of coil ing and is not lim ited sharply from The rib bing is thin and dense. The ma jor ity of ribs near the um bi li - the flanks. The great est whorl width oc curs at ap prox i mately mid- cus are sim ple, with a subset bi fur cat ing at the umbi li cus. On the height. From there, the whorl in clines very grad ually to the venter. flanks, fur ther rib bi fur ca tion can be seen. This bi fur ca tion is less The venter is nar row, flat and smooth and it is sharply sep a rated distinct be cause at close to half the whorl height, the rib bing is from the flanks. weak ened con sid er ably, ex tend ing nearly to the venter. On the The or na ment con sists of S-shaped, al to gether prorsiradiate tran si tion of the flanks to the venter, the ribs again be come dis - ribs. The ribs start in distinctly as simple ribs in the vicin ity of the tinct. On the venter near the ap er ture, the ribs are in ter rupted in a line of coil ing. In the area of the um bil i cal wall, the ribs are curved quite wide smooth zone. In a short sec tion on the op po site side, con vexly. At ap prox i mately mid-height, the ribs bi fur cate. At the near the ap er ture, the orig i nal shell is pre served. In that sec tion, the point of bifur ca tion, there is a lateral tuber cle, which is initially ribs run across the venter in a wide arc with out in ter rup tion. weak and lon gi tu di nally elon gated; sub se quently, tu ber cles be - Mea sure ments: In view of the in complete ness of the shell, the come somewhat stron ger. The orig i nal simple ribs con tinue as an - mea sured val ues are only ap prox i mate. At a D of ap prox i mately te rior bi fur cated ribs. The pos te rior ribs are rursiradiate but prorsi- 39 mm, H = c. 17.8 (0.47) and U = c. 9.9 (0.26). ra di ate there af ter. In the ven tral re gion, be tween the bi fur cated Re marks: The present shell is close in terms of size to T. allobro- ribs, one rib is occa sionally in serted. These short ribs efface near gensis. How ever, it dif fers in ex hib it ing mark edly weak ened rib - the point of rib bi fur ca tion. On the bound ary be tween the flanks bing early on the flanks of the ul ti mate whorl, sim i lar to the Cri - and venter, the ribs end abruptly in the in di cated ventrolateral tu - mean spec i mens assigned by Arkadev and Bogdanova (2012, pl. 9, bercles. The venter is rela tively nar row and flat, with a smooth, figs 1, 2) to Tirnovella occitanica (Pictet, 1867). One of their spec - rel a tively wide, shal low siphonal fur row. The fi nal por tion of the i mens (pl. 9, fig. 1) conforms to the de scrip tion of “Hop lites” ul ti mate whorl ex hib its a ma trix-cov ered ven tral re gion, and thus, occitanicus by Retowski (1893, pl. 11, fig. 9). The spec i mens de- it is not clear whether the smooth zone in the siphonal re gion scribed by Retowski (1893) later became the type ma terial of reaches the max i mum pre served shell di am e ter. Pseudoneocomites retowskyi. This has a chequ ered no mencla ture Measure ments (in mm) and ratios are in Table 3. hav ing been re ported by Sarasin and Schöndelmayer (1901, p. 72), Re marks: For the slightly de formed holotype, Retowski (1893) as Hop lites Retowskyi, then Neocomites retowskyi by Mazenot re corded the fol low ing di mensions: D = 57.0 mm, H/D = 0.37, (1939, p. 211) and Druschic (1960, p. 289) and, subse quently, U/D = 0.39 and B/D = c. 0.18. D. consanguinea is, in contrast to Pseudoneocomites retowskyi (see Hoedemaeker, 1982, p. 68; other delphinellids, char ac ter ised by a quite wide umbi li cus, a rel - Klein, 2005, p. 265). Bogdanova et al. (1999) ar gued that Hoede- a tively wide, smooth zone in the siphonal re gion and early oc cur - maeker (1982) did not de scribe the type spe cies in de tail and thus, rence of lat eral tuber cles. it was diffi cult to deter mine the species concept of P. retowskyi Dis tri bu tion: So far, this spe cies has been reported ex clu sively and the con tent of the ge nus Pseudoneocomites. However, the Cri - from Crimea and south east France. The ge nus Delphinella is spe- mean spec i men re ferred to above should not be at trib uted to the in - ciose, es pecially in Crimea, specif i cally in the ammonite Berria- dex species, Tirnovella occitanica. Addi tional material is needed sella jacobi Zone (Arkadev and Bogdanova, 2012). Al though from Crimea for clarification. those au thors did not men tion the Cri mean spe cies D. consangui- The weak en ing of the ribs on the flank of our spec i men is evi - nea in their mono graph, it can be as sumed that its oc cur rence is dently con nected to its preser va tion as an in ter nal mould. A simi lar confined to the Berriasella jacobi Zone as well. BERRIASIAN AMMONITES FROM THE ŠTRAMBERK LIME STONE 339

Ta ble 3 Measure ments (in mm) and ratios of Delphinella consanguinea (Retowski, 1893)

D H U B B/H H/B UR VR PL 4157 42.8 15.8 (0.37) 17.2 (0.40) 36.7 13.4 (0.365) 13.1 (0.35) c. 8.0 (0.25) 0.60 1.67 16 39

Oc cur rence: Layer 3, near exit road from level 3 to level 4, at Oc cur rence: Layer 6, along the exit road from level 4 to level 3 at Kotouè Quarry. Kotouè Quarry. An other, sub stan tially better-pre served, spec i men of Ps. grandis, from level 6 at that quarry, was de scribed by Vašíèek et al. (2013). Ge nus Pseudosubplanites Le Hégarat, 1971 Type spe cies: Pseudosubplanites berriasensis Le Hégarat, 1973 (Berriasian, southeast France), by origi nal desig na - Ge nus Malbosiceras Grigorieva, 1938 tion of Le Hégarat (1971, p. 850). Vašíèek et al. (2013) Type spe cies: Ammonites malbosi Pictet, 1867 (Berriasian, have recently presented their concept of the genus France), by origi nal desig na tion of Grigorieva (1938, Pseudosubplanites p. 102)

Pseudosubplanites cf. grandis (Mazenot, 1939) Malbosiceras cf. asper (Mazenot, 1939) Fig. 5Q Fig. 5R * 1939 Berriasella grandis n. sp. – Mazenot, p. 133, pl. 22, *1939 Berriasella aspera n. sp. – Mazenot, p. 84, pl. 9, figs figs 3a, b, 6a, b. 2a–c, 4a, b. 1973 Pseudosubplanites grandis (Mazenot) – Le Hégarat, 2005 Malbosiceras asper (Mazenot) – Klein, p. 207 [cum p. 38, pl. 2, figs 3, 4; pl. 37, fig. 9. syn.]. 2005 Berriasella (Pseudosubplanites) grandis (Mazenot) – Ma te rial: A small frag ment of a whorl (PL 4159) of a quite large Klein, p. 158 [cum syn.]. shell, pre served as a com posite mould, of which nei ther the um bil - 2012 Pseudosubplanites grandis (Mazenot) – Arkadev and ical area nor the venter are preserved. Bogdanova, p. 174, pl. 12, figs 1, 2; text-fig. 64-6 De scrip tion: The whorl is slightly arched with the maxi mum [cum syn.]. width at mid-height; maxi mum whorl height is 32 mm. Only three 2013 Pseudosubplanites grandis (Mazenot); Vašíèek et al., promi nent pri mary ribs are pre served. They be gin simple, are p. 463, figs 3, 4. straight, high, rela tively thin and distinctly prorsiradiate. At mid- Ma te rial: An im perfectly pre served fragment of fairly large size height, quite strong tri an gu lar tu ber cles oc cur. From the tu ber cles, (PL 4158), rep re sent ing ap prox i mately one quar ter of the ul ti mate the ribs oc cur in pairs or triplets. When there are two ribs, a third whorl and half of the pen ul ti mate whorl, with recrystallised orig i - rib that reaches as far as the level of the tuber cles is inserted. The nal shell, plus its exter nal mould. split ribs are blunter than the simple ones. De scrip tion: The shell is semi-evolute to evolute, without high Re marks: The char ac ter of the ribbing is simi lar to that ob served whorls. The whorls are only weakly arched. The greatest whorl in both M. asper and the re lated M. chaperi (Pictet, 1868). In view width oc curs in ap prox i mately the lower third. The um bil i cal wall of the prorsiradiate in cli na tion of the lower part of ribs of the for - which passes con tin uously to the flanks of the ul ti mate whorl, is mer species, we re fer it to that species, albeit with a query. low and in clines obliquely to the line of coil ing. Dis tri bu tion: Ac cord ing to Nikolov (1982), M. asper occurs in The ribs are quite strong and prorsiradiate. They be gin as simple the lower Berriasian (Berriasella jacobi Zone) in France, Bulgaria ribs at the um bil i cal wall above the line of coiling. In a short sec- and pos si bly Spain. tion, the ribs are curved con cavely. In ap prox i mately the lower Oc cur rence: Layer 3, along the exit road from level 3 to level 4, at third-to-half of whorl height, the ribs bi fur cate nar rowly. In the the Kotouè Quarry. lower part of the ulti mate whorl, the ribs are thin initially and fur- ther strengthen in creas ingly. In the outer half of the whorl, they are quite wide and flat. One of the ribs pre served on the ul ti mate whorl re mains simple along its en tire length. On the pen ul ti mate whorl, DIS CUS SION the points of rib bi fur ca tion are visi ble. The max i mum height of the ul ti mate whorl is ap prox i mately 23 The Berriasian, as the oldest stratigraphic stage of the mm, which could cor re spond to a shell di ame ter of ap proxi mately Cre ta ceous Sys tem, is cur rently sub di vided (Reboulet et al., 60 mm. 2011) into three inter na tion ally accepted ammonite zones. Re marks: Pseudosubplanites grandis is charac terised by fairly The low est of these, with Berriasella jacobi and corre - large di am e ters. The ma jor ity of spec i mens, il lus trated in the lit er- sponding to the lower Berriasian, is not further subdi vided ature, are rela tively fragmentarily pre served [one excep tion being into subzones. However, Arkadev and Bogdanova (2012) that de scribed by Le Hégarat (1973, pl. 37, fig. 9)]. A spec i men re - did use a tri par tite sub di vi sion of the Berriasella jacobi ported by Mazenot (1939, pl. 22, fig. 3) is morpho log i cally clos est Zone in Crimea: a lowest subzone, with B. jacobi; a middle to frag ment in the pres ent study, but in view of the in complete na - ture of the find, the au thors here use open no mencla ture. subzone with Pseudosubplanites grandis and an upper with Dis tri bu tion: Ac cord ing to Arkadev and Bogdanova (2012), Malbosiceras chaperi. In earlier studies, an inde pend ent Pseudosubplanites. grandis occurs in the lower Berriasian (Berria- Grandis Zone has occa sion ally been indi cated in strata over- sella jacobi Zone) in the Cri mean Mountains, Cau ca sus, Bul garia, lying the Berriasella jacobi Zone (e.g., Le Hégarat, 1973; Ro ma nia, Hun gary, south east France and Tu ni sia. Hoedemaeker, 1982). The authors accept the opinion of 340 Z. VAŠÍÈEK & P. SKUPIEN

Arkadev and Bogdanova (2012) that P. grandis ap pears in ings, usu ally only the ap prox i mate lo ca tion, “Štramberk”, the higher part of Berriasella jacobi Zone. appears on museum labels. Such speci mens are depos ited in In the mid dle Berriasian, a zone with Tirnovella occi- many regional muse ums. The prevail ing opinion is that tanica has been recog nised in re cent studies (Reboulet et these ammonites are of Tithonian age. al., 2011; Arkadev and Bogdanova, 2012). However, in the The newly collected ammonites from the Kotouè Qua- paper by Reboulet et al. (2011), T. occitanica is in di cated as rry, de scribed in the pres ent account, come from a block of Subthurmannia occitanica. The Occitanica Zone con tains limestone that, in con trast to the ma jor ity of mas sive three subzones in this case. The one with Fauriella boissieri Štramberk lime stones, ex hib its strongly de vel oped bed ding. corre sponds to the upper Berriasian. Reboulet et al. (2011) Among this rich collec tion of ribbed ammonites, belong ing also assigned this species to Subthurmannia. to seven spe cies, Berriasella jacobi (index spe cies of the From a stratigraphic point of view, in the new ammo- lower Berriasian) and Pseudosubplanites grandis are of nite asso ci a tion from Kotouè Quarry, berriasellids (Berria- great est strati graphic im por tance. Nei ther pre vi ously was sella jacobi, B. oppeli, Tirnovella allobrogensis, T. cf. allo- con nected to the Štramberk area. An early Berriasian age is brogensis, Delphinella consanguinea, Pseudosubplanites also suggested by the other spe cies asso ci ated with them. cf. grandis and Malbosiceras cf. asper) are the most sig nif i - However, in spite of these early Berriasian ammonites, it re - cant taxa. The in dex species, B. jacobi, occurs in large num- mains valid to conclude that the major ity of older ammonite bers, whereas the oth ers are rarer. Other spe cies found only finds in the Štramberk Limestone are of Tithonian age. spo rad i cally ei ther are long-rang ing (Protetragonites qua- Ammonites post dating the early Berriasian have never drisulcatus d’Orbigny, Haploceras elimatum Oppel) or been found in the Štramberk Limestone. This suggests that their strati graphic po si tions ac cord ing to the lit er a ture are depo si tion of the Štramberk Limestone in the Štramberk not known pre cisely (i.e., small-sized fragments and juve - area came to an end dur ing that time. nile shells of Spiticeras). The follow ing taxa are known: Ac knowl edge ments layer 1 – Berriasella jacobi; layer 2 – Berriasella oppeli, ?Neocosmoceras sp. (PL 4160), The au thors thank K. Mezihoráková (Ostrava) for tak ing the Punctaptychus punctatus; pho to graphs of the ammonites de scribed, the manage ment of the layer 3 – Spiticeras blancheti, Spiticeras sp. juv., Berria- Kotouè Quarry for al lowing them ac cess to the quarry and to the sella jacobi, Tirnovella allobrogensis, T. cf. allobro- re view ers of the pa per, J. W. M. Jagt (Natuurhistorisch Mu seum Maastricht, Nether lands), B. Ko³odziej (Jagiellonian Univer sity, gensis, Delphinella consanguinea, Malbosiceras cf. Kraków, Poland) and M. Machalski (Pol ish Acad emy of Sci ences, asper, Protetragonites quadrisulcatus, Haploceras eli- Warszawa, Po land), for construc tive re marks and sugges tions. We matum (PL 4161), Lamellaptychus beyrichi; thank the first re viewer and F. Simpson (Univer sity of Windsor, Ca- layer 4 – Berriasella jacobi, Spiticeras sp. juv., Protetrago- nada) for lin guis tic help. The research of P. Skupien was sup ported nites quadrisulcatus (PL 4162), Punctaptychus punc- by Grant SP2013/36 from the Min is try of Ed u ca tion, Youth and tatus; Sports of the Czech Re pub lic and by the pro ject RVO 68145535. layer 5 – Berriasella jacobi; layer 6 – Pseudosubplanites cf. grandis; layer 7 – Punctaptychus punctatus, Lamellaptychus beyrichi. REF ER ENCES With regard to the observed succes sion of identi fi able ammonites (which can not al ways be found in the higher Arkadev, V. V. & Bogdanova, T. N., 2004. Ge nus Berriasella part of the sec tion and identi fied with cer tainty, i.e., Pseu- () and ammonoid zonation in the Berriasian of the Cri mean Mountains. Stra tig ra phy and Geo log i cal Cor re- dosubplanites cf. grandis and Malbosiceras cf. asper), the la tion, 12: 367–379. ammonite as soci a tion of the sec tion, described in the pres- Arkadev, V. V. & Bogdanova, T. N., 2012. Golovonogie molljuski ent study, can be consid ered to belong to the upper part of (Ammonity). In: Arkadev, V. V. & Bogdanova, T. N. (eds), the ammonite Berriasella jacobi Zone. Guzhikov, A. J., Lobacheva, S. V., Myshkina, N. V., Pla- There are no data to indi cate a younger age of the tonov, E. S., Savel’eva, J. N., Shurekova, O. V. & Janin, B. T. Štramberk Limestone in the Štramberk area. How ever, as Berrias Gornogo Kryma. Izd. LEMA, Sankt-Pe ters burg, pp. noted on the basis of foraminifera from Štramberk-type 123–224. [In Rus sian]. limestones in the Outer Carpathians (exotic blocks in flysch Arnould-Saget, S., 1953. Les ammonites pyriteuses du Tithonique depos its) and biostratigraphic data on the substrate drilled in supérieur et du Berriasien de Tunisie centrale. Annales des the Carpathian Foredeep (all in Poland), shallow-wa ter sed- Mines et de la Géologie, 10 (for 1951): 1–133. imen ta tion in the northern segment of the Carpathian ba sin Blanford, H. F., 1864. On Dr. Gerard’s col lec tion of fos sils from the Spiti Val ley, in the Asiatic Soci ety’s Museum. Jour nal of persisted locally into the Valanginian (Ivanova and Ko³o- the Asiatic Soci ety of Bengal , 32 (for 1863): 124–138. dziej, 2010, and literature therein). Blaschke, F., 1911. Zur Tithonfauna von Stramberg in Mähren. Annalen des kaiserlich-königlichen naturhistorischen Hof- museums, 25: 143–222. Bogdanova, T. N., Kalacheva, E. D. & Sey, I. I., 1999. O prisustvii CON CLU SIONS zony Tirnovella occitanica (nizhnij mel, berrias) v feodo- sijskom razreze Vostochnogo Kryma. Re gional’naya Geo- In his tor i cal col lec tions of ammonites from the Štram- logiya i Metallurgiya, 1999 (9): 27–32. [In Rus sian]. berk Limestone at Štramberk and its im medi ate surround - Djanélidzé, A., 1922. Les Spiticeras du sud-est de la France. BERRIASIAN AMMONITES FROM THE ŠTRAMBERK LIME STONE 341

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