1Ч. Jb. Geol. Palâont. Abh. 265/3, 249-274 Article ШStuttgart, September 2012

Taxonomy of Late Jurassic - Early Cretaceous aptychi from Bulgaria

Zdenék Vasicek, Neda Motchurova-Dekova, Antoaneta llcheva, and Lubomir Metodiev With 11 figures

In memory of July Stefanov (1932-1966), one of the most talented Bulgarian palaeontologists, on occasion of his 80th anniversary

V a s ic e k , Z., M otchurova-Dekova, N., I lc h e v a , A. & M e t o d ie v , L. (2012): Taxonomy of Late Jurassic - Early Cretaceous aptychi from Bulgaria. - N. Jb. Geol. Palàont. Abh., 265: 249-274; Stuttgart.

Abstract: This paper presents the taxonomic revision of Late Jurassic and Early Cretaceous aptychi housed at the National Museum of Natural History, Sofia (Bulgaria). The aptychi came from the pelagic, flysch-like and flysch deposits of Central and North Bulgaria, and most of them have been initially described by S t e f a n o v (1961). The taxonomic revision has led to the determination of 23 species and subspecies of six genera. A new species named Mortilletilamellaptychus stefanovi is de­ scribed. The collection contains also the holotype of Didayilamellaptychus filicostatus (Stefanov), which was previously elevated from a variety to species level by M ë c h o v à et al. (2010). A valve of Didayilamellaptychus cf. didayi preserved in the body chamber of a Neolissoceras supports the hypothesis that lamellaptychi and perhaps punctaptychi belong to the natural system of ammonites of family Haploceratidae Z i t t e l , 1884. It became evident that the state of preservation may strongly affect the taxonomic determination of the aptychi. Only in exceptional cases (occurrence of both valves in a pair with different state of preservation) this preservation phenomenon can be revealed and erroneous determination of any of the valves could be possibly avoided.

Key words: Late Jurassic, Early Cretaceous, Bulgaria, aptychi, ammonites, parataxonomy, preserva­ tion, new species.

1. Introduction collection was housed at the Geological Institute of the Bulgarian Academy of Sciences (GI-BAS) and an­ This paper presents a taxonomic revision of collec­ other, smaller part - in the Palaeontological Museum tion of Late Jurassic and Early Cretaceous aptychi of Sofia University. The National Museum of Natural housed at the National Museum of Natural History, History, Sofia (NMNHS) was later (1974) restored as Sofia. Most of this collection, 54 specimens of Early an autonomous institution within the Bulgarian Acad­ Cretaceous aptychi from Bulgaria, were initially de­ emy of Sciences. On that occasion, only part of the scribed by Ju l y S t e f a n o v in 1961. S t e f a n o v was one geological collections from GI-BAS was moved to of the talented and most promising Bulgarian palae­ NMNHS in the years following 1974. The reason of ontologists of the 20th century. Unfortunately, he died this selective moving is unknown. To our knowledge, tragically in 1966 at the age of 34. At the time of the the aptychi collection of July S t e f a n o v was one of the publication (S t e f a n o v 1961), one part of the aptychi few important palaeontological collections that were

©2012 E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart, Germany www.schweizerbart.de DOI: 10.1127/0077-7749/2012/0257 0077-7749/2012/0257 $ 6.50 250 Zdenëk Vasfcek et al.

Fig. 1. Map showing the occurrences of Late Jurassic - Early Cretaceous aptychi in Bulgaria. The approximate exposures of the Upper Jurassic and the Lower Cretaceous rocks are plotted on the simplified tectonic framework of Bulgaria (after Z agorchev et al. 2009).

moved to the NMNHS. The specimens of this col­ of certain problematic specimens for which open no­ lection, described earlier by Stefanov, have succes­ menclature is used here. sive museum registration numbers NMNHS F-30809 One of the reasons to revise the Bulgarian collec­ trough to 30876. tion were also the new results reported in the Ph.D. Nineteen Late Jurassic and Early Cretaceous speci­ thesis of L. K ratochvilovâ (2004), where she pro­ mens of aptychi not described earlier are also included posed a serious rearrangement of the taxonomy of in this taxonomic account. They have older museum Early Cretaceous aptychi. The results of her thesis numbers previously assigned (random numbers be­ served as a basis for the suggested new classification tween NMNHS F-335 and NMNHS F-9928) and are (M ëchovâ et al. 2008, 2010). marked with an asterisk in the text. As a result of the At the time of publication (1961), S tefanov’s spec­ taxonomical reappraisal of the Bulgarian material imens were one of the few significant collections of (completed by Z.V.), one new species is erected, named Early Cretaceous aptychi alongside the collections of in honour of J uly Stefanov. Unfortunately, some T rauth (1927-1938) and GдsIOROwsкI (1959, 1962). twelve specimens from the collection remain undeter­ The paper of Stefanov (1961) and the present contribu­ mined because of poor or fragmentary preservation. tion, where new material is added, extend considerably This study aims to facilitate further investigation the knowledge about the distribution of aptychi from of the collection. Future research could focus on the Western and Central to Eastern Europe. Stefanov shell microstructure and/or on the systematic position (1961) had described the collection accurately and in Taxonomy of Late Jurassic - Early Cretaceous aptychi 251 accordance with the aptychi classification at that time. alternation with turbidity sandstones, Tithonian), Since then, however, the knowledge on Early Creta­ Cherniosam Formation (limestones-shales-siltstones- ceous ribbed aptychi has evolved significantly to war­ graywackes in a repeated rhythmic alternation, Titho­ rant a review. nian - Valanginian), Kamchyia Formation (irregular As part of the present taxonomic reappraisal, when alternation of silty marls, siltstones and sandstones, possible, the stratigraphie level of the aptychi occur­ Berriasian - Hauterivian), Hanevtsi Formation (silty rences was critically reviewed using information from marls with interbeds of siltstones, Berriasian - Hau­ the museum labels and the paper by Stefanov (1961). terivian), Salash Formation (irregular marl-limestone The official lithostratigraphie units, which in most alternation, Valanginian - Hauterivian), and Gorna cases were introduced after 1961 (T enchov 1993) are Oryahovitsa Formation (marls with rare interbeds of provided here as well. We consider the revised age de­ sandstones and siltstones, Hauterivian) [for details see terminations confident enough since we have not ob­ Z agorchev et al. (2009), and references cited therein]. served evidence of redeposition and also because the There is a general agreement that these sequences revised ages conform to the range of occurrences of were deposited in wide pelagic environments during the respective taxa as described in the literature with the Late Jurassic times which were replaced from the just one exception (specimen NMNHS F-30841, see end of the Tithonian onwards by long-living foredeep Appendix 1). depositional setting. The data about the localities and horizons as well as Regionally, the rocks which yielded the aptychi Stefanov’s original determinations are summarized in associate largely with the most prominent and most Appendix 1. The original determinations of Stefanov elevated element of the Balkan orogenic system (the are given for each specimen in order to facilitate future Balkan Zone) and less with its foreland (the Moesian research and correlation with his personal archive and, Platform) in Bulgaria (Z agorchev et al. 2009). Just one most importantly, with later Bulgarian publications in locality is in the inner parts of the orogen (the Sred- which these taxa are cited. nogorie Zone). In terms of its geographic distribution, the localities of the studied aptychi display an unequal density of occurrence. The main share of the account 2. Geological setting that follows refers to the localities from a region of the The bulk of the aptychi described in this paper were Central Bulgaria, between the towns of Teteven and collected from Lower Cretaceous rocks, by number of Elena (Lovech and Gabrovo districts). Outside this specimens in ascending order, from the Beriassian to the area, a smaller number of localities in North-eastern Hauterivian. Little portion of our material, only 7 speci­ Bulgaria (Targovishte, Shumen, and Varna districts), mens, came from Upper Jurassic outcrops (Kimmerid- and Western Bulgaria (Sofia, Montana and Vratsa dis­ gian and Tithonian). A few of the aptychi studied have tricts) exist. The localities, ages and stratigraphie posi­ poorly known stratigraphie positions, but fall into the tions of the aptychi are listed for each sample in Ap­ stratigraphie framework outlined. Commonly, the apty­ pendix 1. A map showing the approximate geographi­ chi have scattered occurrences of randomly dispersed cal and geological position of the main localities of examples and data about other associated macrofossils aptychi and the distribution of the Upper Jurassic and preserved are missing. They occur within the pelagic, Lower Cretaceous rocks in Bulgaria is given in Fig. 1. flysch-like and flysch deposits, of great thicknesses that are widely exposed in Central and Northern Bulgaria 3. Aptychi and their taxonomy and crop out on a minor scale in Western Bulgaria. The sedimentary successions of Late Jurassic to The pairs of calcareous and ribbed valves of aptychi Early Cretaceous age that yielded the studied here have already been reported in the literature for almost aptychi are subdivided into several formal lithostrati- 200 years. The relationship between aptychi and am­ graphic units, which alternate gradually one after the monites was suggested already before the mid-19th other. The lithostratigraphic background taken into century. Opinions about their systematic position, consideration for this study includes: Gintsi Forma­ classification, function, and position in the soft body tion (nodular limestones, am m onitico rosso-type, and in the living chamber of ammonites have changed, Kimmeridgian - Tithonian), Glozhene Formation however, several times. The valves of aptychi are con­ (regularly-bedded, mainly micritic limestones, Titho­ sidered to be either opercula of ammonite shells (e.g., nian - Berriasian), Ticha Formation (marl-limestone T rauth 1927-1938; S chindew olf 1958) or the man- 252 Zdenëk Vasîcek et al.

dible of a jaw apparatus (e. g. L ehm ann 1972). Some parataxa exclusively as a common designation. researchers suggest that aptychi had both functions On the basis of further abundant findings of latest (L ehm ann & K ulicki 1990). E ngeser & K eupp (2002) Jurassic/Early Cretaceous aptychi in the Western Car­ summarized the biological function and phylogenesis pathians and Eastern Alps and data from the literature, of Jurassic and Cretaceous aptychi and ammonites. M ëchovâ et al. (2010) concluded that T urculet’s sub­ Those of the Neoammonoidea, in which calcareous generic division fails to cover the range of variability aptychi have been reported; they referred to a new in­ in ribbed aptychi, so it would be perhaps suitable to formal taxonomic unit, Aptychophora. determine some other new subgenera. However, this The calcitic ribbed aptychi occur in the paleon­ would cause an expansion of the old genus Lam ellap­ tological record most frequently as isolated valves. tychus into a disproportionate number of subgenera. Rarely, both valves are found together, and only ex­ Therefore, M ëchovâ et al. (2008, 2010) suggested el­ ceptionally, within the living chamber of an ammo­ evating Turcu let’s subgenera to the level of independ­ nite. Consequently, aptychi classification is entirely ent genera. These authors proposed to group them into artificial. For more details see M ëchovâ et al. (2010: two new families: Punctaptychidae and Lamellapty- 225-239). Therefore, the aptychi belong to the category chidae. Thus, the category of subgenus remains poten­ of artificial groups, similarly to other fossils such as tially open for the future. holothurian ossicles, conodonts, coleoid hooklets, and In this paper, the classification of ribbed aptychi as others. proposed by M ëchovâ et al. (2008, 2010) is followed. The first artificial classification of aptychi was wor­ This classification uses taxa on the levels of families, ked out by T rauth (1927-1938). For the Mesozoic ap­ genera, species and subspecies taken from natural tax­ tychi, he created 14 genera. The genera were divided onomy, but which are in fact artificial taxa (parataxa). into species, which were often subdivided into vari­ Such parataxa could also be possibly designated by eties (var.). T rauth also used the designation ‘forma terms, such as ‘form family’, ‘form genus’ and ‘form typica’ (f. typ.) after the species name when introduc­ species’, but we avoid them in the text that follows in ing subspecies. order to simplify it. Thus, for the ribbed aptychi of Late Jurassic and E ngeser & K eupp (2002) collected arguments, Earliest Cretaceous age (genera Punctaptychus and known earlier, that perfectly calcified thin to thick- Lamellaptychus) the classification based on the no­ walled valves with well developed ribs (lamellaptychi menclature of T rauth (1927-1938) was widely used in and punctaptychi) belonged to ammonites of the fami­ the last century. A huge amount of new, not yet known ly Haploceratidae Z ittel, 1884. This is well confirmed findings of latest Jurassic/Early Cretaceous aptychi by the same stratigraphie range of both fossil groups of both genera, collected in the framework of Deep (Kimmeridgian - Hauterivian; see e.g., W right et Sea Drilling Project (R en z 1972-1983) or in Romania al. 1996). The classification of the lamellaptychi and (T urculet 1994-2000) led to certain turnover in the punctaptychi to a single family of ammonites is also elaboration of the classification of ribbed aptychi based supported by the fact that they are characterised by on the nomenclature of T ra u th . An almost deadlock in shape uniformity of valves, which is marked also by the description and conception of other new species the ratio of valve width (Lat) to valve length (L). The (and largely subspecies) following the rules of T ra uth , value Lat/L moves in a narrow range from 0.43 to heading towards quadrinominal nomenclature (for de­ 0.56. However, some representatives of Thorolamel­ tails see M ëchovâ et al. 2010), urged Turculet (1994) laptychus (Lat/L = 0.57-0.60) and Didayilamellapty- to the definition of subgenera. For the general desig­ chus didayi (Lat/L about 0.65) are an exception. nation of taxa he explicitly used the term parataxa. He proposed the original para-genus Lamellaptychus 4. Systematic palaeontology T ra uth, 1927 to be divided into four para-subgenera: Beyrichilamellaptychus, Lamellosuslamellaptychus, Most of the taxa treated in the present paper were de­ Thorolamellaptychus and Diday ilamellapty chus. scribed recently in detail and their critical synonymy These para-genera were subdivided in para-species was given by M ëchovâ et al. (2010). When these au­ and para-subspecies. thors compiled their manuscript, they did not have at However, none of the previous and subsequent their disposal the extensive monograph by Turculet authors used the prefix “para-“ as a classification cat­ (2000) on Romanian calcareous aptychi. In his mono­ egory (e.g. para-species). They understood the term graph Turculet (2000) summarises his lifelong re­ Taxonomy of Late Jurassic - Early Cretaceous aptychi 253 search on Late Jurassic (mostly) and Early Cretaceous a mapping site, [= точка in Bulgarian]; NMNHS F- - prefix aptychi. The classification he uses is based on his di­ of the numeration in the collections in the National Museum of Natural History, Sofia. vision of the genera Lamellaptychus and Punctapty­ chus into subgenera (T urculet 1994). However, some of the subgenera determined by him do not comply Family Punctaptychidae M ëchovâ, H ousa & Vasicf;к. with nomenclatorial rules for the category of typical 2008 subgenera. Not all illustrations of Turculet (2000) are of satisfactory quality to assess the justification Diagnosis: Ribbed aptychi with valves composed of four of some of his specific determinations. For this rea­ calcareous layers. Outer lamellar layer overlain by porous son, our concept of species in the taxonomic part is (punctate) layer in apical area. supplemented only with the synonymy of those Early Cretaceous species which are described and illustrated Genus Punctaptychus T ra uth , 1927 clearly enough by T urculet (2000), but which were not included in the paper of M ëchovâ et al. (2010). Type species: Aptychuspunctatus Z i t t e l , 1868, p. 52. For all the taxa recently described by M ëchovâ et al. (2010) which we recognised in the Bulgarian Diagnosis: Thick-walled valves with simple ribbing. Ribs material without any doubts, we provide an abridged subparallel to the symphysal margin. Most of ribs end at synonymy only, i.e. the primary reference of papers, outer margin, a few at symphysal margin. where the type material was described and illustrat­ ed. In M ëchovâ et al. (2010) full synonymy and other Punctaptychus punctatus (Z ittel, 1868) relevant information can be found. Thus, for the taxa discussed already in detail by M ëchovâ et al. (2010), 1868 Aptychus punctatus Voltz. - Z ittel, p. 52, pl. 1, fig. only a brief description supplemented in some cases by 15a, b. remarks and measurements of the Bulgarian material 2000 Punctaptychus (Beyrichipunctaptychus) punctatus is presented. punctatus (Voltz). - Turculet, p. 129, pi. 10, figs. 2-3, 6, ? 1, 4-5, 7-9; pi. 11, figs. 1, 3, ? 2, 4-9. Three species of aptychi, including one new tax- 2010 Punctaptychus punctatus (Zittel). - Mëchovâ et al., on (Mortilletilamellaptychus stefanovi n. sp., Tho- p. 231, fig. 8 A (cum syn.). rolamellaptychus cf. aplanatus, and Laevaptychus longus) that were not discussed in the monograph of Material: Three poorly preserved specimens, NMNHS M ëchovâ et al. (2010) are here presented with full syn­ F-30839, 30841, and 337* (see Appendix 1 for details). onymy and described in more details. Measurements: Specimen NMNHS F-30841: L = 27.6 Part of S tefanov’s published material was re-pho­ mm, Lat = 15.0 mm, S - 25.8 mm, Lat/L = 0.54. tographed and is here illustrated with improved qual­ ity. All figured specimens with the exception of one Description: Small to large-sized valves. The outer margin (Fig. 8.10) were coated with ammonium chloride be­ is wide and rounded. Ribs are simple and slightly bent. Near fore photographing. the symphysal margin in the terminal area, ribs are thin and run close to each other. The punctate layer is preserved only The details of geographical and geological setting partly. in Bulgaria, as well as reference data on particular specimens, are to be found in the Appendix 1. Thus, Distribution: Upper Jurassic to Upper Berriasian of the under the sections “Distribution” only the occurrences Mediterranean area (M ë c h o v â et al. 2010). outside Bulgaria are mentioned.

Measurements and abbreviations: The basic morphology Punctaptychus diver gens T ra uth, 1935 of valves of aptychi and measured values is presented on Fig. 3.1 Fig. 2. All the well preserved specimens are measured. The values taken from fragmental specimens are denoted by 1935 Punctaptychus punctatus (V o ltz ) var. n. diver gens. apostrophe (L', Lat') to indicate an uncertainty of the meas­ - T r a u t h , p. 321, text-fig. 1. urement. L - total length of the valve; S - distance between 2000 Punctaptychus (Beyrichipunctaptychus) punctatus the apex and the terminal point; Lat - maximum valve divergens Trauth. - Turculet, p. 132, pi. 12, fig. 2, width; pW - width projection (distance between the termi­ ? 3-5, pi. 13, figs. 1-3, non fig. 4 (-Punctaptychus nal point and the projection of point of maximum width); rectecostatus Cuzzi, 1962). Fm - formation, referring to the official lithostratigraphic 2010 Punctaptychus divergens Trauth. - Mëchovâ et al., unit; t. - in the old museum numbers of specimens refers to p. 234, fig. 8C (cum syn.). 254 Zdenëk Vasicek et al.

inner Inner er facet edge Apical margin 2 edge Umblical projection

Symphysal facet Lateral margin

Symphysal facet 1 Lateral facet

Lateral Harmonic margin edge (Symphysis)

Terminal edge Terminal P°int Outer Outer Outer edge facet margin

Fig. 2. Basic morphology of valves of aptychi (1) and measured values (2) (after M ë c h o v à et al. 2010).

Material: One specimen, NMNHS F-336* (see Appendix Diagnosis: Ribbed aptychi, with valves formed by three 1 for details). calcareous layers. Valve surface with conspicuous ribs.

Description: Valve large in size. The ribs in the terminal area are arranged in a fan-like pattern. Genus Beyrichilamellaptychus Turculet, 1994

Distribution: Newer data place the earliest occurrence Type species: Aptychus beyrichi O p p el, 1865, p. 547. of the species in the Middle Tithonian of Eastern Alps (B o o r o v â et al. 2000). P. divergens was also proven in the Diagnosis: Majority of curved ribs converge along signifi­ lowermost Berriasian of Spain (Vasicek & Hoedemaeker cant section of symphysal margin, where ribs become thin­ 1997). ner and sub-parallel striped pattern condenses. In late evolu­ tionary forms ribbing pattern may be characterized by some of end ribs being discordant to previous ribs. Punctaptychus sp. Fig. 3.2 Beyrichilamellaptychus beyrichi (O ppel, 1865) Material: Four specimens, NMNHS F-335*, 338*, 1890*, 1891* (see Appendix 1 for details). 1865 Aptychus Beyrichi Opp. - Oppel, p. 547. 1868 Aptychus Beyrichi Opp. - Z i t t e l , p. 54, pl. 1, figs. 16, Remarks: Due to poor preservation the determination of 17 (type), 18, ?19. these specimens was possible to a genus level only. 2000 Lamellaptychus (Beyrichilamellaptychus) beyrichi beyrichi (O ppel) em. T r a u t h , /. typ. T r a u th . - T u r c u l e t , p. 85, pl. 1, figs. 2-3, ?figs 4-15, pi. 2, fig. 2, ?figs. 1, 3-4, pi. 24, fig. 9. Family Lamellaptychidae M ëchovâ, H ousa & 2010 Beyrichilamellaptychus beyrichi (Oppel). - Va sic ek, 2008 M ë c h o v â et al., p. 239, fig. 9B (cum syn.). Taxonomy of Late Jurassic - Early Cretaceous aptychi 255

Fig. 3. Representatives of the genus Punctaptychus. 1 - Punctaptychus divergent T r a u t h , NMNHS F-336; Shipkovski Mineral Baths. Lovech District, Tithonian, Cherniosam Fm. 2 - Punctaptychus sp., NMNHS F-335; Erden Village, Mon­ tana District, Tithonian, Ginci Fm - Glozhene Fm. Valve with not preserved punctate layer.

Material: Two fragmental specimens, NMNHS F-9738*, Beyrichilamellaptychus studeri (O o ster, 1857) 9740* (see Appendix 1 for details). Two other specimens, Fig. 4 (one poorly preserved specimen NMNHS F-30825 and one negative imprint NMNHS F-9737*) are referred to as Bey­ 1857 Trigonellites Studeri Ooster. -O oster, p. 26, pi. 7, fig. richilamellaptychus cf. beyrichi in Appendix 1. 4 (type), non figs. 1-3,6 (= Mortilletilamellaptychus breggiensis Renz & Habicht, 1985), non fig. 7 (= M. Description: Small to medium-sized valves, slightly ex gr. mortilleti Pictet & de Loriol, 1858), non fig. vaulted, without a keel or a lateral depression. Along the 5 (= Thorolamellaptychus sp.). symphysal margin in the terminal part of valves the ribs 1961 Lamellaptychus mortilleti (Pictet & Loriol) var. are straight, thin and densely spaced. They converge to the radiata Stefanov. - Stefanov, p. 219, 225-6, 227- symphysal margin at a relatively acute angle. 228, pi. 3, fig. 3. 2010 Beyrichilamellaptychus studeri (Ooster). - Distribution: The stratigraphie interval and area of distri­ M ë c h o v â et al., p. 242, fig. 9D (cum syn.). bution of B. beyrichi is very wide. T I j r c u l e t (2000) reported it from the Oxfordian to the Berrriasian. Material: One pair of valves, NMNHS F-30843 (see Ap­ pendix 1 for details).

Description: Small to medium-sized valves. Juvenile ribs are closely spaced. They converge almost subparallel along the symphysal margin. Adult (peripheral) ribs follow the shape of valves and are discordant to the juvenile ribs. 256 Zdenëk Vasîcek et al.

Genus Mortilletilamellaptychus M échovâ, H ousa & VaSi'Cek, 2010

Type species: Aptychus Mortilleti Pictet & de Loriol, 1858, p. 50.

Diagnosis: Thin-walled valves with thin ribs. In the close vicinity of the symphysal margin the majority of ribs bend towards the terminal point and end at the symphysal mar­ gin. Adult ribs end at the outer margin. In the stratigraphi- cally youngest representatives of this genus, the last few ribs may bend in a complicated way.

Mortilletilamellaptychus mendrisiensis mendrisien- sis (R enz & H abicht, 1985) Figs. 5.1a, b, 5.2

1961 Lamellaptychus mortilleti (Pictet & Loriol) var. longa Trauth. - Stefanov, p. 218-219, pi. 3, fig. 2. 1961 Lamellaptychus submortilleti Trauth. - Stefanov, p. 219-220, pi. 3, fig. 6. 1985 Lcimellptychus mendrisiensis new form; R e n z & H a b ic h t, p. 411, pi. 4, fig. 9. 2010 Mortilletilamellaptychus mendrisiensis mendri­ siensis (Renz & Habicht). - Mëchovâ et al., p. 249, fig. 9N (cum syn.). Fig. 4. Beyrichilamellaptychus studeri ( O o s te r ) , NMNHS F-30843; east of Todorchetata Village, Gabrovo District, Upper Valanginian, Kamchia Fm. Material: One single valve, NMNHS F-30824 and one pair of aptychi, NMNHS F-30834 (see Appendix 1 for details).

Measurements:. NMNHS F-30824: L = 14.5 mm. Lat = about 7 mm, Lat/L = 0.48.

Remarks: S te f a n o v (1961) referred the specimen to as Description: The valves are with a conspicuous keel and a Lamellaptychus mortilleti var. radiata. The thin, dense ribs lateral depression. The ribs between the keel and the sym­ along its symphysal margin, which were regarded by S t e ­ physal margin are quite straight and run obliquely towards fa n o v as radial lines, represent in fact the terminal parts of the symphysal margin. In its vicinity, they are S-shaped and fine juvenile ribs of the “beyrichi” type. converge along a rather longer section of the margin. Sev­ eral ribs end straight at the outer margin. Distribution: According to M è c h o v â et al. (2010) the strati­ graphie range of this species is from the Late Tithonian to Remarks: The right valve of NMNHS F-30834 is strongly the lower part of the Early Valanginian. deformed onto the bedding plane, incompletely preserved

Fig. 5. Some representatives of the genus Mortilletilamellaptychus. 1 - Mortilletilamellaptychus mendrisiensis men­ drisiensis (Renz & Habicht), NMNHS F-30834; Mecha polyana Village (not existing any more), Svoge municipality, Sofia District, Upper Valanginian, Salash Fm; a - note the left valve favourably illuminated; b - note the right valve favourably illuminated. The left valve belongs undoubtedly to M. m. mendrisiensis while the right valve, deformed into the plane of bedding with poorly visible symphysal area could be erroneously identified as M. mortilleti (Pictet & de Loriol). 2 - Mortilletilamellaptychus mendrisiensis mendrisiensis (Renz & Habicht), NMNHS F-30824, Kapinets Village, Antonovo municipality, Targovishte District, Lower Hauterivian, Hanevtsi Fm. 3 - Mortilletilamellaptychus bicurvatus (R e n z & H a b ic h t), NMNHS F-30812; south of the hamlet Hristova mahala, Troyan municipality, Lovech District, Upper Valangin­ ian, Cherniosam Fm. Taxonomy of Late Jurassic - Early Cretaceous aptychi 257 258 Zdenëk Vasicek et al.

Fig. 6. Mortilletilamellaptychus stefanovi n. sp., NMNHS F-30833, holotype with copy of the original label of Stefanov; Makotsevo Village, southeast of the peak Ademitsa, Gorna Malina municipality, Sofia District, Berriasian, Cherniosam Fm (Tithonian - Berriasian flysch).

in the symphysal area; its left valve, however, bears charac­ Remarks: The specimen NMNHS F-30827 previously teristic ribbing near the symphysal margin of terminal area, determined as Lamellaptychus mortilleti ( S te fa n o v 1961, what supports its reference to M. m. mendrisiensis (see fur­ pi. 3, fig. 4) is undeterminable because the valve is broken ther notes in the chapter 'Preservation, palaeoecologic and along the symphysal margin. taphonomic notes’). Distribution: Upper Valanginian of the Western Carpathi­ Distribution: Upper Valanginian of Switzerland (R e n z & an, Northern Calcareous Alps and Switzerland (M ë c h o v â H a b ic h t 1985) and Upper Valanginian to ?Lower Hauteriv­ et al. 2010). ian of Central Western Carpathians (V asîC ek et al. 1994).

Mortilletilamellaptychus stefanovi n. sp. Mortilletilamellaptychus bicurvatus (R enz & Fig. 6 H abicht, 1985) Fig. 5.3 non 1938 Lamellaptychus sub-mortilleti n. n. var. n. retroflexa. - T r a u t h , p. 201, pi. 14, fig. 6 (= 1938 Lamellaptychus suh-mortilleti n. n. var. n. ret- Mortilletilamellaptychus bicurvatus R e n z & roflexa. - T r a u t h , p. 201, pi. 14, fig. 6 (type), H a b ic h t, 1985). pars 1961 Lamellaptychus mortilleti (Pictet & Loriol). 1961 Lamellaptychus submortilleti T r a u t h var. - Stefanov, p. 217, pi. 3, fig. 1, non fig. 4 (= retroflexa Trauth. - Stefanov, p. 220, pi. 3, fig. undeterminable). 7. 1985 Lamellaptychus bicurvatus new name, R e n z & ?pars 1985 Lamellaptychus bicurvatus new name. - R e n z H a b ic h t, p. 409, pi. 3, figs. 25-28. & Habicht, pi. 3, fig. 27 only (non figs. 25-26, 2010 Mortilletilamellaptychus bicurvatus (R e n z & 28 = Mortilletilamellaptychus bicurvatus). Habicht). - Mëchovâ et al., p. 250, fig. IOC (cum syn.). Etymology: In honour of July Stefanov.

Material: One specimen, NMNHS F-30812 (see Appendix Holotype: Valve NMNHS F-30833 (35, t. 237 (M. Hr.) de­ 1 for details). scribed and illustrated by S te f a n o v (1961, p. 220, pi. 3, fig. 7: and re-illustrated herein in Fig. 6), and referred to Lamel­ Description: The juvenile ribs converge along the sym­ laptychus submortilleti var. retroflexa T r a u t h . physal margin. Adult ribs at first broadly bend back to the apex, and then are short, S-shaped in the vicinity of the Type horizon and locality: Berriassian (as part of the Ti- symphysal margin. thonian-Berriasian flysch) near Makotsevo Village, south­ Taxonomy of Late Jurassic - Early Cretaceous aptychi 259

east of the peak Ademitsa, Gorna Malina municipality, So­ cf. 1994 Lamellaptychus aplanatus aplanatus (Gilliéron). fia District. - V a sic e k et al., p. 76, pi. 23, fig. 6.

Material: The holotype only. Material: A single, relatively well preserved valve, NMNHS F-30865 (see Appendix 1 for details). Measurements: L = 14.2 mm, S = 13.0 mm, Lat = 9.5 mm, Lat/L = 0.67. The high value of the index is influenced by Measurements: NMNHS F- 30865: L = 19.3 mm, Lat = 9.4 the considerable deformation of the valve. mm, Lat/L = 0.49.

Diagnosis: Dense and thin ribs are simple on the flanks. Description: The valve is strongly convex, without a keel Double undulation of ribs in immediate vicinity of sym­ and without a depression. Simple, thin, dense ribs follow physal margin. the shape of the valve. Juvenile, imperfectly preserved ribs in the symphysal area are reminiscent somewhat of ribbing Description: The valve is small in size, secondarily flat, of the “beyrichi” type. Similar ribbing can also be seen in apparently wide. Ribs are thin and dense. The juvenile part specimens that are illustrated under the name Lamellapty­ of the valve is poorly preserved. Juvenile ribs are probably chus aff. aplanatus by Renz & Habicht (1985, pi. 3, figs. of the “mortilleti” type. Adult ribs are complicated in the 4, 7). Such juvenile ribbing differs somewhat from typical close vicinity of the symphysal margin, where they are S- representatives of T. aplanatus, and consequently, the speci­ shaped (“zig-zag bends” according to Stefanov). Along the men is left here in open nomenclature. symphysal margin, the ribs run to the terminal apex. The convexity of the valve is strongly affected by deformation Remarks: This is most probably the specimen mentioned onto the bedding plane. by S te f a n o v (1961: 223), assigned tentatively to Lamellap­ tychus herthae var. laevadsymphy salis, but which he did Remarks: Stefanov (1961) identified this valve with the not describe. Trauth’s subspecies Lamellaptychus submortilleti var. retroflexa. L ater R enz & H abicht (1985) selected Trauth’s Distribution: According to Renz & Habicht (1985) typi­ specimen as holotype of new species Lamellaptychus bi- cal specimens of Th. aplanatus come from Valanginian of curvatus. However, the specimen of Stefanov differs from Mediterranean area in Europe and of Atlantic area (Deap the valve of Trauth (1938), as well as from the prevailing Sea Drilling Project). majority of valves of R enz & H abicht (1985) by having a very short portion with S-shaped ribs and by not having the first bend of ribs widely convex. Only a single, incomplete Thorolamellaptychus trauthi (R enz & H abicht, 1985) valve, illustrated by them (R enz & H abicht 1985, pi. 3, fig. Fig. 7.2 27) seems to be close to the Bulgarian species. Because of its markedly higher stratigraphie position, it is referred here 1985 Lamellaptychus trauthi new form. - R e n z & to the new species with a question mark. H a b ic h t, p. 399, pi. 2, figs. 12 (type)-13. 2010 Thorolamellaptychus trauthi (Renz & Habicht). - Distribution: The holotype is from the Chernosiam For­ M ë c h o v â et al., p. 256, fig. 10K (cum syn.). mation, Berriasian of Bulgaria. The specimen of R e n z & H abicht (1985, pi. 3, fig. 27) comes from the Upper Valangin- Material: One specimen, NMNHS F-30853 (see Appendix ian of the Breggia section (Maiolica Formation), Switzerland. 1 for details).

Description: In the juvenile part the simple ribs are di­ Genus Thorolamellaptychus T u r c u l e t , 1994 rected towards the symphysal margin at an angle of about 45°. Above the keel, the adult ribs bend in a broad arch-like Type species: Aptychus thoro O p p e l, 1863, p. 250. manner back towards the apex. Near the symphysal facet, they bend sigmoidally. The end branch of the ribs points Diagnosis: Thin and dense ribs in the juvenile and the adult towards the terminal edge. stage follow the shape of the valves. All or almost all ribs end on the symphysal margin. More complicated ribbing at Distribution: The species is reported from Upper Berria­ final developmental stages. sian to Upper Valanginian of Switzerland, Western Car- parthians and in the Alps (for details see M ë c h o v â et al. 2010). Thorolamellaptychus cf. aplanatus (G illiéro n, 1873) Fig. 7.1 Genus Didayilamellaptychus T u r c u l e t , 1994 cf. 1873 Aptychus aplanatus Peters. - Gilliéron, p. Ill, pi. 10, figs. 3a-c, 4. T^pe species: Aptychus didayi C o q u a n d , 1841, p. 389. ? 1961 ? Lamellaptychus herthae ( W in k le r ) var. laevadsymphysalis Trauth. - Stefanov, p. 223. Diagnosis: Usually thick-walled, medium- to large-sized 260 Zdenëk Vasîcek et al.

1 cm

Fig. 7. Representatives of the genus Thorolamellaptychus. 1 - Thorolamellaptychus cf. aplanatus (Gilliéron), NMNHS F-30865; east of Makotsevo Village, Gorna Malina municipality, Sofia District, Berriasian. Cherniosam Fm. 2 - Thoro­ lamellaptychus trauthi (Renz & Habicht), NMNHS F-30853; north of Beli Osam Village, Troyan municipality, Lovech District, Upper Valanginian, Cherniosam Fm.

Fig. 8. Some representatives of the genus Didayilamellaptychus. 1 - Didayilamellaptychus didayi (Coquand), NMNHS F-30844; inner surface of valve with growth lines, close to the outer margin with serpulid encrustations, southwest of Chernevo Village, Varna District. Lower Hauterivian, Gorna Oryahovitsa Fm. 2 - Didayilamellaptychus didayi (Coquand) with sessile serpulids and bryozoan. NMNHS F-9925; countryside Teke dere near Shumen, Hauterivian, Gorna Oryahovitsa Fm. 3 - Didayilamellaptychus seranonis (Coquand), NMNHS F-30861, Zlatitsa river, Gaganitsa Village, Montana District, Upper Valanginian, Salash Fm. 4 - Didayilamellaptychus cf. seranonis (Coquand, 1841), NMNHS F-30818; corroded surface of valve, strong corroded ribs with pervading growth lines, Chernevo Village, Varna District, Lower Hauterivian, Gorna Oryahovitsa Fm. 5 - Didayilamellaptychus angulodidayi (T rau th ), NMNHS F-30850; west of Balgarski izvor Village, Lovech District, Lower Hauterivian. Gorna Oryahovitsa Fm. 6 - Didayilamellaptychus angulocostatus (Peters), NMNHS F-30831 with radial lines; southwest of Chernevo Village, Varna District, Lower Hauterivian, Gorna Oryahovitsa Fm. 7 - Didayilamellaptychus cf. angulocostatus (P eters), juvenile specimen NMNHS F-30840; Balgarski izvor Vil­ lage, Lovech District, Upper Valanginian. Gorna Oryahovitsa Fm. 8 - Didayilamellaptychus angulicostatus (P ic te t & de Loriol), NMNHS F-30816; Gaganitsa Village, Montana District, ?Hauterivian, Salash Fm. 9 - Didayilamellaptychus cristobalensis (O'Connell), NMNHS F-30817; southwest of Chernevo Village, Varna District, Lower Hauterivian, Gorna Oryahovitsa Fm. 10 - Didayilamellaptychus renzi Mëchovâ, Vasîcek & HouSa. NMNHS F-30856 with radial lines; Krap- chene Village, Montana District. Lower Hauterivian, Salash Fm. Specimen not coated with ammonium chloride before photographing. Taxonomy of Late Jurassic - Early Cretaceous aptychi 261 262 Zdenëk Vasicek et al. valves with strong ribs, only infrequently with thin ones. Adult ribs bend back in a curved to angular manner. In some specimens curved and angular ribbing occurs simultane­ ously. The last few ribs can be incomplete.

Didayilamellaptychus didayi (C oquand, 1841) Figs. 8.1, 8.2

1841 Aptychus Didayi (nobis). - C o q u a n d , p. 389, pi. 9, fig. 10. pars 1961 Lamellaptychus didayi (Coquand). - S te f a n o v , p. 216, pi. 2, figs. 1-3, 5-7, non fig. 4 (=Didayilamellaptychus cf. didayi). 1961 Lamellaptychus subdidayi Trauth. - S te f a n o v , p. 217, pi. 2. figs. 8-10. 2000 Lamellaptychus (Didayilamellaptychus) didayi didayi (Coquand). - Turculet, p. 121, pi. 22, figs. 14-18. 2010 Didayilamellaptychus didayi (Coquand). - M ë c h o v â et al., p. 257, fig. 11A (cum syn.). Fig. 9. Didayilamellaptychus cf. didayi (Coquand), Material: Fourteen specimens, NMNHS F-30810, 30811, NMNHS F-30847 in the body chamber of Neolissoceras 30813, 30828, 30837, 30844, 30852, 388*, 30864, 30835, grasianum (d'Orbigny), NMNHS F-30848, from north of 9925*, 9926*, 30822, 30820 (see Appendix 1 for details). the hamlet Enchovtsi, Troyan municipality, Lovech District, Lower Hauterivian, Kamchia Fm. Measurements: Specimen NMNHS F-30852 is the largest among the ribbed aptychi: L = about 31.5 mm.

Description: The valves are wide, with a prominent keel and a shallow lateral depression. The strong and widely spaced ribs begin to bend in a broad arch back towards the poorly preserved. Because of the considerable imperfec­ apex, approximately, at the area of lateral depression. The tion of preservation of the symphysal area, the width of the width of the valves is notable. In the lateral depression area, valve, rough ribbing and marked (fractal) bend of ribs on the ribs are bent in a sigmoidal pattern. the flank of the valve, the determination of this specimen is uncertain. We tentatively assign it into group of Didayil­ Distribution: Lower Valanginian to uppermost Lower Hau- amellaptychus didayi (C o q u a n d ). terivian in the Mediterranean area (for details see M ë c h o v â et al. 2010). Didayilamellaptychus seranonis (C oquand, 1841) Fig. 8.3 Didayilamellaptychus cf. didayi (C oquand, 1841) Fig. 9 1841 Aptychus Seranonis (nobis). - C o q u a n d , p. 390, pi. 9, fig. 13. pars 1961 Lamellaptychus didayi (Coquand). - Stefanov, pars 1961 Lamellaptychus angulocostatus ( P e te r s ) var. p. 216, pi. 2, fig. 4, non pi. 2, figs. 1-3, 5-7 atlantica (H e n n ig ), transition to Lamellaptychus (=Didayilamellaptychus didayi). seranonis (Coquand). - Stefanov, p. 215, pl. 1, 1961 Lamellaptychus beyrichi (O ppel) var. fig. 12, non fig. 8 (=Didayilamellaptychus cf. fractocosta Trauth. - Stefanov, p. 221, pi. 3, angulocostatus ( P e t e r s , 1854). figs. 8-9. 1972 Lamellaptychus I. - T h o m so n , p. 35, fig. 2a, c, ?b. Material: Three specimens, NMNHS F-30851, 30829, 2010 Didayilamellaptychus seranonis (Coquand). — 30847 (see Appendix 1 for details). M ë c h o v â et al., p. 258, fig. 1 IB (cum syn.).

Remarks: Stefanov (1961, pi. 3, fig. 9, see also here Fig. Material: One specimen, NMNHS F-30861 (see Appendix 9) illustrated the aptychus N M N H S F-30847 occurring in 1 for details). the body chamber of an ammonite determined correctly as Neolissoceras grasianum (d ’O rb ig n y ). Neolissoceras be­ Description: Slender valves, with a keel and a lateral de­ longs to the family Haploceratidae. The body chamber of pression. In maturity, the ribs are strong and spread apart. the ammonite and especially the valve of the aptychus are The ribs between the keel and the symphysal margin bend Taxonomy of Late Jurassic - Early Cretaceous aptychi 263 back in a curved to subangular manner towards the apex. 2000 Lamellaptychus (Didayilamellaptychus) angu­ In the lateral depression, the ribs may be slightly to sigmoi­ locostatus angulocostatus (Peters) f. typ. dally bent. Trauth. - Turculet, p. 126, pi. 23, figs. 7, 9-10, 17, ? figs. 11, 13, 15-16, non fig. 12 (= D. renzi Distribution: From the uppermost Lower Valanginian to M ë c h o v â et al.), non fig. 14 (= D. angulicostatus the Lower Hauterivian of the Mediterranean area (for more Pictet & de Loriol), pi. 24, figs. 2, 4-5, non figs. details see M ë c h o v â et al. 2010). 3, 6 (= D. renzi M ë c h o v â et al.). 2000 Lamellaptychus (Didayilamellaptychus) angu­ Didayilamellaptychus cf. seranonis (C oquand, 1841) locostatus longus Trauth. - Turculet, p. 126, Fig. 8.4 pi. 24, fig. 1. 2010 Didayilamellaptychus angulocostatus ( P e te rs ). - M ë c h o v â et al, p. 260, fig. 1 IF (cum syn.). 1961 Lamellaptychus seranonis (C o q u a n d ) var. radiata n. var. - Stefanov, p. 221, 226, 228, pi. 3, fig. 5. Material: Seven specimens, NMNHS F-30809; 30814, 30819, 30830, 30831, 9905*, 9906*. (see Appendix 1 for Material: Two specimens, NMNHS F-30818, 30842 (see details). Appendix 1 for details). Description: Valves with a keel but without a lateral de­ Remarks: Specimen NMNHS F-30818 is a corroded frag­ pression. Ribs on the flank of valves run subparallel to the mental valve, which S te f a n o v (1961: 221) described as a symphysal margin. In the area of the keel, the ribs are an­ new variety of the species seranonis. We think, however, gular. that the differences, pointed out by the author, are caused by corrosion and do not justify defining a new variety. Remarks: In specimen NMNHS F-30819 the right valve is deformed by lateral pressure, so it would belong to var. “longa ”; while the left valve is with a broken arch, pressed Didayilamellaptychus angulodidayi (T r a u th, 1938) into the bedding plane, so it would belong to var. “lata”. Fig. 8.5 These observations suggest that the varieties longa and lata are not supported in taxonomy and are result of deforma­ 1849 Aptychus Didayi Coquand. - Quenstedt, p. 314, pi. tion. The specimen NMNHS F-30831 shows radial lines 22, fig. 21a, b. (Fig. 8.6). 1938 Lamellaptychus angulo-didayi n. n. f. typ. - T r a u t h , p. 212, pi. 14, figs. 28-29. Distribution: According to Gasiorowski (1962), this spe­ 2010 Didayilamellaptychus angulodidayi (Trauth). - cies and other morphologically related forms occur in a M ë c h o v â et al., p. 260, fig. 11E (cum syn.). wide stratigraphie range - from the Berriasian to the Bar- remian, while according to M ë c h o v â et al. (2010) the spe­ Material: One specimen, NMNHS F-30850 (see Appendix cies is known only from the Late Hauterivian. 1 for details).

Description: Valve with a prominent keel and lateral de­ Didayilamellaptychuscf.angulocostatus(PETERS,\^54) pression. In the juvenile part of valve, the ribs are angular; Fig. 8.7 whereas subangular arch-shaped ribs are observed in the adult part of the valve, which end on the symphysal facet. pars 1961 Lamellaptychus angulocostatus (Peters). - S te f a n o v , p. 212, pl. 1, fig. 4. Distribution: The author of this species placed the type pars 1961 Lamellaptychus angulocostatus ( P e te r s ) var. material in the Neocomian. M ë c h o v â et al. (2010) reported atlantica (Hennig). - Stefanov, p. 215, pl. 1, fig. a distribution from the Upper Valanginian to the Lower 8, non fig. 12 (=Didayilamellaptychus seranonis Hauterivian in the Western Carpathians. (C o q u a n d , 1841).

Material: Three specimens, NMNHS F-30832, 30840, Didayilamellaptychus angulocostatus (P ete r s, 1854) 30845 (see Appendix 1 for details). Fig. 8.6 Remarks: Specimen NMNHS F-30840 is a juvenile valve, 1854 Aptychus angulocostatus Peters. - Peters, p. with a well preserved terminal area. Specimen NMNHS 441. F-30845 is also a juvenile valve, very poorly preserved. pars 1961 Lamellaptychus angulocostatus (Peters). - Specimen NMNHS F-30832 is a poorly preserved fragment. S te f a n o v , p. 212, pl. 1, figs. 1-3, ? 4, non fig. 6 (=Didayilamellaptychus angulicostatus ( P i c t e t & de Loriol, 1858). Didayilamellaptychus angulicostatus (P ictet & de 1974 Lamellaptychus angulocostatus (Peters). - L orio l, 1858) H o u s a , p. 31, pi. 7, figs. 1-4 (type), 5-7. Fig. 8.8 264 Zdenëk Vasîcek et al.

1858 Aptychus angulicostatus. - Pictet & de Loriol, p. 46, pl. 10, fig. 3 (type), ?6-12, non figs. 4-5 (= Didayilamellaptychus cf. renzi M ë c h o v â , VaSi'Cek & Housa, 2010). 1910 Aptychus angulicostatus P ic t. and de Loriol. - K ilia n , pl. 5, fig. 2b. non fig. 2a (= D. cf. renzi). pars 1961 Lamellaptychus angulocostatus (Peters). - Ste­ fa n o v , pl. 1. fig. 6. non figs. 1-3 (-Didayilamel- laptychus angulocostatus ( P e te r s , 1854), non fig. 4 (=Didayilamellaptychus cf. angulocostatus). 1961 Lamellaptychus angulocostatus ( P e te r s ) var. fractocosta Trauth. - Stefanov, pl. 1, fig. 5. 2000 Lamellaptychus (Didayilamellaptychus) angu­ locostatus ( P e te r s ) /. typ. T r a u t h . - T u r c u l e t , pl. 23, fig. 14. 2010 Didayilamellaptychus angulicostatus ( P i c t e t and Loriol). - Mëchovâ et al., p. 262, fig. 11G. H (cum syn.).

Material: Four specimens, NMNHS F-30815, 30816, 30826, 30846 (see Appendix 1 for details).

Measurements: Among all specimens, NMNHS F-30846 is a typical one: L = 26.6 mm, Lat = 12.0 mm, Lat/L = 0.45.

Description: Valves with a prominent keel and lateral de­ pression. In the region between the keel and the symphysal margin, the ribs are angularly bent and run back towards the apex. In the depression area, the ribs are gently inflected to sigmoidally bent.

Distribution: Upper part of Upper Hauterivian of whole Mediterranean area (Mëchovâ et al. 2010).

Didayilamellaptychus cristobalensis (O'C o n nell, 1921) Fig. 8.9 Fig. 10. Didayilamellaptychus filicostatus (Stefanov), NMNHS F-30836, holotype with copy of the original label of S te f a n o v ; Ropot Village (west of Komshtitsa Village), 1921 Aptychus cristobalensis, new species. - O'Connell, p. 7, figs. 7 (type)-8. Godech municipality, Sofia District, Lower Hauterivian, Salash Fm. 1961 Lamellaptychus angulocostatus ( P e te r s ) var. cristobalensis (O’Connell). - Stefanov, p. 213, pi. Lfig. 11. 2010 Didayilamellaptychus cristobalensis (O’Connell). - M ë c h o v â et al., p. 263, fig. 11J (cum syn.).

Material: Two specimens, NMNHS F-30817, 9904* (see Appendix 1 for details). Didayilamellaptychus filicostatus (Stefanov, 1961 ) Fig. 10 Description: The valves are with keel and a faint lateral de­ pression. The juvenile ribs are bent angularly in the area of 1961 Lamellaptychus angulocostatus ( P e te r s ) var. keel. The adult ribs become arch-shaped to rounded. Their filicosta n. var. - S te f a n o v , p. 214, 225, 227, pl. 1, arrangement between the keel and the symphysal margin is fig. 9. complicated by less distinct to evident crenulation. 2010 Didayilamellaptychus filicostatus (Stefanov). - M ë c h o v â et al., p. 266, fig. 1 IL (cum syn.). Distribution: The type material comes from Caribbean area and its stratigraphical position is not clear. In the Slo­ Material: One incomplete specimen, the holotype NMNHS vakian Carpathians the species is known from the Upper F- 30836, from Ropot Village, Godech municipality, Sofia Hauterivian (M ë c h o v â et al. 2010). district (see Appendix 1 for more details). Taxonomy of Late Jurassic - Early Cretaceous aptychi 265

Measurements: L= 13.5 mm.

Description: The valve is of medium size with an indistinct keel and without a lateral depression. Closely spaced and relatively thin ribs on the flank run subparallel to the sym­ physal margin. In the area of the keel, the juvenile ribs are angularly fractured at an acute angle. The angle between the branches of angular fractured ribs is almost symmetric. The adult ribs become gradually less symmetric. The last few ribs in the terminal area are rounded.

Distribution: The holotype is from the Lower Hauterivian ( S te f a n o v 1961). The species is also reported from the Up­ per Hauterivian of Spain and the Carpathians ( M ë c h o v â et al. 2010).

Didayilamellaptychus renzi Mëchovâ, Vasicek & H o u sa 2010 Fig. 8.10

1961 Lamellaptychus angulocostatus ( P e te r s ) var. radiata Trauth. - Stefanov, p. 213, pl. 1, figs. 7, 10; text-figs. 1-2. 1972 Lamellaptychus angulocostatus atlanticus (H e n - nig). - Renz, p. 617, pi. 4, figs. 2a, b, 3 (type), ?4. 2000 Lamellaptychus (Didayilamellaptychus) angulo­ Fig. 11. Laevaptychus longus (v. Meyer), NMNHS F-339; costatus angulocostatus ( P e te r s ) /. typ. T r a u t h . - Cherni Osam Village, Troyan municipality, Lovech Dis­ T u r c u l e t , pi. 23, fig. 12, pi. 24 figs. 3, 6. trict, Tithonian, Cherniosam Fm. 2010 Didayilamellaptychus renzi Mëchovâ, Vasi'Cek & Housa. - Mëchovâ et al., p. 266, fig. 11M , N (cum syn.).

Material: Four specimens, NMNHS F-30856, 30857, 30858, 30859 (see Appendix 1 for details).

Measurements: NMNHS F-30856: L = 23.4 mm, Lat = 12.5 mm, S = 22.0 mm, Lat/L = 0.53. (1960: 91) subdivided the Jurassic representatives of the ge­ nus Laevaptychus into 7 subgenera. He followed the original Description: The valves are with prominent keel and, usu­ idea of T r a u t h (1931). The most suitable parameters meas­ ally, with a shallow lateral depression. The ribs are thin, ured on the valves seem to be: L, S, W (= Lat in Lamellap- closely spaced, and bend at an acute angle. They reach sym­ tychidae), pW, and ratios of these values (S/L, pW/L, W/L), physal margin at an angle of about 10°. The last few ribs as illustrated e.g. in text-fig. 2 in Turculet & Grigore 2006. usually lose the angular character. The patterns are similar to the parameters illustrated on Fig. 2 herein. However, Gasiorowski (1960) did not give proper Distribution: According to M ë c h o v â et al. (2010) D. renzi descriptions of his subgenera. Gasiorowski’s classification, occurs in the Hauterivian across a relatively vast area (from however, without closer analyse, was used, subsequently, by the Caribbean and the Atlantic to Europe (Bulgaria, the Turculet & Grigore (2006). The genus Laevaptychus has Western Carpathians, Switzerland and Spain). not been revised for the last 50 years, thus its family affilia­ tion remains unclear.

Family uncertain Laevaptychus longus (v. M e y er, 1831) Genus Laevaptychus T ra uth, 1927 Fig. 11 Type species: Aptychus meneghinii d e Z ig n o , 1870, p. 11. 1831 Aptychus laevis longus. - V. M e y e r, p. 128, pi. 59, Diagnosis: Thick-walled calcareous aptychi. Cellular struc­ figs. 6-7. ture observed on otherwise smooth surface. 2006 Laevaptychus (Latuslaevaptyclius) longus longus (Meyer). - Turculet & Grigore, p. 30, pl. 1, figs. Remarks: On the basis of the shape of valves Gasiorowski 4-6 (cum syn.). 266 Zdenëk Vasicek et al.

Material: A single, complete left valve, NMNHS F-339* terminal apex is visible. Using this feature only, the (For details see Appendix 1). valve could be determined as Mortilamellaptychus mortilleti (P ictet & de L oriol). The opposite valve, Measurements: NMNHS F-339 is the largest in the col­ lection. The measured parameters and calculated indexes, however, has preserved ribs in the terminal area that which are designated partly differently from those of lamel- run along the symphysal margin in a section longer laptychi, are based on the schematic illustration of T u r c u l e t than in the case of Mortilamellaptychus mortilleti & G r ig o r e (2006, text-fig. 2). The length L = 52.2 mm, the (Fig. 5.1a). Because of this feature, the valve is here distance between the apex and the terminal point S = 44.1 referred to Mortilletilamellaptychus mendrisiensis mm, the maximum valve width W = 35.3 mm, the distance mendrisiensis (R enz & H abicht). between the terminal point and the projection of point of In summary, both maximum width pW = 33.0 mm; W/L = 0.67, pW/L = 0.63, valves belong to M. m. mendrisiensis. This example S/L = 0.84. emphasizes the importance of preservation of the ter­ minal and symphysal area of the valves for correct de­ Description: The valve is of large size, thick-walled, with termination. elongated outline. The venter is apparently smooth; how­ Another problem connected with the preservation ever, its cellular structure is perceptible on the surface and is easily seen under stereomicroscope. of valves can be the partial dissolution of the valve calcareous surface layer. This is well demonstrated, Remarks: Valve morphology and calculated indexes cor­ for instance, in the otherwise well preserved speci­ respond to L. longus. According to Gasiorowski’s classifica­ men NMNHS F-30841, on which the punctate layer tion L. longus belongs to subgenus Latuslaevaptychus. is well preserved only in a small area. Such partial Distribution: L. longus is known from many European dissolution is even more obvious on the incomplete, countries. Trauth (1931) reported it from the Oxfordian to large valve of specimen NMNHS F-335, which lacks Tithonian in autochthonous deposits of England, France, almost entirely the punctate layer (Fig. 3.2). In the lat­ Switzerland, Germany, and in nappes of the Western and ter specimen, when only the ribbing of the symphysal Eastern Alps and the Carpathians. Turculet & G rigore area (with non-preserved punctate layer!) is consid­ (2006) reported ?Oxfordian to Lower Tithonian occurrenc­ es in the area of Svinita in Romania. ered, the valve would be assigned to Lamellaptychus rectecostatus (P eters, 1854). However, with regard to the strength of ribs and the size of the valve, it is rather 5. Preservation, palaeoecologic and close to Punctaptychus rectecostatus Cuzzi, 1962. Fi­ taphonomic notes nally, because of the dissolution phenomenon and the The preservation of the studied aptychi varies consid­ incompleteness of the valve, specimen NMNHS F-335 erably. Both valves of the pair are preserved together is here tentatively determined as Punctaptychus sp. very rarely (only in three cases). In the two of the paired In lamellaptychi the corrosion of valves causes specimens both valves are similarly preserved. In one sometimes the growth lines between the ribs to be case, however, because of preservational deformation, more conspicuous. However, deeper corrosion may af­ the valves of the pair differ (NMNHS F-30834, see fect also the ribbing so that the shape of ribs becomes Fig. 5.1a, b, and Stefanov 1961, pl. 1, fig. 3 and pi. apparently variable (see Fig. 8.4). Thus, a higher de­ 3, fig. 6); one of the valves of the pair has a broken gree of dissolution may cause the ribbing to appear arch and is pressed into the bedding plane; whereas the ostensibly simpler than they actually were, similarly to other valve has a keel that is more distinct as a result the simplification of suture lines in strongly corroded of the deformation. The compressed valve is secondar­ ammonite shells. ily widened. If they have been found separately, these The inner surface of calcareous aptychi, if ex­ two valves of one species, according to e.g. T ra u th’s posed, exhibits dense growth lines. If this side is well (1938) classification, would be referred to two different preserved, as in NMNHS F-30844 (Fig. 8.1), non- taxa; the wider form - to the variety lata, whereas the uniform growth lines are observed. The same speci­ narrower one - to the variety longa. men, NMNHS F-30844, was photographed from the The pair of valves illustrated on Fig. 5.1a, b differs opposite side by S tefanov (1961, pi. 2, fig. 2). Multiple also in the preservation of symphysal margin in the thin lines alternate with single thicker lines at regular terminal area. The contact zone between both valves intervals, which is interpreted as periodicity in animal in the flat valve is slightly submerged below the other growth. valve and is corroded. Consequently, only the bend of Some of the specimens possess on the outer surface ribs in close vicinity of symphysal margin towards the of valves radial lines, having a form of very thin ribs Taxonomy of Late Jurassic - Early Cretaceous aptychi 267

that can be photographed only under suitable illumina­ Acknowledgements

tion (e.g., Fig. 8.6). (The same valve was illustrated by We are grateful to Georgi Mandov, Todor Nikolov and S tefanov (1961, pl. 1, fig. 2), but without obvious radial Platon Tchoumatchenco (Sofia) for their help to clarify lines). In some specific cases, these fine radial lines do and update the information about the provenance of some of not take any clear morphological effect, thus whiten­ the specimens and for help in deciphering the old museum ing with ammonium chloride makes them invisible in labels. H risto Stoynov (Bellevue, Washington, U S A ) kind­ ly improved the English of an earlier version of this paper. photographs. Because of their black colour (probably We acknowledge the valuable help of the photographer K. due to the remains of original, now carbonized organ­ Mezihorâkovâ (Ostrava) who prepared all photographs with ic matter), such lines can be seen on the photographs the exception of the illustrated specimens on Fig. 9 (made only if the specimen is not whitened (Fig. 8.10). by N .M -D .). We are grateful to Jim Kennedy (Oxford) and If, after the death of the living organism (the host Jozef M tchalik (Bratislava) for their helpful comments on earlier versions of the manuscript. ammonite), the isolated valves lay on the muddy bot­ tom for a sufficiently long period before burial, they could be colonized by sessile organisms as document­ References ed on a valve in Fig. 8.2. In this case serpulids and a bryozoan are attached to the valve. A similar, but less B oorovâ, D., L obitzer, H., Skupien, P. & Vasicek, Z. (2000): Biostratigraphy and facies of Upper Jurassic-Lower clear example is a side of the valve with growth lines Cretaceous pelagic carbonate sediments (Oberalm-, illustrated in Fig. 8.1. Schrambach- and Rossfeld Formation) in the Northern Calcareous Alps, South of Salzburg. - Abhandlungen der Geologischen Bundesanstalt, 56: 273-318. 6. Conclusions C o q u a n d , M. (1841): Mémoire sur les Aptychus. - Bulletins As a result of the present taxonomic reappraisal of de la Société géologique de France, 12: 376-391. Cuzzi, G. (1962): Osservazioni sul genere Punctaptychus e the Late Jurassic and Early Cretaceous aptychi from sulla specie Punctaptychus punctatus ( V o ltz ) f. typ. - Bulgaria, 23 taxa from the species group, which are Bolletino délia Società paleontologica Italiana, 1: 43-51. assigned to six genera within the families Punctapty- E n g e s e r , T. & K eupp, H. (2006): Phylogeny of the aptychi chidae, Lamellaptychidae and one uncertain family, possessing Neoammonoidea (Aptychophora nov., Ce­ were recognised. One new species Mortilletilamel- phalopoda). - Lethaia, 34: 76-96. Gasiorowski, S.М. (1959): Succession of Aptychi faunas laptychus stefanovi was erected. Apart from the holo­ in the Western Tethys during the Bajocian-Barremian type of M. stefanovi sp. nov., the Bulgarian collection time. - Bulletin de Г Académie polonaise des Sciences, contains also the holotype of Didayilamellaptychus Série des Sciences de géologie et géographie, 7 (9): 715- filicostatus (Stefanov), which was previously elevated 722. from a variety to a species level by M ëchovâ et al. G a s io r o w s k i, S.M. (1960): Remarques sur les laevaptychi. - Rocznik Polskiego Towarzystwa geologicznego, 30: (2010). Our revision has shown that J uly Stefanov’s 59-97. taxonomic work was precise enough and he had proc­ G a s io r o w s k i, S.M. (1962): Sur les Aptychi a côtes. - Rocz­ essed the collection accurately in accordance with the nik Polskiego Towarzystwa geologicznego, 32: 227-280. aptychi classification at that time. G il l i é r o n , V. (1873): Aperçu géologique sur les Alpes de A valve of Didayilamellaptychus cf. didayi occur­ Fribourg en général et description spéciale du Monsal- ring in the body chamber of an ammonite Neolissocer­ vens. - Matériaux pour la géologie de la Suisse, 12: 1-232. as grasianum (see Stefanov 1961, pi. 3, fig. 9, and Fig. H o u s a , V. (1974): Los Apticos de Cuba I: Lamellaptychus 9 herein) is of special interest. Neolissoceras belongs angulocostatus (P e t.). - Academia de Ciencias de Cuba, to the family Haploceratidae. This finding is a further Serie Geologica, 14: 1-57. confirmation of the hypothesis that both lamellaptychi K ilia n , W. (1910): Erste Abteilung - Unterkreide (Palaeocre- and ?punctaptychi belong to the family Haploceratidae tacicum), Lieferung 2: Das bathyale Palaeocretacicum im südostlichen Frankreich; Valendis-Stufe, Hauterive- Z ittel , 1884 in the natural system of ammonites. Stufe, Barreme-Stufe, Apt-Stufe. - In: F r e c h , F. (Ed.): The state of preservation may strongly affect the Lethaea Geognostica, II, Mesozoikum, Band 3 (Kreide) correct determination of aptychi. Only in exceptional (1907-1913), 169-288; Stuttgart (Schweizerbart). cases like the illustrated occurrence of both valves in Kratochvilovâ, L. (2004): Spodnokndové zebrované ap- a pair with different state of preservation (Fig. 5.1a, tychy - jejich klasifikace, fylogenetické vztahy a stra- tigrafické vyuziti (vnëjsf a centrâlnf Zâpadm Karpaty b) this preservation phenomenon can be revealed and a Severnf vâpencové Alpy). - Unpublished Ph.D. The­ wrong determination of any of the valves could be sis, 155 pp.; Ostrava (VSB - Technickâ univerzita). [in possibly avoided. Czech], 268 Zdenëk Vasîcek et al.

L e h m a n n , U. (1972): Aptychen als Kieferelemente der Am- S te f a n o v , J. (1961): Amonitni operkulumi (aptichi) ot dol- moniten. - Palâontologische Zeitschrift, 46: 34-48. nata kreda na Bâlgarija. [Ammonoid operculums (apty­ L e h m a n n , U. & K u lic k i, C. (1990): Double function of chi) from the Lower Cretaceous of Bulgaria]. - Travaux aptychi (Ammnoidea) as jaw elements and opercula. - sur le géologie de la Bulgarie, Série Paléontologie, 3: Lethaia, 23: 325-331. 209-235 [in Bulgarian with Russian and English sum­ M ë c h o v â , L., H o u s a , V. & V a s îc e k , Z. (2008): Contribution maries]. to the systematics of Lower Cretaceous ribbed aptychi. T e n c h o v , Y. (1993): Glossary of the formal lithostratigraph- - Abstracts of the 9th Palaeontological Conference, 10- ic units in Bulgaria (1882-1992). - 937 pp.; Sofia (BAS 11 October 2008, 57-59; Warszawa (Polish Academy of editorial house) [in Bulgarian], Sciences, Institute of Paleobiology). T h o m so n , M.R.A. (1972): Lower Cretaceous Lamellapty­ M ë c h o v â , L., V a s îc e k , Z. & H o u s a V. (2010): Early Creta­ chus (Aptychi, Ammonoidea) from the south-eastern ceous ribbed aptychi - a proposal for a new systematic Alexander Island. - Bulletin of the British Antarctic classification. - Bulletin of Geosciences, 85: 219-274. Survey, 30: 35-40. M e y e r , H. v. (1831): Das Genus Aptychus. - Verhandlungen T r a u t h , F. (1927): Aptychenstudien I. Über die Aptychen der kaiserlich Leopoldinisch-Carolinischen Akademie im Allgemeinen. - Annalen des Naturhistorischen Mu­ der Naturforscher, 15: 125-170. seums in Wien, 41: 171-259. O’Connell, М. (1921): New species of ammonite opercula T r a u t h , F. (1931): Aptychenstudien VII. Die Aptychen des from the Mesozoic rock of Cuba. - American Museum Malm und der Unterkreide. - Annalen des Naturhistori­ Novitates, 28: 1-15. schen Museums in Wien, 45: 22-136. O o s te r , W.A. (1857): Catalogue des Céphalopodes fossiles T r a u t h , F. (1935): Die Punctaptychi des Oberjura und der des Alpes suisses, II partie: Céphalopodes d ordres in­ Unterkreide. - Jahrbuch der Geologischen Bundesan- certains. - Nouveaux Mémoires de la Société helvétique stalt, 85: 309-332. des sciences naturelles: 14-32. T r a u t h , F. (1938): Die Lamellaptychi des Oberjura und der O p p e l, A. (1863): Über jurassische Cephalopoden. - Pa- Unterkreide. - Palaeontographica, (A), 88: 118-240. laeontologische Mittheilungen aus dem Museum des T u r c u le t , I. (1994): Asupra oportunitâtii separârii de pa- koeniglich Bayerischen Staates, 1 (3): 127-266. rasubgenuri in cadrul paragenelui Lamellaptychus Oppel, A. (1865): Die tithonische Etage. - Zeitschrift der (Cephalopoda, Ammonoidea). - Studii §i Cercetâri de Deutschen geologischen Gesellschaft, 17: 535-558. Geologie, Geofizicâ, Geografie, Serie Geologica, 39: Parkinson, J. (1811): The Fossil Starfish, Echini, Shells, 119-126. Insects, Amphibia, Mammalia & c. - The Organic Re­ T u r c u l e t , I. (2000): Aptihii din România. - 178 pp.; mains of a Former World, 3: 479 pp.; London (Sher­ Bucuresti (Editura Academiei Române). wood, Neely & Jones). T u r c u l e t , I. & G r ig o r e , D. (2006): New Upper Jurassic ap­ P e t e r s , K. (1854): Die Aptychen der ôsterreichischen tychi from the Svinita area (South Carpathians, Roma­ Neocomien- und oberen Juraschichten. - Jahrbuch der nia). - Muzeul Oltenei Craiova. Studii §i comunicâri, kaiserlich-koniglichen geologischen Reichsanstalt, 5: §tiinjele Naturii, 22: 28-39. 439-444. Vasîcek, Z. & Hoedemaeker, PJ. (1997): Aptychi from the P i c t e t , F.J. & Loriol, P. de (1858): Description des fossiles Lower Cretaceous strata along Rio Argos (Caracava, SE contenus dans les terrains néocomiens des Voirons. - Spain). - Scripta Geologica, 115: 29-45. Matériaux pour la paléontologie de la Suisse, 2: 1-64. Vasîcek, Z., M ichalik, J. & R e h â k o v â , D. (1994): Early Cre­ Quenstedt, F.A. (1845-1849): Petrefactenkunde Deutsch- taceous stratigraphy, palaeogeography and life in West­ lands. Erste Abteilung, erster Band. Die Cephalopoden. ern Carpathians. - Beringeria, 10: 1-170. - 580 pp.; Tübingen (Fues). W r ig h t, C.W., Callomon, J.H. & H o w a r th , M.K. (1996): R e n z , O. (1972): Aptychi (Ammonoidea) from the Upper Cretaceous Ammonoidea. - In: K a e s l e r , R.L. (Ed.): Jurassic and Lower Cretaceous of the western North At­ Treatise on Invertebrate Paleontology, part L, Mollusca lantic (Site 105, Leg 11, DSDP). - In: H ollister, C.D. & 4 Revised. - 1-362; Boulder & Lawrence (Geological E w in g , J.I. (Eds.): Initial Reports of the Deep Sea Drill­ Society of America & University of Kansas Press). ing Project, 17: 607-629. Zagorchev, I., Dabovski, H., & N ik o lo v , T. (2009): Geology R e n z , O. (1983): Early Cretaceous Cephalopoda from the of Bulgaria. Vol. II. Mesozoic Geology. - 766 pp.; Sofia Blake-Bahama Basin (Deep Sea Drilling Project, Leg (Prof. M. Drinov Academic Press) [in Bulgarian, with 76, Hole 534A) and their correlation in the Atlantic and English summary]. southwestern Tethys. - In: S h e r id a n , R.E. & Gradstein, Z ig n o , A. d e (1870): Annotazioni paleontologiche. - Memo- F.M. (Eds.): Initial Reports of the Deep Sea Drilling rie del Reale Istituto Veneto di Scienze, Lettere ed Arti, Project, 76: 639-644. 15: 1-27. R e n z , O. & H a b ic h t, K. (1985): A correlation of the Tethys Z i t t e l , K.A. (1868): Palaeontologische Studien ueber die Maiolica Formation of the Breggia section (southern Grenz-Schichten der Jura- und Kreide-Formation im Switzerland) with Early Cretaceous coccolith oozes of Gebiete der Karpathen, Alpen und Apenninen. Abthei- Site 534A, DSDP Leg 76 in the western Atlantic. - Ec- lung I. Die Cephalopoden der Stramberger Schichten. - logae geologicae Helvetiae, 78: 383-431. In: O p p el, A. & Z i t t e l , K.A. (Eds.): Die Cephalopoden Schindewolf, O.H. (1958): Über Aptychen (Ammonoidea). der Stramberger Schichten. - Palaeontologische Mitt­ - Palaeontographica, (A), 111: 1-46. heilungen aus dem Museum des koeniglich Bayerischen Staates, 2: 1-118. Z ittel, К.A. v. (1884): Cephalopoda. - In: Z ittel, K.A. v. (Ed.): Handbuch der Palaontologie, Band 1, 2 (3): 329- 522; München & Leipzig (Oldenbourg).

Manuscript received: March 9th, 2012. Revised version accepted by the Stuttgart editor: May 28th, 2012.

Addresses of the authors:

Z denëk Vasicek, Institute of Geonics of Academy of Sci­ ences of the Czech Republic, Studentskâ 1768, 708 00 Os­ trava - Poruba, Czech Republic; e-mail: [email protected] N eda M otchurova-D ekova (corresponding author), Natio­ nal Museum of Natural History, 1, Tsar Osvoboditel Bvd, Sofia 1000 & Department of Geology and Palaeontology, University of Mining and Geology, Sofia 1700, Bulgaria; e-mail: [email protected] A ntoaneta Ilcheva, National Museum of Natural History, 1, Tsar Osvoboditel Bvd, Sofia 1000, Bulgaria; e-mail: [email protected] L ubomir M etodiev, Geological Institute, Bulgarian Aca­ demy of Sciences, Acad. G. Bonchev Str., Bl. 24, 1113 Sofia, Bulgaria; e-mail: [email protected] Taxonomy of Late Jurassic - Early Cretaceous aptychi 269 270 Zdenëk Vasicek et al.

Appendix

Appendix 1. Annotated list of all studied material of aptychi.

Museum Names appearing number on the old museum NMNHS F- Locality Horizon Illustrated in Remarks labels and/or in (original old S t e f a n o v (1961) number)

Punctaptychus punctatus (Z ittel, 1868) 30839 (7087, t. Lamellaptychus Borushtitsa River, Gabrovo Berriasian, Kamchia Fm 7371) inflexicosta T r a u t h District 30841 (1269) Northwest of Stubel Village, Unknown, Salash Fm Montana District 337* Punctaptychus Shipkovski Mineral Baths, Tithonian, Cherniosam punctatus V o l t z Lovech District Fm

Punctaptychus divergens T rauth, 1 9 3 5 336* Punctaptychus Shipkovski Mineral Baths, Tithonian, Cherniosam This paper, punctatus V o l t z Lovech District Fm Fig. 3.1. Punctaptychus sp. 338* Punctaptychus sp. Barlya Village, Sofia District Tithonian, Ginci Fm - (BAS/5653) Glozhene Fm 1890*, 1891* Aptychus sp. Bov Village, Sofia District, at Kimmeridgian, Ginci the foot of the peak Yavorets Fm 335* Lamellaptychus Erden Village, Montana Tithonian, Ginci Fm - This paper, sparsilamellosus District Glozhene Fm Fig. 3.2. (G u e m b .)

Beyrichilamellaptychus beyrichi (O ppel, 1 8 6 5 ) 9738* (1478 Lamellaptychus Veselina River, south of Berriasian, Hanevtsi Fm (205a)) beyrichi var. longa Todyuvtsi Village, V. T r a u t h Tarnovo District 9740* (1478 Lamellaptychus Veselina River, south of Berriasian, Hanevtsi Fm (205)) beyrichi (Opp.) Todyuvtsi Village, V. Tarnovo District

Beyrichilamellaptychus cf. beyrichi (O ppel, 1 8 6 5 ) 9737* (1375, t. Lamellaptychus South of the hamlet Lipov Valanginian or negative imprint 3729) beyrichi (Opp.) kat, Varna District Tithonian, ?Ticha Fm 30825 (6882) Lamellaptychus North of Shipkovo Village, Berriasian, Cherniosam beyrichi (O p p e l) var. Lovech District Fm seranonoides T r a u t h

Beyrichilamellaptychus studeri (O oster, 1 8 5 7 )

30843 (7083, t. Lamellaptychus east of Todorchetata Village, Upper Valanginian, S t ef a n o v holotype of 7362) mortilleti ( P i c t e t & Gabrovo District Kamchia Fm (1961, pi. 3, fig. Lamellaptychus L o r i o l ) var. radiata n. 3); this paper, mortilleti var. var. Fig. 4. radiata S t e ­ fanov

Mortilletilamellaptychus mendrisiensis mendrisiensis (R enz & H abicht, 1 9 8 5 ) 30824 (3048, t. Lamellaptychus Kapinets Village, Lower Hauterivian, S t e f a n o v 3095) mortilleti ( P i c t e t & Targovishte District Hanevtsi Fm (1961, p i. 3, fig. Loriol) var. longa 2); this paper, T r a u t h F ig . 5.2. 30834 (12872) Lamellaptychus Mecha polyana Village (not U p p e r Valanginian, S t e f a n o v submortilleti T r a u t h existing any more), Svoge Salash F m (1961, p i. 3, fig. municipality, Sofia District 6); this paper, F ig s 5.1 a , b.

Mortilletilamellaptychus bicurvatus (R enz et H abicht, 1 9 8 5 )

30812 Lamellaptychus south of the hamlet Hristova Upper Valanginian, S t efa n o v mortilleti ( P i c t e t & Mahala, Troyan municipality, Cherniosam Fm (1961, pi. 3, fig. L o r i o l ) 1858 Lovech District 1); this paper, Fig. 5.3. Taxonomy of Late Jurassic - Early Cretaceous aptychi 271

Museum Names appearing number on the old museum NMNHS F- Locality Horizon Illustrated in Remarks labels and/or in (original old S t e f a n o v (1961) number) Mortilletilamellaptychus stefanovi n. sp. 30833 (35, t. Lamellaptychus Makotsevo Village, southeast Berriasian, Cherniosam S te f a n o v HOLOTYPE 237 (M. Hr.)) submortilleti T r a u t h of the peak Ademitsa, Sofia Fm (Tithonian - (1961, pi. 3, fig. var. retroflexa T r a u t h District Berriasian flysch) 7); this paper, Fig. 6.

Thorolamellaptychus cf. aplanatus (G illiéron, 1873) 30865 Lamellaptychus east of Makotsevo Village, Berriasian, Cherniosam This paper, this is perhaps (37, t. 237) herthae ( W i n k l e r ) var. Sofia District Fm Fig. 7.1. the specimen laevadsymphysalis mentioned by T r a u t h S te f a n o v (1961, p. 223)

Thorolamellaptychus trauthi (R enz & H abicht, 1 9 8 5 ) 30853 (10194, Lamellaptychus north of Beli Osam Village, Upper Valanginian, This paper, t. 10194) mortilleti ( P i c t e t & Lovech District Cherniosam Fm Fig. 7.2. L o r i o l ) var. longa T r a u t h

Didayilamellaptychus didayi (C oquand, 1841) 30810 Lamellaptychus didayi Balgarski izvor Village, Lower Hauterivian, S te f a n o v ( C o q u a n d ) Lovech District Gorna Oryahovitsa Fm (1961, pi. 2, fig. 5) 30811 Lamellaptychus didayi Balgarski izvor Village, Lower Hauterivian, S te f a n o v ( C o q u a n d ) Lovech District Gorna Oryahovitsa Fm (1961, pi. 2, fig. 6) 30813 Lamellaptychus didayi Chernevo Village, Varna Lower Hauterivian, S te f a n o v (Coquand), District Gorna Oryahovitsa Fm (1961, pi- 2, (transmission to fig- 7) subdidayi) 30828 (67, t. Lamellaptychus didayi north of Stolat Village, Lower Hauterivian, S t e f a n o v 231) ( C o q u a n d ) Gabrovo District Hanevtsi Fm (1961, pi. 2, fig. 3) 30837 (62, t. Lamellaptychus south of Gradnitsa Village, Lower Hauterivian S te f a n o v 241) subdidayi T r a u t h Gabrovo District (1961, pi. 2, [transition to fig. 8) Lamellaptychus didayi (C o q u a n d )] 30844 Lamellaptychus didayi southwest of Chernevo Lower Hauterivian, S t e f a n o v (C o q u a n d ) Village, Varna District Gorna Oryahovitsa Fm (1961, pi. 2, fig. 2); this paper, Fig. 8.1. 30852 Lamellaptychus didayi west of Balgarski izvor Lower Hauterivian, S te f a n o v ( C o q u a n d ) Village, Lovech District Gorna Oryahovitsa Fm (1961, pi. 2, fig. 1) 388* Aptychus sp. Omurtag, Targovishte Hauterivian, Kamchia District Fm 30864 Lamellaptychus southwest of Chernevo Lower Hauterivian, S t e f a n o v subdidayi T r a u t h Village, Varna District Gorna Oryahovitsa (1961, pi. 2, Fm(the upper part of the fig. 9) section) 30835 (2185) Lamellaptychus didayi Debnevo Village (Lozarski Valanginian, Kamchia juvenile ( C o q u a n d ) dol), Lovech District Fm specimen 9925* (332) Lamellaptychus sp. Teke dere, near Shumen Hauterivian, Gorna This paper, Oryahovitsa Fm Fig. 8.2. 9926*(333) Lamellaptychus sp. Teke dere, near Shumen Hauterivian, Gorna Oryahovitsa Fm 272 Zdenëk Vasîcek et al.

Museum Names appearing number on the old museum NMNHS F- Locality Horizon Illustrated in labels and/or in Remarks (original old S t e f a n o v (1961) number)

30822 Lamellaptychus Palilula Village, Vratsa Upper Valanginian, S te f a n o v 1961, Fragment subdidayi T r a u t h District Salash Fm pi. 2, fig. 10 [transition to Lamellaptychus didayi ( C o q u a n d )] 30820 Lamellaptychus didayi D. Vlahovska hamlet, Troyan Hauterivian, Kamchia Fragment ( C o q u a n d ) municipality (to the north of Fm Troyan), Lovech District

Didayilamellaptychus cf. didayi (C oquand, 1841) 30851 Lamellaptychus didayi west of Balgarski izvor Lower Hauterivian, ( C o q u a n d ) Village, Lovech District Gorna Oryahovitsa Fm 30829 (425) Lamellaptychus didayi Shipkovo Village, Lovech Berriasian, Cherniosam S t e f a n o v ( C o q u a n d ) District Fm (1961, pi. 2, fig- 4) 30847 (1016, t. Lamellaptychus north of the hamlet Lower Hauterivian, S t e f a n o v aptychus occur­ 1492) beyrichi (Oppel) var. Enchovtsi, Troyan Kamchia Fm (1961, pi. 3, figs ring in the body fractocosta T r a u t h municipality, Lovech District 8 and 9); this chamber of paper Fig. 9. Neolissoceras grasianum (d ’O r b ig n y )

Didayilamellaptychus seranonis (C oquand, 1841) 30861 Lamellaptychus Zlatitsa River, Gaganitsa Upper Valanginian, S t e f a n o v angulocostatus Village, Montana District Salash Fm (1961, pl. 1, fig. ( P e t e r s ) var. atlantica 12); this paper, ( H e n n ig ) , transition Fig. 8.3. to Lamellaptychus seranonis ( C o q u a n d )

Didayilamellaptychus cf. seranonis (C oquand, 1841) 30818 Lamellaptychus Chernevo Village, Varna Lower Hauterivian, S t e f a n o v holotype of seranonis ( C o q u a n d ) District Gorna Oryahovitsa Fm (1961, pi. 3, fig. L. seranonis var. radiata n. var. 5); this paper, radiata Fig. 8.4. S te f a n o v 30842 northwest of Stubel Village, unknown, Salash Fm only fragment (1269) Montana District of valve

Didayilamellaptychus angulodidayi (T rauth, 1 9 3 8 ) 30850 Lamellaptychus west of Balgarski izvor Lower Hauterivian, This paper, angulicostatus P e t . n. Village, Lovech District Gorna Oryahovitsa Fm Fig. 8.5. var. S te p h .

Didayilamellaptychus angulocostatus (P eters, 1854) 30809 Lamellaptychus south of Balgarski izvor Hauterivian, Gorna angulocostatus P e t t . Village, Lovech District Oryahovitsa Fm 30814 (194/t. Lamellaptychus Gaganitsa Village, Montana ?Hauterivian, Salash Fm note from the 1257) angulocostatus ( P e t e r s ) District original label: “Association Phyllopachyc- eras eichwaldi (Karakasch), Hauterivian - Barremian”.

30819 (239, t. Lamellaptychus southeast of Komshtitsa Lower Hauterivian, S t ef a n o v 982) angulocostatus ( P e t e r s ) Village, Sofia District Salash Fm (1961, pl. 1, fig. 3) 30830 Lamellaptychus southwest of Chernevo Lower Hauterivian, S te f a n o v angulocostatus ( P e t e r s ) Village, Varna District Gorna Oryahovitsa Fm (1961, pl. 1, fig- 1) Taxonomy of Late Jurassic - Early Cretaceous aptychi 273

Museum Names appearing number on the old museum NMNHS F- Locality Horizon Illustrated in Remarks labels and/or in (original old S t e f a n o v (1961) number)

30831 Lamellaptychus southwest of Chernevo Lower Hauterivian, S t e f a n o v angulocostatus ( P e t e r s ) Village, Varna District Gorna Oryahovitsa Fm (1961, pi. l,fig. 2); this paper, Fig. 8.6. 9905*, 9906* Lamellaptychus sp. Railway station Kaspichan, Hauterivian, Gorna Shumen District Oryahovitsa Fm

Didayilamellaptychus cf. angulocostatus (P eters, 1854) 30840 (14, t. Lamellaptychus Balgarski izvor Village, Upper Valanginian, S t e f a n o v 162) angulocostatus ( P e t e r s ) Lovech District Gorna Oryahovitsa Fm (1961, pl. 1, fig. var. atlantica ( P e t e r s ) 8); this paper, Fig. 8.7. 30832 Lamellaptychus Gaganitsa Village, Montana Berriasian, Salash Fm angulocostatus ( P e t e r s ) District var. fractocosta P e t e r s 30845 (5361, t. Lamellaptychus Borima Village, Lovech Lower Hauterivian, S te f a n o v juvenile speci­ 5361) angulocostatus ( P e t e r s ) District Gorna Oryahovitsa Fm (1961, pl. 1, men, note from fig. 4) the original label: “Asso­ ciation of Phyl- lopachyceras infundibulum and Lytoceras sp.”

Didayilamellaptychus angulicostatus (P ictet & de L oriol, 1858) 30815, 30816 Lamellaptychus Gaganitsa Village, Montana ?Hauterivian, Salash Fm This paper, note from the (194, t. 1257) angulocostatus ( P e t e r s ) District F ig . 8.8. original label: v a r. longa T r a u t h “Association Phyllopachyc- eras eichwaldi (Karakasch), Hauterivian - Barremian”. 30826 Lamellaptychus Zlatitsa River, Gaganitsa Berriasian, Salash Fm Stefanov angulocostatus ( P e t e r s ) Village, Montana District (1961, p l. 1, fig. 6) 30846 Lamellaptychus Gaganitsa Village, Montana Berriasian, Salash Fm Stefanov angulocostatus (P e te rs ) District (1961, p l. 1, v a r. fractocosta T r a u th fig. 5)

Didayilamellaptychus cristobalensis (O’C onnell, 1921) 30817 Lamellaptychus southwest of Chernevo Lower Hauterivian, S te f a n o v angulocostatus ( P e t e r s ) Village, Varna District Gorna Oryahovitsa Fm (1961, pl. 1, fig. var. cristobalensis 11); this paper, ( O ’C o n n e l l ) Fig. 8.9. 9904* Lamellaptychus sp. Railway station Kaspichan, Hauterivian, Gorna Shumen District Oryahovitsa Fm

Didayilamellaptychus filicostatus (S tefanov, 1961)

30836 (155, t. Lamellaptychus Ropot Village (west of Lower Hauterivian, S t ef a n o v HOLOTYPE 882) angulocostatus ( P e t e r s ) Komshtitsa), Sofia District Salash Fm (1961, pi. l.fig. var. filicosta n. var. 9); this paper, Fig. 10.

Didayilamellaptychus renzi M ëchovâ, V asicek & H ousa, 2010

30856 Lamellaptychus east of Krapchene Village, Lower Hauterivian, S t ef a n o v angulocostatus ( P e t e r s ) Montana District Salash Fm (1961, pl. 1, fig. var. radiata T r a u t h 7), this paper, fig. 8.10. 274 Zdenëk Vasicek et al.

M useum Names appearing num ber on the old museum N M N H S F- L ocality H orizon Illustrated in R em arks labels and/or in (original old Stefanov (1961) number)

30857 Lamellaptychus east of Krapchene Village, Lower Hauterivian, Stefanov angulocostatus (P e te rs) Montana District Salash Fm (1961, pl. 1, fig. var. radiata T ra u th 10) 30858 Lamellaptychus east of Krapchene Village, Lower Hauterivian, Stefanov angulocostatus (P e te rs) Montana District Salash Fm (1961, text var. radiata T ra u th fig. 2) 30859 Lamellaptychus east of Krapchene Village, Lower Hauterivian, Stefanov angulocostatus (P e te rs) Montana District Salash Fm (1961, text var. radiata T ra u th fig- 1) Laevaptychus longus (v. M eyer, 1831) 339* Punctaptychus sp. Cherni Osam Village, Tithonian, Cherniosam This paper, (BAS/5354) Lovech District Fm Fig. 11. Undeterminable specimens 30862,30863 Tithonian (or (2387a) N eocom ian) 30849 Lamellaptychus south of the hamlet Hristova Upper Valanginian, on same piece mortilleti ( P ic te t & mahala, Lovech District Cherniosam Fm of rock as # Loriol) 1858 30812 (Morti- lletilamellapty- chus bicurva- tus), illustrated in Stefanov 1961, pi. 3, fig. 1 and herein, Fig. 5.3. 2426* (1303 Lamellaptychus Veselina River, south of Berriasian, Hanevtsi Fm (212)) beyrichi var. longa Todyuvtsi Village, V. T ra u th Tarnovo District 30823 (10324) Lamellaptychus west of Ribno Village, Berriasian - Neocomian subdidayi T ra u th Troyan municipality, Lovech D istrict 30838 (11240) northeast of Krapchene Valanginian, Salash Fm juvenile Village, Montana District fragm ent 30827 Lamellaptychus Vidima River, south of the Upper Valanginian, Stefanov in association (2419) mortilleti ( P ic te t & hamlet Skandaloto, Troyan K am chia Fm (1961, pi. 3, w ith Spiticeras Loriol), transition municipality, Lovech District fig. 4). sp. to Lamellaptychus aplanatus (Gilliéron). 30860 Lamellaptychus Zlatitsa River, Gaganitsa Upper Valanginian, juvenile seranonis (Coquand) Village, Montana District Salash Fm specim en var. longa T ra u th 30821 Lamellaptychus n. sp. south of Balgarski izvor Upper Valanginian, note o f J. Village, Lovech District Gorna Oryahovitsa Fm S tefanov: “A s­ sociation of N. neocomiensis and Thurman- nia. B eautiful specimen, pos­ sibly new spe­ cies?” 30855 (10194) north of Beli Osam Village, Upper Valanginian, Lovech District Cherniosam Fm 9927*, 9928* Lamellaptychus sp. Teke dere, near Shumen Hauterivian, Gorna Oryahovitsa Fm