A Review of Gastropods from a Callovian (Middle Jurassic) Glacial Drift at Luków,⁄ Eastern Poland

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N. Jb. Geol. Paläont. Abh. 2008, vol. 247/2, p. 161–176, Stuttgart, February 2008, published online 2008

A review of gastropods from a Callovian (Middle Jurassic) glacial drift at Luków,⁄ Eastern Poland

Andrzej Kaim, Warszawa & Tokyo With 4 figures and 2 tables

KAIM, A. (2008): A review of gastropods from a Callovian (Middle Jurassic) glacial drift at Luków,⁄ Eastern Poland. – N. Jb. Geol. Paläont. Abh., 247: 161– 176; Stuttgart.

Abstract: This paper reviews all available data on gastropods from an abandoned Callovian clay pit in Luków⁄ Lapiguz⁄ located on the large glacial drift. A total of 51 species of gastropods are reported from this locality in contributions by MAKOWSKI (1952), SCHRÖDER (1995), and KAIM (2004). The fauna of the sideritic concretions is dominated by aporrhaid gastropods while the clay fauna is dominated by the alleged eumetulid Cosmocerithium and maturifusids. The latter association is characteristic for Jurassic sunken-wood assemblages and might originate also in such a setting. The PCA and cluster analyses groups well this distinctive association and clearly delimit from other gastropod associations in that region. A new genus and species Makowskispira lapiguzensis is described and attributed to Ampullinidae. The new genus differs from other members of the family by distinct spiral ornamentation.

Key words: gastropods, Jurassic, Callovian, paleoecology, Poland.

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1. Introduction material in the Museum of Faculty of Geology, Uni- versity of Warsaw. The collection consists of seven The outcrop of the glacial drift in Luków⁄ is a well specimens (Tab. 1; Fig. 1) including a specimen of known locality due to excellently preserved Late “Pleurotomaria buchiana D’ORBIGNY, 1845” men- Callovian ammonites (MAKOWSKI 1952, 1962; DZIK tioned but not illustrated by MAKOWSKI (1952). 1990). Three-dimensionally preserved shells from that According to GERASIMOV (1992) this species is a locality were investigated by MAKOWSKI (1962) in younger synonym of Bathrotomaria reticulata order to support his ideas on sexual dimorphism in (SOWERBY, 1821). ammonites. The other groups of fossils from Luków⁄ Some material from Luków⁄ was accessible to have drawn much less attention of palaeontologists. SCHRÖDER (1995) who illustrated five species, three MAKOWSKI (1952) was the first to provide preliminary of them left in open nomenclature. The most com- systematic descriptions of the most common fossils prehensive treatment of gastropods from Luków⁄ is occurring in Luków.⁄ His paper (MAKOWSKI 1952) provided in my monograph on Jurassic-Cretaceous concerned mostly molluscs including 12 gastropod gastropods from Poland (KAIM 2004) in which I species (Table 1). Unfortunately, three of them were described 44 species from Luków⁄ and 13 of them not illustrated and are therefore rather difficult to were new. Unfortunately the clay-pit was already revise. I could trace part of the MAKOWSKI’s (1952) abandoned at the time when I started these invest-

DOI: 10.1127/0077-7749/2008/0247-0161 0077-7749/08/0247-0161 $ 4.00 © 2008 E. Schweizerbart’sche Verlagsbuchhandlung, D-70176 Stuttgart 509-gültig-kaim-alles 26.02.2008 14:30 Uhr Seite 162 (Schwarz Bogen)

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Fig. 1. Two species of gastropods from Callovian of Luków⁄ (block in glacial drift), Podlasie, Poland not illustrated by KAIM (2004). a-c. Bathrotomaria reticulata (SOWERBY, 1925), MWG 004250, mentioned but not illustrated by MAKOWSKI (1952) as “Pleurotomaria buchiana D’ORBIGNY, 1845”. d-f. Makowskispira lapiguzensis n. gen and sp., holotype MWG 004244, illustrated and described by MAKOWSKI (1952) as “Natica crythea D’ORBIGNY, 1850”.

igations and the clay samples collected from a heap town. The brick-pit exploited large blocks of glacial yielded no gastropods. Therefore, I relied exclusively drift until the late 1970s (C. KULICKI, pers. comm. on material from museum collections that were collec- 2007). The outcrop exposed Late Callovian black clay ted by several researchers under variable circum- with ammonite-rich limestone concretions (see review stances. The aim of this paper is to summarise all data in OLEMPSKA & BLASZYK⁄ 2001). The concretions on gastropods from Luków⁄ and their comparison to were ascribed by DZIK (1990) to the Quenstedtoceras the other Middle Jurassic gastropod-bearing localities henrici Subzone of the Quenstedtoceras lamberti in this part of Europe. The facies dependency of the Zone. The rocks at Luków⁄ are from a large glacial Luków⁄ and other Middle Jurassic to Early Cretaceous erratic (so-called glacial drift), which originated most gastropod faunas is analysed and discussed. probably from the bottom of the Baltic Sea, north of Gda´nsk (OLEMPSKA & BLASZYK⁄ 2001). Autochtho- nous Late Callovian strata reached by deep boreholes 2. Geological setting in the Luków⁄ area represent platform carbonate facies The locality called Luków⁄ is an abandoned brick-pit (DAYCZAK-CALIKOWSKA & KOPIK 1976). at Lapiguz,⁄ a suburb in the southern part of Luków⁄ 509-gültig-kaim-alles 26.02.2008 14:30 Uhr Seite 163 (Schwarz Bogen)

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Table 1. List of gastropods described and/or illustrated so far from Callovian of Luków.⁄ Data from MAKOWSKI (1952), SCHRÖDER (1995), and KAIM (2004). Species asterisked are represented in the MAKOWSKI’s (1952) collection in the Museum of Faculty of Geology, University of Warsaw.

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Table 1 Cont.

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Table 1 Cont.

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3. Material and methods rently lost. All specimens here examined were already extracted from the sediment so I was unable to make The largest part of the material described by KAIM direct taphonomic and sedimentological observations. (2004) and analysed herein has been sieved from the The specimens extracted from the concretions could sediment by GWIDON JAKUBOWSKI from the Museum be identified by remnants of the rock matrix attached of Earth, Polish Academy of Sciences, Warsaw (ab- to the shells. The available gastropod shells were breviated MZ) and deposited therein. Some other identified and counted for the taxonomic purposes specimens were collected in the field, removed from of my previous work (KAIM 2004). Best preserved the sediment attached to the sideritic concretions, and representatives of each species were coated with extracted from the concretions by JANUSZ BLASZYK⁄ , platinum and photographed under Philips XL-20 JERZY DZIK, and MARCIN MACHALSKI, all from scanning electron microscope at the SEM laboratory Institute of Paleobiology, Polish Academy of Sciences, of ZPAL. The statistical analyses were carried out Warsaw (abbreviated ZPAL), and deposited therein. using PAST version 1.36 software (HAMMER et al. Part of MAKOWSKI’s (1952) material is housed at the 2001). For details of used statistic and plotting tech- Museum of Faculty of Geology, University of Warsaw niques the reader is referred to HAMMER & HARPER (abbreviated MWG) while its remaining part is appa- (2006). 509-gültig-kaim-alles 26.02.2008 14:30 Uhr Seite 166 (Schwarz Bogen)

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4. The taxonomic diversity of gastropods important part of the Middle Jurassic assemblages at Luków⁄ from Tethys (UHLIG 1881; CONTI & FISCHER 1982, 1983; CONTI & MONARI 2002; CONTI et al. 2004; The gastropods from Luków,⁄ available for this study, SZABÓ 1979, 1981, 1982) and more shallow-water were collected under different circumstances and over settings of its marginal Subboreal basins (GRÜNDEL & long period mostly in the 1960s and 1970s. Therefore, KAIM 2006). The most common vetigastropods at they do not constitute a uniform sample. The main Luków⁄ are Eucycloscala (Fig. 2d) and Torallochus distinction can be done based on the lithology of (Fig. 2c). Eucycloscala is a eucyclinin gastropod the sediment still attached to the shells. I could widespread in the Jurassic of Europe and found in distinguish two basic lithologies: sideritic concretions several outcrops in Poland, Russia, Germany, France, and soft black clay. The specimens from the con- and Great Britain (D’ORBIGNY 1850; HUDLESTON cretions are heavily dominated by aporrhaids with 1887-1896; GERASIMOV 1992; GRÜNDEL 2000; KAIM some contribution of cerithiids. The rest of the gastro- 2004). Torallochus is a barrel-shape turbinid known pods have been obtained by sieving the soft clay with only from a few occurrences including the Late Creta- water. This assemblage is dominated by cerithiids and ceous of Spain (KIEL & BANDEL 2001) and the maturifusids with some admixture of gastropods from Middle Jurassic of Poland (KAIM 2004). The species other groups e.g., eumetulids, polygyrinds, rissoids, of Torallochus seem to occur mostly in outer shelf purpurinds, mathildids, and cylindobullinids (Tab. 1; settings (Faustianka and Luków⁄ in Poland). The Figs. 1-3). specimens at Torallola in Spain were found in an Taking into account the accidental collecting of the olisthostrome containing a mixture of gastropods from material the gastropods were quite diversified and different settings (KIEL & BANDEL 2001); therefore represented by most of families thriving on Middle the original environment of Torallotrochus is un- Jurassic marine soft bottoms in central Europe. certain. Skeneids (Eudaronia and Aequispirella) are Altogether I counted 51 species (Tab. 1; Figs. 1-3) rare but these small sized gastropods (Fig. 2 a-b) are cited in MAKOWSKI (1952), SCHRÖDER (1995), and commonly overlooked by collectors. A detailed study KAIM (2004). of the Bathonian section in Gnaszyn (Central Poland) Pleurotomariids are uncommon in Luków⁄ and are by KAIM (submitted) has revealed that, although not represented by a single specimen of Bathrotomaria numerous, the Eudaronia shells are constantly present reticulata collected by MAKOWSKI (1952). The speci- in the majority of the samples. men is a fully grown shell (Fig. 1a-c). Such patterneschweizerbartxxx of The most common caenogastropods in Luków⁄ are appearance (rare but adolescent or adult specimens) cerithiids, aporrhaids, and maturifusids. Cerithiids are is in accordance with similar pattern observed by represented by six species of Cryptaulax (Fig. 2g-l). KAIM (2004) in Middle Jurassic (Bajocian/Bathonian) Three of them (C. quenstedti, C. echinata, and C. black clays in Central Poland. Relatively rare are also undulata) are exceeding 50 specimens in the col- other vetigastropods (Fig. 2 a -f), which constitute an lections and it seems that most of them were extracted

Fig. 2. Callovian gastropods from Luków⁄ (block in glacial drift), Podlasie, Poland. a. Eudaronia pusilla (GRÜNDEL, 2000), ZPAL Ga. 9/104; b. Aequispirella jurassica KAIM, 2004, ZPAL Ga. 9/107; c. Torallochus lukovensis KAIM, 2004, ZPAL Ga. 9/108; d. Eucycloscala sp. 1 sensu KAIM (2004), ZPAL Ga. 9/113; e. Calliotropis sp. sensu KAIM (2004), MZ VIII Mg 4232/1; f. Gerasimovcyclus gerasimovi (KAIM, 2004), MZ VIII Mg 4233/1; g. Cryptaulax muricata (SOWERBY, 1825), ZPAL Ga.9/142; h. Cryptaulax quenstedti (WALTHER, 1951), ZPAL MZ VIII Mg 4236/1; i. Cryptaulax echinata (BUCH, 1831), MZ VIII Mg 4231/3; j. Cryptaulax undulata (EUDES-DESLONGCHAMPS, 1842), ZPAL Ga.9/145; k. Cryptaulax sp. 3 sensu KAIM (2004), ZPAL Ga.9/167; l. ?Cryptaulax mutabilis (GERASIMOV, 1955), ZPAL Ga.9/168; m. Cosmocerithium kosmai KAIM, 2004, MZ VIII Mg4244/1; n. Cosmocerithium lukovensis KAIM, 2004, MZ VIII Mg 4245/1; o. Cosmo- cerithium polonicum KAIM, 2004, MZ VIII Mg 4246/1; p. Teutonica calloviana GRÜNDEL, 2001, MZ VIII Mg 4226/1; q. Teutonica jakubowskii KAIM, 2004, MZ VIII Mg 4225/1; r. Dicroloma cochleata (QUENSTEDT, 1858), ZPAL Ga.9/29; s. Pietteia subbicarinata (MÜNSTER, 1853), ZPAL Ga.9/33; t. Pietteia pellati (PIETTE, 1891), ZPAL Ga.9/32; u. Bralitzia acuminata (GRÜNDEL, 1999), MZ VIII Mg 4237/1; v. Bralitzia obtusa (LYCETT, 1850), MZ VIII Mg 4242/3; w. Bralitzia striatissima (GRÜNDEL, 1998), MZ VIII Mg 4243/1; x. Buvignieria calloviana GRÜNDEL, 1998, MZ VIII Mg 4238/1: please note that the captions of figs. 66 and 67 in KAIM (2004) were misplaced; y. Buvignieria studenckae KAIM, 2004, MZ VIII Mg 4239/1. 509-gültig-kaim-alles 26.02.2008 14:30 Uhr Seite 167 (Schwarz Bogen)

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Fig. 3. Callovian gastropods from Luków⁄ (block in glacial drift), Podlasie, Poland. a. Buvignieria piserai KAIM, 2004, MZ VIII Mg 4241/1; b. Knightella sp. sensu KAIM (2004), ZPAL Ga.9/178; c. Caenogastropod gen. indet. sp. 1 sensu KAIM (2004), ZPAL Ga.9/179; d. Caenogastropod gen. indet. sp. 2 sensu KAIM (2004), ZPAL Ga.9/181; e. Astandes conspicuus (EICHWALD, 1868), MZ VIII Mg 4227/4; f. Astandes kostromensis (GERASIMOV, 1955), MZ VIII Mg4228/3; g. Purpurina coronata HÉBERT & EUDES-DESLONGCHAMPS, 1860, MZ VIII Mg 4229/1; h. Purpurina formosa (EICHWALD, 1868), MZ VIII Mg 4230/3; i. Ebala varsoviensis KAIM, 2004, ZPAL Ga.9/222; j. Mathilda schmidti (WALTHER, 1951), un- registered MZ specimen; k. Mathilda makowskii KAIM, 2004, ZPAL Ga.9/273; l. Mathilda angulata (GRÜNDEL, 1997), ZPAL Ga.9/238; m. Mathilda podlasiensis KAIM, 2004, ZPAL Ga.9/267; n. Mathilda striatissima (GRÜNDEL, 1999), MZ VIII Mg 4248/1; o. Promathildia gruendeli KAIM, 2004, MZ VIII Mg 4251/1; p. Clathrobaculus sp. 2 sensu KAIM (2004), ZPAL Ga.9/228; q. Bandellina miniperforata (GRÜNDEL, 1998), MZ VIII Mg 4247/1; r. Cylindrobullina schneideri GRÜNDEL, 1997, MZ VIII Mg 4252/1; Sinuarbullina sp. sensu KAIM (2004), MZ VIII Mg 4253/1.

from the soft sediment. However, the majority of are adult individuals with fully developed expansions aporrhaids represented by one species of Dicroloma of the outer lip while the specimens from clays are (Fig. 2r) and two species of Pietteia (Fig. 2s-t) came juveniles. Maturifusids are represented by two species from the concretions. The specimens from concretions (Fig. 3e-f) and one of them (Astandes conspicuus) 509-gültig-kaim-alles 26.02.2008 14:30 Uhr Seite 169 (Schwarz Bogen)

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exceeds 100 specimens in the collections. It is worth Jurassic cerithiids of the genus Cryptaulax are most to mention that all specimens of these three groups are similar to the Recent genera Argyropeza and Varico- large- to middle-sized and easy to collect by surface peza. Gastropods of these genera live in deep waters collecting in the field. This might be a reason of (up to 914 m deep) and are microphagous detritivores their overrepresentation in the collections. Less com- or perhaps filter feeders (HOUBRICK, 1980a, b). It mon middle-sized gastropods are purpurinids (Fig. seems likely that Cryptaulax had the same mode of 3g-h). live. The small-sized specimens are dominated by tiny Recent aporrhaids are infaunal or seasonal bur- ptenoglossans including three species of the eumetulid rowers (BARNES & BAGENAL 1952; PERRON 1978) and Cosmocerithium (Fig. 2m-o) and two species of the graze on diatoms and decaying remains of macro- polygyrinid Teutonica (Fig. 2p-q). The other well algae. It seems that Jurassic Pietteia and Dicroloma diversified (six species) group but with fewer in- also fed on decaying organic matter as suggested by dividuals is Rissoidae represented by Bralitzia (Fig. KAIM (2001, and submitted). 2u-w) and Buvignieria (Figs. 2x-y, 3a). The hetero- Maturifusids and purpurinids are extinct groups branchians are represented by mathildids (nine usually considered as ancestor and/or sister group to species, Fig. 3j-p), cylindrobullinids (two species, neogastropods (RIEDEL 2000; KAIM 2004; KAIM & Fig. 3r-s), Ebalidae (one species, Fig. 3i), and Bandel- BEISEL 2005). Their mode of life is not fully resolved lina (one species, Fig. 3q). The most common are yet. KAIM (2006) reported a distinctive association of mathildids with the dominant species Mathilda these gastropods together with eumetulid Cosmoceri- schmidti (Fig. 3j). All those gastropods are known thium from Middle Jurassic sunken-wood assemblage from clay facies of Middle Jurassic of adjacent areas (see discussion below). Cosmocerithium (and its (WALTHER 1951, GRÜNDEL 1997, 1998a, b, 1999, junior synonym Prisciphora) is usually classified as 2001; GERASIMOV 1992; SCHRÖDER 1992, 1995; Eumetulidae (NÜTZEL 1998; KAIM 2004) although GUZHOV 2002, 2004; KAIM 2004). this assignment seems to be still disputable (BANDEL 2006). However, if that assumption is right then these gastropods, as all triphoroideans, could be primarily 5. Autecology of the gastropods from sponge feeders having juveniles feeding also on sur- Luków⁄ face films (WELLS 1998). Polygyrinidae is a poorly Modern pleurotomariids are spongivores (HICKMAN known extinct group usually placed close to zygo-

eschweizerbartxxx 1984; HARASEWYCH et al. 1988), but it is not certain if pleurids (BOUCHET et al. 2005). Its mode of life their Jurassic counterparts already utilised this source remains unknown. of food. My own observations from fossil collections The rissoids are usually marine shallow-water suggest that Jurassic pleurotomariids are more gastropods living in mid to lower littoral zones but common in silty facies and rather seldom in the clay. with some species known also from continental slope They may not tolerate both high sedimentation rate (PONDER & KEYZER 1998) and deep water settings and too soft substrate. (WARÉN 1996b). The Recent rissoids occur commonly Eudaronia is the most common skeneid gastropod on algae or beneath the rocks PONDER & KEYZER in Jurassic black clays. This gastropod is still present 1998). They feed on surface films or foraminifers in deep waters of Recent oceans (COTTON 1945; (WARÉN 1996a; PONDER & KEYZER 1998). WARÉN 1991) but the anatomy and behaviour of Although Mathildidae is one of the most common, Recent Eudaronia are still poorly known (WARÉN abundant in number of individuals, and well diversi- 1991). Judging from the radula it seems that they are fied groups of Mesozoic gastropods (e.g., KAIM 2004) deposit feeders on very fine sediment (HICKMAN nevertheless its biology and ecology is still poorly 1998; KAIM submitted). The other vetigastropod know as it is rather a relic group in the Recent oceans. group in Luków⁄ is Eucyclinae. Eucyclins form most As far as it is known most of the Recent species live in probably the most primitive branch of Trochidae deep water (BIELER 1995) and feed on coelenterates (MCLEAN 1982; KAIM 2004). They are distributed (BANDEL 1995), chiefly on corals (HEALY 1998). worldwide in the Recent oceans, mostly at bathyal However, corals have never been reported from and abyssal depths on the fine-grained sediment Mesozoic mathildid-rich assemblages (KAIM 2001, (HICKMAN 1998). Eucyclins are selective deposit 2004, and submitted). feeders (HICKMAN 1981). 509-gültig-kaim-alles 26.02.2008 14:30 Uhr Seite 170 (Schwarz Bogen)

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Fig. 4. PCA and cluster analyses of Middle and Upper Jurassic and Lower Cretacous gastropod assemblages from Poland and Russia. a-c. Biplots of scores and loadings of principal component analysis (PCA) on the correlation matrix with singular value decomposition (SVD). PC1 explains 20.1 %, PC2 17.2 %, and PC3 13.6 % of the variation respectively Dashed lines indicate loadings for particular taxa groups used in the analysis. Convex hulls indicate the smallest convex polygon containing all points of a given set. The association from Alpat’evo has been excluded from the convex hull for sand/sandstone as differing too much from the others. d. Results of cluster analysis; pair-group algorithm with Rho similarity coefficient. Note good clustering of sunken wood samples (SW1 and SW2) with Cosmoceritium and/or maturi- fusid rich samples. Data sourced from KAIM (2001, submitted, and unpublished data), GUZHOV (2004, 2006), and GRÜNDEL & KAIM (2006); see also Table 2. Analyses performed due to PAST v.1.36 software (HAMMER et al. 2001). Open diamonds indicate sunken-driftwood and other Cosmocerithium-rich assemblages; open circles indicate assemblages collected from sand and sandstone; shaded circles indicate assemblages from grey clay; solid circles indicate assemblages collected from black clay.

Cylidrobullinids are common tiny gastropods on similar behaviour. The best known Recent species of Mesozoic marine soft bottoms and most probably they Bullinidae, Bullina lineata, lives in tropical and warm are ancestors of acteonoideans (KAIM 2004). The temperate shallow waters and feed on sedentary poly- Recent species of Acteonoidea (e.g., Acteonidae and chaetes (BURN & THOMPSON 1998). Bullinidae) feed on polychaete annelids (BURN & Bandellina is a heterostrophic gastropod of unclear THOMPSON 1998) and cylindrobullinids could have affinities. It is referred to as Valvatoidea by BANDEL 509-gültig-kaim-alles 26.02.2008 14:30 Uhr Seite 171 (Schwarz Bogen)

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Table 2. Data matrix (in %) used for PCA and cluster analyses. Samples SW1 and SW2 from Gnaszyn were collected from sunked-wood assemblage, SWL is a sample from about 1m laterally from the SW1 sunken-wood. Sample “Gnaszyn all” contains 38 samples from whole section accessible at Gnaszyn, exclusive of SW1, SW2, and SWL. For details see KAIM (submitted).

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(1996, 2002) but this assumption remained a matter of in clays.” Outside of such accumulations both controversy (BIELER et al. 1998). In such a situation Astandes and Cosmocerithium are rather uncommon the autecology of this gastropod is also unclear. It both in Russia (GUZHOV 2004) and Poland (KAIM appears, however, in several muddy bottom settings 2006, submitted). The ubiquity of Astandes and Cos- from Late Triassic to Early Cretaceous (BANDEL 1996, mocerithium in Luków⁄ may suggest that some part of 2002; GRÜNDEL 1998b; KAIM 2004). the material has been derived from sunken driftwood assemblage similar to that reported by KAIM (2006) from Bathonian of central Poland or at least from an 6. Discussion association thriving on the secondary hard substrate. The knowledge on Middle Jurassic to Early Creta- The preliminary cluster and PCA analyses (Fig. 4) ceous Subboreal gastropods in Europe has recently support grouping the sample from Luków⁄ together been significantly improved. Rich gastropod assem- with Upper Oxfordian Cosmocerithium-rich samples blages from Germany and NW Poland (e.g., GRÜNDEL from Egor’evskii Quarry 7-2 bis (GUZHOV 2004) and 2003, GRÜNDEL & KAIM 2006 and references therein), two sunken-wood samples from Middle Bathonian of Central Poland (SCHRÖDER 1995; KAIM 2001, 2004, Gnaszyn (KAIM 2006, and unpublished data). and submitted), and Russia (e.g., GERASIMOV 1992; The PCA analysis of Middle-Upper Jurassic GUZHOV 2004, 2006) were described in a series of assemblages provides also some interesting insights papers and monographs. The Middle Jurassic assem- into relation between gastropod composition and their blages described in the aforementioned papers re- host sediment (Tab. 2, Fig. 4a-c). Gastropod asso- present rather uniform series of populations differing ciations from three different lithologies (sand/sand- mostly in contribution of particular taxa to the stone, black clay, and gray clay), although with some assemblage (GUZHOV 2004, 2006). The majority of overlapping, group relatively well (Fig. 4a-b). The them can roughly be attributed to the outer shelf en- cluster analysis (Fig. 4c) is not so unequivocal; how- vironment and muddy bottom (KAIM, submitted). ever, some groupings clearly call for further invest- GUZHOV (2006) attributed differences in assemblage igations. Such detailed analysis of sunken-wood and composition to varying climatic conditions. It seems, other associations will be provided elsewhere. however, that more important are bottom conditions The Jurassic Cosmocerithium and maturifusids and the sedimentation rate expressed, among others, occur also in other parts of Eurasia (D’ARCHIAC by a presence of hiatus concretions (KAIM, submitted). 1843; COSSMANN 1906; BEISEL 1983; SZABÓ 1983;

eschweizerbartxxx KAIM (submitted) showed that the clay sediment in the SCHRÖDER 1992, 1995; GRÜNDEL 1998a, 2001) but vicinity of such concretions contains assemblages, the numerical data are not provided for such occur- which differ from the ones extracted from the sedi- rences, therefore, they are difficult to discuss in the ment away of the concretions. current context. Great importance for gastropod distribution has also a presence of localized hard substrates (e.g. sunken driftwood). It is disputable if such assemblages 7. Selected taxonomy are analogous to the Recent sunken driftwood asso- Class Gastropoda CUVIER, 1797 ciations (KAIM 2006; for description of typical fossil Subclass Prosobranchia MILNE EDWARDS, 1848 sunken driftwood associations see KIEL & GOEDERT Order Caenogastropoda COX, 1959 2006) but undoubtedly they appear in Middle Jurassic Superfamily Campaniloidea DOUVILLÉ, 1904 as distinctive associations. The Jurassic sunken drift- Family Ampullinidae COSSMANN, 1918 wood associations preliminarily reported by KAIM (2006; detailed description is in preparation) are Genus Makowskispira n. gen. dominated by two gastropod genera: Astandes (known also as Maturifusus, both classified to Maturifusidae), Type species: Makowskia lapiguzensis n. sp. (see and Cosmocerithium (allegedly Eumetulidae). Such below). associations are described also by GUZHOV (2004), however, with no clear connection to the wooden Etymology: After the late Professor HENRYK MAKOWSKI, who was the first to describe gastropods from Luków.⁄ substrate. Such associations composed of hundreds- to-thousands of specimens, according to GUZHOV Diagnosis: Ovate ampullinid-type shell with numerous (2004: p. 549) occur in “…form of pocket and lenses spiral grooves. 509-gültig-kaim-alles 26.02.2008 14:30 Uhr Seite 173 (Schwarz Bogen)

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Remarks: The shell of Makowskispira is most similar in 8. Conclusions shape to shells of Jurassic Pictavia and Oonia. Neither of these genera, however, has spiral ornamentation. MAKOWSKI The gastropod assemblage in Luków⁄ indicates lower (1952: 20) mentioned the presence of a “planispirally” littoral zone (or even slightly deeper) in a region of coiled protoconch on one of the specimens at his hand. This high input of fine-grained clastics. The assemblage specimen is apparently lost but his description corresponds extracted from sideritic concretions is dominated by well with some ampullinids, e.g., protoconchs of Naricop- sina illustrated by GRÜNDEL (2001). Therefore, Makowski- aporrhaids and apparently indicating periods of lower spira is classified in Ampullinidae until new material is sedimentation rate. The assemblage recovered from found. soft clay displays close affinities to the sunken-wood associations from Middle Bathonian of Gnaszyn (KAIM 2006) and also some assemblages of uncertain Makowskispira lapiguzensis sp. nov. taphonomy from Upper Oxfordian of Russia. All these Fig. 1d-f assemblages are characterized by high abundances of Cosmoceritium and maturifusids. The analysis of 1952 Natica crythea D’ORBIGNY. – MAKOWSKI, p. 20, pl. 2, fig. 4 non D’ORBIGNY (1850). available data on Middle-Jurassic gastropod associations calls for attention and detailed taphonomic Holotype: MWG 004244, moderately well preserved, observations when mass occurrences of Cosmoceri- adult or adolescent shell. thium and maturifusids are discovered.

Type locality: Luków,⁄ Podlasie region, Poland.

Type horizon: Quenstedtoceras lamberti Zone, Late Acknowledgements Callovian, Middle Jurassic. The following persons and institutions provided the gastropod material for this study: J. BLASZYK⁄ , J. DZIK, and Etymology: After the type locality Luków⁄ Lapiguz.⁄ M. MACHALSKI, (all from the Institute of Paleobiology PAS, Warsaw, Poland); G. JAKUBOWSKI and B. STUDENCKA Material: Holotype only. The remaining material (Museum of the Earth PAS, Warsaw, Poland), and M. mentioned by MAKOWSKI (1952) is apparently lost. NIECHWEDOWICZ (Museum of the Faculty of Geology, Warsaw University, Poland). Their contribution is gratefully Dimensions: The holotype, consisting of about six acknowledged. M. NIECHWEDOWICZ is also acknowledged teleoconch whorls, is 44 mm high and 23 mm wide. for the photographs of MAKOWSKI’s (1952) material. The paper greatly benefited from peer reviews of M. A. CONTI eschweizerbartxxx Occurrence: Late Callovian (Middle Jurassic) of Luków⁄ (Rome), S. KIEL (Leeds), and A. NÜTZEL (München). The Lapiguz⁄ (block in glacial drift). SEM micrographs were taken in SEM laboratory of the Institute of Paleobiology (Warsaw) using Philips XL-20 Diagnosis: Ovate shell with rounded whorls (1.91 as high scanning microscope. This research was supported by as broad). Protoconch orthostrophic with low-spired early Instytut Paleobiologii PAN, the Japan Society for the whorls. The teleoconch ornamented by numerous (22 on the Promotion of Science (JSPS) Postdoctoral Fellowship for last preserved whorl) spiral grooves. Early whorl with Foreign Researchers, and JSPS research grant number ramp. 17.05324 (project number 050500000614).

Description: Protoconch almost planispiral consisting of two whorls and no clear demarcation with teleoconch (after References MAKOWSKI 1952). Shell ovate, whorl rounded, ornamented by shallow grooves, which are sometimes displaced at ARCHIAC, A. DE (1843): Description géologique du départe- enhanced, weakly sigmoidal growth lines. Suture incised. ment de l’Aisne. – Mémoires de la Société Géologique Early teleoconch whorls having relatively wide ramp de France, série 1, 5: 129-418. demarcated from lateral flank by rounded rib. Later in BANDEL, K. (1995): Mathildoidea (Gastropoda, Hetero- ontogeny the ramp disappears. Transition between base and stropha) from the Late Triassic St Cassian Formation. – lateral flank gradual. Aperture oval. Scripta Geologica, 111: 1-83. – (1996): Some heterostrophic gastropods from Triassic Remarks: Most similar is Valanginian (Early Cretaceous) St. Cassian Formation with a discussion on the classi- Ampullinidae gen. et sp. indet. 2, Early Valanginian (Early fication of the Allogastropoda. – Paläontologische Zeit- Cretaceous) reported by KAIM et al. (2004) from Yatrija schrift, 70: 325-365. River, Western Siberia, Russia which most probably also – (2002): About the Heterostropha (Gastropoda) from the belongs to Makowskispira. The Siberian gastropod, how- Carboniferous and Permian. – Mitteilungen aus dem ever, is poorly preserved and difficult to classify to the Geologisch-Paläontologischen Institut der Universität species level. Hamburg, 86: 45-80. 509-gültig-kaim-alles 26.02.2008 14:30 Uhr Seite 174 (Schwarz Bogen)

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