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HISTORY OF THE STUDY. OF CLATHROBACULUS. In 1912 Cossmann established the section Clathro- baculus within the genus Promathildia Andreae, 1887.
ISSN 0031-0301, Paleontological Journal, 2007, Vol. 41, No. 4, pp. 382–394. © Pleiades Publishing, Ltd., 2007. Original Russian Text © A.V. Guzhov, 2007, published in Paleontologicheskii Zhurnal, 2007, No. 4, pp. 35–45.

Systematics and Evolution of the Genus Clathrobaculus Cossmann, 1912 (Gastropoda, Mathildidae) A. V. Guzhov Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya ul. 123, Moscow, 117997 Russia e-mail: [email protected] Received April 7, 2006

Abstract—The morphology and ontogeny of Jurassic gastropods those have been included in the genera Clathrobaculus Cossmann, 1912 and Gordenella Gründel, 1990 are discussed. Based on the ontogenetic research of their shells, several types of ontogeny are established in the species of these genera. The suggestion is made that the genus Gordenella should be considered as a junior synonym of Clathrobaculus within the family Mathildidae. The protoconchs of Clathrobaculus species are studied to reveal their morphological variability and a number of distinguishing characters in which they differ from the protoconchs of other mathildid genera. Four species of Clathrobaculus, of which two are new (C. medidilatatus and C. inconstantiplicatus), are described from the Jurassic deposits of the European part of Russia. The composition of C. fahrenkohli (Rouillier, 1846) and C. krantzi (Rouillier, 1849) is emended, and the protoconchs of these species and of the species C. inconstantiplicatus are described for the first time. DOI: 10.1134/S0031030107040041

HISTORY OF THE STUDY OF CLATHROBACULUS In 1912 Cossmann established the section Clathrobaculus within the genus Promathildia Andreae, 1887 with the type species Cerithium zigzag Eudes-Deslongchamps, 1842, based on the following diagnosis: “Taille assez grande; forme turriculée, étroite, subcylindracée; spire longue, probablement pointue au sommet, croissant regulièrement sous un angle apical de 8 à 12° environ; tours très nombreux, peu convexes, multicarénés, à sutures plus ou moins distinctes; entre les trois carènes principales s’intercalent généralement des cordonnets spiraux, et l’ensemble est décussé par des lignes axiales, disposées en chevrons, qui ne forment pas d’aspérités ni de nodules à leur intersection avec les carènes spirales. Dernier tour peu élevé, habituellement muni d’une carène supplémentaire à la périphérie de la base qui est plane, ornée de cordonnets concentriques et de nombreuses stries rayonnantes. Ouverture arrondie, subanguleuse en avant; labre incurvé; columelle lisse, peu calleuse.” [The shell is fairly high; turriculate, slender, subcylindrical; the spire is long, perhaps pointed at the apex, increasing at regular rate, with apical angle of about 8°–12°; the whorls are numerous, slightly convex, with numerous carinae, with more or less distinct sutures; the three major carinae are commonly intercalated with spiral threads, and they all are intersected by axial lines arranged in chevron patterns and forming no irregularities or pustules at their intersection with the spiral carinae. The last whorl is slightly elevated, usually supplied with one addi-

tional carina on the periphery of the base, which is flat, and sculptured with concentric threads and numerous radial striae. The aperture is rounded, slightly angular anteriorly; the labrum is inflected; the columella is smooth and slightly calloused.] (Cossmann, 1912, pp. 7–8). Cossmann placed the following species into Clathrobaculus: Cerithium zigzag Eudes-Deslongchamps, 1842, C. sinemurense Martin, 1859, C. subcolumnare d’Orbigny, 1850; Turritella (Mathilda) abbas Hudleston, 1892, T. (M.) strangulata Hudleston, 1892, T. legayi Rigaux et Sauvage, 1867, T. clapensis Terquem et Jourdy, 1869, T. subulatissima Hebert et Deslongchamps, 1860, Promathildia multilirata Cossmann, 1912, P. donsieuxi Cossmann (in litt.), and Mathilda bacillaris Cossmann. In my opinion, some of these species cannot belong to Clathrobaculus. This is true of T. legayi, which resembles the genus Rigauxia Cossmann, 1885 in morphology. The type material of C. subcolumnare has never been illustrated, and those shells which have been assigned to this species by others authors most probably belong to other mathildids, since the two well-developed keels that divide the lateral side of the whorl into three equal parts extend the entire shell length. No other spiral sculpture is developed. Clathrobaculus already has some ontogenetic changes in sculpture (see below). T. (M.) strangulata and P. multilirata would be more properly treated as being within the genus Turritelloidea Walther, 1951. In his subsequent publications (Cossmann, 1913, pp. 228– 234; 1915, p. 256) Cossmann extended the scope of Clathrobaculus to include a number of new species.

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ONTOGENY OF THE SHELLS OF THE MEMBERS OF THE GENUS CLATHROBACULUS

S SR

UK SAK LK

LS

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SBK BPA B

Fig. 1. Terminology concerning the shell of Clathrobaculus. Abbreviations: (BPA) basal-palatal angulation, (LS) lateral side, (UK) upper keel, (LK) lower keel, (SAK) surface above the keel, (B) base, (SBK) surface below the keel, (SR) subsutural rib, and (S) suture.

Subsequently, Wenz (1939) treated Clathrobaculus as the subgenus of the genus Promathildia, and Gründel raised it to the rank of genus (Gründel, 2000). The composition of Clathrobaculus has not been revised after Cossmann’s works. In this paper I revise the composition of this genus based on data on the morphology of its type species (using literature sources) and on the morphology and ontogeny of the species that I place into the genus Clathrobaculus based on my own material. Thus, the species composition of the genus is revised and discussed below.

Four species of the genus Clathrobaculus were encountered in clayey sediments of the Callovian and Oxfordian of the European part of Russia: C. fahrenkohli (Rouillier, 1846), C. krantzi (Rouillier, 1849), C. inconstantiplicatus sp. nov., and C. medidilatatus sp. nov. All these species have two keels at the beginning of the teleoconch that divide the lateral side of the first whorl into three equal parts. Each keel has one strong rib. In the discussion of the morphology of whorls, the following abbreviations are used: the part of the lateral side above the upper keel (UK) is referred to as the surface above the keel (SAK), and that located below UK is referred to as the surface below the keel (SBK) (Fig. 1). The spiral sculpture that develops from the beginning of the teleoconch will be referred to as the primary sculpture (in Clathrobaculus this sculpture consists of three ribs located on the keels and basal-palatal angulation), and the sculpture that forms later will be referred to as the secondary sculpture. Different species show different changes in the sculpture of the whorls during the shell growth. In all these species the original pattern of sculpture develops in the first whorl of the teleoconch, on which two identical keels develop to divide the lateral side into three equal parts. The smallest developmental changes are observed in C. medidilatatus (see Fig. 2b): the upper keel drops lower within a few whorls by approximately 0.5–0.6 of the height of the lateral side of the whorl, and

SR UK

SR UK

LK

LK

(a)

(b)

UK

(c)

(d)

Fig. 2. Changes in the morphology of the later whorls in the genus Clathrobaculus: (a) hypothetical shell of the presumed ancestor; (b) C. medidilatatus sp. nov.; (c) C. krantzi (Rouillier); (d) C. fahrenkohli (Rouillier). Abbreviations: (UK) upper keel, (LK) lower keel, and (SR) subsutural rib. Lines connect homologous elements of sculpture in shells of different species. PALEONTOLOGICAL JOURNAL

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the lower keel is located in the middle of the surface below the keel. Thus, the surface above the keel expands at the expense of the surface below the keel. In addition, the lower keel becomes weaker than the upper keel and flattens to disappear to the middle of the teleoconch. The lower keel gives way to a strong spiral rib. The upper keel flattens to disappear on the later whorls. The surface above the keel bears either one or two secondary ribs, of which the subsutural rib (SR, see Fig. 1) is better developed, runs close to the suture, and is the earliest secondary rib on the lateral surface. These characteristic changes are not restricted to C. medidilatatus. The other species of Clathrobaculus (hereafter the original names of species are used) that show similar developmental changes are Promathildia (Teretrina) aculeata Haas, 1953 (Rhaetian); Turritella circinnula Ammon, 1896 (Rhaetian–Lower Hettangian); Cerithium hille d’Orbigny, 1850 (according to Thevenin, 1908; Lower Jurassic); Procerithium westhayense Cox, 1936 (Hettangian–Sinemurian); Cerithium sinemurense Martin, 1859 (Hettangian); Cheminitzia fistulosa Stoliczka, 1860 (Lias); Turritella clapense Terquem et Jourdy, 1869 (Bathonian); Turritella subulatissima Hebert et Deslongchamps, 1860 (Callovian); and Mathilda bacillaris Cossmann (Eocene). However, in contrast to Clathrobaculus medidilatatus, the upper keel of these species does not smooth out. This group of species includes the type species of the genus Clathrobaculus, C. zigzag (Eudes-Deslongchamps, 1842) from the Sinemurian of Normandy. The other species studied, Clathrobaculus krantzi (Rouill.), has a more complex ontogeny than that of C. medidilatatus. This is reflected in the greater change in the morphology of whorls (Fig. 2c). The following changes occur in addition to those in C. medidilatatus. The upper keel shifts to a still lower level, 0.35 of the height of the lateral side of the whorl, and does not smooth out. The rib that takes the place of the lower keel becomes weaker, and the surface below the keel on which it is located becomes strongly concave because of the overhanging upper keel. The secondary sculpture is the same as that of C. medidilatatus. The subsutural rib is well-developed and strongest. In C. krantzi both the amount and the rate of the ontogenetic changes increased. The upper keel of C. krantzi (even on the middle whorls) is located on the lateral side of the shell at a lower level than the upper keel of C. medidilatatus (even on the last whorls); i.e., C. krantzi passes through the ancestral stage of ontogeny at a higher rate. The developmental pattern characteristic of C. krantzi occurs in a few species such as Cerithium zezii Gemmellaro, 1878 (Hettangian–Sinemurian), Cerithium amoenum Eudes-Deslongchamps, 1842 (Bajocian), Turritella eucycla Hebert et Deslongchamps, 1860 (Callovian), and Cerithium ? nerinoides Andreae, 1887 (Oxfordian). Further change in the morphology of whorls occurs in Clathrobaculus fahrenkohli and C. inconstantiplicatus (Fig. 2d). The upper keel drops to the level of 0.15

of the height of the lateral side of the whorl, and the rib left after the lower keel continues to weaken and disappears. The upper keel assumes the form of an angular process below the whorl. Because of the low position of the upper keel, the surface below the keel becomes very narrow and strongly concave, and the suture becomes slit-shaped. In these species the subsutural rib runs over the spiral band of the whorl; thus, the surface above the keel between the subsutural rib and the upper keel becomes steep, flat, or concave. In C. fahrenkohli and C. inconstantiplicatus the morphology of the early whorls changes at a very high rate. By the third whorl, they have a morphology corresponding to the adult C. medidilatatus, and on the fifth whorl their morphology corresponds to the adult C. krantzi. The following species of Clathrobaculus pass through ontogenetic changes resembling those of C. fahrenkohli and C. inconstantiplicatus: Promathildia (Clathrobaculus) subulata Haas, 1953 and ?P. (C.) n. sp. (Haas, 1953) (Rhaetian), Turritella pizzolarii Meneghini, 1880 (Sinemurian–Pliensbachian), Promathildia catenifera Cossmann, 1913 (Bajocian), Mathilda atava Cossmann, 1885 (Bathonian), Turritella (Torcula) osanni Andreae, 1887 (Oxfordian), Gordenella wehrlandia Gründel, 2000 (Middle Callovian), Cerithium ? pommeranum Schmidt, 1903 (Upper Oxfordian), and Cerithium ? deeckei Schmidt, 1903 (Kimmeridgian). Thus, the species of Clathrobaculus may be divided into three groups differing in the developmental patterns of the teleoconch. The first whorl of the teleoconch is identical in all species of Clathrobaculus. The subsequent whorls vary in the amount of change. The minimum amount of change occurs in the species resembling C. medidilatatus (Group 1), the species resembling C. krantzi (Group 2) show a greater amount of change, and the maximum amount of change occurs in the species resembling C. fahrenkohli and C. inconstantiplicatus (Group 3). The increased amounts of developmental change are correlated with increased rates of change. In C. krantzi the region in which the teleoconch has whorl morphology resembling that of C. medidilatatus, is reduced to the first four whorls, i.e., the rate of ontogeny has increased several times over. In C. fahrenkohli and C. inconstantiplicatus the ontogenetic changes that persist in C. krantzi through the entire length of the shell occur within the same first four whorls. It is worth noting that in all species considerable changes in the morphology of whorls occur only on the early whorls, in the subsequent whorls the morphology changes only slightly. Thus, the morphology of the representatives of these groups may be characterized as follows: Group 1. The upper keel persists along the entire shell or, rarely, the rib smooths out. It runs in the middle of the whorl or, rarely, drops slightly lower (up to 0.4 of the height of the lateral side of the whorl). The lower keel smooths out to give way to a strong rib running in the middle of the surface below the keel or slightly lower. The surface above the keel is flat or slightly conPALEONTOLOGICAL JOURNAL

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cave, the surface below the keel is concave. One to three weak secondary ribs appear on the surface above the keel. The subsutural rib varies from weak to strong. The whorls are convex and keeled or, more rarely, the last whorls are not keeled. The suture is deep and angular. Group 1 contains species with the most slender shells, the incremental angle of the teleoconch is 6°–12°. Group 2. The upper keel is strong and drops to the level of 0.35–0.30 of the height of the lateral side of the whorl. The lower keel is reduced, its place is occupied by a strong or weak rib. There are two to four secondary more or less strong spiral ribs on the surface above the keel. The surface above the keel is wide and slightly concave, and the surface below the keel is narrow and strongly concave. The suture is deep and angular. The whorls are keeled in the lower part and are shaped like a pagoda. The incremental angle of the teleoconch is usually no less than 15°. Group 3. The upper keel drops to the level of 0.25– 0.15 of the height of the lateral side of the whorl and becomes weak. Occasionally the keel smooths out. The rib left by the lower keel assumes the shape of a thin thread or disappears. The subsutural rib is usually stronger than the other spiral ribs and is quite often located on a spiral elevation. The spiral elevation is occasionally highly developed, and its combination with the rib takes the shape of a disk. In addition to the subsutural rib, the surface above the keel bears two to four secondary spiral ribs, which are usually fairly strong. The surface above the keel is flat or slightly concave, the surface below the keel is very strongly concave. The suture becomes narrow and slit-shaped, the whorls become flatter and slightly keeled or not keeled in the lower part. The incremental angle of the teleoconch is usually 14°–19°. The evolutionary trend in the ontogeny of the teleoconch of the genus Clathrobaculus has been apparently toward elaboration, as may be seen from the stratigraphic range of its species. Eight species of Group 1, one species of Group 2, and two species of Group 3 have been described from the Lower Jurassic; two species of Group 1, two species of Group 2, and six species of Group 3 have been described from the Middle Jurassic; and one species of Group 1, three species of Group 2, and ten species of Group 3 have been described from the Upper Jurassic. Even though some of the previously described species are synonymous, it is obvious that the species of Group 1 are more common in the Lias; and the species of Group 3, in the Middle and Upper Jurassic. It seems likely that this is caused no only by the increasing diversity of Group 3 but also by its increasing importance in fossil communities. The species of Group 1 are more diverse and more numerous in older sediments. Thus, Group 1 is considered to be ancestral to Groups 2 and 3. Details of the phylogenetic relationships between species within the genus remain, however, unclear. It remains unknown whether these groups developed PALEONTOLOGICAL JOURNAL

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independently after they had split up or Group 1 repeatedly evolved forms with more complex ontogeny, characteristic of the species of Group 2 or, even, Group 3. The earliest occurrences of both of these extreme types of ontogeny are dated to the Triassic. The two earliest species, which in my opinion belong to Clathrobaculus, were described from the Rhaetian of Peru (Haas, 1953). One species (C. aculeatus) belongs to Group 1, the other species (C. subulatus) belongs to Group 3. One can see that the species C. subulatus, as well as C. fahrenkohli, shows the morphology characteristic of Group 3 even on the early whorls. Therefore, the genus Clathrobaculus evolved a teleoconch with the most complex type of ontogeny as early as the Triassic. As to the phylogeny of the species of the genus, data on the rate of the change in the morphology of the early and middle whorls in different species may provide fresh insight into it. Thus, judging from Cossmann’s data (Cossmann, 1913, p. 230, pl. 9, figs. 23–27), the species C. amoenum from the Bajocian of France occupies an intermediate position between C. medidilatatus and C. krantzi in the type of ontogeny. This species has a very prolonged stage in which the whorls have a morphology characteristic of the species of Group 1 (no less than ten whorls), and only at the late stage the appearance of its whorls approximates that of the species of Group 2. This fact suggests the possibility that the species of Groups 2 and 3 derive from the species of Group 1. The progressive elaboration of ontogeny shows a trend toward the disappearance of keels. The first keel to disappear is the lower keel in the species of Groups 1 and 2, the second keel to disappear is the upper keel in the species of Group 3, especially in the most recent C. pommeranus and C. deeckei (Upper Oxfordian and Upper Kimmeridgian, respectively). All species of the genus Clathrobaculus have the same morphology of the first whorl of the teleoconch. Thus, it seems reasonable to assume that the high-turriculate shell with convex whorls and two keels dividing the outer surface into three equal parts (see Fig. 2a) may be considered as an archetype of Clathrobaculus. Clathrobaculus sinemurensis (Martin, 1859) most closely resembles this archetype. ON THE COMPOSITION OF THE GENUS CLATHROBACULUS Gründel (1990) established a new genus, Gordenella, based on the material that he identified as Gordenella ? krantzi (Rouillier, 1849), designated Cerithium ? pommeranum Schmidt, 1905 as the type species of this genus, and placed the high-turriculate mathildids with two primary and several secondary ribs into this genus. In the type species the upper keel shifts to the lower part of the whorl, and the lower keel becomes thinner and disappears. The whorls become flat and not keeled. Gründel also proposed that such species as Mathilda atava Cossmann, 1885;

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Cerithium ? deeckei Schmidt, 1905; Turritella complanata Brösamlen, 1909; T. sauvagea Buvignier, 1852; T. (Torcula) osanni Andreae, 1887; T. (T.) fahrenkohli Rouillier, 1846; and Promathildia catenifera Cossmann, 1913 should be placed into this genus. All these species I place into the genus Clathrobaculus as species of Group 3 (T. sauvagea and T. complanata are included in the synonymy of Clathrobaculus fahrenkohli, see below). It would appear premature to establish the groups of Clathrobaculus, particularly Groups 2 and 3, as separate genera and subgenera. It is still unclear whether the species of Groups 2 and 3 form separate phylogenetic lineages or they are collective groups of species that comprise, at least partly, descendants of species that evolved at different times from Groups 1 and 2 and feature more complex ontogeny than that of their ancestors. Thus, it would appear reasonable that species of Groups 1 and 2 repeatedly were able, following a similar scenario, to give rise to species with more complex ontogeny, as in the species of Groups 2 and 3, respectively. For this reason the placement of Gordenella and Clathrobaculus into different families (Gründel, 2000) seems to be even more unfounded. Gründel established a new family, Gordenellidae, and placed Gordenella (= Groups 2 and 3 of Clathrobaculus) and Proacirsa into this family, whereas he left Clathrobaculus s. s. (= Group 1) outside the family Mathildidae (Gründel, 2000). Moreover, he expressed doubts that Gordenellidae belong to the superfamily Mathildoidea (Gründel, 2003). In my opinion, the genus Clathrobaculus (with the composition proposed here) should be better left in the family Mathildidae, and the genus Gordenella should be regarded as a synonym of Clathrobaculus. MORPHOLOGY OF PROTOCONCHS IN THE GENUS CLATHROBACULUS Makowski (1952, p. 20, text-fig. 4) illustrated a protoconch and the early whorls of a teleoconch and identified them as Turritella krantzi Rouillier, 1849. In the same paper he figured an adult shell (Makowski, 1952, pl. 5, fig. 8) and attributed it to the same species. In my opinion, however, text-fig. 4 and pl. 5, fig. 8 show shells belonging to different genera. Slightly earlier Walther (1951, p. 72, pl. 3, figs. 17a, 17b; pl. 5, fig. 10/10) figured the early stages of ontogeny in the shell of the species Promathildia (Turritelloidea) opalina (Quenstedt, 1858). The shell shown by Makowski in text-fig. 4 belongs to the same group of gastropods as P. (T.) opalina in Walther’s paper. The latter species is characterized by a medioaxial heterostrophic protoconch, and the early whorls of its teleoconch are rounded and bear several spiral ribs. Based on this species, Walther established the subgenus Promathildia (Turritelloidea) (see Walther, 1951). Thus, the specimen shown by Makowski in text-fig. 4 should be better placed into Turritelloidea. Gründel (1973, p. 949) stated the same opinion. The shell shown by Makowski in pl. 5, fig. 8

belongs to Clathrobaculus. It may be identified as C. aff. fahrenkohli Rouillier, 1846. Kaim (2004) described the protoconch of Clathrobaculus, which consists of two whorls, is almost smooth, only umbilical view shows rudimentary collabral plicae. Gründel (1997, pl. 3, figs. 46–47; pl. 4, fig. 48) figured the protoconch that he identified as Tricarilda plana Gründel, 1973. The protoconch is smooth, about 1.5 whorl long. However, it is hardly possible that this protoconch belongs to the genus Tricarilda. In morphology it resembles the protoconchs that I studied in C. fahrenkohli, C. inconstantiplicatus, and C. krantzi. According to my observations the protoconchs of C. fahrenkohli is transaxial, 1.3–1.5 whorl long, and is usually covered by collabral plica on the apical and umbilical sides. The protoconch of C. inconstantiplicatus is 1.5 whorl long and is covered by collabral plicae on the umbilical side, and the protoconchs of C. krantzi are smooth, about 1.3 whorls long. All these protoconchs are transaxial. Of particular interest is the protoconch of the only specimen of C. aff. fahrenkohli (Pl. 5, fig. 4), which differs from the other protoconchs of Clathrobaculus in having a single thick whorl devoid of sculpture. In addition, the spire of the protoconch overhangs the first whorl of the teleoconch. Thus, the protoconch of the genus Clathrobaculus clearly differs from that of the other mathildids in a number of characters: its spire rises above the entire teleoconch, with its last whorl either resting on or even not touching the initial whorl of the teleoconch. In all the other mathildid genera the protoconch is enclosed to a greater or lesser extent by the first whorl of the teleoconch. In addition, the protoconchs of Clathrobaculus are almost planispiral, in contrast to the protoconchs of many other mathildid genera. MATERIAL In the “Material” sections, information on the localities is provided in abbreviated form; full information is given in the list of localities at the end of the paper. A schematic map of these localities was presented in my previous paper (Guzhov, 2004). The material figured in the paper is housed in the Paleontological Institute of the Russian Academy of Sciences (PIN, coll. nos. 4814 and 4863). SYSTEMATIC PALEONTOLOGY S u p e r f a m i l y Mathildoidea Dall, 1889 Family Mathildidae Dall, 1889 Genus Clathrobaculus Cossmann, 1912 Promathildia (Clathrobaculus): Cossmann, 1912, p. 7 (pars). Gordenella: Gründel, 1990, p. 770; 2000, p. 257.

T y p e s p e c i e s. Cerithium zigzag Eudes-Deslongchamps, 1842 from the Sinemurian of France. D i a g n o s i s. Shells multispiral, high-turriculate, of medium size. Protoconch transaxial, sinistral, almost PALEONTOLOGICAL JOURNAL

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planispiral, consisting of one to two whorls, smooth or covered by collabral plicae on apical and umbilical sides. Protoconch rises above the entire teleoconch, with its spire resting on or even not touching the first whorl of teleoconch and with its end flaring. Protoconch-teleoconch boundary distinct, with commissure, marked by transition from sinistral to dextral shell. Teleoconch consists of 20 (? or more) whorls, with extent of involution of about 30%, and with surface of initial whorls of teleoconch divided by two keels bearing one rib each into three equal parts. Ontogeny of teleoconch varies from species to species. Shell morphology varies with the type of ontogeny. After minimum ontogenetic changes, adult whorls become convex, with the lower keel or the rib left by the smoothed keel running in the middle, with the rib of the lower keel running in the middle of the surface below the keel, with one to three secondary ribs on the surface above the keel, and with a deep, open suture. After maximum ontogenetic changes, adult whorls become flattened, with upper keel dropping to suture, occasionally resulting in whorls slightly keeled adaperturally. The surface below the keel becomes very narrow and~stronglyconcave and devoid of spiral sculpture. Two to four welldefined secondary ribs on the surface above the keel, with subsutural rib usually strongest. Suture becomes narrow and slit-shaped. In all species collabral sculpture of shell consists of slender ribs or striae. Last whorl in species of Groups 1 and 2 with lateral side and base separated by angulation with rib on it, that of the species of Group 3, with no such angulation. Shell base very low, flat to slightly convex, covered by spiral threads, numerous and fine or stronger but less numerous. In all shells examined aperture broken. Inner section of whorl almost circular, slightly elongated dorsoventrally. Growth lines opisthocyrt on the lateral side of the whorl, slightly prosocyrt at the base of the shell, and sharply curved backward against columella. S p e c i e s c o m p o s i t i o n. Twenty-nine species: C. aculeatus (Haas, 1953) from the Rhaetian of Peru, C. subulatus (Haas, 1953) from the Rhaetian of Peru, C. ? collenoti (Martin, 1859) from the Hettangian of France, C. sinemurensis (Martin, 1859) from the Hettangian-Sinemurian of France, C. unicarinatus (Quenstedt, 1858) from the Hettangian-Sinemurian of Germany, C. zigzag (Eudes-Deslongchamps, 1842) from the Sinemurian of France, C. j?stulosus (Stoliczka, 1860) from the Lower Jurassic of Austria, C. hille (d'orbigny in Thevenin, 1908) from the Lower Jurassic of France, C. ? sidae (d'orbigny in Thevenin, 1908) from the Lower Jurassic of France, C. westhayensis (Cox, 1936) from the Lower Jurassic of England, C. zezii (Gemmellaro, 1878) from the Lower Jurassic of Italy, C. amoenus (Eudes-Deslongchamps, 1842) from the Bajocian of France, C. cateniferus (Cossmann, 1913) from the Bajocian of France, C. clapensis (Terquem et Jourdy, 1869) from the Bathonian of France, C. atavus (Cossmann, 1885) from the Bathonian of France, 9 (Hebert et Deslongchamps, 1860) PALEONTOLOGICAL JOURNAL

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Fig. 3. Clathrobaculus fahrenkohli (Rouillier, 1846); PIN, specimen no. 48631170 (shell 15 mm high); abandoned quany "Kamushki", Moscow; Middle Callovian.

from the Callovian of France, C. ornatus (Quenstedt, 1858) from the Callovian of Germany, C. subulatissimus (Hebert et Deslongchamps, 1860) from the Callovian of France, C. wehrlandius Grundel, 2000 from the Middle Callovian of Germany, C. pizzolarii (Meneghini, 1880) from the Middle ? Jurassic of Italy, C. fahrenkohli (Rouillier, 1846), C. donsieuxi (Cossmann, 1913) from the Oxfordian of France, C. inconstantiplicatus sp. nov., C. krantzi (Rouillier, 1849), C. medidilatatus sp. nov., C. nerineoides (Andreae, 1887) from the Middle Oxfordian of France, C. osanni (Andreae, 1887) from the Middle Oxfordian of France, C. pommeranus (Schmidt, 1905) from the Upper Oxfordian of Poland, C. deeckei (Schmidt, 1905) from the Kimmeridgian of Poland, and C. bacillaris (Cossmann) from the Eocene of France. C o m p a r i s o n. This genus differs from all the other mathildids in the very long and slender multispiral shells with two primary ribs on the teleoconch and also in the planispiral protoconch rising above the entire initial whorl of the teleoconch. R e m a r k s. In C. collenoti, the keels do not get weaker or change their position. The subsutural rib, which is present in all the other species, is also absent, as well as any other spiral sculpture. Thus, this species represents the type of shell morphology that could be characteristic of the ancestor of the genus Clathrobaculus. C. sidae is represented by a rather poorly preserved shell; thus, we cannot say for certain that it belongs to the genus Clathrobaculus.

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The comparison of the species described below with other species has been hampered by the lack of data on their morphology and ontogeny. The close resemblance in shell morphology between heterochronous species within Groups 1 and 3 has plagued efforts to establish the species composition of the genus Clathrobaculus. Many species described by different authors from deposits of different ages have proved to be too similar to distinguish them at the species level. However, the revision of the species composition should take account of the rate of the ontogenetic changes in the shell, what shells (adult or juvenile) were used in the description of the species, etc. In addition, almost all species were established in the 19th century, when the descriptions were illustrated by drawings, which do not necessarily closely resemble the original specimen. Thus, only morphologically identical coeval species are placed in the synonymy of the species described below, whereas similar species that come from different stratigraphic ranges have been left unrevised. Clathrobaculus fahrenkohli (Rouillier, 1846) Plate 5, figs. 1 and 2; Plate 6, figs. 8–14

Turritella fahrenkohlii: Rouillier, 1846, pl. C, fig. 4; 1848, p. 269; Sokolov, 1912, p. 104, pl. 2, figs. 5–6, 8a; Gerasimov, 1955, p. 183, pl. 40, figs. 18 and 19. Turritella sauvagea: Buvignier, 1852, p. 40, pl. 32, figs. 15 and 16. Turritella (Torcula) fahrenkohlii: Lahusen, 1883, p. 38, pl. 3, figs. 10 and 11. Turritella divisa: Ilovaisky, 1903, p. 261, pl. 10, fig. 4. Turritella complanata: Brösamlen, 1909, p. 276, pl. 21, figs. 2 and 3. Turritella krantzi: Sokolov, 1912, pl. 2, fig. 8a, non Rouillier, 1849. Promathildia (Clathrobaculus) bigoti: Cossmann, 1913, p. 228, pl. 9, fig. 12. ? Nerinella subtricincta: Dmoch, 1971, p. 19, pl. 2, fig. 3. Turritella isjumensis: Makarenko, 1990, p. 113, pl. 32, fig. 19. Turritella (Torquesia) divisa: Gerasimov, 1992, p. 89, pl. 25, figs. 4, 12; pl. 26, figs. 4, 10–15.

L e c t o t y p e. The specimen figured by D.N. Sokolov (1912, pl. 2, figs. 6a–6c) under no. 737/2 from the collection of Fahrenkohl, prepared by Rouillier (1846); TsNIGR museum, coll. no. 10941; Moscow Region, Krasnogorsk district, left bank of the Moskva River near the village of Gal’evo; Upper Oxfordian.

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D e s c r i p t i o n (Fig. 3). The shell more than 18 mm high (presumably up to 28 mm). The protoconch consists of 1.3–1.5 whorls and usually bears collabral plicae on the umbilical and/or apical side. The teleoconch have more than 17 whorls, its incremental angle is 13.5°–16°, rarely up to 18°. The whorls are flattened, slightly angular in the lower part, and attain their maximum width on the upper keel. The suture is angular on the early whorls to become subsequently slitshaped. At the sixth whorl, the upper keel drops to the level of 0.2 of the height of the lateral side of the whorl. Subsequently the position of the keel changes only slightly. The lower keel smooths out at the beginning of the third whorl of the teleoconch, and the rib to which it gives way disappears on the fifth to seventh whorl. The surface above the keel is flat or slightly concave or, rarely, strongly concave, with two or three spiral ribs in addition to the subsutural rib. The surface below the keel is strongly concave. Altogether, the lateral side has from three to six fairly strong ribs and up to two fine threads. The collabral sculpture consists of coarse growth lines, which are fairly regularly arranged and closely spaced. The outer section of the whorl is quadrangular. C o m p a r i s o n. This species differs from C. inconstantiplicatus in the teleoconch without collabral plicae, it differs from C. medidilatatus in the flattened whorls, low position of the upper keel, slit-shaped suture, numerous secondary ribs, and it differs from C. krantzi in the keel being weaker and occupying lower levels, not keeled whorls, and better developed secondary ribs. R e m a r k s. Fahrenkohl’s collection, the material from which was described in Rouillier’s papers of 1845–1849, is apparently completely lost. Only that part of the collection which comes from the village of Gal’evo was saved and revised by Sokolov (1912). Since the quality of line illustrations in Rouillier’s paper is insufficient, Sokolov was not able to determine which shell was illustrated as C. fahrenkohli. Sokolov speculated that “paratype no. 737/37 may have served as a model”; however, he did not provide an illustration. Only specimen no. 737/2 is illustrated. The latter comes from the type series, and I here designate it as lectotype. In contrast to the drawing in Rouillier’s work (Rouillier, 1846), it has five rather than six ribs on the lateral surface of the whorl.

Explanation of Plate 5 Figs. 1 and 2. Clathrobaculus fahrenkohli (Rouillier, 1846); bank of the Oka River near the village of Nikitino; Middle Oxfordian; (1) PIN, specimen no. 4814/149: (1a) abapertural view of the shell (shell 3.35 mm high), (1b) apical view of the protoconch, (1c) umbilical view of the protoconch; (2) PIN, specimen no. 4814/150: (2a) abapertural view of the shell (shell 5 mm high), (2b) apical view of the protoconch, (2c) umbilical view of the protoconch, (2d) morphology of the early whorls (fragment 4 mm high), and (2e) character of collabral sculpture. Fig. 3. Clathrobaculus krantzi (Rouillier, 1849); PIN, specimen no. 4814/151: (3a) morphology of the early whorls (fragment 3.9 mm high), (3b) umbilical view of the protoconch; quarry at the town of Shchurovo; Middle Oxfordian, tenuiserratum Zone. Fig. 4. Clathrobaculus aff. fahrenkohli (Rouillier, 1846); specimen was lost after taking photographs: (4a) apertural view of the shell, (4b) apical view of the protoconch; Egor’evsk phosphorite mine, quarry no. 7-2bis; Upper Oxfordian, serratum Zone, serratum Subzone. Fig. 5. Clathrobaculus medidilatatus sp. nov.; PIN, holotype no. 4814/152 (shell 15.9 mm high): (5a) apertural view of the shell, (5b) abapertural view of the shell; quarry at the town of Shchurovo; Middle Oxfordian, tenuiserratum Zone. PALEONTOLOGICAL JOURNAL

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Shells of C.fahrenkohli from different localities differ in the character of collabral sculpture. Thus, shells from the village of Tyrnovo (Ryazan Region) and those

from the Upper Oxfordian clays of Egor'evsk phosphorite mine (Moscow Region) have a collabral sculpture that is more regular, stronger, and more widely spaced

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than in the shells from the Middle Oxfordian clays of the town of Shchurovo (Moscow Region), village of Nikitino (Ryazan Region), Unzha River basin, and others. In Rouillier’s paper (Rouillier, 1846) a C. fahrenkohli with a coarser collabral sculpture was described. Apparently that is the reason why Gerasimov (1992) erroneously identified the species which is here described as C. inconstantiplicatus as C. fahrenkohli. However, the collabral sculpture of C. fahrenkohli consists of scattered coarse growth lines, and that of C. inconstantiplicatus consists of groups of closely spaced coarse growth lines separated by spaces where the shell is covered with very weak growth lines. O c c u r r e n c e. Middle Callovian–Upper Oxfordian of the central part of European Russia; Middle Oxfordian of the lower Volga region; Oxfordian of the Donets Basin; Middle–Upper Callovian of Germany; perhaps, Middle–Upper Callovian of Poland; Oxfordian of France. M a t e r i a l. Eight hundred and seventy-nine specimens: Middle Callovian, quarry “Kamushki” (2 specimens); Upper Callovian–Middle Oxfordian, village of Tyrnovo (26 specimens); Lower Oxfordian, village of Nikitino (7 specimens); Lower–Middle Oxfordian, villages of Nikitino (4 specimens) and Chevkino (4 specimens); Middle Oxfordian, Afanas’evskii quarry (2 specimens), villages of Vasil’kovo (9 specimens) and Mikhalenino (52 specimens), open pit “Severnyi Makar’ev” (2 specimens), settlement of Novoselki (1 specimen), and the villages of Nikitino (137 specimens + 16 specimens with a protoconch) and Novoselki (1 specimen); Middle Oxfordian, densiplicatum Zone, open pit “Severnyi Makar’ev” (1 specimen); Middle Oxfordian, densiplicatum Zone, densiplicatum Subzone, village of Mikhalenino (2 specimens) and the open pit “Severnyi Makar’ev” (13 specimens + 2 specimens with a protoconch); Middle Oxfordian, tenuiserratum Zone, open pit “Severnyi Makar’ev” (40 specimens + 4 specimens with a protoconch), village of Mikhalenino (21 speci-

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mens + 1 specimen with a protoconch); town of Shchurovo (Zarech’e sector) (410 specimens + 1 specimen with a protoconch); town of Shchurovo (cement plant) (6 specimens), and Mt. Ulagan (4 specimens); Middle Oxfordian, tenuiserratum Zone–Upper Oxfordian, alternoides Zone, village of Konstantinovo (4 specimens); Upper Oxfordian, village of Nikitino (1 specimen); Upper Oxfordian, alternoides Zone, ilovaiskii Subzone, village of Mikhalenino (2 specimens); Upper Oxfordian, alternoides Zone, alternoides subzone, village of Mikhalenino (62 specimens + 5 specimens with a protoconch); Upper Oxfordian, serratum Zone, serratum Subzone, Egor’evsk phosphorite mine, quarry no. 7-2bis (15 specimens) and quarry no. 10 (1 specimen); and the Upper Oxfordian, ravni Zone (beds with Amoeboceras gerassimovi), village of Mikhalenino (1 specimen). Clathrobaculus krantzi (Rouillier, 1849) Plate 5, fig. 3; Plate 6, figs. 2–4

Turritella krantzi: Rouillier, 1849, p. 378, pl. L, fig. 95; Sokolov, 1912, p. 106, pl. 2, figs. 7, 8c (non 8a); Gerasimov, 1955, p. 106, pl. 40, fig. 2. ? Turritella praecursor: Andreae, 1887, p. 22, pl. C, figs. 19 and 20. Turritella bicostata: Ilovaisky, 1903, p. 262, pl. 10, figs. 5 and 6. Turritella (Torquesia) krantzi: Gerasimov, 1992, p. 93, pl. 25, figs. 1–3, 6; pl. 26, figs. 2 and 9.

L e c t o t y p e. The specimen, figured by Sokolov (1912, pl. 2, fig. 8c (non 8a = (C. fahrenkohli))) under no. 737/4 from the collection of Fahrenkohl, described by Rouillier (1849); TsNIGR museum, coll. no. 10941; Moscow Region, Krasnogorsk district, left bank of the Moskva River near the village of Gal’evo; Upper Oxfordian. D e s c r i p t i o n. The shell no less than 16.5 mm high (presumably up to 20.5 mm). The protoconch is smooth, consists of 1.3 whorls. The teleoconch has more than 17.5 whorls, its incremental angle is 9.5°–14°. The whorls are keeled in the lower part and flattened above

Explanation of Plate 6 Fig. 1. Clathrobaculus medidilatatus sp. nov.; specimen PIN, no. 4814/153 (shell 9.6 mm high), abapertural view of the shell, quarry at the town of Shchurovo; Middle Oxfordian, tenuiserratum Zone. Figs. 2–4. Clathrobaculus krantzi (Rouillier, 1849); quarry at the town of Shchurovo; Middle Oxfordian, tenuiserratum Zone; (2) PIN, specimen no. 4814/154 (shell 15.8 mm high): (2a) apertural view of the shell, (2b) abapertural view of the shell; (3) PIN, specimen no. 4814/155 (shell 16.4 mm high), abapertural view of the shell; and (4) PIN, specimen no. 4814/156 (shell 13.2 mm high), abapertural view of the shell. Figs. 5–7. Clathrobaculus inconstantiplicatus sp. nov. Middle Oxfordian. (5) PIN, specimen no. 4814/157 (shell 12.2 mm high), abapertural view of the shell; quarry at the town of Shchurovo; tenuiserratum Zone; (6) PIN, holotype no. 4814/158 (shell 17 mm high): (6a) abapertural view of the shell, (6b) apertural view of the shell; quarry at the town of Shchurovo; tenuiserratum Zone; (7) PIN, specimen no. 4814/159 (shell 17.3 mm high), abapertural view of the shell; Oka River bank near the village of Nikitino. Figs. 8–14. Clathrobaculus fahrenkohli (Rouillier, 1846). (8–13): quarry at the town of Shchurovo; Middle Oxfordian, tenuiserratum Zone; (8) PIN, specimen no. 4814/160 (shell 18.2 mm high), abapertural view of the shell; (9) PIN, specimen no. 4814/161 (shell 12.9 mm high), abapertural view of the shell; (10) PIN, specimen no. 4814/162 (shell 18.3 mm high), apertural view of the shell; (11) PIN, specimen no. 4814/163 (shell 8.6 mm high), abapertural view of the shell; (12) PIN, specimen no. 4814/164 (shell 11.5 mm high), abapertural view of the shell; (13) PIN, specimen no. 4814/165 (shell 15.3 mm high), abapertural view of the shell; (14) PIN, specimen no. 4814/166 (shell 8.4 mm high), abapertural view of the shell; alluvium of the Kazach’ya River near the village of Tyrnovo; Upper Callovian–Middle Oxfordian. Fig. 15. Clathrobaculus inconstantiplicatus sp. nov.; PIN, specimen no. 4814/167, character of collabral sculpture; quarry at the town of Shchurovo; Middle Oxfordian, tenuiserratum Zone. PALEONTOLOGICAL JOURNAL

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the keel and attain their maximum width on the upper keel. The suture is deep and angular. On the fourth and fifth whorls of the teleoconch, the upper keel runs along a line of 0.43–0.47 of the height of the lateral side, and subsequently drops to the level of 0.35 of its height. The lower keel reduces to leave a strong or, more rarely, weak rib on the surface below the keel. The surface above the keel is slightly concave and bears below the subsutural rib one more rib and one or two threads. The surface below the keel is concave. Altogether, the lateral surface has three or four fairly strong ribs and one to three threads. The collabral sculpture consists of slender striae and threads. The outer section of the whorl is trapezoidal. C o m p a r i s o n. See Comparison sections for other species. R e m a r k s. The shells of C. krantzi from the Upper Oxfordian have a coarser and more widely spaced collabral sculpture than specimens from other stratigraphic ranges. O c c u r r e n c e. Middle and Upper Oxfordian (except for the ravni Zone) of the central part of European Russia. M a t e r i a l. One hundred and twenty-five specimens: Middle Oxfordian, villages of Vasil’kovo (1 specimen) and Mikhalenino (3 specimens), open pit “Severnyi Makar’ev” (1 specimen), and the village of Nikitino (14 specimens + 1 specimen with a protoconch); Middle Oxfordian, densiplicatum Zone, open pit “Severnyi Makar’ev” (1 specimen); Middle Oxfordian, densiplicatum Zone, densiplicatum Subzone, village of Mikhalenino (1 specimen with a protoconch) and the open pit “Severnyi Makar’ev” (2 specimens); Middle Oxfordian, tenuiserratum Zone, open pit “Severnyi Makar’ev” (11 specimens), village of Mikhalenino (3 specimens), and the town of Shchurovo (Zarech’e sector) (73 specimens + 2 specimens with a protoconch); Upper Oxfordian, alternoides Zone, alternoides subzone, village of Mikhalenino (2 specimens + 5 specimens with a protoconch); Upper Oxfordian, serratum Zone, koldeweyense Subzone, village of Mikhalenino (1 specimen); and the Upper Oxfordian, serratum Zone, serratum Subzone, EFR, quarry no. 7-2bis (4 specimens). Clathrobaculus medidilatatus Guzhov, sp. nov. Plate 5, fig. 5; Plate 6, fig. 1

E t y m o l o g y. From the Latin medium (middle) and dilatatio (dilatation). H o l o t y p e. PIN, no. 4814/152; Moscow Region, Kolomna district, town of Shchurovo (Zarech’e sector); Middle Oxfordian, tenuiserratum Zone. D e s c r i p t i o n. The shell is no less than 16 mm high. Only fragments of the teleoconch have been found, their incremental angle is 9°–10°. The early whorls are keeled, the later whorls are convex, attaining their maximum width on the rib of the upper keel. The

suture is angular and deep. The upper keel is smoothed on the later whorls and runs slightly below the middle of the whorl. The lower keel is reduced and gives way to a strong rib running in the middle of the surface below the keel. The surface above the keel is flat of has two strong spiral ribs, the upper of which is subsutural. A very weak rib may occasionally develop above the subsutural rib, immediately adjacent to the suture. The surface below the keel is slightly concave and bears two ribs, the lower of which is usually weak. Altogether, the lateral surface of the whorl has four or five fairly strong ribs. The collabral sculpture consists of fine striae and ribs. The outer section of the whorl is hexagonal or rounded-hexagonal. C o m p a r i s o n. This species differs from the other species described in the convex adult whorls with smoothed keels. M a t e r i a l. Six specimens from the town of Shchurovo (Zarech’e sector). Clathrobaculus inconstantiplicatus Guzhov, sp. nov. Plate 6, figs. 5–7, 15

Turritella (Torquesia) fahrenkohlii: Gerasimov, 1992, p. 91, pl. 25, figs. 5, 7–11; pl. 26, figs. 3 and 16.

E t y m o l o g y. From the Latin inconstans (inconstant) and plica (fold). H o l o t y p e. PIN, no. 4814/158; Moscow Region, Kolomna district, town of Shchurovo (Zarech’e sector); Middle Oxfordian, tenuiserratum Zone. D e s c r i p t i o n. The shell is no less than 17.5 mm high (presumably up to 22.5 mm). The protoconch consists of 1.5 whorls, is smooth on the apical side, and has rudimentary collabral plicate on the umbilical side. The teleoconch has no less than 14 whorls, its incremental angle is 15°–19°. The whorls are flattened, slightly angular in the lower part. The whorl attains its maximum width on the upper keel. The suture is angular on the early whorls to become subsequently narrow and slit-shaped. By the sixth whorl, the upper keel drops to the level of 0.2 of the height of the lateral side of the whorl. Subsequently the position of the keel changes only slightly. The lower keel smooths out at the beginning of the third whorl of the teleoconch, and the rib to which it gives way disappears on the fifth to seventh whorls. The surface above the keel is flat or slightly concave and has two or three spiral ribs in addition to the subsutural rib. The surface below the keel is strongly concave. Altogether, the lateral side has four or five fairly strong ribs and up to two fine spiral threads. The collabral sculpture appears to consist of plicae, which in fact are formed by several closely spaced coarse growth lines. The spaces of the shell covered with coarse growth lines are separated by spaces where growth lines are very weak. The outer section of the whorl is quadrangular. The peculiar collabral sculpture develops as follows: The first 4–4.5 whorls of the teleoconch have closely spaced, but single growth lines (= threads). PALEONTOLOGICAL JOURNAL

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Then groups of two closely spaced ribs appear and begin to predominate on the next whorl. Subsequently threads are arranged in groups of three or four and closely approach each other to form plicae. The plicae form flat thickenings at their intersections with spiral ribs. The reverse process occurs on the later whorls (specimen no. 4814/159, see Pl. 6, fig. 7): the groups of ribs begin to disintegrate. However, the ribs remain coarse and very closely spaced. Thus, the whorls appear rough, and the collabral sculpture appears imbricated. This reverse process does not occur on the teleoconch before the 15th whorl. C o m p a r i s o n. On the later whorls C. inconstantiplicatus closely resembles C. fahrenkohli, but it differs from the latter in the dense imbricated collabral sculpture. O c c u r r e n c e. Upper Callovian?, Lower–Middle Oxfordian of the central part of European Russia. M a t e r i a l. Thirty-four specimens: Upper Callovian–Lower Oxfordian, quarry Peski (1 specimen); Lower Oxfordian, village of Nikitino (2 specimens); Middle Oxfordian, open pit “Severnyi Makar’ev” (1 specimen) and the villages of Mikhalenino (5 specimens) and Nikitino (6 specimens + 1 specimen with a protoconch); Middle Oxfordian, densiplicatum Zone, village of Vasil’kovo (1 specimen) and the open pit “Severnyi Makar’ev” (1 specimen); and the Middle Oxfordian, tenuiserratum Zone, town of Shchurovo (Zarech’e sector) (16 specimens).

Village of Novoselki—Ryazan Region, Rybnoe district, right bank of the Oka River near the upstream end of the village of Novoselki. Nikitino—Ryazan Region, Spassk-Ryazanskii district, right bank of the Oka River, 1 km downstream of the village of Nikitino. Peski—Moscow Region, Kolomna district, quarry south of the urban-type village Peski. “Severnyi Makar’ev”—Kostroma Region, Makar’ev district, right bank of the Unzha River near the northern end of the town of Makar’ev, open pit “Severnyi Makar’ev”. Tyrnovo—Ryazan Region, Pronsk district, bottom of the Kazach’ya River near the upstream end of the village of Tyrnovo, 1 km away from the confluence of the Kazach’ya and Ist’ya Rivers. Ulagan—Volgograd Region, Elton (Pallasovka) district, abandoned quarries on Mt. Ulagan, located near the eastern shore of Lake Elton. Chevkino—Ryazan Region, Spassk-Ryazanskii district, right bank of the Oka River downstream of the village of Chevkino. Shchurovo (Zarech’e sector)—Moscow Region, Kolomna district, quarries southwest of the Zarech’e district of the town of Shchurovo. Shchurovo (cement plant)—Moscow Region, Kolomna district, dump west of the cement plant of the town of Shchurovo.

LIST OF LOCALITIES Afanas’evskii quarry—Moscow Region, Kolomna district, quarry south of the village of Ratchino. Vasil’kovo—Kostroma Region, Makar’ev district, right bank of the Unzha River downstream of the village of Vasil’kovo. Egor’evsk phosphorite mine (EFR), quarry no. 7-2bis—Moscow Region, Voskresensk district, Egor’evsk phosphorite mine, quarry no. 7-2bis, located 5 km east of the village of Baranovskoe. Egor’evsk phosphorite mine, quarry no. 10—Moscow Region, Voskresensk district, Egor’evsk phosphorite mine, quarry no. 10, located between the villages of Novocherkasskoe and Elkino. “Kamushki”—Moscow, within the Fili sector. Konstantinovo—Ryazan Region, Rybnoe district, right bank of the Oka River near the upstream end of the village of Konstantinovo. Mikhalenino—Kostroma Region, Makar’ev district, right bank of the Unzha River 1 km upstream of the village of Mikhalenino. Settlement of Novoselki—Ryazan Region, Ryazan district, right bank of the Oka River downstream of Novoselki settlement of summer cottages near the mouth of the tributary of Dyat’kovskii, 1 km upstream of the village of Novoselki.

ACKNOWLEDGMENTS This work was supported by the Russian Foundation for Basic Research (project no. 04-04-48703a “Morphogenesis and Ecogenesis of Bivalve and Gastropod Mollusks”).

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REFERENCES 1. A. Andreae, “Die Glossophoren des Terrain à Chailles der Pfirt,” Abh. Geol. Spez. Elsaß–Lothringien 4 (3), 1–45 (1887). 2. R. Brösamlen, “Beitrag zur Kenntnis der Gastropoden des schwabischen Jura,” Palaeontographica 56 (4), 177– 321 (1909). 3. A. Buvignier, Statistique géologique, minéralogique, minérallurgique et paléontologique du département de la Meuse. Atlas (J. B. Bailliere, Paris, 1852). 4. M. Cossmann, Essais de paléoconchologie comparée (L’auteur et J. Lamarre & Cie, Paris, 1912a), Vol. 9. 5. M. Cossmann, “Contribution à la paléontologie française des terrains jurassiques. 3. Cerithiacea et Loxonematacea,” Mém. Soc. Géol. France, Paléontol. 20 (46, Fasc. 1–2), 89–264 (1913). 6. M. Cossmann, Essais de paléoconchologie comparée (L’auteur, Paris, 1915), Vol. 10. 7. I. Dmoch, “Osady i fauna górnojurajska z K l e¸bów na Pomorzu Zachodnim,” Stud. Soc. Sci. Torunensis, Sec. C, Geogr. Geol. 7 (4), 1–60 (1971).

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PALEONTOLOGICAL JOURNAL

Vol. 41

No. 4

2007