Ammonites and Stratigraphy of the Upper Bajocian Garantiana Garantiana Zone in the Interfluve Between the Kuban and Urup Rivers (Northern Caucasus) V

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ISSN 0031-0301, Paleontological Journal, 2019, Vol. 53, No. 11, pp. 1188–1202. © Pleiades Publishing, Ltd., 2019.

Ammonites and Stratigraphy of the Upper Bajocian Garantiana garantiana Zone in the Interfluve between the Kuban and Urup Rivers (Northern Caucasus) V. V. Mittaa, b, * aBorissiak Paleontological Institute, Russian Academy of Sciences, Moscow, 117647 Russia bCherepovets State University, Cherepovets, 162600 Russia *e-mail: [email protected] Received March 12, 2019; revised March 29, 2019; accepted April 1, 2019

Abstract—This paper presents the results of the study of the Upper Bajocian Garantiana garantiana Zone (Middle Jurassic) and characteristic ammonites in sections of the basin of the Kuban River (Karachay-Cher- kessia). The assemblage contains species of the genera Garantiana, Pseudogarantiana, Paragarantiana, Djanaliparkinsonia (all family Stephanoceratidae) and Vermisphinctes (family Perisphinctidae). A section of the Garantiana Zone on the Kyafar River contains (from bottom to top) Beds with Djanaliparkinsonia alanica (also recognized on the Kuban River), Beds with Garantiana subgaranti and Beds with Paragarantiana, approximately corresponding to the Dichotoma, Garantiana, and Tetragona subzones of the standard scale. Garantiana subgaranti Wetzel and Vermisphinctes martiusii (d’Orbigny) are described. Prorsisphinctes Buck- man, 1921 is proposed as a junior subjective synonym of Vermisphinctes Buckman, 1920.

Keywords: Upper Bajocian, ammonites, Garantiana, Paragarantiana, Djanaliparkinsonia, Vermisphinctes, biostratigraphy, Northern Caucasus DOI: 10.1134/S0031030119110066

INTRODUCTION lished records of representatives of Garantianinae in the Garantiana Zone in the Northern Caucasus The Garantiana garantiana Zone is located (Ob”yasnitelnaya…, 1973; Yura…, 1992) are not sup- between the Upper Bajocian Strenoceras niortense and ported by figures or collections, and are mostly based Parkinsonia parkinsoni zones of the standard scale and on field identifications. Thus, the Garantiana Zone in is subdivided into three subzones: Pseudogarantiana the Northern Caucasus is one of the least studied dichotoma, Garantiana garantiana [=Garantiana sub- Upper Bajocian intervals, and the same applies to its garanti, by some authors] and Garantiana tetragona. In ammonites. New data on the ammonites and stratig- Western Europe, especially in Germany, France, and raphy of the Garantiana Zone of the basin of the England, the Garantiana Zone is characterized by a Kuban River are presented below. high taxonomic diversity of ammonites. Ammonites of the subfamily Garantianinae, family Stephanocer- atidae are key elements for identification and infra- MATERIAL zonal subdivision of this zone. Representatives of Peri- The paper is based on material collected by the sphinctidae, Lissoceratidae, Oppeliidae, and some author during field studies in the Zelenchuk and Ust- other families are of secondary importance. Dzheguta districts of the Karachay-Cherkessia In the Northern Caucasus, the subfamily Garan- Republic in 2014–2018. The figured specimens are tianinae appears in the Niortense Zone where it is rep- housed in the Borissiak Paleontological Institute resented by diverse species of the genera Baculatoceras (PIN), coll. no. 5546. [M], Orthogarantiana [M], Strenoceras [m], and Kep- pleritiana [M, m]). Up the section, in the Garantiana Zone, occurrences of Garantianinae are extremely DESCRIPTIONS OF SECTIONS rare in this region. Until recently, only one finding was In the Kuban–Urup interfluve, the Upper Bajo- published, from the upper subformation of the Dzhor cian is represented by terrigenous, mainly argillaceous Formation of the Khodskiy Mountain Pass (North rocks, with numerous interbeds of concretions of var- Ossetia), identified as Hlawiceras aff. subgaranti (Wet- ious shapes and compositions, and less commonly, by zel) (Besnosov and Mitta, 1993; 1998). Other pub- large lenses of siltstone and interbeds of argillaceous

1188 AMMONITES AND STRATIGRAPHY OF THE UPPER BAJOCIAN 1189

50 km N SEA OF AZOV Udobnaya Kuban Krasnodar Stavropol Novorossiisk Belaya Laba Kuban Urup Malyi Zelenchuk Kuban Bolshoi Zelenchuk BLACK SEA Ispravnaya Ust-Dzheguta Sochi (a) 44 00 II 19 1751 Pregradnaya 5(10) 1560 Krasnogorskaya Kyafar Djangura Urup Storozhevaya Zelenchukskaya Mountain Urup Kardonikskaya

Malyi Zelenchuk Dausuz

Urup Bizhgon Aksaut Karachayevsk

2130 Marukha Kardonik Kyafar

0 5 10 km Kuban Bolshoi Zelenchuk (b) 43 40 41 00 41 30 E42 00

Fig. 1. Schematic map showing the sections studied in the Upper Bajocian Garantiana garantiana Zone in the interfluve between the Kuban and Urup rivers: (a) general and (b) detailed. limestone. These deposits belong to the lower part of and casts of underwater desiccation cracks (Fig. 3b). the upper subformation of Djangura Formation, This interbed, subdividing the section into two almost which is in general dated Bajocian–Lower Bathonian equal parts, appears to represent a short-term discon- (Besnosov, 1967). One of the most complete Upper tinuity of deposition. Apart from riverbank outcrops, Bajocian sections, in some intervals is relatively well the Garantiana Zone was sometimes observed in the characterized by ammonites, is currently exposed on episodically drying temporary channel of the Kyafar the banks of the Kyafar River (right tributary of the (locality 36a: Fig. 4), which allowed extensive collec- Bolshoi Zelenchuk River) in the Zelenchuk District of tion owing to a large surface of natural exposure. Karachay-Cherkessia. The siltstone concretions across the entire section Outcrops of the Garantiana Zone are on the right of the Garantiana Zone have a particular shape. They bank of the Kyafar downstream of the Storozhevaya are mostly flat discoid, reaching 10–15 cm in diame- (Fig. 1). The steep river gorge shows a member of ter, and are almost devoid of fossils. The lower half of dark-gray argillite-like clay for a distance of 450 m, the section (Bed 1), unlike its upper part, large (up to with scattered gray (on the surface yellowish-brown) 0.4 m in diameter) contains infrequent nodules of dark siltstone concretions, forming numerous layers (local- gray hard limestone, often with remnants of bivalve ities 5, 10, 36) (Figs. 2, 3). The middle part of the shells and small-sized ammonites usually identifiable member contains a thin (10–20 cm) layer of argilla- only to genus. The total thickness of the Garantiana ceous limestone concretions (Fig. 3a) with a distinct Zone on the Kyafar River, taking into account the dip cone-syneresis texture (combination of cone-in-cones of the beds (5° to NNE) reaches 50 m.

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m Lithology Ammonites Bed Substage Chronozone Beds with fauna

45 Paragarantiana with sp. [M] 40

2 sp. [M] [m] Paragarantiana [M]

35 Garantiana [M] [M, m] Garantiana subgaranti Garantiana 30 with Pseudogarantiana minima Pseudogarantiana Garantiana subgaranti subgaranti Garantiana ? sp.

25 martiusii Vermisphinctes Djanaliparkinsonia alanica Garantiana Garantiana Upper BajocianUpper Garantiana garantiana Garantiana 20

1 15

10 Djanaliparkinsonia alanica 1 with 2

5 3

Fig. 2. Section of the Garantiana garantiana Zone on the Kyafar River. Explanations: (1) clay argillite-like, (2) siltstone concretions, (3) limestone with cone-in-cone-syneresis, (4) large limestone concretion.

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(b) (a)

Fig. 3. Exposure of the Garantiana garantiana Zone, Kyafar River, (a) middle part of outcrops with an interbed of argillaceous limestone with texture of cone-in-cone and syneresis cracks (indicated by arrows) and (b) a fragment of a surface with syneresis texture, washed by the river.

Bed 1 contains frequent Djanaliparkinsonia, Vermi- In the adjacent Ust-Dzheguta District on the left sphinctes, and small incomplete shells, apparently of bank of the Kuban River downstream of the village of the genus Garantiana; Bed 2 contains fewer fossils; Krasnogorskaya, there is another outcrop of the representatives of Garantiana, Pseudogarantiana, Garantiana Zone (locality 19). Here, the towpath at Paragarantiana have been collected here. low water (from October to April) exposes argillite- like clay, similar to Kyafar clay, with layers of concre- Clays of the Garantiana Zone in the valley of the tions (Fig. 5a), and large lenses of siltstone in the basal Kyafar River are overlain (with an interruption in part. The basal 2–2.5 m of clay yielded Djanaliparkin- observation) by a member of dark-gray argillites with sonia and Vermisphinctes. These beds are basal for the rare interbeds of argillaceous limestone and numerous Garantiana Zone, overlying the sandstones with sideritic concretions, both scattered, and forming lay- numerous Zoophycos (Fig. 5b); based on the occur- ers. Besides numerous Rarecostites spp., the beds con- rence of Baculatoceras sp. these sandstones belong to tains occasional Spiroceras obliquecostatum (Quenst- the Niortense Zone. edt), Patrulia karachaica Mitta (Mitta, 2017a, 2017b; Mitta and Sherstyukov, 2018; Mitta et al., 2018). The argillite member belongs to the lower part of the Par- kinsoni Zone, Rarecostites subarietis Subzone. AMMONITES, THEIR TAXONOMIC COMPOSITION Clays of the Garantiana Zone are underlain (also AND DISTRIBUTION IN THE SECTION an interrupted observation) by a series of dark gray arenaceous clay, with interbeds of siltstone and car- As mentioned above, until recently ammonites of bonate concretions. The upper part of this series con- the Garantiana Zone from the Northern Caucasus tains species of the genus Baculatoceras characteristic were poorly studied and were mainly published in lists of the standard Niortense Zone (its upper Baculatum and general reviews. As shown by our field studies, this subzone), including the index zonal species B. bacula- is due to two factors. Firstly, the zone in this region is tum (Quenstedt), and also Strenoceras, Sphaeroceras, very poorly characterized by macrofossils. In ten field Infraparkinsonia, Spiroceras, Lissoceras, Oecotraustes, trips to the basin of the Kuban River, more than once etc. (Mitta, 2017a). I had to leave sections of the Garantiana Zone without

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(a) (b)

Fig. 4. Dried temporary channel of the Kyafar River (locality 36a), exposing the surface of the lower part of the Garantiana garantiana Zone (a), and a shell of an adult macroconch Vermisphinctes martiusii (d’Orbigny), found here in situ (b).

(b) (a)

Fig. 5. Lower part of the Garantiana garantiana Zone on the Kuban River (a), and a fragment of the surface of the underlying sandstone of the Strenoceras niortense Zone with Zoophycos (b).

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10 mm

* (a) (b)

Fig. 6. Djanaliparkinsonia alanica Mitta, specimen PIN no. 5546/250, shell of a young macroconch with a part of a body chamber, (a) ventral view, (b) lateral view; Karachay-Cherkessia, Ust-Dzheguta District, Kuban River downstream of the village of Krasnogorskaya, locality 19; Upper Bajocian, Garantiana garantiana Zone, Beds with Djanaliparkinsonia alanica; collected by the author in 2018. finding fossils. Secondly, the taxonomic diversity of labeled it as from an adjacent locality. Ammonites of ammonites is very poor in the Garantiana Zone of the the Garantiana Zone in the Kyafar River have a char- region, in comparison, for example, with the underly- acteristic form of preservation with externally red- ing Niortense Zone. Without taking into account occa- brownish siltstone nodules, usually containing a body sional shells of the order Phylloceratida, suborder chamber with a phragmocone imprint, and the rock Phylloceratina (transit species of Adabofoloceras, matrix covering the umbilicus may have slickensides. Phylloceras, and Holcophylloceras), the zone contains Hence, according to the emended data, D. alanica four or five genera of two different family rank taxa exclusively characterizes the lower part of the Garanti- (Fig. 2). The subfamily Garantianinae (superfamily ana Zone. Stephanoceratoidea, family Stephanoceratidae) is represented by the genera Garantiana, Pseudogaranti- Garantiana Mascke, the type (macroconch) genus ana, Djanaliparkinsonia, and Paragarantiana. of the subfamily, is represented by a larger number of species but fewer specimens. A fragment of a small Macroconchs and microconchs of the genus Djanaliparkinsonia Kutuzova have recently been shell of Garantiana ? sp., which could not be more pre- described as one species, D. alanica Mitta (Mitta, cisely identified, comes from the upper part of Bed 1 2018), from two localities on the Kyafar River and on the Kyafar River; two fragments of Garantiana sp. Kuban River. Material collected on the subsequent are found in the interval 7–8 m below the top of the trips (Figs. 6, 7) did not add anything significant to the section. And, finally, the middle part of Bed 2 on the characterization of the species. However, the speci- Kyafar River yielded two specimens described below mens figured in the original description as coming as G. subgaranti Wetzel (Fig. 8). The latter species in from the Subarietis Subzone of the Parkinsoni Zone of Western Europe characterizes the middle subzone of Kyafar (Mitta, 2018, p. 382, Fig. 3), most likely from the Garantiana Zone (Rioult et al., 1997; Dietze et al., the lower part of the Garantiana Zone. M.P. Sherstyu- 2002; Chandler et al., 2014), and many authors use it kov who donated this material, probably erroneously as the index species of this subzone (Pavia and Sturani,

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(a) * (c)

* 10 mm

(b) (e) (f) (d)

Fig. 7. Djanaliparkinsonia alanica Mitta: (a, b) slightly distorted shell of an adult microconch, specimen PIN no. 5546/198, (a) lateral view, (b) ventral view; (c, d) shell of a microconch with an imprint of the inner whorls, specimen PIN no. 5546/199, (c) lateral view, (d) ventral view; (e, f) shell of a young macroconch with part of a body chamber, (e) lateral view; Karachay-Cher- kessia; (a, b) Kuban River downstream of the village of Krasnogorskaya, locality 19; (c–d) Kyafar River downstream the village of Storozhevaya, locality 36a; Upper Bajocian, Garantiana garantiana Zone, Beds with Djanaliparkinsonia alanica; collected by the author in 2018.

1968; Fernández-López, 1997; Rioult et al., 1997; the developed aperture with lappets, this an adult shell O’Dogherty, 2006). of a representative of the genus Pseudogarantiana A small-sized shell (22 mm in diameter) comes Bentz (Fig. 9). The ornamentation is well-developed, from 10 m below the top of Bed 2, but, judging from represented by bipartite and fewer individual ribs,

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10 mm

(a) (b) (c)

Fig. 9. Pseudogarantiana minima (Wetzel), shell of an adult macroconch with a complete body chamber and an aperture with lappets, specimen PIN no. 5546/178, * (a) ventral view, (b) lateral view, (d) apertural view; Kara- (a) (b) chay-Cherkessia, Zelenchuk District, Kyafar River downstream of the village of Storozhevaya, locality 5(10); Upper 10 mm Bajocian, Garantiana garantiana Zone, Beds with Garan- tiana subgaranti; collected by the author in 2016.

upper subzone of the Garantiana Zone; while the genus characterized the entire Zone (Dietze et al., 2002)1. The genus Pseudogarantiana was first established as a subgenus of the genus Garantiana (Bentz, 1928); recently most authors consider these taxa as a dimorph pair Garantiana [M]/Pseudogarantiana [m] (Dietze et al., 2002; Fernández-López, Pavia, 2016). M.P. Sherstyukov found another ammonite of the subfamily Garantianinae represented by half of a body chamber (Fig. 10) in the interval 5–6 m below the top of Bed 2 of a section on the Kyafar River. Based on the shell shape (medium-wide whorls, rounded-trapezoid in cross-section with the maximum width near the umbilicus; the umbilicus is wide, with a relatively steep umbilical wall; the umbilical shoulder is rounded) and ornamentation (prominent, relatively widely (c) (d) spaced weakly curved bipartite and tripartite ribs, alternating with intercalating and simple ribs; ribbing coefficient 2.4.), this ammonite belongs to the Garan- Fig. 8. Garantiana subgaranti Wetzel: (a, b) shell of a mac- tiana alticosta Wetzel, 1911 Group (Dietze et al., roconch with an incompletely preserved body chamber, specimen PIN no. 5546/140, (a) apertural view, (b) lateral 2002). This group of species of Garantianinae is char- view; (c, d) slightly compressed body chamber of an adult acteristic of the upper part of the Garantiana Zone macroconch with an imprint of inner whorls, specimen (Tetragona subzone)—lower part of the Parkinsoni PIN no. 5546/139, (c–d) ventral view, (d) lateral view. Zone (Subarietis Subzone), and was considered as a Karachay-Cherkessia, Zelenchuk District, Kyafar River separate genus, named Paragarantiana Gauthier, downstream the village of Storozhevaya, locality 5(10); 2003, with the type species Garantiana longidoides Upper Bajocian, Garantiana garantiana, Beds with Garan- tiana subgaranti; collected by the authors, 2016. The aster- Gauthier et al. (Gauthier et al., 2000; Dietze et al., isk (*) here and further marks the beginning of the body 2002; Gauthier, 2003; Fernández-López and Pavia, chamber. 2016). The Kyafar specimen, judging from the gradual disappearance of the ventral weakening of the ribs (ventral groove or band) is an adult shell. It differs without nodes at the bifurcation point; the ends of the from all other species of Paragarantiana by the domi- branches on the ventral size are considerably enlarged. nance of tripartite ribs (bipartite, intercalating, and Based on these features and considering the size of the adult shell, this microconch belongs to P. minima 1 The hypothesis of N.V. Besnosov (Besnosov and Mitta, 1993) of (Wetzel) (Wetzel, 1911, p. 167, text-fig. 11, pl. 11, the occurrence of a Northern Caucasian species of Pseudogaran- tiana in the Subfurcatum [=Niortense] Subzone, was apparently figs. 11–16; Bentz, 1925, p. 171, pl. 7, fig. 8). In Ger- erroneous, resulting from exceptional rarity of their findings in sec- many, this species is characteristic of the middle and tions, without association with stratigraphically important taxa.

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10 mm

(a) (b)

Fig. 10. Paragarantiana sp., body chamber of an adult macroconch, specimen PIN no. 5546/13, (a) ventral view, (b) lateral view; Karachay-Cherkessia, Zelenchuk District, Kyafar River downstream of the Storozhevaya, locality 5(10); Upper Bajocian, Garan- tiana garantiana Zone, Bed with Paragarantiana; collected by M.P. Sherstyukov. simple ribs appear because of the loss of the connec- Garantiana (Subgarantiana) subgaranti: Bentz, 1928, c. 189, tion with a primary rib). Most likely, this is a new, pl. 16, fig. 1; Dietl, 1974, p. 13; pl. 2, fig. 5. undescribed species, which at this stage should be Garantiana (Garantiana) garantiana: Callomon et al., 1987, p. 8, pl. 1, fig. 2. referred to as Paragarantiana sp. ? Garantiana subgaranti var. garantiana: Dietze et al., 2002, The family Perisphinctidae (superfamily Peri- p. 25, pl. 3, fig. 2; pl. 4, fig. 2; pl. 6, fig. 2 [=?Garantiana garantiana sphinctoidea) is represented in the Garantiana Zone, (d’Orbigny)]. found only in its lower part. Our collection presently ? Garantiana subgaranti var. wetzeli: Dietze et al., 2002, p. 25, contains only macroconchs of this family, although of pl. 6, fig. 6 [=? Garantiana wetzeli Trauth]. non Garantiana (Hlawiceras) subgaranti: Rostovtsev et al., various growth stages. The identification to genus or 1985, p. 154, pl. 42, fig. 5 [=Baculatoceras]. species was difficult because of the chaos in the taxon- Lectotype. Garantiana subgaranti Wetzel, 1911, omy of early perisphinctids still persisting despite the p. 175, text-figs. 20, 21; pl. 13, figs. 3, 4; Germany, efforts of several generations of specialists. I hope to be North Rhine-Westphalia, vicinity of Bielefeld; Upper able to provide to a more detailed discussion of the Bajocian, Zone (“Lower taxonomy of this group; the species from the Garantiana garantiana Par- Garanti- Beds”); designated in Dietze et al., 2002, ana Zone of the basin of the Kuban River is described kinsonia below as Vermisphinctes martiusii (d’Orbigny). p. 25; refigured: Dietze et al., 2002, pl. 6, fig. 3; Howarth, 2017, p. 20, figs. 15c, 15d. D e s c r i p t i o n (Fig. 8). The adult shell, up to SYSTEMATIC PALEONTOLOGY 80–85 mm in diameter, medium-wide. The cross-section is oval with the maximum width in the lower third Superfamily Stephanoceratoidea Neumayr, 1875 of the flanks. The umbilicus is wide, shallow, the Family Stephanoceratidae Neumayr, 1875 umbilical wall is steep, the umbilical shoulder is Subfamily Garantianinae Wetzel, 1937 rounded. The ornamentation is represented by mainly bipartite ribs, prominent, slim, raised on the mold, Genus Garantiana Mascke, 1907 alternating between every three–four ribs with a sparse Garantiana subgaranti Wetzel, 1911 [M] simple rib. When the connection with the primary ribs is lost, intercalating ribs appear. The ribbing coeffi- Garantiana subgaranti: Wetzel, 1911, p. 175, text-figs. 20, 21; pl. 13, figs. 3, 4; Dietze et al., 2002, p. 25, pl. 6, figs. 1, 3; Chandler et cient is 1.8–2. The primary ribs are subradiate, al., 2014, p.145; text-fig. 8.1; Howarth, 2017, p. 20, text-fig. 15 c–g. appearing near the umbilical seam, the bifurcation

PALEONTOLOGICAL JOURNAL Vol. 53 No. 11 2019 AMMONITES AND STRATIGRAPHY OF THE UPPER BAJOCIAN 1197 point is in the mid-flank or slightly nearer the venter. Lectotype. Ammonites Martiusii d’Orbigny, The secondary ribs are noticeably bent anteriorad in 1845, pl. 125, figs. 1, 2 [= Vermisphinctes martiusii the ventrolateral region, and forming an obtuse angle (d’Orbigny); Rioult et al., 1994, pl. 49, fig. 1]; France, in the mid-venter. The ventral groove, readily notice- Calvados, vicinity of Bayeux; Upper Bajocian, Garan- able with age on the phragmocone, becomes a ventral tiana garantiana Zone; designated by Rioult et al. band, and disappears entirely nearer the aperture in (1994, p. 111). the adult shell. D e s c r i p t i o n. The shell is up to 200–250 mm Dimensions in mm and ratios: in diameter, with flattened whorls, oval in cross-section. The umbilicus is very wide and shallow. The Specimen umbilical wall is low; the umbilical shoulder is Dm WH WW UW WH/Dm WW/Dm UW/Dm no. rounded. The shell characteristically has deep con- strictions, following the direction of the ribs; as the 5546/140 67 22.5 20 28 0.34 0.30 0.42 shell grows, their number increases from two to three– 50 18.5 17.5 20.5 0.37 0.35 0.41 four per whorl. The ornamentation of young whorls is represented by subradiate primary ribs and secondary C o m p a r i s o n. A modern comparison with sim- ribs bent orad; on the venter, the ribs are similarly ilar species, in more detail, is given by Dietze et al. noticeably bent orad. The ribs are mainly bipartite, but (2002). there are also simple and intercalating ribs formed R e m a r k s. The lectotype in some papers (Ros- when the connection with the primary rib is lost. The tovtsev, 1985; Besnosov and Mitta, 1993; Howarth, ribbing coefficient is 1.7. The bifurcation point of the 2017) was erroneously cited as the holotype, although ribs is near the mid-flank. At the 50–60 mm diameter the author did not designate a holotype, and the type there are slight nodes, including on the simple prima- series was not monotypic. ries. With age, the ribs become arched orad, more The synonymy of G. subgaranti, cited by Dietze densely spaced and are expressed less. The body et al. (2002, p. 25), is in my opinion, unjustifiably chamber of young specimens is more than one whorl, expanded. Perhaps it is possible to agree with the adult–about 3/4 whorls. The aperture is simple, with assignment to it of transitional specimens found at the a prominent ventral projection. same stratigraphic level as the typical G. subgaranti. Dimensions in mm and ratios: However, ammonites described from Armenia by K.O. Rostovtsev under this species name (by the syn- Specimen Dm WH WW UW WH/Dm WW/Dm UW/Dm onymy above), judging from the description and illus- no. tration (shell up to 40 mm in diameter with raised bifurcation points and ends of branches on the venter) 5546/242 185 52 41 94 0.28 0.22 0.51 do not belong to this species and most likely belong to 5546/243 66 18 16 35 0.27 0.24 0.53 a species of the genus Baculatoceras from the 56 16 13.5 28 0.28 0.24 0.5 Niortense Zone. M a t e r i a l. 2 specimens from locality 5 (10) on C o m p a r i s o n. This species is most similar to the Kyafar River, Bed 2; Upper Bajocian, Garantiana V. p se u d o m a r t i n s i (Siemiradzki), from which it differs, garantiana Zone, Beds with Garantiana subgaranti. judging from the holotype (Perisphinctes pseudomartinsi Siemiradzki, 1899, pl. 22, fig. 27), mainly bipartite ribs in early whorls, with greater ribbing coeffi- Superfamily Perisphinctoidea Steinmann, 1890 cient. Family Perisphinctidae Steinmann, 1890 Remarks. A. d’Orbigny, when establishing this Subfamily Leptosphinctinae Arkell, 1950 species, gave several figures, including illustrations from the flank and venter of a subadult shell Genus Vermisphinctes Buckman, 1920 (d’Orbigny, 1845, pl. 125, figs. 1, 2). S.S. Buckman Vermisphinctes martiusii (d’Orbigny, 1845) [M] established that the illustrations of this ammonite are Plate 16, figs. 1; Plate 17, figs. 1–4; Plate 18, fig. 1 “in fact, synthetographs”, i.e., characters from differ- Ammonites Martinsii: d’Orbigny, 1845, p. 381, pl. 125, figs. 1–4. ent specimens combined in one illustration (micro- Vermisphinctes (Vermisphinctes) martinsi: Galácz, 1980, p. 107, conchs and macroconchs, in modern terminology), pl. 25, fig. 1. and pointed to the typographic error in the name of Vermisphinctes martiusii: Rioult et al., 1994, c. 111, pl. 49, the species name explaining subsequent differing figs.1. spellings (Buckman, 1920, p. 26). Nevertheless, Buck- Vermisphinctes (Prorsisphinctes) sp.: Galácz, 1980, p. 110, man designated this specimen as the holotype, which pl. 29, figs. 1. is not a valid action under the ICZN rules. cf. Vermisphinctes martinsi: Topchishvili et al., 2006, pl. 32, figs. 5. Judging by the description of d’Orbigny, the type non Perisphinctes Martinsii: Borissjak, 1914, p. 21, pl. 2, fig. 13 series included specimens with weakened ribs on the [=? Leptosphinctes sp.]. venter, currently assigned to the ancestral genus Lep-

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Plate 16

10 mm

Explanation of Plate 16 Fig. 1. Vermisphinctes martiusii (d’Orbigny), specimen PIN no. 5546/242, shells of an adult macroconch with a complete body chamber and a preserved aperture in lateral view; Karachay-Cherkessia, Zelenchuk District, Kyafar River, downstream of the village Storozhevaya, locality 36a; Upper Bajocian, Garantiana garantiana Zone, Beds with Djanaliparkinsonia alanica; collected by the author in 2018.

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Plate 17

10 mm

3a 3b

Explanation of Plate 17 Figs. 1–4. Vermisphinctes martiusii (d’Orbigny): (1) specimen PIN no. 5546/242, ventral view, (lateral view in Plate 16, fig. 1); (2) specimen PIN no. 5546/256, inner whorls of an adult macroconch (see Fig. 4b) in lateral view; (3) specimen PIN no. 5546/243, shell of a young macroconch with a complete body chamber, (3a) lateral view, (3b) ventral view; (4) specimen PIN no. 5546/241, shell of a young macroconch with a complete body chamber, lateral view. All from: Karachay-Cherkessia, Zelen- chuk District, Kyafar River, downstream of the village Storozhevaya, locality 36a; Upper Bajocian, Garantiana garantiana Zone, Beds with Djanaliparkinsonia alanica; collected by the author in 2016–2018.

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Plate 18

10 mm

Explanation of Plate 18 Fig. 1. Vermisphinctes martiusii (d’Orbigny), specimen PIN no. 5546/202, shell of an adult macroconch with body chamber and preserved aperture, lateral view; Karachay-Cherkessia, Ust-Dzheguta, District, Kuban River downstream of the village of Kras- nogorskaya, locality 19; Upper Bajocian, Garantiana garantiana Zone, Beds with Djanaliparkinsonia alanica; collected by the author in 2017.

PALEONTOLOGICAL JOURNAL Vol. 53 No. 11 2019 AMMONITES AND STRATIGRAPHY OF THE UPPER BAJOCIAN 1201 tosphinctes from the Niortense Zone, and also younger Zone on the Kyafar River based on the occurrences of specimens from the Garantiana and Parkinsoni zones; Garantiana sp. and Paragarantiana sp. is recognized as both macroconchs and microconchs. Taking all this Beds with Paragarantiana, approximately correspond- into account, the species was for a long time inter- ing to the upper subzone (Tetragona) standard Garan- preted in various ways, although it was proposed as the tiana Zone. type species of Spathia Schindewolf, 1925. In the latest The taxonomic composition of the ammonites revision of d’Orbigny’s materials, Rioult et al. (1994) from the Garantiana garantiana Zone of the Northern supported Buckman’s conclusions and designated a Caucasus at this stage seems less diverse than that of lectotype from 36 syntypes, coming mainly from Nor- the northwestern part of the Tethys. However, the mandy. These actions and published photographs of presence in the basin of the Kuban River of genera and the valid type species martiusii significantly affected some species previously described from Western the existing taxonomy (Galácz, 1980; Donovan et al., Europe suggests similar biotopes with a normal pro- 1981; Besnosov and Mikhailova, 1981; Besnosov, portion of wide-ranging and endemic species. 1982) of early perisphicnctids. Rioult et al. (1996), in my opinion, correctly assigned this species to Vermisphinctes. The lectotype ACKNOWLEDGMENTS of martiusii is only slightly different from the holotype In the fruitful part of the field work on the study of the Perisphinctes pseudomartinsi (Siemiradzki, 1899), Garantiana garantiana Zone, I was joined by my friends and which is the type species of the genus Prorsisphinctes colleagues V.V. Kostyleva (Geological Institute of the Rus- also described from the Upper Bajocian of Normandy. sian Academy of Science, Moscow), I.A. Starodubtseva Evidently, the name Prorsisphinctes Buckman, 1921 (Vernadsky State Geological Museum of the Russian Acad- should be considered as a junior subjective synonym of emy of Science, Moscow), and the paleontology enthusiasts Vermisphinctes Buckman, 1920. Microconchs of Ver- M.P. Sherstyukov (Stavropol), O.V. Trofimov (Rostov-on- misphinctes are ammonites formally assigned to the Don), O. Nagel (Radeberg, Germany), and S. Gräbenstein genus Planisphinctes Buckman, 1922. (Bodelshausen, Germany). S.V. Bagirov (Paleontological M a t e r i a l. Five specimens from localities 36 and Institute of the Russian Academy of Sciences) took the pho- 36a on the Kyafar River and two specimens from tographs of the ammonites. I am grateful to everyone who Locality 19 on the Kuban River; Upper Bajocian, contributed to the collection of the material and the prepa- Garantiana garantiana Zone, Beds with Djanalipar- ration of this work. kinsonia alanica. REFERENCES CONCLUSIONS Bentz, A., Die Garantienschichten von Norddeutschland mit besonderer Berücksichtigung des Brauneisenoolithhor- In Western Europe, species of the dimorph pair izontes, Jb. Preuß. Geol. Landesanstalt, vol. 45, 1924 (1925), Garantiana [M]/Pseudogarantiana [m] occur pp. 119–192. throughout the interval of the Garantiana Zone, while Bentz, A., Über Strenoceraten und Garantianen insbe- the species G. subgaranti Wetzel, 1911 characterizes its sondere aus dem mittleren Dogger von Bielefeld, Jb. Preuß. middle subzone. The genus Paragarantiana appears in geol. Landesanstalt, 1928, vol. 49, pp. 138–206. the upper subzone of the Garantiana Zone. The exact range of Djanaliparkinsonia (species D. bentzi (Wetzel) Besnosov, N.V., Bayosskie i batskie otlozheniya Severnogo Kavkaza (Bajocian and Bathonian of the Northern Cauca- was described from Germany) has not been yet estab- sus), Pistrak, R.M., Ed., Tr. Vses. Nauchno-Issled. Inst. lished in this region. Prirod. 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