P .. I .... nl .. I(J~; •. ul l",,,,,,.,. IM. J9. No, 5. 2005. pp. 4/17-497. Trom.lu't rl/,,,," /'ulro""'lo~;, -hr,'t;i Zilurn<ll, No, 5, 2005. pp_ JO-J/I, Or;~;",,1 RU<-li<ln Tut Cop,,-righi 0 2005 byArLJrl;~ •. ButJ;Jnow •. En~'i.'h Tmn.d",;,,,, C"pyr;~hl 0 2005 by MAIK ·'Nauhl / IMrrprr;orl;cu "(Rus.,;u).

Genus Delphinella (Ammonoidea) from the Berriasian of the Crimean Mountains V. v. Arkadiev* and T. N. Bogdanova**

*St. Petersburg State University, Universitetskaya nab. 719. St. Petersburg, 199034 Russia e-mail: arkad@GG2686.spb.edu

**Karpinsky All-Russia Research Institute of Geology. Srednii pr. 74, St. Petersburg, 199026 Russia Received April 28. 2004

Abstract-Seven species of the genus Delphinella, i.e., D. subehaperi (Retowski), D. erimensis (B urckhardt), D. obtusenodosa (Retowski), D. tresanensis Le Hegarat, D. delphinensis (Ki lian). D. janus (Retowski), and D. peetinata sp. nov., and the species Berriasella berthei (Toucas) from the upper grandis Subzone of thejaeobi Zone of the Berriasian of the Crimean Mountains are described. Their stratigraphic distribution is different from that in the stratotype area,

Key words: new species, Delphinella, Ammonoidea, Berriasian, Crimean Mountains.

INTRODUCTION

Species of the genus Delphinella frequently occur in the Berriasian of the Crimea and are very important for strat igraphy. Along with other Berriasian genera (Pseu­dosubplanites. Berriasella, Fauriella, Tirnovella, etc .) they form stable zonal assemblages eas il y traceable in the sect ions. Delphinella have been known from the Berriasian of the Crimea for a long time, although they received little monographic study except for the works of Retowski (1893) and Druschits (1960). D. delphin­ensis, D. ex gr. obtusenodosa, D. subchaperi, D. cri­mens is. and D. benhei were recorded but not described from the vicinity of Feodosiya (Bogdanova et al., 1984).

. The authors of thi s paper have a large collection (several tens of spec imens) of Delphinella she lls, col­lected over years in the Crimea by v.v. Druschits, BT. Yanin, T.N. Bogdanova, S.v. Lobatscheva, v.A. Pro­zorovskii, T.A. Favorskaya, V.V. Arkadiev, and Yu.N. Save lieva. These she lls were found predomi­nantly in the eastern Crimean Mountains (vicinity of Feodosiya) and, to a lesser extent, in the central Crimean Mountains (basin of the Tonas River), where the clayey carbonate beds of the jacobi Zone are wide­spread (Fig . I).

In the section on Cape St. lIya (Fig. 2), the assem­blage of Delphinella (D. subchaperi, D. crimensis. D. obtusenodosa, D. tresanensis. D. delphillensis, D. janus, and a new spec ies, D. pectinala) was found together with Pseudosubplallites ponticus (Re!.), P gralldis (Maz.), P. lorioli (Zill.), and P. combesi Le Hegarat in the 13-m-thick "Feodosiya Marl" Mem­ber, wh ich is presently dated the grandis Subzone of the jacoiJi Zone (Arkadiev and Savel ieva, 2002; Arkadiev, 2(03). Delphillella that were found in the vic inity of the

vi llages of Nanikovo and Sultanovka also belong to the jacobi Zone (Fig. 2) (Bogdanova et af., 1984).

In the Tonas River basin, infrequent Delphinella sp. were found together with Pseudosubplanites pontieus and P. lorioli (Bogdanova et al., 198 1).

The collection contains molds completely rep laced by rock matrix, so detailed study of the shell ontogeny is impossible .. The collection is housed in TsNIOR Museum (St. Petersburg) (coil. no. 13055). We also studied the collection of Retowski from the Feodosiya Section (TsNIGR Museum, coIl. no. 10916).

HISTORY OF STUDIES OF DELPHINELLA SPECIES

Retowski (1893) described an usual grou p of ammonoids characterized by ornamentation that

BLACK SEA o 25

Fig. l. Location of the sections studied: (1) Cape SI. l1ya. (2) village of Suttanovka. and (3 ) village of Nanikovo. Bar· akol Valley.

487

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g, E

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Sultanovka

; E

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~ Litho)og,cal column

Nanikovo

; '" E

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Litholog,cal column

; Lithological column ~~i"~~z:::"'::~ 1 • PrrutWsubplunilrs luri(J1i ~ .: ~

" '-' £ .!! .... ~ g ~ !i ,!< PrrwJoJubp!Qn;ttJ grandis. P. 8 11 : : : : : : : :

i'-flZ==Z==2jAl Puudo.ubplolli't J lorioli. P. ponli· : : : : : : :: .. CUJ. Btrrio"l/a opprli

Vl N Vl ::;: ~ pom;(uJ, P./orloli. P. combni. BrrriaJtl/o mMsicD, 8. J"beat· _ _ _ _ _ _ _ _ 8uriastllu IJ.(rthti. Dtlphintllo "" jisto, Dtlphinrllo subcbop<ri . r z ~AI OrlphintIlOlrt Jalltl1JiJ.D. jollus.

- - - - - - - - .. RtIOK'JkiCtnJJ ondruSJoK'i

11' . ~ e j ~

. !:! subchopt,i, D. crim.nsis, D. oh· : : : : : : :: .. D, crim.,ul" D. obtu$lIWsosa. 11 8 13 - - - - - - - - • ',uStl.odbJo. D; IrrSD"."sis. D. 1 1 D, ,.,rsontnJiJ. D. dt/finensiJ.

--------, I Jij

1: - - - - - - - - ","UJ. D, JUcronoto D, J'l"U'. R"",,'sktarOJ sp. ;> Z <

(,:J ________ 6 7 ___ _ ___ _

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:: : : : : : : _ _ _ _ _ _ _ _ SpiticrroJ t.' xr. t/rrudusio. • Spiticrras ffuttuS 1 13 _ _ _ _ _ _ _ _ • Ptychophyll<KtrolSrmi.ulctl· - - - - - - - -

-------- 5 12 :::::::: /um.Hop/OCtfflStiimmll'" -------- ,- . . _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ .~ PstudoJubpiamlt. {oTwl" P. poM' _ _ _ _ _ _ _ _ GP ell', Ptychopltyllocrros stmlsu/ra· :::::::: _~ __ ~ __ ~ 4 13 rum. Hap/(Juraulimmum

6110\=-------- 4 12 ________ . Pr(JrrfrOR{miluW"ric"s =======-:::::::: • /{up{oc,rase{.caracJuh,i, -------- 3 10 :-'!': ::.

s I 17

4 I 8

3 I 13

2 13.6

--------

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Btrrio .. 1l0 c/wmrrDctnJiJ. fa,,' • ri<lla c/.j1oquinrnsi., PlyeM'

pnyllocuos t:/. .rmisu/r:alum

----------------3 12 .. ' .. ... 1 ...... .

2 1212 2 1. JlupJ(JuruuiimUlum

~ Spili,r rasori,nrol •. £upnyl/a.

- - - - - - - - • euus .. rum, Pf)lehuphJII<XY· 1 12 ra, .,,,,isui<'mum, P .... 8r. in·

- - - - - - - - (JrdinOlum, /{oploctroJ rarorn· ________ Ih.i.

2 18

20

• • •

--------I 0 I •

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--------.. ' m __ '.' .. * ..

_ Pra/tlr08o,.,irtJ lauricu., Hapl/Kt . .. ru. carochlhtis. 11 . • Iimalum

[::::lJ §2 §3 ~4 ~5 B6 [!]7

Fig, 2. Corre lation of the Berriasian sequences of the Eastern Crimea. Explanation: (I) Clay, (2) mar], (3) limestone, (4) conglomerate limestone, (5) calcareous sandstone. (6) stf3tigraphic unconfonnity. and (7) levels of ammonoid occurrences .

A ~ ~

> '" " >-" ~ 8 " > Z o ~

GENUS DELPHINELLA (AMMONOIDEA) FROM THE BERRIASIAN 489

becomes smooth on the living chamber and by the pres· e nce of umbilical and lateral nodes from the Be rriasian section near Feodosiya (Hoplires calislo vac kaffae, H. calislo var. berrhei, H. calislo var. deJphinensis, H. janus, H. obrusenodosus, H. consonguinells, and H. sllbchapen). Later Mazenot (1939) described and assigned to the group Berriasella delphinemis the fol· lowing ammonites from the Beniasian of France: B. deiphinensis. B. gamieri. B. aff. janus, 8. sp. ind ., B. moravica, B. oblllsenodosa, B. aff. oblusenodosa, and B. alpillensis. Mazenot ( 1939, p . 67) ind icated that "thi s group includes BerriaseJla of small and medium size and flat shape in which the ribbed oma· mentation ... smoothens o n the body chamber begin· ning from the ex.ternal third of the flanks, spreading more or less completely to the more external and inter­nal portions of the shell ."

Based on the group of species B. delphinensis described by Mazenot, Le Hegarat (1971) described the genus Delpllinella and gave it the following diagnosis (p. 852) : "Ammonites with rather evolute coiling. The ornamentation consists of simple and bifurcating rings, di scontinued on the venter, with thickened termina· tions. The cross section is trapezoid in the first whorls to become ell iptical later. The siphonal band is stable. The ornamentation shows a trend to smoothe ning in the body chamber." Le Hegarat assigned I1 species to Del­phinella (D. autollensis, D. boisseli. D. berthei, D. del­phinensis. D. gamieri. D. obtusenodosa. D. sllbcha­peri. D. Iresanensis. D. ellenica, D. sevenieri, and D. crimensis). Later Le Hegarat (1971 ) indicated that four of the II species have bundles at the beginning of the ribs (D. aUl.Onensis. D. crimensis. D. subchaperi, and D. rresane'l sis), while these spec ies should proba· bly be recognized as a separate subgenus.

Latcr changes were made regarding the rank of Il}embers of Delphinella. Hoedemacker (1982) placed them as a subgenus in the genus Berriasella. Tavera (1985) did not accept the genus Delphinella and ass igned the species berlliei, delpilinensis. gamieri. and obtusenodosa to Berriasella (Berriasella), and the spec ies subchaperi to the genus l abronella. Partly, without indicating exactly which species he meant, he included the genus DeJphinella into the synonymy of the genera Eienaella, Fallrieila, and 7imovella.

In the new edition of the American list o f Cretaceous Ammonites. Delphinella is considered as a junior syn· onym of the subgenus Elennella Nikolov, 1966 of the genus Berriasella (Wright el al., 1996). At the same time Nikolov (1982) regarded Elellaella as a subgenu s of the genus Dalmasiceras. This opi nion was supported by Howarth ( 1992), when he desc ribed Dalmasiceras from the Chia Gara Formation of Iraq for the volume of Treatise.

In our o pinion , the gen us Delphinella sensu Le Hegarat ( 197 1) includes species with different sets of morpho logical characters. On the one hand. the list

PALEONTOLOGtCAL JOURNAL Vol. 39 No. 5 200S

includes microconches (species berthei and delphinell­sis) with s imple ornamentation (paired and sing le ribs), and, on the other hand, the list includes macroconches (species crimensis. janus, and sllbchapert) with com­ple.lt ornamentation (triple and bidichotomous ribs) and umbilical and. occasionally, lateral nodes. Neverthe~ less, we follow Le Hegarat and accept this genus. We consider the smoothening of the ornament on the body chamber and clear discontinuity of ribbing on the ven~ ter to be the main diagnostic characters of this tax.on. Because o f this, we assign the species berthei. in which the ribs are not smoothened, to the genus Berriasella.

SYSTEMATIC PALEONTOLOGY

Family Berriasellidae Spath. 1922

Genus Delphinella Le Hegarat. 1971

Type s p ec ie s. Hoplites delphinensis Kilian. 1889; Berriasian of southeastern France.

o i a g n os i s. Shells discoidal, evolute, micro~ and macroconches. Umbilicus wide and stepped. Omamen· talion consisting of e ither paired or s ingle ribs, or has triply bundled, or bidichotomous ribs. In the bifurcation point, ribs thickened , sometimes forming umbilical and lateral nodes. On body beginning from mid~height , ornamentation smoothened , although sharply increas~ ing near venter. Ribs discontinuing on venter. Suture ammonitic.

Species compo s ition. Fifteen species: D. delphilleflSis (Kilian), D. gamieri (Mazenot), D. oblusenodosa (Retowski), D. subchaperi (Retowski), D. ellenica (Nikolov), D. crimens;s (Burckhardt), D. janus (Retowski), D. aUl.(mensis Le Hegarat, D. boisseli Le Hegarat. D. /resanensis Le Hegarat, D. sevenieri Le Hegarat, D. mollovi Nikolov, D. vesse­linens;s Nikolov, D. consanguinea (Retowski), D. pec­li"ata sp. novo from the Berriasian of western Europe, Crimea, Caucasus, and Tunisia.

Co m par i so n. The genus Delphinella closely connected with the genus Berriasella, and in many cases the separation of the two genera is difficult. Large shells o f DeJphinella differ sharply from Berriasella in two ways. smoothening of the ribbing and the presence of lateral nodes. Some paleontologisls consider mem· bers of the genus Deiphinella as macroconches and those of BerriaseJla as microconches. By the smooth· ening of the ribbing, which is also observed in Dalma­siceras. Deiphinella differ from this genus in the absence of umbilical nodes and by the presence of pro· nounced lateral nodes. Delphinella is very similar to tubercular berriasianellins, for e.ltample. Malbosiceras s.1. (Wright el al., 1996, p. 50), from which it is distin~ guished by the presence of a sharp discontinuity of the ribs on the venter with the rib terminations thickened on the ventrolateral shoulder, by the absence of umbilical nodes, and by the thi nner ( less robust) ribs.

41)(l ARKADIEV, BOGDANOVJ\

Plalc 5

1',\ J.EON I{lI.Otj lt ' Al, JOUk NI\1. Vu!. III Nil,.' 2011.'

GENUS fJELPI-IINELLA (AMMONOIDEA) r:1~OM THE BERRIASIAN 4lJ I

f) e/phinella )'"bclraperi (I{clo .. '.~ ki, 11I9J)

PI~le .'\. figs. )\ ilnd 9

1foJllile.t subdwperi: Retowski. 1893. p. 64, pI. 4, figs. 3 and 4; Simioncseu, 1899, p_ 5. pI. 1. fig_ 2.

Olllmasiam:; ? suhchaperi: Grigorieva. 1938. p. 118. pI. 7. figs. 2a and 2b.

Berriaselfa Subc1wpf'ri: Luppov el al .. 1949. p. 220. pI. 63. fig. I; non Berriaselfll $uhchaperi: Drusehils. 1960, p_ 276, pi. 21, fig. I (= Relowskiceras tlntirus.wwi (Ret. ).

Oelphinelfa subcJllIperi: Le Hcgarat, 1971. p. I 12, pI. 13, fig. 12. pI. 42. fig. 10: Bogdanova el al.. 1984. pI. 4. figs. 4 and 5: Khimshiashvili. 1989, p. 17, pI. 4 . fig. 1

Oelphinella oblusellOdosa: Nikolov. 1982. p. 86. pI. 20, fig. 5.

Jabrollellll subchaperi: Tavera. 1985 . p. 304. pI. 46. fig. I. text~ fig.2]/A.

S h e ll s ha pe. She ll s are large, d iscoidal, wi th slowly ex pand ing whorls. Flan ks are wide, almost flat. The venler is ftauened. The umbilicus is shallow, mod­erately wide, with a narrow, slopi ng umbilical wall. The umbil ical shoulder is gently sloping. The whorl cross section is rectangul ar to oval, compressed.

Orname nt ation. The whorls less than 50-60 mm in d iameter have very th in and dense ribbing. Pai red ribs dichotomi zing at mid¥f1 an k or slight ly above predominate. Very rarely the bi furcation occurs near the umbilical shoulder with fu rther dev iati ng of both ribs. S ingle intercalating ribs are present. In the adul t whorl two pairs of thickenin gs appear on the ribs: umb ilica l and lateral (at the point of rib bifu rcation). Umbilical swellings virtually do not develop into nodes bu t remain as e longated sharp ridges. These ridges give rise to indi sti nct dic hotomous ly branch ing ri bs. Approximately in the midd le of the whorl, these branches are connected by nodes, whic h in some spec­imens are elongated and poimed, and are orie nted pe r­pendicular to the ribs. After the lateral node in the upper part of the whorl wi th in the pair of ribs, a th ird ri b appears. Two or three sing le intercalating ribs are present between the bund les. These ribs begin from the mid-whorl or slightly lower. Near the lateral nodes the ribs are smoothened and indistinct. Near the venter the ri bs are distinct, wide, and ri bbon-like. They are inter­rupted on the ve nter by a smoot h band. The rib te nnina­tions on the ventrolatera l shoulder are directed slight ly forward.

Co m pa r i son. This spec ies is distinguished from the si milar species D. obwsellodosa by the less robust

ornamentation in ear ly whorls, more pronounced umbilical ridges, and by the lateral nodes in adults.

R e m ark s. Following Grigorieva (1938) we include the spec imen figured by Simionescu, 1899, pI. I, fig. 2 in the synonymy list of this species. although it differs somewhat from the typ ica l subcha­peri in the longer (reaching the umbilicus) intercalating ribs, we th ink that thi s is just variability. The specimen figured and desc ri bed by Nikolov (1982, p. 86, pI. 20, fig . 5) as D. obtusenodosa has st rongly pronounced umbilical nodes. We assign thi s spec imen to D. subcha­peri. The spec imen fi gured by Druschi ts (1960, pI. 21. fig . 1) has coarse ribs (wit hout smoothening) and sharpl y pronounced lateral nodes and most likely should be assigned to Relowskiceras andrussovi.

o c cur r e n C e. Berriasian. jacobi Zone of the Crimea (graf!di.~ Subzone), Caucasus, Bulgari a, south­eastern France, Spain; Berrias ian of Switzerland.

M a t e r i a I. Three specimens (nos. 1113055-3113055) from Cape St. lIya and the vi ll age of Sultanovka; coil. by T. N. Bogdanova.

Delphinella crimt niis (Burckhardl, 1912)

Plate 6, figs. 7-9

Hopliles calislO d'Orbigny var. de/phillensis: Retowski. 1893. p. 57. pl.]. fig. 4.

HopUles occiJaniclIs: Retowski. 1893, p. 60, pI. 3. figs. 7 and 8. non fig. 9.

Sleueroceras cr;mense: Burckhardt, 1912. p. 165. De/phinefla crim€me: Le Hcgarat. 1971. p. 10]. pI. 42. figs. 6

and 8: l3ogdanova el al .. 1984. pI. 2. fig. 6; pI. ]. fig. 6.

S h e II s h ape. She ll s are medium-s ized. wi th dis­coida l, moderate ly low and weakly expand ing whorl s. The flanks are strongly flattened . The venter is narrow, very weakly convex. The whorl cross section is high and rectangular to ova1. The umbilicus is moderately wide and shallow. The umbi lica l shoulder is smooth. The umbilical wall is narrow and gently slopi ng.

Or n a m enta ti on. On the inner whorls (up to Dm = 30-35 mm), the ornamentation consists of very th in , densely spaced, weakly curved ribs. Tri ply bun­dled ri bs with low bifurcation points at the umbilical shoulder and high bifurcation poi nt s of one of the branches in the mid-flank dominate . In addition, doubly bundled ribs wit h bifurcation al the umbilica l shoulder and paired ri bs dichotomizi ng at the mid-whorl are

Explanation of Plale 5 In all cases Ihe size is natural. figs. 1-3. OelphilreflajanllS (Relowski): (I) specimen no. 35/ 1305: (I a) laler,1I view. (I b) ventral view: village of Sultanovka; gratl­d;s Subwne: (2) specimen no. 3311 3055. later::t! view: village of Nanikovo: grandis Subwne: (3) specimen no. 32/ 1 3055: (3a) lal­eral view. Ob) ventral view: Cape SI. Ilya: gramlis Subzone. Figs. 4 and 5. Del/,/rinel/(I /reswrmsis Le Hcgarnt: (4) specimen no. 20113055. l,lIer;!1 view: village of Nanikovo; gmntlis Subzolle; (5) specimen no. 19/13055. lateral view: l3:lrakol Valley: gml1dis Subzone. Fi~~ . 6 and 7. Dell,/riwlla O/JllI.\·/'rr"dll,wr (Retowski): (6) specimen no. 16/U055. lateral view: Cape St. Ily:l: .~m/l{lis Suhzone: (7) ~pecitnen no. 1.'1/13055,laler,lI view: the same ,lge ;uullocality. Fil!-~' Hand 9. Ild"iJilldlu .m/x:/IUI",ri (RelOwski ): (8) spcdl11en no. 1/13055. 1:I(cr;11 vie\\': village of Sultamwb: .~rmrd;J Subz(lllc: (')1 sll\;(imen no . . V I 3055: (9a) lillcr:11 view. (9b) VC1llr,rl vie\\'; thc .~amc age and lo .. :alil'!.

1';\I.I :() ,'.:- rOI.OlilCAl. JOUI{ 01AI. Vpl. 1') Nu. 5 20()5

ARKADIEV, BOGDANOVA

Plalc6

I'i\LEONTOLO(; ICi\\, JOURNi\L Vul. ,I'J Nu, ~ 200~

GENUS IJELPNlNELLII (AMMONOIDEA) fROM THE BERRIASIAN 493

present. Rare ly, bidichotomous ribs and . very rarely. si ngle ribs arc present. No pattern in a lternat ion of the rib types is present. In the body chamber at WH = 23-25 mm, the whorl s increase more slowly in he ight. while the ribbin g sll100thens in the mid-flank . At thi s stage. thin single crestlike ribs are seen runnin g from the umbilica l shoulder to one-third of the who rl he ight. Further, the ribs smoothen to the extent that the pl aces of their bifurcation become very indi stinct. After branching, the ornamentation consists of bundles of two or three ribs, which are sharpl y pronounced near the ventro lateral shoulder. In the inner whorl s, the ribs are separated on the venter by a smooth band that dis­appears in the body chamber because ribs continue onto the venter.

Dim e n s i o n s in mm, ratio s, a nd num ­b er of whorl s p e r half whorl:

E Number of ribs Speci- E E per half whorl

men no. ~ e ~ ~ E '" " '" inner outer

Cl ~ ~ ::> ~ ~ ::> 4/13055 68 24 10 23 0.35 0 . 15 0.34 34 64

30 25 5/ 13055 50 23 14 0 .46 0.28

28 20 4S 6/ 13055 71 32 18 0,45 0.25

7113055 46.5 19 12.5 0.41 0.27 24 49

Co m par i so n. Thi s species diffe rs from other species in the di versity of ribs and their considerab le smoot hening in the body chamber.

R e mark s. Burckhardt (19 12) only named the spec ies crimeflsis, without describing o r figuring it. Le Hegarat ( 1971) was the first to describe the species. This author placed th is spec ies in the synonymy list of Rop/ires calisto d 'Orbigny var. de/phinensis, described by Retowski ( 1893). We studied Retowskii 's co llecti on and concl uded that the variety delphifleflsis, similar to specimens described by Le Hegarat, is characterized by the development of various ribs: triple (bundled), bidi­choto mous, paired, and single.

o c c ur r e n c e . Berriasian, jacobi Zone of the Crimea (gralldis Subzone), southeastern France, Spain.

M at e r i a I. Nine specimens (nos. 4/1 3055- 12113055) from Cape SI. l1 ya, Barakol Valley. and the village of Sultanovka: coli. by Y.Y. Drusc hits, BT. Yanin . T.N. Bogdanova, and vv. Arkadiev.

Delphinella obluu nodosa (Rcl(lwski. 11193)

Plale S. figs. 6 and 1: Plate 6. fig. 6

Hoplites obtusenodosa: Retowski. 1893. p. 62. pI. 3, figs. 10 and 1 \.

IJerriasel/u obtu.renodosa: Ma7cnol. 1939. p. 73, pI. 8. figs. 3 and4; Druschits. 1960, p. 275. pI. 22. fig. l.

lJerriasd/a n. sp. ind. (gr. n. delphinellsis ): ,\ i3z.cnot. 1939. p. 71. pI. 6. fig. 19.

non Berriasella aff. oblusellodosa: Maz.cnot. 1939. p. 72, pI. 7. figs . 6 and 7 (= De/phinel/a el/enica (Nikolov).

non Benimel/a oblUsenodosa: Amould-Sagct, 1953. p. 46. pI. 5. fig. I (=? Berriasella).

? Berriasella aff. obltlsenodosa: Lc Hcgarat and Rcmane. 1968. pI. 4. fi g. 2.

non Oelphinella oblusenodosa: Khimshiashvili, 1976. p. lOO. pI. 16. fig. 6.

non Berriruelia (De/phinel/a) obtusenodosa: Sapunov. 1979. p. 178. pI. 56. fig. 8 (= Delphinellajanus (Rctowski ).

De/phinel/a obtusenodosa: Le Hegarat. 1971. p. 109. pI. )3. fi gs. 10. 11 , 13. and 14; pI. 41. figs. 6 and 7 (non fig. I): Nikolov. 1982. p. 86. pi. 21. fig. 1 (non pi. 20, fig. 5): Bogdanova et al . . 1984. pI. 4. fig. 2.

Sh e II s h ap e. She ll s are medium-sized to large, wi th di scoidal, low, moderately expandi ng whorls. The whorl cross secti on is high and rectangular. The umbi ­licus is shallow and mode rately wide. The umbilical wall is narrow, poorly de lineated, and gently sloping.

Or n a men tat i on. Flanks on the inner whorls up to Om = 25- 30 mm are covered by thin bifurcating strai ght ribs. They begin on the umbilical wall , lean weakly backwards on the umbilical shoulder, and after that sli ghtly slanting orad run across the flank. Some­what above the mid-flank , the ribs branch into two sim­ilarly pronounced ribs, with the posterior lean ing slightly backwards. On the venter, the ribs are inter­rupted by a smooth narrow band. The ornamentation of ad ult whorls at Dm = 50 mm consists of paired and. more rarely, single ribs. Weak crests develop on the umbilical shoulders of the body chamber. Swel lings develop in the bifurcation points. These swellings sometimes become indi sti nct e longated nodes. Imme­diately after the swellings, ribs smoothen. This effect disappears near the venter. The ribs on the venter of the

E xplanati o n of P la te 6 In all cases the size is natural. Figs. I and 2. Delphillel/a peetillata sp. nov.: (t) holotypc no. 3911 )05.'i . lateral view : Cape SI. tlya: gnmdis Subzonc; (2) sp..:ci men no. 40/1 3055. lateral v\cw: the same age and locality. Figs. J and 4. Berriaulla bertlrei (Toucas): (3) specimen no. 421 13055: (3a) vent ral view. (3b) lateral view; village of Sultanovka: grmrdis Subzone: (4) .~pccimen no. 41/13055. lateral view: Barakol Valley: /i,mrdis Subzonc. Fig. 5. De/phincl/a dcll'iri.rl'lrsis ( Ki lian): specimen no. 28/ 13055. lateral view; village of Nanikovo; gramlis Subzone. Fig. 6. Delplrind!a ohtUSI'/I(){!OSll (Retowskil; specimen no. 1311)05): (6a) ventral view. (6h) aper1ural view. (6c) lateral view: Cape SI. Ilya: gmndis Subznl1e. Figs. 7- 9. Ol"philrel/a uil/J/'lIsix (Burckhardt ): (7) specimen no . . ~I t 3(5). lateral view: Cap..: SI. Ilya: gmm/iI SUblOlle; 8) .~pccimen 110. 4/1305.'i: (ga) ventral view. (Xh) brteral view ; Barakol Valley: ,~mlldi.{ Sub/.one; (9) specimen no. 6/ 130.'i). lateral view: village of Suhanovka: g/"lllllfi.\· Suh/.onc . • ' il-: ' tU. {)d/J llindlajllllus (Retowskil; spec imen no. 341130.'i5. !:ttcral view: Cape SI. lIya: gmlldix Suhl.llne.

!'A1.EONTOl.(l(iICM . J(ll IRNA I. V"I. .N Np . .'i ::! 1I0~

494 ARKAD1EV. 130GDANOVA

body chamber arc separated by a smooth band. Rib ter­minat ions are curved antcriorly.

Dim e n s ion s in mm, ratio s, and num ­ber o f who rl s p e r half wh o rl:

E E Number of rihs

Specimcn E per half whorl

" " " 00.

" - ii ii E '" " '" inner outer

" " " => " " => 13/1305551 20.5 9.5 16.50.400.19 0.32 22 42

141!3055 26.5 I I 7 0.42 - 0.26 25 48

Co m par i so n. This spec ies differs from D. sub­chaperi in the less stron gly pronounced umbili cal ridges and weakly deve loped lateral nodes.

R e m ark s. Khimshiashv ili (1976) described D.obruSelZodosa from the Northern Caucasus. The attentive study of hi s paper shows that the only speci ­men from the Caucasus possesses tripl ed ribs and has no smoothening of the o rnamentation, femures that are not characteristic of D. obtuseflodosa. Because of this, the authors of this paper do not place th e spec imen from the Northern Caucasus in the synonymy list.

O cc urr e nce. Berriasian, jacobi Zone of th e Cri mea (gralldis Subzone); Bulgaria, southeastern France, and Spain.

M a t e r i a I. Six specimens (nos. 13/13055- 18/1 3055) from the Cape St. Il ya and from the vi llage of Sul­tanovka; col I. by T.N. Bogdanova.

D<.>lphinellll IrtSlln t nsis Le Hcgaral , 1911

Plate 5. figs . 4 and 5

f/op/iles occilanicus: Relowski. 1893. p. 60. pI. 3. fig. 9. non figs. 7 and 8.

De/phinel/a lresanensis: Le Hegarat. 1971. p. 113. pI. 13. fig. 15; pI. 42. figs. 1 and 2.

Berriasella (Deiphinel/a) lresanensis: Hocdemaeker. 1982. pI. 1. fig. 4.

, S h e 11 s h ape. Shell s are medium-sized, flat­tened, wi th high moderate ly expanding whorls. The flanks are weakly convex. slightly diverging toward the umbilicus. The venter is narrow, flat , and smooth. The umbilicus is shallow, moderatel y narrow, with low sloping wall s. The umbil ica l shoulder is very rounded .

Or n a m e n tat i o n. The ribbing is thin and dense. The ribs are weakly curved, paired. tripled (bundled), and occasionally single. In the tripled rib. the bottom branching occurs somewhat above the umbilical shoul + der. One of the branches again fork into two branches slightly below the middle of the flank. Paired ribs, branc hing on the umbilical shoulder or subdividing approximately in the mid-flank are present. Sparse sin ­gle ribs, appearing near the umbilical shoulder, arc presen t. Whorls about 60 nUll in diameter show weak­ening of the ribbing in the mid-flank. which is clearly vis ible in the spec imen fi gurcd by Retowsk i ( 1893. pI. 3. fig . 9). All ribs arc flattened toward the pcriphery and arc interrupted 011 the vcntrolateral shoulder. It.::av­ing thc vcntcr smoot h.

Dim e n s io n s in m m. rat i 0 s, and n u m -b e r o f whorls p e r half wh o rl :

E E E Number of ribs

Speci men

" " " per hal r whorl

00.

'" 3 3 'i' ii ii E inner " " " => 3 3 => ouler

t9113055 63 28 18.5 0.44 0.29 27 60

201 13055 62 J2 13 10.5 0.52 0.2 1 0.17 24 55

21113055 39 18 11 0.46 0.28 24 48

22113055 27 14 8 0.52 0.30 2J 47

Co m par i s 0 n. This species differs from the sim­ilar, densely ribbed spaces D. crimensis in the sparser and straighter ribs in the inner whorls (up to Om = 25-30 mm) and also, to a lesser extent, in the smoothening of the ribbing on the body chamber.

o c c ur r e n c e. Berriasian, jacobi Zone of the Crimea (gralldis Subzone), southeastern France, and Spain.

M a t e r i a I. Nine specimens (nos. 19/ 13055-27113055) from Cape St. Il ya, Barakol Valley, and the villages of Nanikovo and Sultanovka; co il. by y.y. Druschits, T.N. Bogdanova, and VY. Arkadiev.

Delphinella delphinensiJ (Kilian, 1889)

Plate 6. fig. 5

Hop/iles de/phinensis: Kilian. 1889. p. 662. fig. !.

Hop/iles ca/islo: Rctowski. 1893, p. SS. pI. 3, fig. 1, non Hop­lires calislo var. de/phinensis: Rctowski. 1893. p. 57, pI. 3. fig . 4 (= D. crime/uis).

Berriasella de/phinensis: Ma:tenot. 1939. p. 67. pI. 6 . figs. 14 and 15; Amou ld-Sagel. 1953. p. 45. pI. 4. figs. JO and Il.!cx[-fig. 14.

De/phi/Jella de/plrinensis: Le Hl!garal. 1971. p. 104, pt. 13. figs. 7 and 8; pI. 42. figs. 3 and 9: Nikolov. 1982, p. 86. pI. 20. figs. 2-4. non Berriasella (DelphinellaJ cf. de/phinensis: Sapunov. 1977, pI. 6, fig. 5; 1979. p. 177. pI. 56. fig. 7 (= De/phil/el/a mol/ovi Nikolov).

She 11 s h a p c. She ll s are small (mi croconches). with slowly ex panding whorl s. The flanks are flattened . The venter is flat. The umbilicus is shallow and wide. The umbilical wall is narrow. indistinct, and gently slop­ing. The whorl cross secti on is high, rectangular to oval.

o rn a men ta t i on. The flanks are covered by straight or weakly curved bifurcating or. less com­monly, single ribs. The whorls 20-30 mm in diameter possess weak swe lling and flattening of ribs. A general smoothening of the ribbi ng is deve loped above the mid­flank . The ribs do not cross the venter, on which a wide smooth band is devel oped.

Dimen s i o n s in mm. ratio s, and num ­b e r of whor l s p e r half whorl :

E E Number of rihs

Specimen E per half whorl

" " " no.

" " ~ 'i' ii ii E imlcr

" ~ " => ~ " => ()u[cr

1Xf1 .~05 .~ 41 15.5 11 .~ (Ut-: 0,:\0 25 41i

GENUS OELI)/IINEtLA (AMMONOIDEA) FROM THE 3ERRIAS IAN 49,

Co III par i so n. This species differs from most spec ies in the smaller size (microconches) and early srnootheni ng of the ribbing. The latter character is very weakl y developed .

R em a rk s. The study of Retowski 's collection showed that the specimen of Hoplifes caliSlO figured by him (RelOwski . 1893. pI. 3. fig. I) has the ribbi ng smoothened on the body chamber and . hence. should be assigned to D. deipliillensis. Prev iously Arkadiev and Bogdanova (2004 ) identifi ed thi s spec imen as Berria­sella ca/isto. The variety delphillellsis. described by RelOwski. 1893. shows various types of ribs (t riple. bid­ichOlomous, etc.). which does not agree with the diag­nosis of the species deiphillellsis given by Kil ian (1889). Triple ribs are present in B. (D.) cL delphillell­sis. described by Sapunov ( 1977. 1979). which appar­entl y instigated Nikolov ( 1982) to recognize it as a new spec ies. D. moflovi.

o c c ur r e n c e. Berriasian. jocohi Zone of the Crimea (gral/dis Subzone). Bulgaria. southeastern France. Spain , and Tunisia.

M ate r i a I. Three specimens (nos. 28/13055-3011 3055) from the Cape SI. lI ya, Barakol Va lley, and lhe village of Sultanovka; coli . by V. V. Druschits and T.N. Bogdanova.

Dtlphintllajanus (Relowskl . 1893)

Plate S. figs. 1-3; Plme 6. fig . 10

Hoplittsjmuu: RClowski , 1893, p. 59, pI. 3, figs. 5 and 6.

Berriastlla aff.jmUls: MazenOl, 1939. p. 70, pi . 6. figs. 20 and 21.

Delpilinellajamu: Nikolov. 1982. p. 88. plo 21. fig. 5.

Delphinella dd(llzinensis : Bogdanova et al., 1984. pI . 2. fig. 3.

S h e II s h a p e. The she ll s are discoidal and medium-sized. The whorls are high. weakl y overlap­ping. and slowly ex pandi ng. The fianks are very weakly c-onvex. The venter is narrow and weakl y convex. The umbil icus is wide and shallow. The umbi lical shoulder is rounded. The umbili cal wall is narrow and steep.

Orna m e nt a ti o n. In juvenile whorl s (up to Dm = 40 mm), the fiank s possess vari ous ribs. mai nly paired and trip led (bundled) and. less commonl y. bidi­chotomous and single. The ribs beg in in the middle of the umbilica l wall. The ir terminations are directed sharply forward. On the umbilical shou lder. they form an arc di rected backward, whereas on the fi anks again slant forward. The ribs run straight across the flanks. The first branching occ urs on the umbilical shoulder; the upper branching is in mid-fl ank . In the upper pall of the whorl . a ll ribs become flattened. and soon after crossing the vent rolateral shoulde r di scontinue to leave a smoot h band on the venter. The shape of the ribbing on the body chamber at Om = 50- 60 mm changes . Paired ribs prevail . and single ribs arc less common. In the Illiddle part. the ribs smoothCIl: hence. the tipper br.mching is indistinct. The frequency of ribbings com­pared to younge r Slages is reduced. The ribs become th icker and sparser.

D ime n s i o n s in mm. r at io s, and n um ­ber o f w horl s per h a Ir whorl:

~ E E

Numhcr ofribs Specimen 0 0 per half whorl

'0. ~ :> -E ~

~ '" ~ 0 ~ ~ ::> ~ ~ ::> inner oUler

3 1113055 66 25.5 ]I 21.5 0.39 0. 17 0033 28 60 32113055 " 18.5 9 - 0.36 0. 17 25 " )3113055 " 19 9.218 0.36 0. 17 0.34 25 53

Co m par i so n. This spec ies differs from D. del­phillensis in the larger size and more intense ribbing, wh ich become smoother later in ontogeny. It is distin­guished from the closely related D. crimensis by the less dense and more crestlike ribs on the body chamber.

o c c ur r e n c e. Berrias ian, jacobi Zone of the Crimea (grandis Subzone). Bulgaria , and southeastern France.

M ate r i a I. Eight spec imens (nos. 3111 3055-38113055) from Cape SI. lIya, Barakol Valley. and the vi llages of Sultanovka and Nan ikovo; colI. by T. N. Bogdanova, Y.Y. Drusch its, and V.V. Arkadiev.

Ihlphilltlla ptclinata Arkadie .. et 8ogdnnD~·II. sp. no ...

Platt 6. figs. 1 and 2

E l y m 0 l og y. From the Latin pectinarus (pecti­nate).

H o l o t Y p e . TsNIGR Museum, no. 39113055; Feodosiya, Cape SI. lIya. Berriasian, jllcobi Zone, gramlis Subzone.

She I I s h a p e. Judg ing from two fragments of adult whorls. Ihe shells are discoida l, with wide flal ­tened flanks and narrow ven ter.

Or n a III e n t at i on. Two fragments of a quarter of a whorl possess a distinctive ornamen tation. Six inner ri bs arc present. They begin on the umbilical shoulder wi th small spiky nodes and slan t orad. Somewhat above one-third of the height of the whorl a crestlike swelling on the fl anks is present. Ribs near nodes are naITOW and sharp and, afte r entering the swe lling. become fiat­tened. wide. and indistinctly bifu rcate. The swelling subdivides the fl ar:k in to two unequal pans. Before the swell ing. the thickness of the shell increases to become slightly reduced later. After the swel li ng. a wide depres­sion is present on the shel l. Hardly noticeable ribs run across the depress ion, slanting backwards. Near the vent rolateral shoulder, the ends or the ribs become sharp. distinct. and thin . They are then turned fo rward again . An additional two or three ribs appear belween the bundles of ribs. The number of exte rnal ribs on a quarter of the whorl reaches 25. On the venter, the ribs are intermpted by a narrow smooth band .

Co III par i so n. Th is spccie$ di ffers from DeI­phillel/a in the very distincti ve orn amen tation.

o c c tI r r e n c e. Berrias ian . jac:o/Ji Zone, Kramlis Suhzont! of the Crimea.

496 ARKADIEV, BOGDANOVA

M ate r i a I. Two specimens (nos. 39113055 and 40113055) from Cape SI. lIya: coli . by v.Y. Arkildiev.

Genus Herriaselfa Uhlig, 1905 Rtrri.o.stlla btrtht'; (Toucas.. 1890)

Plate 6. fi gs. 3 and 4

lIoplites callislO ... ar. her/hei: Toucas. 1890, p. 601, pI. 17. figs. 6 arid 7: Rctowsld. 1893. p. 55, pI. 3, fi g. 3.

Berriasdla /x!rtirei: Mazcnot, 1939. p. 48. pI. 2. figs. 9 and 10. non figs . 11 and 12 (= D. sevtl1ieri): Amould-$aget. 1953. p. 26. pI. 2. I1gs. 12 and 13; pI. 3. fi g . 3: Le HcgarJl and Remanc. 1968. pI. 5. fig . 5.

De/p/rinella bertllt!i: Le Hcgarat . 1971. p. 99. pI. 13. figs. 3-5: pI. 42. figs . I1 and 12: Nikolo .... 1982. p. 87. pI. 21. figs . 2-4.

Berriasdla (BerriasellaJ herthe;: Ta ... era, 1985, p. 248. pt. 34, fi gs. I arid 2. text.fig. I9IH.

5 h e ll s h a p e. The shells are small (micro-conches), fl attened, wit h high, moderate ly ex pandi ng whorls. Flanks are weakl y convex . The venler is fl at­tened. The umbi licus is wide, with low, gently or steeply sloping wa ll.

Or n a me n ta t ion. The flanks possess relati ve ly coarse, straight. mainl y bifurcating ribs. Branc hing occurs in the middle part of the flanks. Sparse single ribs beg inning from the umbilicus are present. On the body chamber, al the place where ribs are branching, crestl ike swellings are developed. The ribs become coarser, fl atter, and sparser. The rib tenninations on the ventrolateral shoulder are weakly curved forward. On the venter the ribs arc interrupted by a narrow smoOlh band.

Dime n s i o n s i n mm, rati os, and num­b e r of who rl s p e r ha lf w h o rl :

E Number of ribs Specimen E E per half whorl e " " "0. ,. ~ ~ E '" ,. '" inner outer

" ,.

'" => ,. ,. => 4 tll 3055 45 17.5 - 16 0.39 0.36

421 13055 Jl 12 6.5 9 0.39 0.21 0.29 20 39

Co rn par i so n. This species is dist in gui shed from ot her spec ies by the type of ri bbing near the venter and slight swell ings at the points of rib bifurCal ion.

R e in ark s. This species has a conflicting set of characters. On the one hand. it shows a weak thickening and flatteni ng of the ribs on the body chamber, which is characteristic of the genus Delphinella; on the other hand , the ribbing (pai red and sing le ribs) and the absence of smoothen ing of the ri bbing suggest it s affin­ity 10 the genus Berriasella. Such a combinat ion of characters certain ly indicates a close connect ion between these gene ra. Wc decided to leave the species berrhei in the genus Berriasella. However, its precise diagnost ics will be possible onl y when new dala on the morphogenesis of the shell s and the sUlura l out line become ava ilable.

o c cu r r e n cc. Bcrri ' l" ian.jllcohi- occ:ifllllic{I Zones (.~III}{/lpi"(I Subzone) soutllc;tstc rn Franc\!, Bu lgaria ;

Bcrriasiiln , jacoh; Zone of the Crime;] (grandiJ Sub­zone), Spain , and Tuni sia.

Materia l. Two specimens (nos. 41 / 13055-42/ 13055) from Cape St. llya and from the Barakol Val ­Icy; coil. by T N. Bogdanova.

CONCLUS IONS

As a resuh of th is study, a rich assemblage of Del· phinella species (seven spec ies, includi ng D. subcha· peri, D. crimensis, D. obtllsenodosa, D. lres(lnensis, D. deipliinensis, D. janus. and D. pecliflOta) has been described for the fi rst time from the Berriasian of the Crimean Mountains. Th is assemblage a ll ows the recog­nition of the standard jacobi Zone in the section and correlation with southwestern France (Le Hegarat, 197 1), Spain (Ta vera, 1985), Bulgaria (Nikolov, 1982), CaucaslI s (Khimshi ashvili, 1976, 1989), and other areas of the Mediterranean region.

The cllOmeracellsis Subzone, recogn ized by Ark­adiev in the Feodosiya section in the lower part of the jacobi (Arkadiev and Save lieva, 2002; Arkadiev, 2003), is presen tly characterized only by members of the gen­era Berriasella and Fauriel/a. De/phinella have not been fou nd in thi s subzone. In the C rimea all records of thi s ge nus are restricted to the grandis Subzone, where they have been found together with Pselldosubp/anites, Berriasella, and RelolVskiceras. The study by Arkadiev of the section of the jacobi Zone in the bas in of the Tonas Ri ver in 2003 showed that very few Delphiflella occur with Pseudosllbp/aflires grondis and other Pse/ldoplmllllites, i.e., they are a lso characteri stic onl y of the grandis Subzone. Furthermore, in the lower part of the Tonas section, below the levels wi th Pse/ldosllb­p/anites grandis, the species Berriasella jacobi was found . This allowed the subdivision of thejacobi Zone into two subzones. jacobi and grandis, rather than chomeracellsis and gram/is, as was suggested prev i­ously (A rkadiev, 2003).

For southeastern France Le Hegarat ( 197 1) noted that D. crimensis, D. delphillensis, D. oblllsenodostl, D. subchaperi, and D. Iresanensis are characteri stic only of the jacobi Zone in Le Hegarat's terminology. i.e., fo r the lower part of the jacobi Zone in its modem understanding (Hoedemaeker and Bulot. 1990). No separation of the finds of De/phinel/o and Pseudosub­p/anites have been so far found in the Crimea, perhaps, because a ll the sections stud ied (Cape SI. lIya, Su l­tanovka, and Nan ikovo) are incom plete. They do nOl ex pose depos its covering thejacobi Zone in a complete sec tion. Hence, wc can correlate the Crimean gram/is Subzone onl y with the en tirejacobi Zone. However, the analysi s of ammonites from Retowski 's co llection a llows the recogni tion of the overlyi ng occitallica Zone in thl! Feodosiya Secti on (Bogdanova er (11., 1999). Most likel y. depos its of th is zone arc presentl y in the bu il! up part of the town and are not exposed.

I'ALt:ON']'OI.(XiICAL JOURNAL vU!. W Nu. S 200S

GENUS fJELPIIINEU..II (AMMONOlDEA) FROM THE BE RRIAS IAN 497

We emended the di agnos is of the genu s De/phille/lo, accordi ng to w hich the s pecies bertltei is assigned to the genus Berriasellll.

A c lose connection of the Be rrias ian marine bas in of the C rimean Mountains ex.isted w ith the western Euro­pean bas ins. Of 15 s pec ies of the genus De/phinel/o known now, seven a re recorded from the C rimea, of whic h five arc sha red wi th France.

ACKNOWLEDGM ENTS

The study was supported by grant E02-9.0- 111 of the M inistry of Education o f R ussia in the Field of Nat­ural Sciences.

REFERENCES

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16. W. Kilian, "Mission d'Andalousie. 2. Etudes paleon­tologiques sur lcs terrai ns sc<:ondaire et tertiaires de l' Andalousie," Mem. pres. par d ivers savants Acad. Sci. Inst. Francc 30, 601 - 75 1 ( 1889).

17. G. Le Hegarat, "Perisphi nctidae et Beniasellidae de la lim ite Jurassique Crctace. Genres nouveaux et rcvision crit iquc de quelques defi nitions taxonomiques anterieures," C. R. Acad. Se i. Pari s. Scr. D 273 (10), 850-853 (197Ia).

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