Cainozoic Research, 7(1-2), pp. 109-117, April 2010

A in between striking convergence conchological morphology

Oligocene-Miocene lottiids (, Patellogastropoda) from

the North Sea Basin and the Paratethys

Olga+Yu. Anistratenko¹Adri+W. Burger² & Vitaliy+V. Anistratenko³

1 Institute of Geological Sciences ofNationalAcademy ofSciences ofthe Ukraine, O. GontcharaSir., 55-b, 01601, Kiev,

Ukraine. E-mail: [email protected] 2 P. Soutmanlaan 18, NL-1701 MC Heerhugowaard, The Netherlands. E-mail: [email protected]

3 1.1. SchmalhausenInstitute ofZoology ofNationalAcademy ofSciences ofthe Ukraine, B. Khmelnitsky Str., 15, 01601.

Kiev, Ukraine. E-mail: [email protected]

Received 24 February 2009; revised version accepted 14 February 2010

The protoconch and teleoconchmorphology of Patella compressiuscala Karsten, 1849 (North Sea Basin, Chattian - Langhian) is de- The Boreobliniais established for this which is characterized indicative of scribed and illustrated. new species by a protoconch a

lecithotrophic typeof early development, lacking even a short free-swimming larval stage. The distinctness ofBoreobliniagen. nov. from

other Patellogastropoda such as Tectura,Patella, and Helcionexhibiting the typical patellogastropodprotoconch types is supported also

by its unusual shell structure. An amazing heterochronous convergence ofboth protoconch and teleoconch morphology between the new

species from the North Sea Basin and Miocene Flexitectura subcostata (Sinzow, 1892)from the Sarmatianofthe Paratethys is shown.

Detailed descriptions and SEM images of species involved are presented.

KEY WORDS:- , Lottidae, Miocene, Europe, protoconch, new genus

Introduction marine gastropods with a planktotrophic larva in their on-

togeny {e.g., Bandel, 1982, 1991; Sasaki, 1998; Kahn,

2004). The characters of embryonic, larval, and juvenile Oligocene and Miocene patellogastropods from the Medi- shells can be used to reconstruct the phylogeny of some terranean and Paratethys as well as from the North Sea the of the the gastropods. In case patellogastropods proto- Basin have usually been classified in different families, conch characters can reveal the size of the eggs and the such as PatellidaeRafmesque, 1815, Acmaeidae Carpenter, mode of embryonic development, e.g., the presence of a 1857, or Tecturidae Gray, 1847. The small-shelled repre- free swimming larva. sentatives ofthis group occur regularly but not abundantly For the first time a comparative study is presented between in various shallow to deeper marine and brackish water the minute Patella-like Patellacompressiuscala Karsten, habitats and were studied by many authors (Eichwald, 1849 that lived in the North Sea Basin from Chattian until 1830-1853; Karsten, 1849; von Koenen, 1882; Sinzov,

Langhian time (Late Oligocene - Middle Miocene), and 1892; Friedberg, 1928); Kolesnikov, 1935; Jekelius, 1944; similar species known from the later Badenianand Sarma- Anderson, 1959;Baluk, 1975; Janssen, 1984; Il’ina, 1993; tian (Middle Miocene) from the Paratethys. The former Anistratenko, 2000, 2001; Harzhauser and Kowalke, exhibits unusual of which indicates an type protoconch, a 2002). of This of lecithotrophic type early development. type pro- Correct taxonomic identificationofthese species is some- toconch has been discovered before only in several species times difficult because the restricted number of concho- ofSarmatian lottiids from the Paratethys (see Anistratenko characters available and insufficient of logical knowledge also and discuss & Anistratenko, 2007). We present new the range of intraspecific morphological variation. Fur- data on the shell structure ofthe studied species. thermore, few data on protoconch morphology and type of For the Oligocene/Miocene species the new genus name early development ofthese gastropods have beenobtained Boreoblinia is proposed. Although the generic level of

so far {e.g., Anistratenko et al., 2006; Anistratenko & taxonomic attributionofthe patellogastropods investigated Anistratenko, 2007). here requires further confirmation we assign them to the Protoconch ornamentation and shape are now generally family Lottiidae Gray, 1840 (for more detailed discussion useful taxonomic in accepted as characters, particularly see below under“”). - no-

Figure I.Stratigraphic correlation chartof the standard scale (MediterraneanStages) with local stagesof the North Sea Basin, the Cent- ral and the Eastern Paratethys (Mediterranean Stages after Gradstein etal, 2004; CentralParatethys regional Stages afterHarzhauser

& time in Mandic, 2008). The ranges ofPatellogastropods with a differenttype ofprotoconch shown vertical lines.

— Figure 2. Localities, where studied materials were collected. 1 Miste, near Winterswijk, the Netherlands (local stage Hemmoorian,

lower part ofthe Langhian); 2 - Pinnow, in the neighborhoodof Schwerin, Germany (“StembergerGestein”, Neochattian); 3 - Heist-

op-den-Berg, located between Antwerp and Aarschot, Belgium (Zonderschot Sands, late Hemmoorian,early Langhian). - Ill -

Material and methods Nomenclatural notes and taxonomy

Our study is based on material from three localities in the Originally the species discussed was described as Patella

North Sea Basin (Figures 1,2): compressiuscala and subsequently has been attributed to

1. Miste near Winterswijk (Gelderland province, the differentgenera: Acmaea Eschscholz in Rathke, 1833(e.g.

Netherlands). It concerns a Middle Miocene fauna (local Kautsky, 1925), Scurria Gray, 1840{e.g. Anderson, 1959),

stage Hemmoorian, lower part ofthe Langhian) from the Lepetella Verrill, 1880 {e.g., Janssen, 1984). According to

Breda Formation that is described by Janssen (1984). At Waren(1972) the species belongs to the Cocculinacea and

this lot of have been made for the Acmaeidae. locality a pits especially not to In our opinion Patellacompressius- fossil Six collecting. specimens came from the pit, made in calaKarsten, 1849 should be assigned to the family Lotti-

the other six from several between idae it is 1968; come pits, dug Gray, 1840 as characterized by the sametype and 1968 and 1977. All provided by A.W. Burger. morphology of protoconch as Blinia Anistratenko, Bandel

2. Six specimens ofPatellacompressiuscala from theNeo- & Anistratenko, 2006 and Flexitectura Anistratenko, 2000 chattian “Sternberger Gestein” kindly provided by mr (more details see below).

A.F.J. Jansen could be studied, collected near Pinnow in Since the publication ofKarsten (1849) the incorrectspell-

the neighborhood of Schwerin (Schleswig-Holstein, Ger- ing “compress iuscula” has been generally used {e.g. Kaut-

many). sky, 1925; Anderson, 1959; Janssen, 1984; Wienrich,

3. Also A.F.J. Jansen 18 “ provided by mr were specimens 2001), although the correct original spelling is com- from a” Heist-op-den-Berg (Brabant Province, Belgium), pressiuscal (Karsten, 1849: 12), and there is no nomen-

where also several have been for fossil clatural it. in this risk pits dug collecting. reason to change Moreover, case the

The material originates from the Zonderschot Sands, also of homonymy with Pileopis compressiuscula Eichwald, of late Hemmoorian(early Langhian) age. 1830 (though it is attributed to another genus - Tectura) is

Adult shell characters were studied with an optical stereo- fortunately removed.

microscope. Standard dimensionsfor shell characters were The distinct similarity ofthe protoconch and teleoconchof

used. Morphological features of protoconchs such as P. compressiuscala with Flexitectura subcostata (Sinzov,

shape, size, sculpture and character of boundary with the 1892) from the Middle SarmatianofUkraine and Moldova teleoconch were examined by means of scanning electron might be considered as evidence for their congenerity or

of microscope (SEM). Altogether thirty-six specimens Pa- even conspecifity. However, there are some strong argu-

tellacompressiuscala were analysed. Four specimens were ments against this attribution.First of all this Middle Sar-

partially broken for SEM examination of the shell micro- matian lottiidwith a pancake-like protoconch is assumed to

“ ” structure. be derived from the Early Sarmatian Tectura and might

Most of the SEM obtained in the Institute of of of endemic mollusc images were represent one a large group genera Geological Sciences National Academy ofSciences ofthe of the Middle Sarmatian Paratethys basin (Kolesnikov,

Ukraine (Kiev, Ukraine). Shells were mounted on stubs, 1935; Paramonova, 1994; Harzhauser & Kowalke, 2004;

with and then documented sputter-coated gold using a lonesi et al., 2005; Anistratenko & Anistratenko, 2007).

JSM-6490 Scanning Electron Microscope. A few addi- Noteworthy, the Early Sarmatian basin was already land-

tional SEM micrographs ofshell structure were performed locked and immigrants from the Mediterranean or the

in the Instituteand of North Basin Geological-Paleontological Museum Sea could not penetrate. The last occurrence

the University of Hamburg (Germany); here the digital ofP. compressiuscala in the North Sea Basin predates the

SEM LEO 1455 VP was used. All figured specimens are first occurrence ofBlinia andFlexitectura about2.5 Ma. If

housed at the Institute of Geological Sciences National P. compressiuscala from the North Sea Basin and F. sub-

of Sciences of the Ukraine would be attributed the Academy (Kiev, Ukraine), costata to a single genus mono- abbreviated IGS of as NANU. phyly the genus Flexitectura would become question-

able. Hence, in spite ofa striking resemblance we treatthe

Morphological terms for the shell description used in text similarities between Patellacompressiuscala and Flexitec-

subcostata and for the (Figure 3): tura as a convergence, we suggest

AP - i.e. distanceof from frontal apex position, apex edge, Oligocene/Miocene lottiidfrom the North Sea Basin a new

- diameter - DP greater of a protoconch, HS height of genus name: Boreoblinia.

shell, LS - length of shell, WS - width of shell.

Shell microstructure characters and its taxonomic im-

plications

Some patellogastropod taxa are characterized by having

both calcitic and aragonitic shell layers and this is inter-

preted as the most primitive gastropod shell structure. Cal-

citic layers in patellogastropod shells include foliatedand

Figure 3. A sketch of studied lottiid shell (schematic); muscle homogeneous structures; aragonitic layers are predomi-

crossed-lamellar. Some families scar on the inner surface indicatedby arrow. See explanation nantly e.g., Patellidae,

for - measurements in the text. possess both foliated and crossed lamellar structures. -112-

4. in Figu re The shell structure of patellogastropods discussed the text. A, B. Boreoblinia compressiuscala (Karsten, 1849). A - Speci-

men (IGSNANU, Miste_l 977-4/2008)fromthe Langhian. Cross-section, showing the dissected aragonitic crossed lamellarstructure

B - (DCL) and the aragonitic porous outer layer ofthe shell. Specimen (IGS NANU, Heist-op-den-Berg_2001-3/2009)from the

Langhian. The same aragonitic crossed lamellar structure as in the specimen of fig. 4A and also the inner prismatic layer(IPL) are

visible. C. Tectura zboroviensis (Friedberg, 1928). Specimen (IGSNANU, 18/2003)from the ChokrakianofYurkino (Crimeapenin-

sula, reveals similar with crossed lamellar and inner D. Ukraine), a composition layer (DCL) prismatic layer (IPL). Blinia sp. Speci-

men (IGS NANU, 34/2004) from the middle Sarmatian ofLetichev, West Ukraine. Section showing the aragonitic crossed lamellar

and of structure (DCL) more rough sets the inner crossed lamellarlayers of the shell, compared withB. compressiuscala. E. Tectura

virginea (Muller, 1776). Recent specimen from the North Sea, Doggerbank, (private collection ofDr. Jens Hartmann)shows devel-

oped inner shell layer as aragonitic basal prismatic layer (BPL), middleshell layercomposed of transversally fractured, aragonitic

crossed lamellar layer (TCL), and outer calcitic layer of inclined sheets (1CA). - 113 -

Figure 5. A-G. Boreoblinia compressiuscala (Karsten, 1849). A - IGS NANU, Miste_1968-4/2008, Langhian; AI - apical view; A2 -

right lateral view; A3 - anterior view; A4 - posterior view; A5 - details ofapical view of the shell. B - IGS NANU, Heist-op-den-

Berg_2001-3/2009, Langhian; B1 - apical view; B2 - detailsofapical view of the embryonic shell. C - IGSNANU, Heist-op-den- - Berg_2001-2/2009,Langhian; Cl - apical view; C2 - detailsof apical view of the embryonic shell.D IGS NANU, Miste_1968-

5/2008, Langhian;apical view of the embryonic shell; the clear demarcationbetween the protoconchand early teleoconch is visible.

E- IGS NANU, Schwerin-1/2008,Chattian; apertural view, the muscle scars are visible as gentle expressive tracts. F - IGS NANU,

Schwerin-2/2008,Chattian; FI - posterior view; F2 - left lateral view; F3 - apical part of the shell viewed from the left. G - IGS

NANU, Schwerin-3/2008, Chattian; G1 - apical view; G2 - right lateral view; G3 - apical partofthe shell viewed from the left. -114-

whereas Lottiidae have shells with thin outer calcitic of the shell is thin as Blinia) and porous, both are ar- underlain homogeneous layers by aragonitic crossed- agonitic. That distinguishes both genera from Patella and lamellar layers (MacClintock, 1967; Lindberg, 1988). its relatives, which have a calcitic outer layer that is usually

The shell structure of Boreoblinia is characterized by a thick and has a complex structure. The shell structure of simple type of aragonitic crossed-lamellar structure, the Boreobliniaand Blinia is similar in general to that of the

“ ” same as is known from the Chokrakian “Tectura” zborovi- Sarmatian Tectura zboroviensis, but the first two differ ensis and Sarmatian Blinia from the from the latter (Friedberg, 1928) as well as from all otherknown patellogas-

Paratethys (Anistratenko et al, 2006). In all these cases tropods (except for Flexitectura and Squamitectura) in most of the shell is of of crossed la- composed one layer having a “pancake’Mike protoconch. The new genus dif- mellae in which the needles ofthe two directions oflamel- fers from Blinia in larger and more globular shape of the lae of the first order commonly intersect (Figure 4). A protoconch, which has no expressed sculpture, wrinkles or similar is known from the Triassic structure patellid Scutel- other specific ornamentationor pit on its surface. lastraea costulata (Munster, 1841) from the St Cassian

Formation ofthe Alps, and from apatellogastropod species Description — The shell is small to moderatein size, thin- of similar shape from the Paleocene of Alabama(Bandel, walled, high-conical in shape without marginal slit, apical

The sets of 1982). crossed lamellae are more delicate in hole or internal septum. It measures up to 4 mm in length,

“ Blinia' shell wall than in Boreoblinia and to s “Tectura” up 2 mm in width and up to 2 mm in height. The apex is zboroviensis. situated eccentric and slightly tilted forward. The lateral

The outer layer of Boreobliniaalso resembles that of the edges of the shell are almost parallel to each other. The

Blinia shell - it is thin and porous, and also aragonitic. teleoconch surface has numerous concentric lines. The

These characters distinguish both Boreoblinaand Blinia protoconch is round to oval “pancake’Mike in shape, quite clearly from Patella and itsrelatives, which have a calcitic bulbous and measures from 0.18 mm to 0.25-0.28 mm in

outer layer with a characteristic layered structure that is maximum diameter. The protoconch surface is apparently

usually quite thick and has a rather complex structure (Fig- smooth, perhaps the ornamentation is not preserved. The ure 4). transition from the embryonic shell to the teleoconch is

the of the Boreoblinia shell is charac- Hence, structure usually clearly marked by a constriction. terized as follows: the outer layer is thin, porous and ar-

most of the shell has an crossed- Discussion — The less agonitic (1), aragonitic more or cap-like or semi globe- lamellar of of structure, composed a single layer coarse shaped embryonic shell of Patella compressiuscala indi- crossed lamellae and thin inner exists (2) a prismatic layer cates a lecithotrophic type of early development and the it is often off, and the inner surface of (3), though peeled absence ofany short free-swimming larval stage following of the adult individuals be this most appears to lacking the yolk-rich embryogenesis (e.g., Bandel (1982)). The specific layer. shape and proportions of a “pancake”-type ofprotoconch

the of may suggest brooding young snails in the maternal adult individuals (as it is in the modernErginus moskalevi

Systematic paleontology (Golikov and Kussakin, 1972) illustratedby Sasaki (1998,

fig. 21a-c). The imprints on the inner surface ofthe shell Class Gastropoda Cuvier, 1797 of Boreoblinia which we interprete as the muscle scars, , Order 1986 Patellogastropoda Lindberg, are horseshoe-like, gentle expressive tracts (Figure 3). The Lottiidae 1840 Family Gray, impressions were SEM-documented as well (Figure 5e).

Genus Boreoblinianov. The combinationof shell features and the structure proto-

conch morphology support that the Oligocene/Miocene

Type species — Patella compressiuscala Karsten, 1849. lottiid from the North Sea Basin discussed here should be

Late Oligocene (Chattian) - Middle-Miocene(Langhian) considered as belonging to an independent genus. ofthe Netherlands, Belgium, NW Germany and Denmark.

“ ”

the — Derived from Etymology of name Borealis (Latin, Boreobliniacompressiuscala (Karsten, 1849)

northern) and “Bliri” (Russian, pancake) for the general (Figure 5) shape ofthe protoconch that is like a thick pancake.

1849 Patella compressiuscala n. sp?; Karsten, p. 12.

— small Diagnosis Conical, relatively patellogastropods 1868 Patella compressiuscala Karsten; Koch & Wiech-

12. with high-conical, laterally compressedshell and “pan- mann, p. 562, pi. 12, fig.

1876 Patella cake’Mike protoconch. Teleoconch ornament ofconcentric compressiuscula Karsten; Koch, p. 165.

1882 Patella Karsten; von ribs; compressiuscula Koenen, p. growth aperture elongated in shape, anteriorand pos- 323. terior edges have distinct flexures. 1925 Acmaea 55. compressiuscula Karsten; Kautsky, p.

1959 Scurria compressiuscula (Karsten, 1849) [non Eich- Differentiation — Differs from SarmatianBliniawith simi- wald]; Anderson, p. 45-46, pi. 2, fig. 2a, b. lar type of protoconch mainly in having larger protoconch 1984 Lepetella compressiuscula (Karsten, 1849); Janssen, and more rough sets ofthe inner crossed lamellar of 2a-c. layers p. 121, pi. 44, fig. the shell, whereas the organization of these layers in both 2001 Lepetella compressiuscula (Karsten, 1849); Wien-

2a-d. taxa is similar. The outer layer ofthe Boreoblinia(as well rich, p. 394-395, pi. 63, fig. -115-

Diagnosis — See diagnosis for genus. ally a little bit smaller than those of specimens from all

other localities (see Table 1).

Material — Twelve from the of the shell few notes shouldbe specimens Langhian Concerning variability a pre-

Miste, Holland, IGS NANU, MisteJ 968-1/2008-6/2008 sented here. We examined more than fifteen specimens and

coll. IGS (ex A.W. Burger); NANU, Miste_1977-l/2008- the limits ofdimensions’ variability and shape ofshell are

6/2008 (ex coll. A.W. Burger); six specimens from the restricted. The protoconch in Bliniacan be with or without

Chattian of Schwerin- small Schwerin, Germany, IGS NANU, pit (a depression in the centralpart ofthe protoconch

1/2008-6/2008(ex coll. A.F.J. Jansen); eighteen specimens surface: Anistratenko & Anistratenko, 2007) whereas the

from the Langhian of Heist-op-den-Berg, Belgium, IGS protoconch in all studied Boreoblinia specimens has no

NANU, Heist-op-den-Berg_2001-1/2009-18/2009 (ex coll. expressed sculpture, wrinkles or other specific ornamenta-

A.F.J. Jansen). tion or pit on its surface.

The specimens from the studiedmaterial generally corres-

Description — Shell small, high-conical, with lateraledges pond well to the original description of Patella

The is tilted forward. subparallel. apex eccentric, slightly compressiuscala (Karsten 1849). Unfortunately, the The from front apical angle (measured the view) varies, original description contains no illustration, whereas the usually about 55-65° in different specimens. The anterior shape, proportions and ornament of specimens of this and the described and illustrated posterior slopes are straight or slightly convex; species, by Janssen (1984,121, pi.

“ ” anterior and posterior edges with distinct flexures. Aper- 44, fig. 2a-2c), correspond fully to the compressiuscala ture oval. The muscle scars are horseshoe-like and are specimens studied here. Due to corrosion, the protoconch- usually clearly visible as gentle expressive tracts (Figures teleoconchtransitionis sometimes not clearly visible even

3, 5). The protoconch is “pancake”-like to semi globe- in SEM images of adult specimens. Morphologically,

shaped, apparently smooth. The teleoconchsurface is co- specimens from the Chattian “Sternberger Gestein” are vered by coarse concentric growth lines alternating with almost indistinguishable from the Langhian specimens and

fine transition from the shell ones. The embryonic to the they shouldbe consideredas a single species with a strati- teleoconch is usually marked by a constriction or a rim. graphic rangeofat least25-15 Ma. There are other species

known which have been living as long as this species -

— See Table modemtaxa in the Mediterranean Atlantic Measurements 1. many and per-

sist from the Late Mioceneor even Oligocene ( Coccu- e.g.,

No LS HS ws AP DP lina rathbuni Dali, 1882; Epitonium clathrus (Linnaeus,

1758), Bittium reticulatum (Da Costa, 1778), Cerithium

vulgatum Bruguiere, 1792,Alvania (Payraudeau, IGS NANU, Schwerin montagui 1/2008 2.2 1.2 1.5 1.2 -0.18 1826)). 2/2008 2.55 1.2 1.6 1.3 0.23-0.25*

3/2008 2.25 1.0 -1.6 1.3 0.18-0.19*

and — This 4/2008 2.75 1.5 1.4 1.6 -0.15 Stratigraphic geographic range species has

5/2008 2.9 1.7 1.76 1.5 0.20 been recorded in fully-marine deposits of the North Sea

6/2008 2.7 1.45 1.7 1.25 - Basin from Late Oligocene (Chattian) untilEarly Serraval-

IGS Miste NANU, lian, where it occurs mostly in low numbers (e.g., Karsten, 1968-1/2008 1.9 1.15 1.1 0.9 -0.16 1849; Janssen, 1984). 1968-2/2008 2.5 1.45 1.4 -1.0 0.19

1968-4/2008 2.3 1.45 1.45 0.95 0.18-0.20*

1968-5/2008 2.35 1.5 1.4 0.95 0.24-0.25*

IGS NANU, Miste Palaeoecological implications

1977-1/2008 2.75 1,85 1.65 1.2 0.28

1977-2/2008 - - - - -0.18 As it is suggested, the modifications in protoconch mor- 1977-3/2008 - - - 0.23-0.27*

phology ofthe Badenian/Sarmatianlottiidsfrom a plankto- IGS NANU, Heist-op-den-Berg trophic to a lecithotrophic one, could have been triggered 2001-1/2009 1.7 0.8 1.2 0.8 0.13

2001-2/2009 2.6 1.4 1.3 1.1 0.18 by decreasing salinities of the water (Anistratenko et al.,

2001-3/2009 3.9 1.8 1.9 1.7 0.16 2006; Anistratenko& Anistratenko, 2007). What was a key 2001-4/2009 18 1.4 1.75 0.8 0.12 for the in in reason change ontogenetic strategy occurring

the Late-Oligocene in fully marine circumstances in the

North Sea Basin? Table 1. Measurements of Boreoblinia compressiuscala. All It is that in the measurements in mm. The asterisk denotes two measure- possible, already Early-Oligocene predesse-

of Boreobliniaexisted with the of ments (biggest and smallest) of a protoconch. sors same type proto-

conch. However that only puts the problem furtherback in

time. One of the possible reasons could be, that properties

Remarks — Specimens of Boreoblinia compressiuscala of bottom waters changed during the Oligocene, caused by both from the Chattian and the Langhian differ from the changes in the palaeogeography, also influenced by the

MiddleSarmatian Flexitectura subcostata in its and of Was it larger development a stronger seasonality. really an

inflated more globular not protoconch. The Langhian B. independent development in the North Sea Basin, that was differs from compressiuscala the Chattian specimens in a repeated during the Sarmatian ofthe Paratethys, or is there

more eccentricic position ofthe apex. The protoconchs of a possibility, that these groups could have a common de-

B. fromthe compressiuscala Heist-op-den-Berg are gener- velopment? -116-

The between the last time-gap occurrence ofBoreoblinia new genus ofpatellogastropod with unusualprotoconch from Miocene of Acta Polonica compressiuscala and the first occurrence ofpatellogastro- Paratethys. Palaeontologica 51 (1), 155-164. pods with a pancake-like protoconch in the Paratethys Baldi, T. 1986. Mid-Tertiary and paleogeographic (Blinia Flexitectura and there is stratigraphy , Squamitectura), suggest 201 evolutionofHungary. AkademiaiKiado, Budapest, pp. no directrelation between both developments. But it seems Baluk, W. 1975. Lower Tortonian gastropods from Korytnica, that there existed possible a, perhaps short-lived, connec- Poland. Part I. Palaeontologia Polonica 32, 1-186. tion between the North Sea Basin and the Paratethys dur- Bandel, K. 1982. Morphologic und Bildung der friihonto- the Middle-Miocene the southern bor- ing highstand along genetischen Gehausc bei conchiferen Mollusken.Facies 7,1-

der ofPoland, ofwhich the deposits have beenremoved by 198.

and erosion. that there is K. 1991. from the subsequent uplift In case, a pos- Bandel, Gastropods brackish and fresh waterof Jurassic-Cretaceous sible closer relationship between Blinia, Flexitectura and transition (a systematic reevaluation). Berliner ReiheA 9- Boreoblinia, and the time-gap becomes artificial. Unfortu- GeowissenschaflenAbhandlungen, 134, 55. nately no paleogeographic evidence is known for such a E. d' 1830. Naturhistorische Skizze Eichwald, von Litthauen, connectionyet. On the other hand it could be hypothesized Volhynien und Podolien in geognostischer, mineralogischer, that a faunalinterchange betweenmentionedbasins existed botanischer und zoologischer Hinsicht. Wilna, 1-256. earlier, e.g. during the Late-Oligocene (Chattian) since the Eichwald, E. d’ 1853. Lethaea Rossica ou Paleontologie de la main connectionofthe with the sea was in Paratethys open Russie. III. Derniereperiode. 1-518 Stuttgart (Schweizerbart). the towards the North Basin west, Sea (e.g. Baldi, 1986; Friedberg, W. 1911-1928. miocehskie ziem Polskich.

Popov et al, 2004). In this case forerunners of the Ba- Czesc I. Slimaki i todkonogi. (MolluscamiocaenicaPoloniae.

“ ” denian Tectura zboroviensis and Sarmatian Blinia Pars I. Gastropoda et Scaphopoda). 1-631.Lwow-Poznah, [in

Polish], probably branched off from a common ancestor of Gradstein, F.M., Ogg, J.G. & Smith, A.G. 2004. A geologic time Boreoblinia and immigrated into the Paratethyan Basin. scale. I-XIX+1-589, Cambridge (Cambridge University And the resemblance in structure of the shell in

“ ” Press). Boreohlinia, Blinia and Tectura zboroviensis seems to Harzhauser, M. & Kowalke, T. 2002. Sarmatian (Late Middle this idea. support Miocene) Gastropod Assemblages of the CentralParatethys. Facies, 46:57-82; pi. 9-13.

Harzhauser, M. & Mandic, O. 2008. Neogene lake systems of Acknowledgements Central and South-Eastern Europe: Faunal diversity, gradients and interrelations. Palaeogeography, Palaeoclimatology,

Palaeoecology 260, 417-434. We express our sincere thanks to Mr A.F.J. Jansen, who Il'ina,L.B. 1993. Handbook for identification ofthe marine Mid- kindly provided six specimens of Patellacompressiuscala dle Miocene gastropods of Southwestern Eurasia. Trudy from the Neochattian of "Sternberger Gestein" (Pinnow, Paleontologiceskogo Instituta AkademiiNauk SSSR 255. 1- Germany) and 18 specimens from the late Hemmoorianof 151 \in Russian], Zonderschot Sands (Heist-op-den-Berg, Belgium). Olga Janssen, A.W, 1984. Mollusken uit het Mioceenvan Winterswijk and Vitaliy Anistratenko were partially the supported by - Miste. KNNV36/NGV/RGM: 1-451; 82 pis.

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