Paleoecology, stratigraphy and taxonomy of the pliocene marine diatoms from central crete (Greece)

R E V U E DE MICROPAL]~ONTOLOGIE

VoI. 42, n ° 4, d 6 c e m b r e 1999, pp. 269-300

PALEOECOLOGY, STRATIGRAPHY AND TAXONOMY OF THE PLIOCENE MARINE DIATOMS FROM CENTRAL CRETE (GREECE) PALEOECOLOGIE, STRATIGRAPHIE ET TAXONOMIE DES DIATOMEES MARINES DU PLIOCENE

DE

CR TE CENTRALE CRECE by

Dimitris

FRYDAS*

ABSTRACT. -- A rich siliceous p h y t o p l a n k t o n assemblage of late E a r l y Pllocene to e a r l y Late Pliocene age, c o n t a i n i n g 31 g e n e r a of m a r i n e diatoms w i t h 55 specific t a x a , was f o u n d in fine l a m i n a t e d diatomite beds in the sections of K a l l i t h e a , M a r a t h i t i s , P r a s s a s and A i t a n i a from the n o r t h e r n c e n t r a l p a r t of the Crete I s l a n d ( P r o v i n c e of H e r a k l i o n , Greece). The sediments of the studied sections belong to the u p p e r p a r t of the S t a v r o m e n o s facies-unit ( E a r l y to Late Pliocene). The d i a t o m associations were m a i n l y i n v e s t i g a t e d for their paleoecology, s t r a t i g r a p h y and taxonomy. The d i a t o m assemblages are c h a r a c t e r i z e d b y species t y p i c a l of low l a t i t u d e s . They belong to the u p p e r p a r t of the Nitzschia jouseae biozone ( E a r l y Pliocene - Late Pliocene) because of the p r e s e n c e of the m a r k e r species N. jouseae, w h i c h is r a r e to f r e q u e n t in the K a l l i t h e a a n d the M a r a t h i t i s sections, and r a r e in the P r a s s a s a n d Aitania sections. The u p p e r p a r t of the N. jonseae biozone c o r r e s p o n d s to the C N l l b (Discoaster asymmetricus) a n d CN12a (D. tamalis) subzones of c a l c a r e o u s nannofossils as well as to the Globorotalia puncticulata a n d G1. bononiensis biozones of p l a n k t i c f o r a m i n i f e r a . Among the diatoms the p l a n k t i c , centric species d o m i n a t e in t e r m s of t h e i r p e r c e n t a g e d i s t r i b u t i o n a n d reflect a succession of two paleoclimate i n t e r v a l s : a s u b t r o p i c a l i n t e r v a l with the w a r m - w a t e r species Azpeitia noduliJ~r, Hemidiscus cuneiJbrmis a n d Thalassiosira leptopus a n d a t e m p e r a t e i n t e r v a l c h a r a c t e r i z e d b y the c o l d - w a t e r species Coscinodiscus marginatus. These two i n t e r v a l s are p r o p o s e d as two new local acme subzones. R~SUME. -- Les diatomites f i u e m e n t lamin6es de la p a r t i e s e p t e n t r i o n a l e de la P r o v i n c e d ' H @ a k l i o n (Cr~te, Grace) ont 6t6 6tudi6es dans les coupes de K a l l i t h e a , de M a r a t h i t i s , de P r a s s a s et d ' A i t a n i a , qui a p p a r t i e n n e n t ~t la p a t t i e s u p 6 r i e u r e de l'unit6 de facies de Stavromenos (Plioc~ne i n f @ i e u r ~ Plioc~ne s u p @ i e u r ) . Elles ont f o u r n i une fiche association de p h y t o p l a n c t o n siliceux c o n t e n a n t 31 genres et 55 esp~ces de Diatom6es m a r i n e s d n Plioc~ne i n f 6 r i e u r t e r m i n a l a u Plioc~ne s u p @ i e u r basal. L a pal6o6cologie, la r 6 p a r t i t i o n s t r a t i g r a p h i q u e et la taxonomie des Diatom6es ont 6t6 analys6es en d6tail. Les Diatom6es examin6es sont des esp~ces c a r a c t 6 r i s t i q u e s des basses latitudes. Elles a p p a r t i e n n e n t ~ la p a r t i e s u p 6 r i e u r e de la biozone a Nitzschia jouseae (Plioc~ne i n f 6 r i e u r - Plioc~ne s u p @ i e u r ) d ' a p r ~ s l'esp~ce guide N. jouseae qui est r a r e ~ f r 6 q u e n t e d a n s les coupes de K a l l i t h e a et de M a r a t h i t i s et r a r e d a n s les coupes de P r a s s a s et d ' A i t a n i a . La p a r t i e s u p 6 r i e u r e de la biozone ~ N. jouseae c o r r e s p o n d a u x sous-zones C N l l b (Discoaster asymmetricns) et CN12a (D. tamalis) d u n a n n o p l a n c t o n calcaire et aux biozones ~ Globorotalia puncticulata et GI. bononiensis des F o r a m i n i f ~ r e s p l a n c t o n i q u e s . P a r m i les Diatom~es, les esp~ces c e n t r i q u e s , p l a n c t o n i q u e s , d o m i n e n t en p o u r c e n t a g e ; elles refl6tent la succession de d e u x intervalles pal6oclimatiques : l ' u n subtropical, avec les esp~ces d ' e a u x c h a u d e s Azpeitia nodulifer, Hemidiscus cuneiformis et Thalassiosira leptopus, et l ' a u t r e temp6r6, caract@is6 p a r l'espbce d ' e a u x froides Coscinodiscus marginatus. Nous p r o p o s o n s de c o n s i d @ e r ces d e u x intervalles comme d e u x nouvelles sous-zones d ' a c m 6 , ~ v a l e u r au moins locale.

-

K e y - w o r d s : M a r i n e diatoms Diatoufite - Neogene - Pliocene - M e d i t e r r a n e a n a r e a - Crete i s l a n d - Paleoecology - S t r a t i g r a p h y T a x o n o m y - New local acme subzones - Q u a n t i t a t i v e d a t a - P a l e o c l i m a t i c i n t e r v a l s .

Mots-el6s : Diatom6es m a r i n e s - Diatomites - N~og~ne - Plioc~ne - M 6 d i t e r r a n 6 e - Cr~te - Pal6o6cologie - S t r a t i g r a p h i e - T a x o n o m i e - Nouvelles sous-Zones d ' a c m 6 locales - Donn6es q u a n t i t a t i v e s - l n t e r v a l l e s pal6oclimatiques.

* U n i v e r s i t y of P a t r a s , D e p a r t m e n t of Geology, Division of M a r i n e Geology a n d G e o d y n a m i c , GR-26500 P a t r a s - R i o n , Greece.

270

DIATOMI~ES PLIOCI~NES DE CRI~TE

C R E T E

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FIG. 1. - L o c a t i o n of the studied sections on C e n t r a l Crete. 1 : K a l l i t h e a , 2 : M a r a t h i t i s , 3 : P r a s s a s , 4 : Aitania. The localities of C h a m a l e v r i (5) a n d A s t e r i (6) h a v e b e e n p r e v i o u s l y investigated. Localisation des coupes dtudides en Cr~te centrale. I : K a l l i t h e a , 2 : Marathitis, 3 : Prassas, 4 : Aitania. Les localitds de C h a m a l e v r i (5) et d'Asteri (6) oat f a i t l'objet de p u b l i c a t i o n s antdrieures.

INTRODUCTION

After the deposition of predominantly terrigenousclastic sediments in several grabens in Early-Middle Tortonian time, the Cretan area was transformed into a relatively stable platform with numerous islands and shoals, separated by shallow seas (Figs. 1-2). This paleogeographic configuration came into existence during the Middle Tortonian and lasted until the end of the 'Middle' Pliocene (Tsapralis, 1976). During the early Late Pliocene, predominantly calcareous muds were deposited in shallow, often fault-bounded basins, that adjoined relative highs with a thin bioclastic limestone cover and/or fringing reefs. Especially in the Prefectures of Rethymnon and Heraklion these paleogeographic configurations can be constructed in great detail. The calcareous muds accumulating in the graben-like depressions are composed of alternations of finely-laminated and homogenous marls, which show large differences in their sedimentary and macrofaunistic features. On the island of Crete, two sedimentation cycles were distinguished, which range corresponds to the Late Miocene and Pliocene time respectively (Christodonlou, 1963). The Miocene and Pliocene macrofossils of central Crete have been investigated by Papapetrou-Zamani (1965) and Symeonidis and Konstandinidis (1970). Deposits of more than 500 m in thickness comprise the above two cycles within the Neogene basins. During the Late Miocene Mediter-

ranean "salinity crisis", rythmic alternations of thick evaporites, sandstones and diatomites were deposited (Frydas 1985, 1987, 1993, 1994b). In turn, the Pliocene succession mainly consists of grey silty clays and yellow marls, which alternate with light grey to white diatomites or diatomaceous marls. The silty clay layers and the fine-laminated diatomites represent the paleoclimatic changes, or the periods of upwelling due to upward movement of cold water currents. From a total of nine formations which have been distinguished in the Crete Island, three were deposited during the Late Miocene (Kissamou Formation, Cherethiana F. and Aghia Varvara F.), and the rest (Tavronitis Formation, Asteri F., Aghios Vlassios F., Finikia F., Stavromenos F. and Faneromeni F.) were deposited during the Early and Late Pliocene (Freudenthal, 1969; Meulenkamp et al., 1979; Frydas, 1990, 1996; Frydas and Keupp, 1992).

THE KALLITHEA, MARATHITIS, PRASSAS AND AITANIA SECTIONS

The Neogene sequence in the Heraklion district is synsedimentary faulted, especially in the northern part (Jonkers, 1984). Two orthogonal systems of NNW-ENE and NNE-WNW trending faults influenced and controlled the paleogeographical evolution of four small, individual basins, which were generally

FRYDAS

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Miocene

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Messiniangypsum

271

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Pliocene

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FIG. 2. - Simplified geological m a p of the n o r t h e r n p a r t of the H e r a l d i o n p r o v i n c e ( J o n k e r s , 1984). I. Finikia-Aghios Vlassios Basin, II. Kallithea Basin, I1~. Prassas Basin, IV. Aitmfia Basin. Investigated sections in this paper. 1 : Kallithea, 2 : Marathitis, 3 : Prassas and 4 : Aitania.

Carte gdologique simplifide de la partie septentrionale du Nome d'Hdraklion (Jonkers, 1984). Bassins sddimentaires : I. Finikiu-Aghios Vlassios, II. KaUithea, Ill. Prassas et IV. Aitania. Coupes examindes clans ce travail. 1 : Kallithea, 2 : Marathitis, 3 : Prassas et 4 : Aitania.

separated by the presence of intrabasinal basement highs (Fig. 2). These four adjacent basins used in our study are Finikia-Aghios Vlassios, Kallithea, Prassas and Aitania Basins. The Neogene sediments of all these basins, which transgressively overlie the Tripoli limestones, mainly consist of homogeneous marls, grey clays with brownish interbeds, fossiliferous yellowish marls and diatomites, with a total thickness of more than 100 m. The present paper concerns the investigation of diatom assemblages for establishing useful local diatom zones, which can be correlated with the zones of silicoflagellates and calcareous nannoplankton (Frydas, 1990, 1996; Driever, 1988). The sediments of the Kallithea, Marathitis, Prassas and Aitania sections (Figs. 3-6) belong to the upper p a r t of the Stavromenos unit (late Early Pliocene to early Late Pliocene), which are found all over the Heraklion province (Jonkers, 1984). The Marathitis and Prassas sections are considered in this p a p e r as representative sections because of their great abundance in both genera and taxa of marine diatoms (Tab. 1).

The investigated sections represent an alternation of yellowish-grey homogeneous marls, sapropel and white-greyish laminated diatomite beds, which generally appear in the u p p e r p a r t of the sections (Fig. 7). A rich diatomaceous assemblage has been observed in the diatomaceous layers from these sections. Submarine andesitic eruptions from the island volcanic arc of the south Aegean Sea probably furnished the silica to coastal waters where it could be used for the skeleton building of the diatoms. The genesis of siliceous sediments in the neighbouring of active island arcs is explained in detail in Siever (1983, p. 15; fig. 3). The planktonic foraminifera from Aghios Vlassios section near that of Marathitis, have been studied by Spaak (1983, fig. 18). He has recognized the Pliocene guide species Globorotalia margaritae and G. puncticulata. The benthic foraminifera and the calcareous nannoplankton from the section Aghios Vlassios have been studied by Jonkers (1984, fig. 45) and Driever (1988, fig. 55), respectively. The diatomite beds in the u p p e r p a r t of both Aghios Vlassios and Marathitis sections were investigated for silico-

272

DIATOM]~ES PLIOCI~NESDE CRI~TE

FIG. 3. - Locatiot~ of the K a l h t h c a section.

FIG. 5.

- -

L o c a t i o n of the P r a s s a s section.

Localisation de la coupe de Kalithea.

Localisation de la coupe de Prassas.

FIG. 4. - L o c a t i o n of the M a r a t h i t i s section.

FIG. 6. - L o c a t i o n of the A i t a n i a section.

Localisation de la coupe de Marathitis.

Localisation de la coupe de Aitania.

FRYDAS

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FIG. 7. Litbostratigraphy, diatom, planktonic foraminifera1 and calcareous nannofossil biostratigraphy, and position of the samples in the investigated sections on Central Crete. - 1 - indicates the correlation level corresponding to the increase in the Coscinodiscus marginatus and Actinocyclus curvatulus abundances. Lithostratigraphie, biostratigraphie d'apr$s les diatom6es, foraminifdres planctoniques et nannofossiles calcaires, et position des 6chautillons darts les coupes ~tudi6es en Cr$te centrale. 1 indique le niveau de correlation correspondant & l'augmentation d'abondance de Coscinodiscus marginatus et d'Actinocyclus curvatulus. -

flagellates as well as f o r c a l c a r e o u s n a n n o f o s s i l s f o r stratigraphical and paleoecological purposes by Frydas (1990, fig. 7, 1996, fig. 2). T h e y b e l o n g to t h e CNllb ( D i s c o a s t e r a s y m m e t r i c u s ) to C N 1 2 a (D.

t a m a l i s ) s u b z o n e s of c a l c a r e o u s n a n n o f o s s i l s as well as t h e D i c t y o c h a b r e v i s p i n a b r e v i s p i n a a n d D. cf. n e o n a u t i c a a c m e - s u b z o n e s of silicoflagellates (Late Pliocene).

274

DIATOMI~ES PLIOCI~NES DE CRI~TE

The lower part of the Prassas section is placed in the Globorotalia pancticulata (Zanclean) zone while the upper part belongs to the G. bononiensis (Piacenzian) zone (Spaak, 1983, fig. 18). The macrofossil assemblage, with the Terebratula group, examined by Georgiades-Dikeoulia (1979), correlated with the microfossil zones, and can be considered as a 'Middle' Pliocene one. The fish fauna mainly consists of numerous skeletons of Bregmaceros albyi (SAUVAGE) and some of Diaphus sp. characterizing the upper mesopelagical part of a subtropical to temperate marine environment (Miiller and Strauch 1994). The silicoflagellates belong to the Cannopilus major and Distephanus boliviensis boliviensis acme subzones (Frydas 1990, 1996).

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MATERIAL AND METHODS

Five to seven distinctly separated diatomaceous horizons have been identified in the upper part of the investigated sections with a thickness of 2.0 and 12.0 m. From each horizon, representative samples were examined containing well preserved diatom valves. For quantitative and qualitative diatom study microscope slides with randomly distributed microfossils were used. The cleaning of the sediment samples, preparation of permanent mounts for light microscopy and SEM investigations were carried out by following standard procedures : In a 250 ml beaker, 10 % HC1 was added to 1 c m 3 o f diatomite until the carbonate reaction ceased ; then it was heated in 10 % H 2 0 2 and washed repeatedly in distilled water. Strewn slides of cleaned material were prepared using a Hyrax mounting medium and cover glaces of 50 × 24 mm dimension. The diatoms were then identified at magnifications of approximately 1200 ×. From each slide 500 undamaged, complete diatom valves were counted.

PALEOECOLOGY

The dominance of planktic diatom species, that represent between 72.0 % and 92.5 % of the investigated assemblages, is given in Fig. 9. Therefore full marine conditions in a polyhalobian, meioeuryhaline environment (Fig. 8) must have dominated in Late Zanelean-Early Piacenzian time in the Crete area. The percentage variation of the diatom species reflects the succession of two paleoelimatie intervals :

FIG. 8,

Salinity tolerance diatom distribution after Pankow (1976). Distribution des diatomdes selon lear tolerance h la salinitd, d'apr~s Pankow (1976). -

-

a subtropical interval and a temperate one.The first one, corresponding to the diatoms of the Kallithea and Marathitis sections, is indicative of warm-water conditions as revealed by the common presence of the warm-water species Azpeitia nodulifer, Nitzschia

marina, Thalassionema nitzschioides, Thalassiosira leptopas and Triceratium balearicum f. biquadrata (Tabl. 1-2). Th6 second interval, indicative of temperate prevailing conditions, corresponds to the diatom assemblages of Prassas and Aitania sections and is characterized by the common occurrence of the cold-water species Actinocyclus curvatulus and Coscinodiscus marginatus (Tabl. 1-3). Warm-water species A. nodulifer, N. marina, T. nitzschioides and T. leptopus are totally missing from the second interval. In addition, similar paleoclimatic changes were found by Frydas (1990, fig. 7; 1996, fig. 2) through the study of the silicotlagellate percentage variation of both Marathitis and Aitania sections. The dominance of Dictyocha brevispina brevispina and D. cf. neonautica in the Marathitis section indicates a warm-water environment (Frydas, 1990), in contrast to the temperate paleoclimate interval revealed by the dominance of Cannopilus major and Distephanus boliviensis boliviensis in the Prassas section (Frydas, 1990). The succession of the silicoflagellate assemblages from subtropical to temperate climates, indicates a climatic instability during the early Late Pliocene in the southeastern Mediterranean Sea (Driever, 1988).

FRYDAS

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FIG. 9. - P e r c e n t a g e d i s t r i b u t i o n of p l a n k t i c (p), m e r o p l a n k t i c (m) a n d b e n t h i c (b) d i a t o m s as well as of the z o n a l m a r k e r the w a r m - w a t e r Thalassiosira leptopus a n d t h e c o l d - w a t e r Coscinodiscus marginatus species.

Nitzschiajouseae,

Distribution des pourcentages de Diatom~esplanctoniques (p), m6roplanctoniques (m) et benthiques (b) ainsi que des esp~ces Nitzschia jouseae (marqueur de biozone), Thalassiosira leptopus (forme d'eau chaude) et Coscinodiscus marginatus (forme d'eau froide).

According to Stradner and Bachmann (1978) low temperatures may have been dominated in the Late Pliocene (NN16) Aegean Sea, with warm intervals inbetween (paleotemperatures derivated from Dictyocha Distephanus ratio in the Aegean Basin, DSDP Leg 42). A warm chmate period must have been dominated in the Mediterranean Sea between 3.42 and 3.18 Ma, whereas a temperate one, accompanied by important seasonal fluctuations, may characterize the time-interval between 3.18 to 2.87 Ma (Driever, 1988). According to Zachariasse et al. (1989), long term changes in surface water temperatures include a warming between 3.38 and 3.18 Ma, followed by a cooling from 3.18 to 3.03 Ma. The warming of Mediterranean surface waters at 3.38 Ma is believed

to be associated with the final closure of the Isthmus of Panama. According to Thunell et al. (1990) the oxygen isotope results for Site 653-0DP (Tyrrhenian Sea) indicate high variable conditions during the late Early Pliocene (Zone MPL3). The interval from 4.7 to 3.5 Ma shows a periodic fluctuation in the 8180 record. This variability is more related to surface water temperature fluctuations of nearly 5 °C, resulting in salinity changes, and not to ice volume changes, because in contrast, most open-ocean studies have considered the Early Pliocene to be a time of very stable climate. For the early Late Pliocene (Zone MPL4) the 8180 record is marked by the stratigraphical distribution of the Globorotalia inflata group. The first occurrence of Globorotalia bo-

276

DIATOMI~ES PLIOCENES DE CRETE

noniensis and G. inflata are associated with coolings while the last occurrences of Globorotalia puncticulata and G. bononiensis coincide with warmings (Zachariasse and Spaak, 1983). Both paleoclimatic intervals in this study are in good agreement with Driever's paleotemperature curve (1988, fig. 55) during Late Zanclean-Early Piacenzian time. On one hand we have the dominance of Thalassiosira leptopus, a warm water species, which is common to abundant in both the Kallithea and Marathitis sections (Tab. 1-2). On the other hand the last occurrence of G. puncticalata in the upper part of the section Marathitis (Fig. 8) during Late Zanclean time coincides also with a warm period (Thunell et al., 1990). The cold water diatom species Coscinodiscus marginatus, which is frequent to common in the Prassas and Aitania sections, in connection with the first appearance of G. bononiensis in these sections, indicates a temperate climate period.

The presence of N. jouseae indicates assignment to the N. jouseae zone or subzone A of the Rhizosolenia praebergonii zone of Burckle (1972). Recently, Baldauf and Iwai (1995)have proposed to replace, where needed, Rhizosolenia praebergonii zone with the Nitzschia marina zone, and have extended the Nitzschia jouseae zone to include the total range of N. jouseae. R. praebergonii is rare outside of the equatorial Pacific, and Baldauf and Iwai (1995) favor the use of a N. jottseae zone based on the total range of N. jouseae (J. Barton, pers. comm., June 30, 1998). The N. jouseae biozone in this work can be correlated more or less with : the middle part of the Chaetoceros sp. zone (Bukry and Foster, 1973), - - the Dictyocha stapedia stapedia biozone of silicoflagellates in the section of Chamalevri in western Crete (Frydas, 1989, tabl. 1) ; Fig. 1, this work, the G. puncticulata/G, bononiensis biozones of planktic foraminifera (Gradstein, 1974 ; Zachariasse, 1975), the CNllb/CN12a calcareous nannoplankton subzones of Okada and Bukry (1980), Frydas, 1990 ; Rio et al., 1990) and the NN14-15B to NN16-17D acme-subzones of I)riever (1988, fig. 53), (- 3.7 to 2.9 Ma), the upper and the lower part of the Dictyocha fibula and D. stapedia stapedia silicoflagellate zones of Bukry (1981). - -

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Diatoms from the Kallithea, Marathitis, Prassas and Aitania sections can be correlated with the well known tropical/subtropical diatom zouation (Burckle, 1972, 1978; Baldauf and Iwai, 1995; Barron, 1985a, b) based on low-latitude or cosmopolitan marker species. The magnetostratigraphic timescale (Fig. 10) is based on biostratigraphic datings given by Berggren et al. (1985).

- -

Moreover, a similar diatom assemblage was described in the uppermost unit of the Caleta Herradura de Mejillones se~ction (lower Late Pliocene), north of Antofagasta in Chile (Koizumi, 1990) as well as in the Onzole (Ecuador) and Pisco (Peru) sections (Koizumi, 1992).

DISCUSSION ABOUT THE NITZSCHIA JOUSEAE ZONE OF BURCKLE (1972) The low-latitude Nitzschia jouseae biozone of Burckle (1972, 1978) and Barron (1985a,b)is defined as the interval from the first occurrence of Nitzschia jouseaeto the first occurrence of Rhizosolenia praebergonii MUCHINA (Early Pliocene to early Late Pliocene, fig. 10). The Nitzschia jouseae zone is equivalent to the upper planktonic foraminiferal N19 zone (Blow, 1969), the CN10c, C N l l , CN12a subzones of the calcareous nannofossils, and the Spongaster pentas z o n e of radiolarians (Sanfilippo et al., 1985). Nitzschia jouseae appears generally in low nunnbers in t h e investigated sections (Tabl. 1-2). On the contrary, Thalassiosira convexa and Th. oestrupii~ two significant species for the Pliocene, are f r e q u e n t to common in botll sections (Tabl. 1-3).

PROPOSITION OF TWO NEW LOCAL ACME-SUBZONES WITHIN THE N. JOUSEAE ZONE As revealed by tables 1 to 3 the investigated sections are characterized by different diatom assemblages. The Kallithea and Marathitis sections, dominated by subtropical diatoms, are proposed as the Thalassiosira leptopus acme subzone based on the frequent occurrence of T. leptopus. In the Prassas and Aitania sections, the subtropical species Azpeitia nodulifer and Thalassiosira leptopus are missing. These sections contain more common to frequent occurrences of the cold-water species Coscinodiscus marginatus and Actinocyclus curvatulus, and are proposed as the Coscinodiscus marginatus acme subzone.

FRYDAS

P

L

Late Zanclean

I

O

C

E

277

N

E

Epoch Age

Piacenzian ~G~ A ~ U •

o

S

S

,~

,

,

Chron Berggren et al. (1985)

Rhizosolenia praebergonii

Nitzschia jouseae

z Discooster asl,mr~letrictts'~

Discoaster tamalis

0"

to

to

2

Reticul. pseudo- _ umbilica ~

~r-)-

~,~

Discoaster surculus

~

0 m

-'

i

DicO:ocha fibula

i i i

, 1

Dictyocha stapedia stapedia

i

r~>.

i

i

t

Sphaero idinellops is subdehiscens

Globigerinoides elongatus

v

GI. puncticulatu

Globorotalia

GI. puncticnlata

Globorotalia

bononiensis

N ~z

~'~- N ~'H .

"-"

Dictyocha Dict. Cannopilus brevispina qf. neo- mqjor brevispim~ nautica Thalassiosira h'ptopus

~C3

Z

O~

crassaformis

Disteph. boliviensis boliviensis

C'o.~'cinodi.~ctt,% murginattts

~

~

FIG. 10. - Correlation of the studied sections in Central Crete based on diatom biostratigraphy and comparison with calcareous nannofossfl, silicoflagellate and planktic f'oraminiferal zonations.

Corr61ation des coupes dtudiges en Crbte centrale d'aprbs les diatomdes, et comparaison avec les biozonations de nannofossiles calcaires, silicoflagellds et foramini~res planctoniques.

278 TABLE ] . -- P e r c e n t a g e

DIATOMI~ES PLIOCI~NES DE CRI~TE v a r i a t i o n o f d i a t o m s i n t h e M a r a t h i t i s a n d P r a s s a s s e c t i o n s . 1-3 % : r a r e , 4 - 1 0 % : f r e q u e n t , a n d 20 % : a b u n d a n t . Variation des pourcentages

MARATIlITIN SECTION Diatom Ilora: (p)planktic, (m) mcroplanktic. (b) b~mthic Diatoms from Marathitls (M) section M8 M10 M14 Actinoc)~clus curvatuhts (p) 4 I 2 A. octonarms (m) 1 2 A. octonarius var. tenella Qn) 5 l Aclinopt)'chus senarius (m) 3 <1 ..I. senarilts var. nllnor (lit) .1 .,1. sldendens (b) Auliscus incertus (m) <1 <1 /lzFeitia noduliler (p) 4 6 Az. vetustJssmm (p) 6 4 <1 Bacteriastrum comosum (m) 1 <1 Biddulphia tuome),J (b) <1 <1 <1 Cestodiscus sp. (13) Chaetoceros ~astridium (p) 3 5 Cocconeis scutellum (b) <1 <1 Coscinodiscus a.steromphalus (p) 8 7 7 C. marginatus (p) 1 1 C., oculus-iridis (p) 21 24 18 C. radiatus (p) I Dimerosramma marinum (b) <1 .. DqJIoneis c£ advena (m) ~1 13. hamhus (b) 1 "1 -1 l). aamarttensis (1t) D. smithii (b) 1 <1 Epithemia sp. (b) <1 Grammatophora ansulosa (b) G. marina (p) <1 G. oceanica var. macilenta (b) 12 ltemidiscus cuneformis (p ) 5 9 <1 Navtcula henned),i (b) c1 Nttzschia,lonseae (p) 3 2 5 N. marina (p) "1 1 PSelldodnnerogramnla elongalunl ". l

(b)

Simonseniella barboi (p) Stephano~, onia aculeata (b) Stephanop)'xis corona (rn) S. turris (m) Stictodlscus parallelus var. parallelus fa. quadrata (b) Surirella ovata (b) 53,nedra.fidgens (b) 5),. tabulata (b) Thalassionema nitzschioides ~p) ThalassJosira convexa (p) Th. eccentrica (p) Th. leptopus (p) 7h. oestrupti (p) Thalassiothrix ct~ lon~issima (p ) Trachyneis aspera (b) 7)'weratium balearicum fa. biquadrata (b)

d e d i a t o m 6 e s d a n s les c o u p e s d e M a r a t h i t i s et P r a s s a s .

M18 2 3 2 <1 I 3 2 2 3 1 2 <1 5 2 16 3 <1 1 l

M21 <1 3 2 2

M23 1

1

l <1 2 2

2 3 4 4 1 4 2 6 2 21 4

3

1

3

1 <1 3 2 3 1 "1

10 2 <1

2

1

1

<1 4 1

10 6

21 9

4 5 <1 14 16

l <1

1

1 1 <1 4 4 "22 2

2

2 <1 3 2 3 2

2

1

<1 <1

2 1

2

<1 <1 l 4

11-20 % : c o m m o n ,

4 1

PRASSAS SECTION l)iatom flora: ~p) planktic, (m) meroplanktie, (b) baathic Diatorrm from Prassas (P)section P5 P10 Actinoc, vclus curvatulus (~) 6 7 A. octonarms (m) 3 6 A. octonarius var. tenella (m) 1 3 Actinoptvchus senarius (m) A. senorius var. minor (m) Aul(scus incertus <1 Azpeitia vetustissima (p) 2 Bacteriastrum comosum (m) 3 2 B. delicatulum (nQ I Biddulphia tuomeyi (b) 2 1 Chaetoceros 8astrJdium (p) 3 3 Cocconeis scutellum (b) Coscinodiscus asteromphalus (p) 6 5 C mar~inatus (p) 12 8 C: 9culus-iridis (p) 18 20 C. radiams (p) 3 Dirnero~,ramma marinum (b) <1 Diploneis e£ advena (in) 1 D. bombus (b) D. octmaruensis (b) 1 Grammatol~hora angulosa (b) 1 Ilemidiscus cunei~ormis (p) 4 5 Navicula hennedyi (b) 2 3 Nitzschia ]ouseae (p ) 3 1 Rhabdonema adriaticum (b) 1 2 Rhizosolenia hebetata fa. semispina (p) 3 2 ? Rhizosolenia sp. Stephanogonia aculeata (b) <1 Stephanopvxis turris (m) 4 3 Surirella ovata (b) Synedra undulata (b) 3 Thalassiosira convexa (p) 8 6 Th. eccentrica (p) Th. oestrutgii (p) Thqlassiothrix eE lon~issima (p ) Triceratium balearicum fa. biquadrata

O)

Trigonium arcticum var. pentagonalis

(b)

3 10 1

4 12 1

P12 6 3 2

<1 2 2 2 6 10 19 2

P14 7 4 2 3 1

PI5 4 6 2 1

3

6 3

5 <1 <1

3 <1 2 i1 17

3 3

9 20

<1 <1 4 3 <1 2 2 3 <1 5
<1 8

4

1 <1 6 <1

2 2 3

<1

2 <1

3

6

9

5 9 1 1

5 11 <1 <1

<1

<1 <1 1 5

<1

4

2 7

11 8

10 6

12 6

<1

<1

I 1

<1

<1

The paleogeographical evolution on Crete Island during the Late Neogene is characterized by the migration of the tectonic activity northwards (Kontopoulos et al., 1996). According to these synsedimentary tectonic movements, a number of individual basins have developed in the studied area with a

different stratigraphical history. Subsequently, the two paleoclimatic intervals recognized by diatom assemblages coincide with the stratigraphical evolution determined by planktonic foraminifera and calcareous nannofossils during Late Zanclean-Early Piacenzian period (Fig. 7).

FRYDAS

279

TABLE2. Percentage variation of diatoms in the Kallithea section. Variation des pourcentages de diatom~es dans la coupe de Kalethea. -

Diatom flora: (p) planktic, (in) mero Diatoms from Kallithea K1 section Actinoclvclus curvatulus (p) 4 .-t. octonarius (m) 4 .'4. octonarius var. tenella 6

KALLITHEA SECTION 31anktic. 09) benthic K2 K3 K4 K5

2 6 7

1 2 <1

5 7 <1 1

3 4 <1 <1

<1 8 5

<1 3 6


<1

1

<1 2

K6

K7

K8

5

2 6

<1 8 2

K9

K10 7 5 <1

(m) .:lctinop@'chus senarius (m) Azpeitia nodulifer (p) .-tz. vetustissima (p) Bacteriastrunz comosum

6 4 <1

<1 <1

<1

4 10 <1 26 3 12 2 4

3 2 2

3 2 3 4

3 3 7

4 4 9

2 32 5 3

2 17 4 4

(m) Biddulphia tuomevi Oo) Chaetoceros ~,astridium (p) Coscinodiscus asteromphalus (p) (7.. mar einatus (p) C. oculus-iridis (p) C. radiatus (p) Hemidiscus cune!formis (p) Navicula hennedvi (19) Nitzschia/ouseae (p) A;. marina (p) Rhabdonema adriaticum (b) Simonseniella barboi (p) Stephanopyxis turris (m) Surirella ovata (b) Thalassi onema nitzschioides (p) Thalassiosira convexa (p) Th. eccentrica (p) Th. leptopus (p) Th. oestmtpii (p) Thalassiothrix cf. Lon~issima (p) Tri ceratium balearicum f~ biquadrata (b)

Thalassiosira

leptopus

20 7 2

4

4 1 21 3

<1

23 <1 9 <1 4 <1

11

7

6 8

1 26


1 27

11 <1

14 <1

16 3

6 <1 <1

1

6 <1 15

9 <1 13 5 1

1

acme-subzone

Definition : Interval where Thalussiosira leptopus is frequent to a b u n d a n t . Localities : The diatomiteo horizons of both Kallithca and Marathitis sections.

1 3 <1 2 1

2

<1 2 <1

1 6 6

21 3 8 2 2 <1 <1

3 1 4

2 1 3

1

4

1

<1

7

I1

8

4

4

23 4

10 2

13 5

20

21 3

_~ <1 14 2

<1

<1

6
5 7 6

<1 24 4 5 <1 <1

2 4 2 5 18

1

<1

<1

Age : Late Zanclean - E a r l y Piacenzian. Regional distribution: Kallithea and Marathitis sections in central Crete, Heraklion Province, as well as Asteri section (Fig. 1) on western Crete, Rethymnon Province (Frydas, 1998).

DIATOM]~ES PLIOC]~NES DE CRI~TE

280

TABLE3. Percentage variation of diatoms in the Aitania section. Variation des pourcentages de diatom~es dans la coupe d'Aitania. -

AITANIA

SECTION

Diatom flora: (p) planktic. (m) meroplanlaik. (b) benthic Diatoms from Aitania section A1 A2 A3 Actinocvclus curvatulus (p) 5 7 8 A. octonarius (m) 2 6 5 A. octonariusvar, tenella (m) <1 <1 2 Azpeitia vetustissima (p) 2 <1 <1 Biddulphia tuomevi 09) <1 2 1 Chaetoceros ~,astridium (p) 4 6 3 Cocconeisscutetlum (19) <1 2 Coscinodiscus asteromphalus 6 14 10

A4 10 1

A5 7 4 7

A6 11 8

A8 9 5

14

A7 7 8 <1 4 <1 6 <1 9

3

<1 <1 2

<1 7 <1 9

3 3 1 13

8 19 <1

!2 28

8 21

9 25

10 29

9 <1 3 8 3

10 3

11 22 <1 <1 5

7

9 <1

7


3 4 7

6

4

4

8

(p) C. mar~znatus (p) C. oculus-iridis (p) Diploneis oamaruensis (b) Epithemia sp. (b) Hemidiscus cuneiformis (p) Ntn,icula hennedvi (b) Nitzschiajouseae (p) Rhabdonema adriaticum (b) Stephanopvxis turris (m) Slictodiscus sp. (b) Thalassiosira convexa (p) Th. eccentrica (p) Th. oestrupii (p) Triceratium balearicum fa. biquadrata 09)

14 28

14

6 4 9 5

10 16 <1 <1 11 <1 2 2 6 2 8 1 4

Assemblage : The diatom nannoflora (Table 1-2) mainly consists of planktic centric (70-90 %), meroplanktic (5-20 %) and benthic species (2-15 %). The most important diatoms are : a) Planktic species - A z p e i t i a nodulifer (0 to 8 %), Coscinodiscus asteromphalus (4-10 %), C. oculus iridis (15-30 %), Hemidiscus cuneiformis (3 to 15 %), Nitzschia jouseae (0-5 %), Thalassiosira convexa (4-10 %), Th. leptopus (10-25 %), Th. oestrupii (0-15 %). b) Meroplanktic species -Actinocyclus octonarius (0-8 %), A. octonarius vat. tenella (0-7 %), Actinoptychus senarius (0-5 %), Bacteriastrum comosum (0-4 %) and Stephanopyxis turris (0-6 %). c) Benthic species - Biddulphia tuomeyi (0-5 %), Navicula hennedyi (0-2 %) and Triceratium balearicum f. biquadrata ( 0 - 1 % ) .

8 <1 8 <1

6 2 <1 5 <1 8

6

3 6 2 4 <1 3 <1

<1 <1 8 <1 14 4

Correlation: The Thalassiosira leptopus acme subzone can be correlated with : - - the silicoflagellate Dictyocha brevispina brevispina and D. cf. neonautica acme - subzones (Frydas, 1990, 1996), which indicate a warm-water paleoclimate interval, --the C N l l b - Discoaster asymmetricus and CN12a - D. tamalis subzones of calcareous nannofossils (Okada and Bukry, 1980), --the NN14-15B calcareous nannofossfl acme subzone in the Kallithea section, and the NN16-17A to NN16-17B in the Aghios Vlassios section (Driever, 1988, fig. 53), - the Globorotalia puncticulata zone of planktic foraminifera in the Kallithea section (Spaak, 1983, fig. 18; Jonkers, 1984, fig. 45).

FRYDAS

Coscinodiscus marginatus a c m e - s u b z o n e Definition : Interval t¥om the level of increase in the Coscinodiscus marginatus and Actinocyclus curvatulus a b u n d a n c e s (sa. P5 : Prassas section and sa. AI : Aitania section), above the last occurrences of Azpeitia nodulifer, Thalassionema nitzschioides and

281

--the Globorotalia crassaformis zone in the Prassas (Spaak, 1983, fig. 18) and Aitania (Jonkers, 1984, fig. 45) sections.

TAXONOMY

Thalassiosira leptopus. Localities : The diatomite horizons of both P r a s sas and Aitania sections. Age : Early Piacenzian. Regional distribution : Prassas and Aitania sections in central Crete, Heraklion Province. Assemblage: The diatom assemblage from both sections (Tabl. 1, 3) consists of planktic (75-90 %), meroplanktic (10-20 %) and benthic species (315 %). The most important diatoms are : a) Planktic species - A c t i n o c y c l u s curvatulus (510 %), Coscinodiscus asteromphalus (2-15 %) C. marginatus (8-15 %), C. oculus-iridis (15-30 %), Hemidiscus cuneiformis (3-15 %), Nitzschia jouseae (0-3 %), Rhizosolenia hebetata fa. semispina (06 %), continuously occurring in Prassas section, but missing in Aitania samples, Thalassiosira convexa (5-15 %) and Th. oestrupii (3-12 %). b) Meroplanktic species : Those of the Marathitis section plus Bacteriastrum delicatulum (0-3 %). c) Benthic species : Those of the Marathitis section. Correlation : The Coscinodiscus marginatus acme subzone can be more or less correlated with : the Aurila convexa emalthiae zone of ostracodes (Tsapralis, 1976), - - the silicoflagellate Cannopilus major and Distephanus boliviensis boliviensis acme - subzones (early Late Pliocene) (Frydas, 1990, 1996), which indicate a temperate paleoclimatic interval, the Dictyocha stapedia stapedia zone of silicoflagellates as well as the G. bononiensis zone of planktic foraminifera in the Chamalevri section on western Crete (Frydas, 1989; F r y d a s and Bellas, 1994, pl. 1, fig. 11-15), the CN12a Discoaster tamalis calcareous nannofossil subzone of Okada and B u k r y (1980), (Frydas, 1990), - - the NN16-17C to NN16-17D calcareous nannofossil acme subzones (Early Piacenzian) (Driever, 1988, fig. 53) for the u p p e r p a r t of the Prassas and Aitania sections, the Globorotalia bononiensis zone of planktic foraminifera (Zachariasse, 1975), -

-

-

-

-

-

-

-

Genus Actinocyclus EHRENBERG (1837) Actinocyclus curvatulus JANISCH in Schmidt (1878) (P1. 1, fig. 15-16) 1874-1900. - Actinocyclus curvatulus JANISCH. -- A. Schmidt, pl. 57, fig. 31. 1973. - Actinocychts curvatulus JANISCtL-- Schrader, pL 19, fig. 2. 1995. - Actinocyclus curvatulus JANISCH.-- Fenner, p. 78, pl. 1, fig. 7.

Remarks: Eocene to Recent. F r e q u e n t pelagic form in recent seas and oceans. Common to r a r e in our samples. Actinocyclus o c t o n a r i u s EHRENBERG (1837) (P1. 2, fig. 1-4; P1. 4, fig. 1-2, 4-5) 1837. - Actinocychts octonarius n. sp. - Ehrenberg, p. 172, pl. 21, fig. 7. 1861. -Actinocyclus ehrenbergii RALFS. - Pritchard, p. 834.

Remarks : Specimens of Actinocyclus octonarius, A. octonarius var. teneUa, Actinoptychus splendens, Coscinodiscus marginatus, C. oculus-iridis and Thalassiosira leptopus belong to the group of species characterized by t u b u l a r fultoportulae and/or stalked lipped rimoportulae, which are usually symmetrically a r r a n g e d on the valve lace or on the valve mantle. The arrangement of the rimoportulae tends to be species - specific (Round et al., 1990). Both their length and their form can be used as an additional morphological criterion for distinguishing the species mentioned above. Distinct formed rimoportulae (the term, labiate processes, has been used by Medlin et al., 1986; Sancetta 1987) are symmetrically a r r a n g e d on the valve-mantle (P1. 4, fig. 1, 4). These small m u s h r o o m - s h a p e d processes have a very short stalk and a length of about 3 p m with a split in their top (P1. 4, fig. 2, 5). They are an important additional criterion for the determination of A. octonaflus. Cosmopolitan in recent seas and oceans. Polyhalobe, meso- to meioeuryhaline (Pankow, 1976). Miocene to Holocene. In Crete, known from the Late Miocene Cherethiana and Panassos I sections (Gersonde, 1980). F r e q u e n t to r a r e in our material.

282

DIATOM]~ES PLIOC}~NES DE CR}~TE

Actinocyclus

octonarius var. teneHa H E N D E Y (1964)

(BRI~BISSON)

(P1. 1, fig. 18-19; PI. 4, fig. 7-9) 1854. - Eupodiscus tenelhts n. sp. - Br6bisson, p. 257, pl. 1, fig. 9. 1930. - Actinocyclus ehrenbergii RALFS var. tenella (BREBISSON). -- Hustedt, part 1, p. 530, fig. 302.

Remarks : The rimoportulae of A. octonarius var. tenella differs from those of A. octonarius through their 60 ° arrangement and their long stalk of about 1,25 llm (P1. 4, fig. 7-9). Miocene to Holocene in recent seas and ocean environments. Meroplankton (neritic), polyhalobe, meso- to meioeuryhaline form (Pankow, 1976).

(Abbott and Andrews, 1979). In the Mediterranean Sea occurs in lesser numbers as in the northern coasts of Middle Europe (Hustedt, 1930). In the western Baltic Sea neritic, polyhalobe, meioeuryhaline (salinity : 35 to 20-17 %~, cosmopolitan (Pankow, 1976). A. senarius has been recorded in the Late Miocene calcareous nannofossil zone NN11a from Merada section, NW-Crete by Bellas and Frydas (1994, tab. 2, fig. 26). Frequent to rare in the here investigated sections.

Actinoptychus senarius var. m i n o r (A. CLEVE) HAJOS (1968) (P1. 1, fig. 14)

Genus Actinoptychus EHRENBERG (1843) 1941. - Actinoptychus undulatus vat. minor. p. 179, pl. 6, fig. 92-94.

Aetinoptyehus senarins ( E H R E N B E R G ) E H R E N B E R G (1843) (P1. 1, fig. 13) 1837. -Actinocyclus senarius n. sp. - Ehrenberg, p. 172, pl. 21, fig. 6. 1843. - A c t i n o p t y c h u s senarius (E~IR.). - Ehrenberg, p. 400, pl. 1, fig. 27. 1930. - Actinoptychus undulatus (BAILEY) RALFS. -- Hustedt, part 1, p. 475, fig. 264.

Remarks : Cretaceous to Holocene. Common occurrence in cool, coastal waters in seas and oceans

Cleve-Euler,

1968. - A c t i n o p t y c h u s senarius (EHR.) EHRENBERG vat. minor (A. CLEVE). -- ttajos, p. 121-122, pl. 30, fig. 6-8.

Remarks : Diameter 20-30 ~am. Margin with 14-15 teeth in 10 lam. Valve divided into 6 sectors. Hexagonal hyaline central area. About 8 areolae in 10 pm. Rare in the Marathitis and Prassas section. This variety is only known fossil from South Lapland and from the upper marine diatomite-earth of the Szurdokpiispi~ki section, Miocene of Hungarn (Hajos, 1968).

PLATE 1 Optical microscopy, x 850 (approximately), except fig. 10 : x 2500 approximatelly. Microscopie optique, x 850 environ s a u f fig. 10 : x 2500 environ. la-c. Coscinodiscus asteromphalus EHRENBERG. 12. Actinoptychus splendens (SHADBOLDT) RALFS. Marathitis, M23. Arrows indicate the hyaline ray in the Marathitis, sample M10 (la) and M14 (1b-c). middle of the sector (see also P1. 3, fig. 4b). Les f ~ c h e s 2a, b. Thalassiosira convexa MUCHINA. indiquent le rayon hyalin au milieu du secteur : voir aussi Marathitis, M23. Pl. 3, fig. 4b. 3-5. Coscinodiscus oculus-iridis EHRENBERG. 13. Actinoptychus senarius (EHRENBERG) EHRENBERG. Marathitis, M18(3), M10(4) and M23(5). Arrows indicate opeMarathitis, M18. nings of rimoportulae. Les fl~ches indiquent les ouvertures 14. Actinoptychus senarius (EHR.) EHR. var. minor (A. CLEVE) des rimoportulae. HA.lOS. 6-8. Thalassiosira leptopus (GRUNOW) HASLE and FRYXELL. Marathitis, M18. Marathitis, M10(6), M18(7) and M23(8). Arrows indicate ope15-16. Actinocyclus curvatulus JANISCH in Schmidt. nings of rimoportulae. Les fl~ches indiquent les ouvertures Marathitis, M8(15) and M18(16). des rimoportulae. 17a-c. Thalassiosira oestrupii (OSTENFELD) PROSCHKINA-LAVRENKO. 9-10. Coscinodiscus radiatus EHRENBERG. Marathitis, M18(17a) ; Prassas, P5(17b, c). Marathitis, M18. 18-19. Actinocyclus octonarius EHRENBERG var. teneUa (BRI~BIS10 : C. radiatus. Detail from central area, with a small hyaline field. Ddtail de la rdgion centrale, & aire hyaline petite. SON) HENDEY. Marathitis, M8(18) and M18(19). 11. Coscinodiscus marginatus EHRENBERG. Marathitis, M23.

20. Cestodiscus sp. Marathitis, M18.

FRYDAS

REVUE DE MICROPALt~ONTOLOGIE, VOL. 42, N ° 4

PLATE 1

284

DIATOMI~ES PLIOCI~NES DE CRt~TE

Actinoptychus splendens (SHADBOLDT) RALFS in Pritchard (1861) (P1. 1, fig. 12; P1. 3, fig. 4); P1. 4, fig. 15)

The oldest known certain occurrence seems to be of Early Pliocene age and it is a common Recent littoral species (Andrews, 1980). Rare in the upper part of the Marathitis section.

1854. - A c t i n o s p h a e n i a splendens n. sp. - Shadboldt, p. 16. 1930. - Actinoptychus splendens (SHADBOLDT)RALFS in Pritchard. - Hustedt, part 1, p. 478-480, fig. 265.

Remarks: The most significant criterion of A. splendens is the narrow hyaline ray (P1. 3, fig. 4b), which extends from the hyaline central area of the valve to the vicinity of the fuhoportulae near the margin of each depressed or elevated sector (P1. 3, fig. 4a). Fultoportulae have been observed around the valve mantle (PI. 4, fig. 15).

Genus Auliscus EHRENBERG (1845) Auliscus incertus SCHMIDT (1886) (P1. 2, fig. 19; P1. 3, fig. 16a, b) 1930. - Auliscus iacertus SCHMIDT,1886. - Hustedt, part 1, p. 522, fig. 296. 1989. - Auliscus incertus SCHMIDT, 1886. - Desikachary and Sreelatha, p. 84, pl. 35, fig. 4.

PLATE 2 Optical microscopy, × 1200 (approximately). - Microscopie optique, × 1200 environ. 23. Stictodiscus parallelus (EHRENBERG) var. parallehts fa. qua1-3 : Marathitis, sample M10(1), M14(2) and M21(3). drata TEMPI~RE and PERAGALLO. 4 : Prassas, P5. Marathitis, M23. 24. SurireUa ovata KOTZING. 5. Trigonium arcticum var. pentagonalis (TEMPI~RE and PERAMarathitis, M18. GALLO) DESIKACHARYand SREELATHA. Prassas, P15. 25. Cocconeis scutellum EHRENBERG. Marathitis, M21. 6. Thalassiosira eccentrica (EHRENBERG)CLEVE. 26-27. Grammatophora angalosa EHRENBERG. Prassas, P14. Marathitis, M21 : (26) girdle view (vue connective). (27) valve 7, 8. Azpeitia nodulifer (A. SCIIMIDT) FRYXELL and SIMS. view (rue valvaire). Marathitis, M8(7), M10(8), 28. Grammatophora marina (LYNGBYE) K[JTZING. 9. Hemidiscus cnneiformis WALLICH. Marathitis, M23 ; valve view (vue valvaire). Marathitis, M10. 29. Epithentia sp. 10-12. Biddulphia tuomeyi (BAILEY) ROPER. Marathitis, M14. 10, 11 : Girdle view, (vue connective) Marathitis, M18(10) and 30. Synedra fttlgens (GREVILLE)W. SMITH. M21(ll). Marathitis, M18. 12 : Valve view, (vue valvaire), Marathitis, M21. 31. Trachyneis aspera (EHRENBERG)CLEVE. 13, Diploneis oamaruensis (CLEVE) MILLS. Marathitis, M18. Marathitis, M18. 32. Pseudodimerogramma elongatum SCHRADER. 14. Diploneis cf. advena (A. SCHMIDT)CLEVE. Marathitis, M18. Marathitis, MI8. 33. Rhabdonema adriaticum K{]TZING. Prassas, P10 : girdle view (vae connective). 15. Diploneis bombus EHRENBERG. Marathitis, M18. 34. Sintonseniella barboi (BRUN) FENNER. Marathitis, M23. 16. Diploneis smithii (BR~mSSON) CLEVE. Marathitis, M8. 35. Rhizosolenia hebetata fa. semispina (HENSEN) GRAN. Prassas, P5. 17. Navicula hennedyi W.SMITH. 36. Thalassiothrix cf. longissima CLEVE and GRUNOW. Prassas, P5. Prassas, P5. 18. Triceratium balearicum CLEVE and GRUNOW fa. biquadrata 37. Synedra undalata (BAILEY) GREGORY. (JAnIscn) HUSTEDT. Prassas, P5. Marathitis, M18. 38. Stephanogonia aculeata PANTOCSEK. 19. Auliscus incertus SCHMIDT. Prassas, P10. Prassas, P5. 39. Bacteriastrum delicatulum CLEVE. 20. Synedra tabulata (AGARDII) K~)TZING. Prassas, P14. Marathitis, M18. 40. Bacteriastrum c o m o s t t n t PAVILLARD. 21. Stephanopyxis turris (GREVILLEand ARNOTT) RALFS. Marathitis, M21. Maratifitis, M8. 41-42. Nitzschia jouseae BURCKLE. 22. Stephanopyxis corona (EHRENBERG) GRUNOW. 41 : Marathitis, M14(41a), M23(41b). 42 : Prassas P5(42a), P15(42b). Marathitis, M8.

14. Actinocyclus octonarius EHRENBERG.

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R e m a r k s : Valve elliptical. Diameter 30 to 50 pro. Surface rising slightly from the central area to ocelli. Central area circular. Valve markings distinct, striate, striae converging to the ocelli, the other diverging, straight along the minor axis, elsewhere convexe towards the ocelli. Two subcirctdar ocelli, 12.5 pm broad (P1. 3, fig. 16b). Oldest known occurrence observed in the Oainaru Diatomite, Upper Eocene of New Zealand (Desikachary and Sreelatha, 1989). Very rare in our material.

1986. - Azpeitia nodulifer (A. SCHMIDT). -- Fryxcll and Sims, in Fryxell et al., p. 19, fig. 17, 18 (1, 2, 4, 5) ; SEM : 30 (3, 4).

R e m a r k s : Valve circular. Diameter 60 pm. Areolae hexagonal but somewhat irregularly arranged, 4-5 in 10 pm, decreasing in size towards the margin. A distinct rounded papille in the centre of the valve (P1.4, fig. 11). Border clear, striate, with 9-10 striae in l0 pm. Miocene to Recent. Planktic, polyhalobe, warm-water species, cosmopolitan. Frequent to rare in the Kallithea and Marathitis sections.

Azpeitia vetustissima (PANTOCSEK) in Fryxell et al. (1986) (P1. 3, fig. 6)

Genus Azpeitia PERAGALLOin Temp~re and Peragallo (1912)

SIMS

1886. - Coscinodiscus vetustissimus n. sp. - Pantocsek, I, p. 71, tabl. 20, fig. 186.

Azpeitia nodtdifer (A. SCHMIDT) FRYXELL and SIMS in Fryxell et al. (1986) (P1. 2, fig. 7-8; P1. 4, fig. 10-11)

1930. - Coscinodiscus vetastissimus PANTOCSEK. -- Hustedt, part 1, p. 412-414, fig. 220.

1878. - Coscinodiscus nodulifer A. S C H M I D T , Atlas der Diatomaceenkunde, 15, pl. 59, fig. 20, 22, 23.

1986. - Azpeitia vetustissima (PANTOCSEK). -- Sims in Fryxell et al., p. 16.

1930. - Coscinodiscus nodulifer A. SCIIMIDT. -- Hustedt, part 1, p. 426-427, fig. 229.

1989. - Azpeitia vetustissima (PANTOCSEK). -- Sims et al., p. 303, pl. 2, fig. 11 13, pl. 3, fig. 10.

PLATE 3 Scanning electron microscope views, x 850 (approximately) except fig. 2a, 3a and 15 : x 2,200 ; fig. 5b and 17 : x 3,500 ; fig. 2b, 3b and 4b : × 10,000. Microseopie 6lectronique ~t balayage, x 850 environ, s a u f fig. 2a, 3a et 15 : × 2 200 ; fig. 5b et 17 : x 3 500 ; fig. 2b, 3b et 4b : × 10 000. 1-3. Thalassiosira leptopus (GRUNOW) HASLE and FRYXELL. 12a, b. Biddulphia tumneyi (BAILEY) ROPER. Marathitis, sample M23. Marathitis, M18(12a) : valve view (rue valvaire); (12b) : 2a, b : Details of the small spinules on the margin. Ddtails girdle view (vue connective). des petites @ines marginales. 13. Thalassiothrix cf. longissima CLEVE and GRUNOW. 3a, b : Arrows indicate the tubular rimoportulae with an Marathitis, M23. anlipped opening on the inner valve mantle. Les fl~ches 14. Bacteriastrum comosum PAVILLARD. indiquent les rimoportulae tabalaires h ouverture arronPrassas, P5. Specimen with nine well prcserved bristles. die sans l~vre, sur la marge interne de la valve. Specimen ~t 9 soies bien conservdes. 4a-b. Actinoptychus splendens ( S H A D B O L D T ) RALFS. 15. Chaetoceros sp. (bristles, soies). Marathitis, M23(4a) Prassas, P5. 4b : Arrow indicates the narrow hyaline ray of each elevated sector. La fl~che montre le mince rayon hyalin de l'un 16a, b : Aaliscus incertas SCtIMIDT. des secteurs surdlevds. Marathitis, M18; 16b : detail of ocellus, 12.5 pm broad. Ddtail d'un ocelle, large de 12,5 ttm. 5a, b. Coscinodiscus radiatus EHRENBERG. Marathitis, M18; 5b : detail for the hyaline central area. 17. Rhizosolenia sp. (?). Detail de l'aire centrale hyaline. Prassas, P14. 6. Azpeitia vetustissima (PANTOCSEK) SIMS. 18. Grantmatophora oceanica (EIIRENBERG) var. macilenta (W. Marathitis, M10. SMiTh) GRUNOW. Marathitis, M18. 7. Navicula hennedyi W. SMtTI-I. Prassas, P5. 19. Synedra undnlata (BAILEY) GREGORY. Prassas, P5. 8. Stephanopyxis corona (EHRENBERG) GRUNOW. Marathitis, M8. 20. Grammatophora attgulosa EHRENBERG.

9. Stephanopyxis turris (GREVILLE and ARNOTT) RALFS. Marathitis, M8. 10-11. Dimerogramma m a r i n u m (GREGORY) RALFS. Marathitis, M8.

Marathitis, M21. 21. Triceratium balearicum CLEVE and GRUNOW fa. biquadrata (JANISEII) H U S T E D T . Prassas, PI0.

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DIATOMI~ES PLIOCt~NES DE CRI~TE

R e m a r k s : Late Oligocene t r o u g h Late Miocene (Sims et al., 1989). Miocene of H u n g a r y , f r o m the u p p e r m a r i n e d i a t o m i t e e a r t h of the locality Sznrd o k p i i s p ~ k i ( H a j o s , 1968). Middle Miocene f r o m South Carolina and Georgia (Abbott and Andrews, 1979) a n d Miocene of the Atlantic Margin ( A b b o t t , 1980). K o i z u m i (1973) gave a range f r o m N9 to N17 f o r a m i n i f e r a l zones, o r t h r o u g h o u t the Middle to Late Miocene. F r e q u e n t to r a r e in the investigated sections. Genus B a c t e r i a s t r u m SHADBOLT (1853) B a c t e r i a s t r u m e o m o s u m PAVILLARD (1905) (PI. 2, fig. 4 0 ; P1. 3, fig. 14) 1973. - Bacteriastrum comosttm PAVILLARD.-

Hajos,

Genus C e s t o d i s c u s GREVILLE (1865) C e s t o d i s c u s sp. (PI. 1, fig. 20) R e m a r k s : S i m i l a r to Actinocyclus octonarius b u t w i t h o u t p s e u d o n o d u l e s . S i m i l a r f o r m s , belonging to the Denticula d i m o r p h a zone (Middle/Late Miocene b o u n d a r y ) a n d to the R h a p h i d o d i s c u s m a r y l a n d i c u s - to P y x i l l a sp.-zone (Earliest Miocene), have been p r e s e n t e d f r o m S o u t h Pacific Ocean b y S c h r a d e r (1976). A d d i t i o n a l l y similar forms were d e s c r i b e d as Cestodiscus sp. 1, C. sp. 3, C. sp. 4 a n d C. sp. 6 b y G e r s o n d e (1980). P l a n k t i c , p o l y h a l o b i a n species. V e r y r a r e in M a r a t h i t i s section.

pl. 6, fig. 2.

R e m a r k s : Valve c i r c u l a r . D i a m e t e r a b o u t 10 lira. 9 to 11 c u r v e d bristles arise f r o m the m a r g i n of the valve, a n d extend o u t w a r d in a d i s t a n c e of a b o u t 3 d i a m e t e r length f r o m the valve m a r g i n . F r e q u e n t in a n u m b e r of i n v e s t i g a t e d sections in Crete. K n o w n at least f r o m the Messinian ( F r y d a s 1985, 1994b). B a c t e r i a s t r u m d e l i e a t u l u m CLEVE (1897) (P1. 2, fig. 39)

Genus C h a e t o c e r o s EHRENBERG (1845) C h a e t o c e r o s g a s t r i d i u m (EHRENBERG) BRIGHTWELL (1856) (P1. 4, fig. 3) 1846. - Chaetoceros didymus n. sp. - Ehrenberg, p. 75. 1973. - Chaetoceros gastridium ( E H R E N B E R G ) BRIGHTWELL. Hajos, p. 962, pl. 6, fig. 5.

--

1897. - Bacteriastrum delicatulum n. sp. - Cleve, p. 298, fig. 15. 1994. - Bacteriastrum delicatulum CLEVE.- - Frydas, pl. 1, fig. 22.

R e m a r k s : T h e ends of the bristles a r e b i f u r c a t i n g at a distance e q u a l to the elliptical k n o b .

R e m a r k s : Valve c i r c u l a r . D i a m e t e r a b o u t 8 to 10 pm. A v a r i o u s n u m b e r of bristles (7 to 11) arise f r o m the m a r g i n of the valve a n d e x t e n d o u t w a r d , b i f u r c a t i n g at a d i s t a n c e of a b o u t 1.3 d i a m e t e r f r o m the valve m a r g i n ( F r y d a s , 1994b). F o u n d in g r e a t a b u n d a n c e in the d i a t o m i t e f r o m S t a v r o m e n o s section, E a r l y P i a c e n z i a n of C e n t r a l C r e t e , which belongs to the D i s t e p h a n u s boliviensis boliviensis a c m e - s u b z o n e of silicoflagellates ( F r y d a s , 1994b). R a r e in the P r a s s a s section.

Oldest k n o w n o c c u r r e n c e o b s e r v e d in the D i a t o m i t e , U p p e r E o c e n e , New Z e a l a n d c h a r y a n d S r e e l a t h a , 1989). C o m m o n in the d i a t o m i t i c deposits of Crete. P o l y h a l o b i a n , r y h a l i n e , cosmopolitan.

Genus B i d d u l p h i a GRAY (1821) B i d d u l p h i a t u o m e y i (BAILEY) ROPER (1859) (P1. 2, fig. 10-12; P1. 3, fig. 12) 1837. - Denticella tridens n. sp. - Ehrenberg, p. 129. 1841a. - Biddulphia tridens n. sp. - Ehrenberg, p. 205. 1844. - Zygoceros tuomeyi n. sp. - Bailey, p. 158, fig. 3,4. 1859. - Biddulphia tuomeyi ( B A I L E Y ) . - Roper, p. 8, pl. 1, fig. 12. R e m a r k s : C r e t a c e o u s to Holocene benthic form. F r e q u e n t in m o d e r n coastal waters of w a r m e r seas. Miocene f r o m Sicily, S p a i n , Algeria a n d Crete (Gersonde, 1980), a n d R e c e n t in the coastal region of the K a l y m n o s i s l a n d , G r e e c e (Foged, 1985b). R a r e to c o m m o n in o u r m a t e r i a l .

Oamaru (DesikaPliocene meioeu-

C h a e t o c e r o s sp. (Pl. 3, fig. 15) R e m a r k s : Complete spores of Chaetoceros a r e f r e q u e n t to r a r e in o u r samples. U s u a l l y two long bristles (P1. 3, fig. 15) a r e j o i n t t o g e t h e r in one elliptical k n o b .

Genus C o c c o n e i s EHRENBERG (1837) C o c c o n e i s s c u t e l l u m EItRENBERG (1837) (P1. 2, fig. 25) 1837. - Cocconeis scutellum n. sp. - Ehrenberg, p. 194, pl. 14, fig. 8. 1959. - Cocconeis scutellum EHRENBERG. - - Hustedt, part 1, p. 337-339, fig. 790. R e m a r k s : Benthic m a r i n e f o r m f r o m Miocene to H o l o c e n e , l i t t o r a l , e p i p h y t i c , cosmopolitan. A polyh a l o b i a n , meio- to p l e i o e u r y h a l i n e ( P a n k o w , 1976).

FRYDAS Genus C o s c i n o d i s c u s EHRENBERG (1837) C o s c i n o d i s c u s a s t e r o m p h a l u s EHRENBERG (1846) (P1. 1, fig. l a - c ) 1846. - Coscinodiscus asteromphalus n. sp. - Ehrenberg, p. 77. 1854. - Coscinodiscus asteromphalus EHR. - Ehrenberg, pl. 18, fig. 45; pl. 33, fig. 15(7). 1930. Coscinodiscus asteromphahts EHR. -- Hustedt, part 2, p. 452-454, fig. 250. R e m a r k s : Original d e s c r i p t i o n b y E h r e n b e r g front the M i d d l e Miocene deposits at R i c h m o n d , Virginia, U n i t e d States. Known in Crete f r o m Miocene a n d Pliocene sections ( G e r s o n d e , 1980). C o m m o n to f r e q u e n t in all the s t u d i e d sections. P o l y h a l o b i a n , meso- to m e i o e u r y h a l i n e , c o s m o p o l i t a n species ( P a n kow, 1976).

C o s c i n o d i s c u s m a r g i n a t u s EtIRENBERG (1843) (P1. 1, fig. 11) 1843. - Coscinodiscus marginatus n. sp. - Ehrenberg, 1843, p. 329, 371. 1854. - Coscinodiscus marginatus EttR. Ehrenberg, pl. 17, fig. 44. 1930. - Coscinodiscus marginatus EnR. Hustedt, part 1, p. 416418, fig. 223. R e m a r k s : C. marginatus a n d C. oculus-iridis belong to the g r o u p of species c h a r a c t e r i z e d f r o m b o t h f u h o p o r t u l a e a n d r i m o p o r t u l a e on the valve face a n d on the valve mantle. C. marginatus is distinguished f r o m C. oculus-iridis by the possession of a b r o a d h y a l i n e b a n d p r o j e c t i n g i n w a r d f r o m the valve edge, which seems like a striate m a r g i n in light m i c r o s c o p y . It is distinguished from Eudictya hungarica HAJOS (1968) b y the possession of a r e o l a e distinctly h e x a gonal a n d n o t r o u n d e d as b y E. h u n g a r i e a , Eocene to Holocene (N. S t r e l n i k o v a , J. B a r r o n , pers. comm.). C o s m o p o l i t a n , p l a n k t o n i c c o l d - w a t e r species, w i d e s p r e a d . In surface sediments, m o r e common (15-25 % ) in the Guff of A l a s k a a n d at a b o u t 45 ° N, b e t w e e n 180 ° a n d 150 ° W ( S a n c e t t a , 1987). More c o m m o n in the P r a s s a s a n d A i t a n i a t h a n in the K a l l i t h e a a n d M a r a t h i t i s sections, w h e r e it is v e r y rare.

C o s e i n o d i s e u s o c u l u s - i r i d i s EHRENBERG (1841b) (P1. 1, fig. 3-5; P1. 4, fig. 12-14) 1841b. - Coscinodiscus ocuhts-iridis n. sp. - Ehrenberg, p. 147. 1854. - Coscinodiscus centralis EHR. -- Ehrenberg, pl. 21, fig. 3. 1930. - Coscinodiscus oculus-iridis EHR. - Hustedt, part 1, p. 454459, fig. 252. R e m a r k s : Small f u h o p o r t u l a e o c c u r at the u p p e r edge of the valve mantle (2 in 10 p m , P1. 4, fig. 1213) as well as small p o r e s , r e p r e s e n t i n g the e x t e r n a l

289

a p e r t u r a l openings of the r i m o p o r t u l a e on the valve face (P1. 4, fig. 14). The functioning of these organelles is not fully elucidated. A c c o r d i n g to Medlin et al. (1986) m o v e m e n t is r e p o r t e d for the first time in a centric diatom. T h e m o v e m e n t is d i r e c t i o n a l , a n d s t r o n g c i r c u m s t a n t i a l evidence f r o m h i s t o c h e m i s t r y a n d e l e c t r o n m i c r o s c o p y s u p p o r t s the p r o p o s a l t h a t motility is achieved by the p r o d u c t i o n of mucopolys a c c h a r i d e s t h r o u g h the p o r t u l a e . T h e y were called mucilage p o r e s (Gallertporen) b y w o r k e r s such as H u s t e d t (1930-1966) in R o u n d et al. (1990). I l l u s t r a t e d u n d e r this n a m e b y E h r e n b e r g (1854, pl. 19, fig. 2) f r o m a Miocene siliceous m a r l f r o m the G r e e k i s l a n d of Aegina in the Guff of S a r o n i k o s , Attica. B u t E h r e n b e r g ' s locality on Aegina was not given ( E h r e n b e r g , 1842). In a r e c e n t w o r k , F r y d a s (1996) s t u d i e d the thin d i a t o m i t e beds of the Aghios T h o m a s section on the i s l a n d of Aegina, which were p l a c e d in the silicoflagellate Mesocena circulus acmes u b z o n e , a n d in the NN14 (Discoaster asymmetricus) c a l c a r e o u s nannofossil zone of M a r t i n i (1971).

C o s c i n o d i s c u s r a d i a t u s EHRENBERG (1841b) (P1. 1, fig. 9 - 1 0 ; PI. 3, fig. 5) 1841b. - Coscinodiscus radiatus n. sp. Ehrenberg, p. 148, pl. 3, fig. la-c. 1986. - Coscinodiscus radiatus EHRENBERG.- Hasle and Sims, p. 310-312, fig. 8-32, 35-39. R e m a r k s : Eocene to H o l o c e n e . C o m m o n species with w o r l d w i d e d i s t r i b u t i o n in m o d e r n oceans. R a r e in M a r a t h i t i s a n d P r a s s a s sections, b u t m o r e common in the u p p e r p a r t of K a l l i t h e a section.

Genus D i m e r o g r a m m a

RALFS (1861)

D i m e r o g r a m m a m a r i n u m (GREGORY) RALFS in P r i t c h a r d (1861) (P1. 3, fig. 10-11) 1959. - Dimerograntma marinum ( G R E G O R Y ) R A L F S . -- Hustedt, part 2, p. 119, fig. 642. 1976. - Denticula marina G R E G O R Y , 1857. - Pankow, p. 138, fig. 275. R e m a r k s : F r e q u e n t in the M e d i t e r r a n e a n Sea ( H u s t e d t , 1959, p a r t 2), a n d in the coastal region f r o m N o r t h e r n E u r o p e ( P a n k o w , 1976). Benthic f o r m , Miocene to Recent, p o l y h a l o b e , cosmopolitan. R e c o r d e d in the Late Miocene M e r a d a section (NWCrete) b y Bellas a n d F r y d a s (1994, tab. 2). R a r e in the M a r a t h i t i s a n d P r a s s a s sections.

290

DIATOMI~ES PLIOCI~NES DE CRI~TE Diploneis oamaruensis (CLEVE) MILLS (1934) (Pl. 2, fig. 13)

Genus Diploneis EHRENBERG ex CLEVE (1894) Diploneis cf. advena (A. SCHMIDT) CLEVE (1894) (P1. 2, fig. 14) 1987. - Diploneis adveeta (A. SCHMIDT) CLEVE. -- Foged, p. 37, pl. 14, fig. 6. Remarks : Valve less constricted in the middle than by D. advena. Central nodule quadrate. Segments elliptical-oblong, angular at their ends. Length o f o b s e r v e d s p e c i m e n s : 6 0 t o 70 p m . L o n g i t u d i n a l canal broad. Transapical s t r i a e 1 0 - 1 2 i n 10 p m . Polyhalobe. Rare in the Marathitis and Prassas sections.

Diploneis b o m b u s EHRENBERG (1845) (P1. 2, fig. 15)

1934. - Diploeteis oamaretensis 1959. - Diploneis oamarueetsis p. 614, fig. 1027d.

--

Mills, p. 622. -- Hustedt, part 2,

MILLS.

Remarks : Valve deeply constricted in the middle. Central nodule quadrate. Longitudinal canal narrow, with inner poroid rows near to the horns, and outer poroid rows adjacent to the alveoles. Alveoles entire, with outer poroid rows homogeneous, and inner ones bigger near the margin. Valve 60-80 pm long, 12-20 p m b r o a d at the constriction, 5-8 transapical costae in 10 pm. Late Eocene of the Oamaru diatomite on the eastern coast of the South island in New Zealand. Extremely rare in the Marathitis, Prassas and Aitania sections.

1845. - Pietetuletria (Diploeteis) bombus n. sp. - Ehrenberg, p. 84. 1959. - Diploneis bombus EHRENBERG. -- Hustedt, part 2, p. 704, fig. 1086a-e.

R e m a r k s : Middle Miocene to Recent. Benthic species in modern coastal waters. Polyhalobe, cosmopolitan. Known in Greece from several modern localities at the islands of Samos, Kos and Kalymnos (Foged, 1985a,b), as well as from the Late Miocene Chairethiana Formation of NW-Crete (Merada section, Bellas and Frydas, 1994). Rare in our samples.

(CLEVE). (CLEVE)

Diploneis smithii (BRI~BISSON) CLEVE (1894) (P1. 2, fig. 16) 1853. - Navicula elliptican, sp. - W. Smith, t. 17, fig. 152/a. 1894. - Diploneis smithii (BRI~BISSON). -- Cleve, I, p. 96. 1959. - D i p l o n e i s smithii (BRI~B.) CLEVE. -- Hustedt, part 2, p. 647, fig. 1051.

R e m a r k s : Miocene to Holocene. Benthic form, polyhalobe, pleiceuryhahne, cosmopolitan. Very rare in the Marathitis section.

PLATE 4 SEM views, approximate magnification : fig. 1, 4, 6, 7, 10, : x 1,000 ; fig. 3, 5, 8, 12 : x 3,500 ; fig. 2, 9, 11, 13-15 : × 10,000. MEB, grossissemeetts approximatifs : fig. 1, 4, 6, 7, 10 : x 1 000 ; fig. 3, 5, 8, 12 : × 3 500 ; fig. 2, 9, 11, 13-15 : x 10 000. 11 : Detail of the valve centre with a distinct rounded papille. 1-2, 4-5. Actinocyclus octonarius EHRENBERG. Ddtail du centre de la valve avec uete papille arrondie Marathitis, sample M21(I); Prassas P15(2). bien distincte. 2, 5 : Detail of the inner mantle area showing mushroom-shaped, lipped rimoportulae of about 3 p m length. DEtail 12-14. Coscinodiscus oculus-iridis EHRENBERG. de la face interne du manteau, montraett des rimoporMarathitis, M18. tulae labi~s en f o r m e de champignon, d'une Iongeur d'h 12 : Internal view showing a symmetrical arrangement of fulpeu pros 3 ~tm. toportulae around the mantle and rimoportulae on the 3. Chaetoceros gastridiunt (EHRENBERG) BRIGHTWELL. valve face. Vue interne de la valve, montrant des fultoPrassas, P5. portulae symEtriquement arrangEes sur le bord du ntanteau et des rimoportulae dispersEes sur la f a c e valvaire. 6. Triceratium balearicum CLEVE and GRUNOW fa. biquetdrata 13 : Detail of the internal view of the mantle showing two (JANtSCn) HUSTEDT. fultoportulae. D~tail de la vue interne du bord du manPrassas, P14. teau, moettrant deux fultoportulae. 7-9. Actietocyclus octonarius EItRENBERG vat. teeteUa (BRI~BISSON) 14 : Detail of the valve face showing a rimoportula with a HENDEY. narrow slit. Ddtail de let Jhce valvaire montraett uete Marathitis, M21. rimoportula h fente Etroite. 8, 9 : Detail of the inner mantle region showing stalked curled rimoportulae with a significant long stalk. DEtail du bord 15. Actinoptychus splendens (SHADBOLDT) RALFS. interne du manteau, moettretnt des rimoportulete ~t long Marathitis, M23. pddoncule et h l~vres incurvdes. Arrows indicate two tubular fnhoportulae on the external valve mantle. Les fl~ches montrent deux fultoportulae tubu10-11. Azpeitia nodulifer (A. S C H M I D T ) FRYXELL and SIMS. laires sur le borcl exterete du manteau. Marathitis, M18.

FRYDAS

REVUE DIE MICROPALEONTOLOGIE, VOL. 42, N ° 4

PLATE 4

292

DIATOMI~ES PLIOCI~NES DE CRI~TE Genus E p i t h e m i a BRI~BISSON (1838) E p i t h e m i a sp. (P1. 2, fig. 29)

R e m a r k s : Concave c e n t r a l side a n d convex d o r s a l side. S u r f a c e of valve with t r a n s v e r s e costae. A c u r ved r a p h e is extending f r o m the ends to an a n g u l a r flexure n e a r the d o r s a l m a r g i n . W o r l d w i d e e p i p h y t i c in R e c e n t f r e s h - w a t e r e n v i r o n m e n t s . V e r y r a r e in the M a r a t h i t i s a n d A i t a n i a sections, c o n t r a r y to the Late P h o c e n e d i a t o m i t e deposits on w e s t e r n P e l o p o n n e s u s recently studied by Frydas and Economou-Amilli (1998), w h e r e the species E p i t h e m i a argus (EHRENBERG): K1JTZING a n d E. g o e p p e r t i a n a HILSE a r e the m a i n p a r t i c i p a n t s of the f r e s h - w a t e r d i a t o m flora.

Genus G r a l n m a t o p h o r a EHRENBERG (1841b) G r a m m a t o p h o r a a n g u l o s a EHRENBERG (1841b) (P1. 2, fig. 26-27; P1. 3, fig. 20) 1841b. - Grammatophora angulosa n. sp. - Ehrenberg, p. 153. 1959. - Grammatophora angulosa EHRENBERG. Hustedt, part 2, p. 39-40, fig. 564. R e m a r k s : Miocene to Recent. Benthic species, p o l y h a l o b e , m e i o e u r y h a l i n e , cosmopolitan. F o u n d in Recent sediments on the G r e e k islands Samos, Kos a n d K a l y m n o s (Foged, 1985a). R a r e in o u r m a t e r i a l . Grammatophora

m a r i n a (LYNGBYE) KUTZING (1844) (P1. 2, fig. 28)

1819. - Diatoma ntarinum n. sp. - Lyngbye, pl. 62A. 1844. - Grammatophora marina (LYNGBYE). -- Kfitzing, p. 128, pl. 17, fig. 24 (1 6) ; pl. 18, fig. 1(1-5). Remarks: polyhalobe, R a r e in the sediments of well as in the b ; 1986).

Miocene to Recent. Benthic species, meso-meioeuryhaline, cosmopolitan. M a r a t h i t i s section. F o u n d in R e c e n t Samos, K a l y m n o s a n d Kos islands as Gulf of Volos (Greece) b y F o g e d (1985a,

G r a m n m t o p h o r a o e e a n i c a (EHRENBERG) v a t , m a c i l e n t a (W. SMITH) GRUNOW in Van H e n r c k (1881) (P1. 3, fig. 18) 1886. - Grammatophora oceanica EHRENBERG v a t . subtilissima BAILEY. - Pantocsek, I, p. 38, tab. 16, fig. 112. 1959. - Grammatophora oceanica ( E H R E N B E R G ) GRUNOW vat'. macilertta (W. SMITH)GRUNOWf. subtilissima (BAILEY).-- Hustedt, part 2, p. 48, fig. 575. 1976. - Grammatophora oceanica ( E H I / . ) GRUNOWvar. ntacilettta ( W . S M I T I I ) G R U N O W ; -- Pankow, p. 120, fig. 204.

R e m a r k s : Cell n a r r o w , long-lancet s h a p e d with b l u n t r o u n d e d e n d s ; between m i d d l e a n d ends distinctly contricted. L e n g t h : 80-120 l~m, w i d t h : 810 p m . Miocene to Recent. Benthic species, neritic, p o l y h a l o b e , mesceuryhaline. S p o r a d i c a l l y o c c u r r i n g in t h e M a r a t h i t i s section.

Genus H e m i d i s c u s WALLICH (1860) H e m i d i s c u s c u n e i f o r m i s WALLICH (1860) (P1. 2, fig. 9) 1930. -Ilemidiscus cttneiformis WALL1CIk--Hustedl, part 1, p. 904, fig. 542. R e m a r k s : Middle Miocene to Pleistocene in the P a n a m a Basin, D S D P Sites 157, 158, ( B u k r y and F o s t e r , 1973). Some v a r i a n t s a r e s t r a t i g r a p h i c a l l y r e s t r i c t e d , like the large a n g u l a r forms of the Miocene D S D P Site 158 ( B u k r y a n d F o s t e r , 1973, pl. 14, fig. 11). P l a n k t i c , p o l y h a l o b e , w a r m - w a t e r species. C o m m o n in all investigated sections.

Genus N a v i c u l a BORY (1824) N a v i c u l a h e n n e d y i W. SMITH (1856) (P1. 2, fig. 17; P1. 3, fig. 7) 1856. - Navicula hennedyi n. sp. - W. Smith, t. 2, p. 93. 1964. - Navicula hennedyi W. SMITH. - Hustedt, part 3, p. 453455, fig. 1516. R e m a r k s : Miocene to Recent. Benthic species, p o l y h a l o b e , m e i o e u r y h a l i n e (salinity 35 to 20-17 %~, c o s m o p o l i t a n ( P a n k o w , 1976). R a r e in o u r m a t e r i a l .

Genus Nitzscllia HASSALL (1845) N i t z s c h i a j o u s e a e BURCKLE (1972) (P1. 2, fig. 41-42) 1972. - Nitzschia jouseae n. sp. - Burckle, p. 240, pl. 2, fig. 1721. 1978. - Nitzschia jottseae BURCKLE- Schrader and Gersonde, p. 157, pl. 5, fig. 1, 4-6. R e m a r k s : 9-11 striae in 10 p m a c c o r d i n g to B u r ckle (1972), 9-10 in o u r m a t e r i a l . It differs f r o m N. cylindrica BURCKLE in its t r a n s a p i c a l swelling a n d the g e n e r a l geometry of the valve. A c c o r d i n g to B a r r o n (1985a), the N. j o u s e a e zone is d e f i n e d as the i n t e r v a l f r o m the first o c c u r r e n c e of N. j o u s e a e to t h e first o c c u r r e n c e of Rhizosolenia p r a e b e r g o n i i MUCtllNA ( E a r l y Pliocene to e a r l y L a t e Pliocene). G e n e r a l l y in low n u m b e r s in the i n v e s t i g a t e d sections ( T a b l . 1-3).

FRYDAS Nitzschia m a r i n a GRUNOW in Cleve a n d G r u n o w (1880) 1880. - Nitzschia marina GRUNOW.- Cleve and Grunow, p. 70. 1973. - Nitzschia marina GRUNOW- Schrader, pl. 4, fig. 17-19. 1986. - Nitzschia marina GRUNOW- Akiba and Yanagisawa, p. 443, pl. 22, fig. 12. R e m a r k s : I n our m a t e r i a l only fragments were observed. Striae in the t r a n s a p i c a l axis very densely a r r a n g e d , a b o u t 13 to 15 in 10 pm. P l a n k t i c , w a r m water species, polyhalobe.

Rare to common in the Kallithea a n d Marathitis sections.

293

R e m a r k s : Valve is a wedge - shaped tube, which ends i n a long, thick spine. K n o w n from the Messin i a n deposits in C e n t r a l (Panassos I, II) a n d W e s t e r n Crete (Cherethiana) (Gersonde, 1980). P l a n k t i c , polyhalobe, m e i o e u r y h a l i n e , cosmopolitan species. I n o u r m a t e r i a l this species only occurs in the Prassas section.

R h i z o s o l e n i a sp. (?) (P1. 3, fig. 17) R e m a r k s : The long, thick spine is more or less p r e s e r v e d , from the illustrated R h i z o s o l e n i a sp. (?).

Genus S i m o n s e n i e l l a FENNER (1991) Genus P s e u d o d i m e r o g r a m m a SCHRADER (1976) S i m o n s e n i e l l a b a r b o i (BRUN) FENNER (1991) (P1. 2, fig. 34)

P s e u d o d i m e r o g r a m m a e l o n g a t u m SCHRADER (1976) (P1. 2, fig. 32) 1976. - Pseudodimerogramma elongata SCHRADER-- Schrader and Fenner, 13. 993, pl. 3, fig. 19-20. 1979. - Pseudodimerogramma elongatum S C H R A D E R . -- Abbott and Andews, p. 249, pl. 5, fig. 5. R e m a r k s : E a r l y to Middle Miocene from the Norwegian Sea ( S c h r a d e r a n d F e n n e r , 1976). Very r a r e in o u r material.

1991. - SimonsenieUa barboi (BRuN) Fenner, p. 108, pl. 3, fig. 1, 3. 1991. - Proboscia barboi (BRUN). - Jordan and Priddle, p. 56, fig. 1,2. R e m a r k s : Very r a r e in the Marathitis a n d Kallithea sections.

Genus S t e p h a n o g o n i a EHRENBERG (1845) S t e p h a n o g o n i a a c u l e a t a PANTOCSEK (1889) (P1. 2, fig. 38)

Genus R h a b d o n e m a KOTZING (1844)

1889. - Stephanogonia aculeata n. sp. - Pantocsek, II, p. 77, pl. 13, fig. 221.

R h a b d o n e m a a d r i a t i c u m K1JTZING (1844) (Pl. 2, fig. 33) 1844. - Rhabdonema adriaticum n. sp. - Kiitzing, p. 126, pl. 18, fig. 7. 1959. - Rhabdonema adriaticum KOTZING. - Hustedt, part 2, 13- 23, fig. 552. Remarks: Late Eocene diatomite e a r t h from O a m a r u , New Z e a l a n d (Desikachary a n d Sreelatha, 1989). Recent benthic form, epiphytic, polyhalobe, m e i o e u r y h a l i n e , cosmopolitan, with p r e f e r e n c e for w a r m - w a t e r e n v i r o n m e n t s ( P a n k o w , 1976). R a r e i n the Kallithea a n d Prassas sections, a n d r a r e to freq u e n t in the A i t a n i a section.

R e m a r k s : K n o w n from the Late Eocene of New Z e a l a n d (Desikachary a n d Sreelatha, 1989). Very r a r e i n the Marathitis a n d P r a s s a s sections.

Genus S t e p h a n o p y x i s EHRENBERG (1845) S t e p h a n o p y x i s c o r o n a (EHRENBERG) GRUNOW in V a n H e u r c k (1882) (PI. 2, fig. 2 2 ; P1. 3, fig. 8) 1854. - Systephania corona n. sp. - Ehrenberg, pl. 33, 15.22 ; 131. 17.16. 1882. - Stephanopyxis c o r o n a ( E H R E N B E R G ) G R U N O W . - Van Heurck, pl. 83c, fig. 10, 11. 1968. - Stephanopyxis c o r o n a ( E H R E N B E R G ) Grunow. - Hajos, p. 87-88, pl. 11, fig. 2, 4.

Genus R h i z o s o l e n i a BRIGHTWELL (1858) R h i z o s o l e n i a h e b e t a t a fa. s e m i s p i n a (HENSEN) GRAN (1908) (P1. 2, fig. 35) 1959. - Rhizosolenia hebetata fa.semispina ( H E N S E N ) Hustedt, part 2, p. 592, fig. 338, p. 575, fig. 327.

GRAN.

--

Remarks: Valve d i a m e t e r : 15-20 llm, h e i g h t : 15 l~m. L e n g t h of the o u t e r spines : 5 to 7 pm. Areolae hexagonal, a r r a n g e d in diagonal crossing rows, 5-7 in 10 1Jm. Marine fossil form. Middle Miocene of M a r y l a n d , a n d Ptiocene of California (Lohm a n , 1938, 1948). Very r a r e in the Marathitis section.

294

DIATOMI~ES PLIOCI~NES DE CRI~TE

S t e p h a n o p y x i s t u r r i s (GREVILLE a n d ARNOTT) RALFS in P r i t c h a r d (1861) (P1. 2, fig. 2 1 ; P1. 3, fig. 9) 1857. - Creswellia turris GREVILLE. - Gregory, p. 538, pl. 14, fig. 109. 1861. - Stephanopyxis turris ( G R E V I L L E and A R N O T T ) R A L F S . -Pritchard, p. 826, pl. 5, fig. 74. 1930. - Stephanopyxis turris ( G R E V I L L E and A R N O T T ) R A L F S . Hustedt, part 1, p. 304, fig, 140. R e m a r k s : A long r a n g i n g m a r i n e , p l a n k t i c spec i e s : C r e t a c e o u s to Recent. M e r o p l a n k t o n i c (neritic), p o l y h a l o b e , oligc~uryhaline (salinity 35-30 %~, c o s m o p o l i t a n , with p r e f e r e n c e for s u b t r o p i c a l to temp e r a t e regions ( P a n k o w , 1976). F r e q u e n t in all stud i e d assemblages.

Genus S t i e t o d i s c u s GREVILLE (1865)

S t i c t o d i s c u s p a r a l l e l u s (EHRENBERG) v a t . p a r a l l e l u s fa. q u a d r a t a TEMPERE a n d PERAGALLO (1914) (PI. 2, fig. 23) 1989. - Stictodiscus parallelus (ErIR.) vat. parallelus fa. quadrata TEMPERE and PERAGALLO.- Desikachary and Sreclatha, p. 240, pl. 103, fig. 2. R e m a r k s : Valve s u b q u a d r a n g u l a r . Sides slightly convex. Valve surface a r e o l a t e , a r e o l a e r o u n d e d . C e n t r a l a r e a wide, with only a few s c a t t e r e d a r e o l a e . N e a r the m a r g i n , a r e o l a e a r r a n g e d in s h o r t p a r a l l e l rows. 4-5 a r e o l a e in 10 pm. K n o w n f r o m the Late Eocene d i a t o m i t e - e a r t h f r o m O a m a r u , New Z e a l a n d ( D e s i k a c h a r y a n d S r e e l a t h a , 1989). V e r y r a r e in the M a r a t h i t i s section.

Genus S u r i r e l l a TURPIN (1828)

Genus S y n e d r a EHRENBERG (1830) S y n e d r a fulgens (GREVILLE) W. SMITH (1856) (P1. 2, fig. 30) 1959. - Synedra fulgens 2, p. 228, fig. 717a.

(GREVILLE)

W. SMITH. -- Hustedt, part

Remarks: Common species along the coasts of E u r o p e , a n d in the C a r i b b e a n (Foged, 1986). Benthic f o r m , p o l y h a l o b e to mesohalobe. V e r y r a r e in the M a r a t h i t i s section.

S y n e d r a t a b u l a t a (AGARDIt) K~TZING (1844) (P1. 2, fig. 20) 1832. - Diatoma tabulatum n. sp. - Agardh, p. 50. 1844. - Synedra tabulata ( A G A R D H ) . - Kfitzing, p. 68, pl. 15, fig. 10 (1-3). 1959. - Synedra tabulata (AGARDH)K~TZING. - Hustedt, part 2, p. 218, fig. 710a-d. R e m a r k s : Valve small, l a n c e t - s h a p e d with thin b i l l - s h a p e d ends a n d slightly r o u n d e d poles. Length : 25-200 pro, w i d t h : 3-5 p m , 10-15 s t r i a e in 10 p m continuous in the middle. P s e u d o r a p h e wide, lancets h a p e d . Miocene to Recent. Benthic f o r m , mesohalobe, e u r y h a l i n e , c o s m o p o l i t a n ( P a n k o w , 1976). R a r e in the M a r a t h i t i s a n d P r a s s i a s sections.

S y n e d r a u n d u l a t a (BAILEY) GREGORY (1857) (P1. 2, fig. 3 7 ; P1. 3, fig. 19) 1959. - Synedra undulata ( B A I L E Y ) G R E G O R Y . -- Hustedt, part 2, p. 224, fig. 714. 1976. - Toxarium undulatum B A I L E Y , 1853. - Pankow, p. 134, fig. 256-257. R e m a r k s : Miocene to Recent. Benthic f o r m , pol y h a l o b e , m e i o e u r y h a l i n e , c o s m o p o l i t a n with pref e r e n c e for w a r m - w a t e r e n v i r o n m e n t s ( P a n k o w , 1976). R a r e in the P r a s s a s section.

Genus T h a l a s s i o n e m a GRUNOW (1881)

S u r i r e l l a o v a t a K1JTZING (1844) (P1. 2, fig. 24)

T h a l a s s i o n e m a n i t z s c h i o i d e s (GRUNOW) HUSTEDT (1959)

1844. - Surirella ovata n. sp. - Kfitzing, p. 62, pl. 7, fig. 1-4. 1968. - Surirella ovata KOTZING. - H a j o s , p. 205-206, pl. 60, fig. 4.

1862. - Synedra nitzschioides n. sp. - Grunow, p. 403. 1881. - Thalassiothrix nitzschioides (GRUNOW). -- Van Heurck, pl. 43, fig. 7, 10. 1959. - Thalassionema nitzschioides (GRt/NOW). -- Hustedt, part 2, p. 244-246, fig. 725.

R e m a r k s : Miocene to Recent. Oligohalobe (indifferent) mesceuryhahne, a l k a l i p h i l e , cosmopolitan. O b s e r v e d in the R e c e n t sediments of the G r e e k islands Samos, Kos a n d K a l y m n o s (Foged, 1985a,b). R a r e in o u r m a t e r i a l .

R e m a r k s : L a t e Miocene to Recent. P l a n k t i c , neritic species, p o l y h a l o b e , meio- to mesceuryhaline, cosmopolitan. R a r e in the M a r a t h i t i s a n d K a l l i t h e a sections.

FRYDAS G e n u s T h a l a s s i o s i r a CLEVE (1873) T h a l a s s i o s i r a c o n v e x a MUCHINA (1965) (PI.1, fig. 2a, b) 1965. - Thalassiosira convexa n. sp. - Muchina, p. 22, pl. 11, fig. 1-2. 1968. - Thalassiosira undulosa (MANN)SIIESIIUKOVA-PORETZKAYA. - Koizumi, pl. 35, fig. 26a-b. Remarks : O c c u r s i n the N o r t h Pacific D i a t o m (NPD) Zones 6 to 11, Leg 18, Site 173, N o r t h e a s t Pacific, t o g e t h e r with N i t z s c h i a j o u s e a e ( E a r l y Pliocene to e a r l y L a t e P l i o c e n e ) ( S c h r a d e r , 1973). F r e q u e n t to r a r e i n the i n v e s t i g a t e d sections.

T h a l a s s i o s i r a e c c e n t r i e a (EHRENBERG) CLEVE (1904) (P1. 2, fig. 6) 1845. - Odontodiscus excentricus n. sp. - Ehrenberg, p. 79. 1854. Coscinodiscus excentricus Eltl~. - Ehrenberg, pl. 18, fig. 32 ; pl. 21, fig. 6. 1972. - Thalassiosira eccentrica ( E I I R E N B E R G ) C L E V E . -- Fryxell and Hasle, p. 300-312, fig. 1-18. R e m a r k s : L a t e Miocene to R e c e n t . O n e of the most c o m m o n d i a t o m s i n m o d e r n n e r i t i c p l a n k t o n . P o l y h a l o b e , m e i o - to meso~uryhaline, c o s m o p o l i t a n ( P a n k o w , 1976). O b s e r v e d i n the lower P l e i s t o c e n e d i a t o m i t e s f r o m A d a m a s section, Milos I s l a n d , Aegean Sea ( F r y d a s , 1994a). F r e q u e n t i n the P r a s s a s section, while r a r e i n the K a l l i t h e a , M a r a t h i t i s a n d A i t a n i a sections.

T h a l a s s i o s i r a l e p t o p u s (GRUNOW) HASLE a n d FRYXELL (1977) (P1. 1, fig. 6 - 8 ; P1. 3, f i g . l - 3 b ) 1840. - Coscinodiscus lineatus n. sp. - Ehrenberg, p. 129. 1854. - Coscinodiscus lineatus EIIR. - Ehrenberg, pl. 18, fig. 33 ; pl. 22, fig. 6a, b ; pl. 35, A.16, fig. 3,17. 1833. - Coscinodiscus lineatus var. leptopus GRUNOW. -- Van Heurck, Synopsis, pl. 131, fig. 5, 6. 1977. - Thalassiosira leptopus (GRUNOW). -- Hasle and Fryxell, p. 20-22, fig. 1-14, 94-96. Remarks : Small rounded rimoportulae with an apertural opening, symmetrically arranged on the i n n e r valve m a n t l e (P1. 3, fig. 3 a , b ) as well as s m a l l s p i n u l e s o n t h e m a r g i n (P1. 3, fig. 2a, b). M i o c e n e to H o l o c e n e . C o m m o n p l a n k t o n i c species i n m o d e r n w a r m - w a t e r o c e a n s . I n the K a l l i t h e a a n d M a r a t h i t i s sections, c o m m o n to a b u n d a n t , while t o t a l l y m i s s i n g i n the P r a s s a s a n d A i t a n i a sections. P r o b a b l y m a r k e r for s e a s o n a l c h a n g e s f r o m w a r m to t e m p e r a t e w a t e r c o n d i t i o n s . Also f o u n d i n g r e a t n u m b e r s i n the A s t e r i section, W e s t e r n C r e t e , R e t h y m n o n p r o v i n c e ( F r y das, 1998).

295 T h a l a s s i o s i r a o e s t r u p i i (OSTENFELD) PROSCHKINA-LAVRENKO (1956) (P1. 1, fig. 17)

1900. - Coscinosira oestrupii n. sp. - Ostenfeld, p. 52. 1956. - Thalassiosira oestruppi ( O S T E N F E L D ) . -- Proschkina-Lavrenko, p. 57. Remarks: E a r l y P l i o c e n e to R e c e n t . T h e first a p p e a r a n c e of T. o e s t r u p i i is n o t e d b y S c h r a d e r (1973) as E a r l y P l i o c e n e i n his N P D Z o n e 9. Also B a r t o n (1976) r e p o r t e d t h e first o c c u r r e n c e of T. o e s t r u p i i i n U n i t 18 of the B o l i v i n a o b l i q u a Z o n e i n C a l i f o r n i a of p r o b a b l e E a r l y P l i o c e n e age. Cosmop o l i t a n , p o l y h a l o b e m a r i n e species c o m m o n i n R e c e n t s u b p o l a r to t r o p i c a l e n v i r o n m e n t s . R e c o r d e d f r o m G u l f of Mexico, t h e A t l a n t i c , P a c i f i c a n d I n d i a n O c e a n s (Foged, 1986). F r e q u e n t to c o m m o n i n the i n v e s t i g a t e d sections.

G e n u s T h a l a s s i o t h r i x CLEVE a n d GRUNOW (1880)

T h a l a s s i o t h r i x cf. l o n g i s s i m a CLEVE a n d GRUNOW (1880) (P1. 2, fig. 3 6 ; P1. 3, fig. 13) 1873. - Synedra thalassiothrix n. sp. - Cleve, p. 22, pl. 4, fig. 24. 1880. - Thalassiothrix longissima n. sp. - Cleve and Grunow, p. 108. 1959. - Thalassiothrix longissima CLEVEand GRUNOW.- Hustedt, part 2, p. 247, fig. 726. R e m a r k s : M i o c e n e to R e c e n t . O c c u r s i n m o d e r n o c e a n s , especially a b u n d a n t i n cold w a t e r s . O n l y a few f r a g m e n t s were o b s e r v e d i n t h e K a l l i t h e a , Mar a t h i t i s a n d P r a s s a s sections.

G e n u s T r a c h y n e i s CLEVE (1894)

T r a e h y n e i s a s p e r a (EHRENBERG) CLEVE (1894) (P1. 2, fig. 31) 1841a. - Pinnularia aspera n. sp. - Ehrenberg, p. 213. 1854. - Navicula aspera ( E I I R E N B E R G ) . -- Ehrenberg, pl. 35, A. fig. 5; pl. 17, fig. 26. 1894. - Trachyneis aspera ( E H R E N B E R G ) . -- Cleve, I, p. 191. R e m a r k s : M i o c e n e to R e c e n t . B e n t h i c f o r m , polyhalobe, meioeuryhaline, cosmopolitan (Pankow, 1976). V e r y r a r e i n the M a r a t h i t i s section.

296

DIATOMt~ES PLIOCI~NES DE CRI~TE Genus Triceratium

EHRENBERG (1839)

T r i c e r a t i u m b a l e a r i c u m CLEVE a n d GRUNOW fa. b i q u a d r a t a (JANISCH) HUSTEDT (1930) (P1. 2, fig. 1 8 ; P1. 3, fig. 2 1 ; P1. 4, fig. 6) 1886. - Triceratium biquadratum JANISCH. -- Schmidt : Atlas, pl. 98, fig. 4-6. 1930. - Triceratium balearicum CLEVE and GRUNOWfa. biquadrata (JANISCII). -- Hustedt, t. 1, p. 815, fig. 477. R e m a r k s : Miocene to Recent. B e n t h i c f o r m , p o l y h a l o b e , l i t t o r a l , w i t h p r e f e r e n c e f o r w a r m - w a t e r env i r o n m e n t s . A l w a y s r a r e in t h e i n v e s t i g a t e d sections.

T r i g o n i u m a r e t i e u m v a t . p e n t a g o n a l i s (TEMPERE a n d PERAGALLO) DESIKACHARY a n d SREELATHA (1989) (P1. 2, fig. 5) 1910. - Triceratium arcticum vat. pentagonalis. - Tempere and Peragallo, p. 210, n ° 393. 1989. - Trigonium arcticum vat. pentagonalis ( T E M P E R E and PERAGALLO) n o v . c o m b . - Desikachary and Sreelatha, p. 272, ph 114, fig. 7; ph 125, fig. 8. R e m a r k s : E o c e n e to R e c e n t . B e n t h i c f o r m , p o l y halobe, littoral, with preference for cool-water envir o n m e n t s . O n l y o b s e r v e d , v e r y r a r e , in t h e s a m p l e P15 of t h e P r a s s a s s e c t i o n .

AKNOWLEDGEMENTS I a m g r a t e f u l to t h e M i n i s t r y f o r D e v e l o p m e n t , G e n e r a l S e c r e t a r y of R e s e a r c h a n d T e c h n o l o g y f o r financial support during my field work. I am deeply i n d e b t e d to M r s . D r . E. F o u r t a n i e r ( C a l i f o r n i a A c a d e m y of S c i e n c e s ) , Mr. D r . J . A . B a r r o n ( U . S . G e o l o g i c a l S u r v e y ) a n d Mrs. D r . M . - C . J a n i n (CNRS, Centre IBT, Troyes) for their many helpful c o m m e n t s a n d v e r y c r i t i c a l r e a d i n g of t h e m a n u s c r i p t . I also t h a n k P r o f . D r . H . K e u p p ( I n s t i t u t e of P a l e o n t o l o g y , F r e e U n i v e r s i t y of B e r l i n ) f o r his assistance concerning the SEM imaging under a common research program and the German Academic E x c h a n g e S e r v i c e ( D A A D ) f o r s u p p o r t i n g m y two m o n t h s r e s e a r c h at t h e F . U . of B e r l i n ( J u n e - J u l y , 1998). T h a n k s a r e d u e to M r s . D r . J. F e n n e r (Bundesanstalt fiir Geowissenschaften und Rohstoffe, H a n n o v e r ) f o r h e r c o m m e n t s on t h e i n i t i a l m a n u s c r i p t . I t h a n k also M r s . P r o f . U. G e i s s l e r (Inst. of S y s t e m a t i c B o t a n y , F . U . B e r l i n ) , M r s . D r . P . A . Sims (Natural History Museum, London) for their inform a t i o n s a b o u t t h e b e h a v i o u r of m o d e r n d i a t o m s , a n d D r . M r . P . A v r a m i d i s ( U n i v e r s i t y of P a t r a s ) f o r t h e presentation of the manuscript. Additional SEM a s s i s t a n c e was p r o v i d e d by M r s . U. H e y e r (Inst. of Paleontology, F.U. Berlin).

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DE CRI~TE

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