New Pliocene and Pleistocene calcareous dinoflagellate cysts from southern Italy and Crete

Review of Palaeobotany and Palynology, 78 (1993): 353-380 Elsevier SciencePublishers B.V., Amsterdam

353

New Pliocene and Pleistocene calcareous dinoflagellate cysts from southern Italy and Crete G e r a r d J.M. Versteegh Laboratory of Palaeobotany and Palynology, University of Utrecht, Heidelberglaan 2, 3584 CS Utrecht, The Netherlands (Received August 8, 1992; revised and accepted December 17, 1992)

ABSTRACT Versteegh, G.J.M., 1993. New Pliocene and Pleistocenecalcareous dinoflagellate cysts from southern Italy and Crete. Rev. Palaeobot. Palynol., 78: 353-380. Samples from the south-Italian Singa section (early Pliocene-early Pleistocene)and from a piston core from an offshore location southwest of Crete (late Pleistocene)have been investigated for their calcareous dinoflagellate cyst content. The assemblages include four new genera, eight new species and one new variety: Bicarinellum tricarinelloides sp. nov., Calciperidinium asymmetricum gen. nov., sp. nov., Calcicarpinum bivalvum sp. nov., C. perfectum sp. nov., Follisdinellum splendidum gen. nov., sp. nov., Melodomuncula berlinensis gen. nov., sp. nov., Pseudopithonella striatula gen. nov., sp. nov., Praecalcigonellum schizosaeptum sp. nov. and Sphaerodinella albatrosiana var. spinosa var. nov. The genus Calcicarpinum Deflandre, 1948, is emended.

Introduction

Calcareous dinoflagellate cysts are qualitatively and quantitatively, significant contributors to the microfossil content of marine sediments from Jurassic times onwards (e.g. Bolli, 1974; Keupp, 1980a, 1981; Dale, 1986). Presently, calcareous dinoflagellates are found in a variety of marine environments and their cysts are therefore a potential source o f (paleo)environmental information (e.g. Wall and Dale, 1968a; Gilbert and Clark, 1983; Dale, 1986; Akselman and Keupp, 1990). Most information on fossil representatives of this group deals with the morphology and taxonomy of Cretaceous material (e.g. Bolli, 1974, 1978; Keupp, 1978; Willems, 1985, 1988; Keupp and Ilg, 1989). There is a general lack of data on Neogene and Quaternary calcareous dinoflagellate cysts. Miocene cysts were described by Deflandre (1947, 1948). Published information concerning Pliocene and Pleistocene material is limited to the work of Ffitterer (1976, 1977a, 1984), Gilbert and Clark (1983) and Keupp and Versteegh (1989), whereas 0034-6667/93/$06.00

(sub)recent data are available from e.g. Wall and Dale (1968a), Wall et al. (1970, 1973), Dale (1977, 1986), Tangen et al. (1982), Akselman and Keupp (1990), and Binder and Anderson (1990). By describing the species observed in samples from southern Italy and offshore Crete, the present study is aimed at a first qualitative inventory of the calcareous dinoflagellate cysts in the Pliocene and Pleistocene of the Mediterranean realm. This inventory may provide the background for subsequent analysis of the paleoenvironmental signal of Pliocene-Pleistocene calcareous dinoflagellate cysts. Material and age assessment

Initially, samples were taken from different lithologies and ages in order to see to what extent the destribution of calcareous dinoflagellate cysts depends on these parameters. However, the amount of new taxa made it necessary to do a purely taxonomical study first. Therefore, no attention has been paid to palaeoenvironment. Samples

© 1993-- ElsevierSciencePublishers B.V. All rights reserved.

354

G.J.M. VERSTEEGH

have been studied from two localities (Fig. 1): (1) the Pliocene to lowermost Pleistocene Singa section in southern Italy, and (2) an uppermost Pleistocene piston core T 87/2/20G taken offshore, south of Crete.

¢

F

l

90 2

i'I

~

.,1 uJ~

b

W

~i

>.

8, O .J

!

The Singa section

The Singa composite section is named after the mountain "Monte Singa" in southern Calabria. The upper part of the section, from which the samples have been taken, has a total thickness of 149 m. It is composed of a homogeneous clay sequence in which some sets of organic-rich layers are intercalated (Fig. 2). For a detailed description of the section reference is made to Hilgen (1987). The Singa section has been extensively studied with respect to lithology (Hilgen, 1987), paleomagnetism (Zijderveld et al., 1986), calcareous nannofossils (Driever, 1988), clay minerals (de Visser, 1991), carbon and oxygen stable isotopes (Gudjonsson 1987; Lourens et al., 1992), planktonic foraminifera (Zachariasse et al., 1990; Lourens et al., 1992), and organic dinoflagellates (Zevenboom and Versteegh, in prep; Versteegh, submitted). The section has been correlated with the astronomical time scale of Berger and Loutre (1991) by Hilgen (1991a,b). The samples from this section that have been examined for their calcareous dinoflagellate cyst content are listed in Table I.

2.1 -

~ "='~ ~

ST

' ~B 5

.5-! ~

, B~

2.2--

B2

1.6

i

2.3

I

' B1 1.7--

2.,1

1,8 n i

2.5 -! ]

C1 CO

J

2.7

2.6~

1.9 --i

!

A1

BB

SAPROPEL

~

CLAY

m

SAMPLE POSITION

Fig. 2. The upper part of the Singa section. Lithology and sapropel codation according to Hilgen (1991). Age calibration according to Lourens et al. (1991). Sample locations are indicated by an asterix. ~7

The T 87/2/20G piston core )~ &V-

-' ~. '.P. : . .~. ( '/-

~

o6 /

B Fig. 1. Sample locations. A =Singa Section, Italy; B = C o r e T87/2/20G, Crete.

The T 87/2/20G piston core was recovered in 1987 from the southern Crete continental margin at 34°.58 ', 4 N, 23°.44 ', 8 E at 707 m waterdepth during cruise Tyro 1987-2 Crete Perimeter of the Dutch research vessel Tyro. It consists of a greybrown homogenous marl sequence in which four grey-green layers, rich in organic matter ("sapro-

NEW PIOCENE AND PLEISTOCENE CALCAREOUS DINOFLAGELLATE CYSTS FROM SOUTHERN ITALY AND CRETE

355

TABLE I Samples from the Singa section Sample

9652 9651 9453 9452 9451

Depth (m from base)

Lithology

137.45 137.30 89.45 88.95 88.45

sapropelic (CI 1) non-sapropelic non-sapropelic non-sapropelic sapropelic (B7)

Age (x lOs a)

~

+ 1.60 + 1.60 + 2.08 _ 2.09 +2.09

<

I.-J

/

$1

~ /

s3

50--

Data source: Lourens et al., 1991.

100-pels"), are intercalated (Fig. 3). From top to bottom these organic rich layers correlate with the first, third, fourth and fifth (respectively S1, $3-$5) sapropelic layers in the Eastern Mediterranean basin (Rohling and Gieskens, 1989). The section has been described in detail by Rohling and Gieskens (1989) who also studied the section on planktonic foraminifera. Furthermore, the core has been investigated on its organic dinofiagellate cyst content (Zonneveld and Versteegh, in prep). Samples, which have been examined for calcareous dinoflagellates cysts, are listed in Table II.

150--

200-m

250-85

~t

Methods

Approximately 1.5 g of each sample was washed over a 20 Ixm (samples 9452, 9453, 9652) or a 10 ~tm (other samples) mesh sieve and then dried for one night at 60°C. Ultrasonic treatment was used to speed up the sieving procedure. The residual fraction was dried and specimens of each calcareous dinofiagellate cyst morphotype were isolated using a binocular (x 200) and a needle. Subsequently, the selected specimens were placed on polished scanning electron microscope (SEM) stubs, or were stuck on SEM stubs covered with double-sided gummed tape. The stubs were then coated with gold-palladium for 240 s. These were examined in a Cambridge-360 Stereoscan. Some specimens were broken and coated again to show the structure of both the inner and outer wall. The terminology of Keupp (1981, 1987) was used to describe the calcareous dinoflagellate cysts.

$4

BB

SAPROPEL

(SS]

StLTYCLAY

m

SAMPLEPOSITION

Fig. 3. Core T87/2/20G. Lithology adapted from Rohling and Gieskens (1987). Sample locations are indicated by an asterix.

TABLE II Samples from the core T/87/2/20G Samples

Depth (cm from top

Lithology

Age ( x 103 a)

$2/87-88 S1/80-81

170-171 264-265

sapropelic ($3) sapropelic ($5)

+76 __+122

Data source: Rohling and Gieskens, 1989.

356

PLATE I

G.J.M. VERSTEEGH

357

NEW PIOCENE AND PLEISTOCENE CALCAREOUS DINOFLAGELLATE CYSTS FROM SOUTHERN ITALY AND CRETE

Taxonomy A considerable number of previously undescribed taxa were found. These are formally described in the following paragraphs. The taxonomic concept used here for the family level and higher levels, is based on both dinoflagellate cysts and thecae (see Bujak et Davies, 1983). For taxonomic levels lower than families, the taxonomic system o f Keupp and Versteegh (1989) is used. This system is based on fossil calcareous dinoflagellate cysts exclusively. The prefix "para", which is in use to indicate morphological features on cysts that are considered homologous to morphological features on thecae and that have otherwise the same name for cysts and thecae, has not been used because it is clear that this study only deals with fossil (cyst) material. The prefix " p a r a " therefore adds no substantial information. The obliquipithonelloid forms (Bicarinellum tricarinelloides sp. nov., Calciperidinium asymmetricum gen. nov, sp. nov., Calcicarpinum bivalvum sp. nov., C. perfectum sp. nov., and Follisdinellum splendidum gen. nov., sp. nov.) have already been figured earlier by Wall and Dale (1968a,b), Wall et al. (1973) and Fiitterer (1977b). Pseudopithonella striatula gen. nov., sp. nov. and Melodomuncula berlinensis gen. nov., sp. nov., have not been reported before. Information on orthopithonelloid cysts obtained

during the present study already led to a revision of the subfamily Orthopithonelloidea (Keupp and Versteegh, 1989). The only new orthopithonelloid form described in this study is Praecalcigonellum schizosaeptum sp. nov.

Description of taxa Division PYRROPHYTAPascher, 1914 Class DINOPHYCEAEFritsch, 1929 Order PL~rDI~_ALESHaeckel, 1894 Suborder PERIDINIrN'EAE (Fott, 1959) Bujak et Davies, 1983 Family CALCIODINELLACEAE(Fott, 1959) Bujak et Davies, 1983 Subfamily OBLIQUIPITHONELLOIDEAEKeupp, 1987

Genus BiearineUum Deflandre, Keupp, 1984

1948

Bicarinellum triearinelloides sp. nov. (Plate I, 1-12; Fig. 4) 1977b Incertae sedis (Rhodophyceae?) Fiitterer, p. 63, 68, pl. 24, fig. 1-6

Derivation of name: Latin, tri, three; carina, keel.

Bicarinellum tricarinelloidessp. nov. (All photographs x I000, except where otherwise stated.) I.

emend.

Type species: Bicarinellum castaninum Deflandre,

PLATEI

2,3. 2. 3. 4, 5. 4. 5. 6. 7, 10, 11. 7. 10. 11. 8. 9, 12. 9. 12.

1948

Cyst T20-S1/80o81/8;lateral view (negative 250). Cyst T20-S1/80-81/20;paratyp¢. Antapical view, sulcus (below), antapical plates (above) (negative66). Detail of cyst wall (negative 67), x 5000. Cyst T20-$1/80-81/14;holotyp¢. Apical view (negative 77), x 750. Ventro-lateral view (negative 75), x 750. Cyst T20-$1/80-81/7;ventral view (negative 23). Cyst T20-S1/80-81/40;note the intratabular areas overgrownby ridges, the opcrculumremains free. Apical-lateral view (negative 131), x 750. Antapical-lateral view (negative 129), x 750. Detail of cyst wall (negative 130), x 5000. Cyst 9651/39; note the intratabular areas partly overgrownby ridges (negative 37). Cyst 9651/40; open cyst. Apical-lateral view (negative 39). Detail of archaeopylemargin (negative40), x 10,000.

358 PLATE I|

G.J,M VERSTEEGH

NEW PIOCENE AND PLEISTOCENE CALCAREOUS DINOFLAGELLATE CYSTS FROM SOUTHERN ITALY AND CRETE

A

B

C

359

D

Fig. 4. Schematic diagram of Bicarinellum tricarinelloides sp. nov. (A) Apical view. (B) Antapical view. (C) Ventral view. (D) Dorsal view. For the pronounced antapical plate margins resemble a third keel.

Holotype: Cyst no. T20-S1/80-81/14, SEM micrographs 74-77 (Plate I, 4, 5). Paratype: Cyst no. T20-S1/80-81/20, SEM micrographs 66-68 (Plate I, 2, 3).

Type locality: Piston core T87-2-20G. Southern continental margin of Crete, 340.58 ', 4 N, 230.44 ', 8E.

Type stratum: Upper Pleistocene, sapropel $5, _ 1 2 x 104a.

Repository: Palynology, Netherlands.

Laboratory of Palaeobotany and University of Utrecht, The

Diagnosis: A species o f Bicarinellum Deflandre, 1948, which is characterised by broad ridges and a well-developed polar horn or, in case the horn is missing, a circular opening instead. From the polar horn to the other side o f the cyst two parallel

(sub)circular transversal ridges are present on the upper half of the cyst. Ridges on the lower half of the cyst completely reflect the plate boundaries of the antapical plates. Two ridges connect the ventral corners of reflected antapical plate margins with the lowermost (sub)circular ring on the upper half of the cyst. Sometimes additional ridges reflect details of tabulation.

Description: Calcareous dinoflagellate cysts having a subspherical endocoel, a subspherical to elongate shape and a single layered wall composed of numerous randomly orientated calcite needles. On the apical side the circular archaeopyle possesses a well developed central protrusion. On the antapical side both antapical plates are completely reflected by broad sutural ridges. Broad ridges elsewhere on the cyst, are on the basis of the reflected antapical tabulation considered to be sutural as well. On the epicyst these ridges form two transversal and parallel (sub)circular closed rings. Two straight ridges connect the ventral corners of the antapical plate sutures with the lowermost epicystal ring. For their position on the cyst these ridges may reflect the margins of the

PLATE II

Calciperidinium asymmetricurngen. nov., sp. nov. (All photographs x 1500, except where otherwise stated.) 1-3. Cyst T20-S1/80-81/106;holotype. 1. Dorsalview (negative 322). 2. Apicalview (negative 317). 3. Detailof dorsal apex (negative 320), x 3000. 4, 7. Cyst T20-S1/80-81/101. 4. Lateralview (negative 273). 7. Detailcyst wall, note the (due to distal expansion) plate-like crystal ends (negative 272). 5. CystT20-S1/80-81/84;closed cyst, dorso-lateral view (negative 262), x 10000. 6, 8. Cyst T20-S1/80-81/107. 6. Ventralview (negative 23). 8. Detailcyst wall, crystal ends lacking distal expansion (negative 311), × 10,000.

360

lower part of the sulcal area. A small intratabular protrusion may be present on the antapical plates. Occasionally, and mostly on the hypocyst, tabulation is indicated by additional sutural ridges. On individual specimens the ridges may vary in width due to the varying amount of overgrowth over the intratabular areas. On specimens with very broad sutural ridges the tabulation is mostly obscure. The calcite crystals on the sides of the ridges and on the intratabular areas are more loosely arranged than on the crests of ridges where a dense tapetum is formed.

G.J.M. VERSTEEGH

Diagnosis: Obliquipithonelloid dinoflagellate cysts with a subspherical endocoel and a peridinioid shape: one apical and two antapical protrusions. The archaeopyle is positioned below the apex of the cysts. It is probably formed by the loss of one or more intercalary plates. Remarks: This genus differs from other genera in having a single apical and two antapical protrusions. Up to now, the type species is the only species known from this genus. Calciperidinium asymmetricum sp. nov. (Plate II,

Dimensions: Holotype: length 52 ~n, width 53 lam. Other specimens: length 40-59 lam, width 34-64 I.tm, n = 13. Occurrence: Rare in the upper Pliocene of the Singa section, common in core T87-2-20G, southern continental margin of Crete, upper Pleistocene. According to FiJtterer (1977b) this species is also present in the Recent sediments of the Persian Gulf and the Adriatic Sea. Comparison: B. tricarinelloides differs from all previously described Bicarinellum species by the welldeveloped broad ridges and the loose arrangement of the calcite crystals on the intratabular areas. The "Bicarinate-type" as described by Wall and Dale (1968a) closely resembles B. tricarinelloides, but differs by lacking an apical protrusion.

Genus Caleiperidinium gen. nov. Type species: Calciperidinium asymmetricum sp. nov.

Holotype of type species: Calciperidinium asymmetricum (Plate II, 1-3). Derivation of name: with reference to the peridinioid shape. Repository: Laboratory of Palaeobotany and Palynology, University of Utrecht, The Netherlands.

1-8) 1973 "Pyriform-type" Wall et al., pp. 25-26, pl. 1, fig. 22

Derivation of name: Greek, a-symmetros, notfitting. With reference to the position of the endocoel. Holotype: Cyst no. T20-S1/80-81/106, SEM micrographs 316-322 (Plate II, 1-3). Type locality: Piston core T87-2-20G, southern continental margin of Crete, 34o.58', 4 N, 230.44 ', 8E. Type stratum: Upper Pleistocene, sapropel $5, _ 1 2 x 104a. Repository: Laboratory of Palaeobotany and Palynology, University of Utrecht, The Netherlands. Diagnosis: Obliquipithonelloid dinoflagellate cysts with a subspherical endocoel and a peridinioid outline. Tabulation is only indicated by the archaeopyle. Description: Calcareous dinoflagellate cysts having a subspherical endocoel (light-microscopic observations), one apical and two antapical lobes, and a wall composed of numerous randomly orientated calcite needles. The subspherical endocoel has an asymmetrical position. In consequence, the lower part of the cyst is solid as is the apex. The archaeopyle is located on the dorsal side of the

361

NEW PIOCENE AND PLEISTOCENE CALCAREOUS DINOFLAGELLATE CYSTS FROM SOUTHERN ITALY AND CRETE

cyst, just below the apex. It is probably formed by the loss of one or more intercalary plates. On the cyst surface, areas with a fine granulate structure (formed by the distal ends of the rodlike wall crystals), may alternate with more coarsely granulate areas (due to horizontal overgrowth and fusion of the crystal ends).

Dimensions: Holotype: length 40 gm, width 28 ttm, height 25 grn. Other specimens: length 36-45 gm, width 23-30 gm. n = 4.

emended to include all 3-5 faced obliquipithonelloid cysts on which the apical face is connected with all other faces. As has been demonstrated by Keupp (1984), the type species of the genus displays an obliquipithonelloid wall structure. The revision of Calcicarpinum by Keupp (1984) leaves only room for species with a tetrahedral outline which is, seen in the light of the forms described herein, a too narrow genus concept. The Cretaceous "Pithonella" usheri Krasheninnikov et Basov, 1983 differs from species of Calcicarpinum in lacking an apical face.

Occurrence: Frequent in core T87-2-20G, southern continental margin of Crete, upper Pleistocene. Also known from recent coastal sediments of South Carolina and the Upper Quaternary of the Black Sea (Wall et al., 1973).

Comparison: This species is the only calcareous dinoflagellate cyst known with a peridinioid outline. Obliquipithonella usheri (Krasheninnikov et Basov, 1983) Keupp, 1987, may also have an asymmetrical archaeopyle but it lacks a peridinioid shape. The structure of the cyst wall, with distal expansion of the calcite crystals on the cyst surface, corresponds with Obliquipithonella patriciagreeleyae (Bolli, 1974) Keupp et Mutterlose, 1984 (see Keupp, 1980a).

Calcicarpinum bivaivum sp. nov. (Plate III, I-8, Fig. 5) 1968a "Keeled-type" Wall and Dale, pp. 1406-1407, text-fig. 3, figs. 14, 15, pl. 172, fig. 23

Derivation of name: Latin, bi, two; valva, valve. Holotype: Cyst no. T20-S1/80-81/79, SEM micrographs 268, 269, 288-291 (Plate III, 1, 2). Type locality: Piston core T87-2-20G, southern continental margin of Crete, 34°.58 ', 4 N, 23°.44 ', 8E. Type stratum: Upper Pleistocene, sapropel $5,

Remark: Due to the position of the archaeopyle,

_+12x 104a.

the apex is easily removed from the rest of the cyst. As a consequence damaged cysts seem to possess an apical archaeopyle.

Paratype: Cyst no. 9651/2, SEM micrographs 65, 72-74 (Plate III, 3, 6, 7).

Genus Calcicarpinum Deflandre, 1948, emend.

Repository: Laboratory of Palaeobotany and

Type

Palynology, Netherlands.

species: Calcicarpinum tetraedricum

Deflandre, 1948, pp. 216, 218; text-figs. 35-37. Holotype: text-fig. 37.

Emended diagnosis: Obliquipithonelloid dinoflagellate cysts with subcircular to elongate endocoel and 3-5 faces. The single apical face contacts all other (2-4) faces of the cyst. Faces contact always at an angle of less than 90 °.

Remarks: All species thusfar included in this genus appear to be single walled. The diagnosis is

University

of

Utrecht,

The

Diagnosis: A species of Calcicarpinum with only three faces. The outline of the apical face is ellipsoid. Both remaining faces, which are of identical size and shape, have a semi-ellipsoid outline with the minor axis forming the straight margin. Each of the margins on either side of the major axis of the apical face, is connected over its full length with the straight margin of a semi-ellipsoidal face. The remaining, curved margins, of the semiellipsoid faces connect each other. As a result a

362 P L A T E III

GJM

V E R S I EEGH

NEW PIOCENE AND PLEISTOCENE CALCAREOUS DINOFLAGELLATE CYSTS FROM SOUTHERN ITALY AND CRETE

A

B

C

363

D

Fig. 5. Schematic diagram of Calcicarpinumbivalvum gen. nov., sp. nov. (A) Apical view. (B) Antapical view. (C) Ventral/dorsal view. (D) Lateral view.

distinct keel is formed of which the edge initiates at the points of strongest curvation o f the apical face. The endocoel is subspherical. The (sub)spherical archaeopyle has a central position on the apical plate and is probably formed by the loss o f plate 3'.

Remarks: The apical face of the single walled cysts is always about planar. Centered on the apical face a planar subcircular depression with a different wall structure may be present. This differently structurated area is interpreted as the operculum. The operculum may be completely missing leaving a circular archaeopyle. Sometimes, however, a hole is present within the operculum through which the organism may have left the cyst. The semi-ellipsoid faces are devoid of pliae. The cyst surface shows a tapetum of differently orientated calcareous crystal rods. Dimensions: Holotype: length 50 Ixm, apical ellipse 43 lam x 65 Ixm. Other specimens: length 47 lam, apical ellipse 65 txm x 42 Ixm. n = 4. Width endocoel 29 l~m. n = 1.

Occurrence: Rare in the upper Pliocene of the Singa section and frequent in core T87-2-20G, southern continental margin of Crete, upper Pleistocene. Also known from Recent sediments of the southeastern part of the Mediterranean Sea and the waters around Bermuda (Wall and Dale, 1968a). Comparison: This species differs from all other species of Calcicarpinum in having only three faces.

Calcicarpinum perfectum sp.

nov. (Plate IV, 1-11,

Fig. 6) 1968a "Wedged-type"Wall and Dale, p. 1406, text-fig.3, figs. 16, 17, pl. 172, fig. 24

Derivation of name: Latin, perfectus, perfect. With reference to the perfectly wedged shape. Holotype: Cyst no. 9453/13, SEM micrographs 33-36, 120-122 (Plate IV, 6, 7, 10).

PLATE III

Calcicarpinum bivalvumsp. nov. (All photographs x 1000, except where otherwise stated.) 1,2, Cyst T20-S1/80-81/79;holotype. 1. Lateral view (negative 289). 2. Apical view, operculum partly removed (negative268). 3,6,7. Cyst 9651/2; paratype. 3. Lateral-apical view (negative 65). 6. Cross-section of cyst wall (negative 73), x 5000. 7. Cross-section of dorso-ventral ridge (negative 74), x 5000. 4,5,8. Cyst T20-S1/80-81/21. 4. Detail of operculum and part of remaining apical face (negative 72), × 10,000. 5. Apical view (negative 70). 8. Dorsal or ventral view (negative 73).

364

PLATE IV

G.J.M. VERSTEEGH

NEW PIOCENE AND PLEISTOCENE CALCAREOUS DINOFLAGELLATE CYSTS FROM SOUTHERN ITALY AND CRETE

A

B

C

365

D

Fig. 6. Schematic diagram of Calcicarpinum perfectum gen. nov., sp. nov. (A) Apical view. (B) Antapical view. (C) Ventral/dorsal view. (D) Lateral view.

Paratype: Cyst no. 9453/53, SEM micrograph 259 (Plate IV, 1).

Type locality: Sample Calabria, south Italy.

9453,

Singa

section,

Type Stratum: U p p e r Pliocene, _ 20 x 105 a.

Repository: Laboratory of Palaeobotany and Palynology, Netherlands.

University

of

Utrecht,

The

Diagnosis: A species of Calcicarpinum with two triangular faces of similar size and three rectangular faces. The faces are orientated in such a way

that a wedge is formed with both triangular faces being parallel. The rectangular apical face is situated at the widest end of the wedge. The archaeopyle is circular and is situated at the center of the apical face. All faces are flat except for the apical face on which the operculum normally possesses a centered protrusion. The endocoel is subspherical.

Remarks: This single-walled obliquipithonelloid dinoflagellate cyst has a subspherical endocoel. The circular archaeopyle is, in analogy with other better tabulated calcareous cyst taxa, considered to be formed by the release of plate 3'. Antapically two rectangular faces connect, forming an "antapical rim". This rim may be interpreted as represent-

PLATE IV

Calcicarpinum perfectum sp. nov. (All photographs x 1000, except where otherwise stated.) I.

Cyst 9453/53; paratype, lateral-antapical view showing the antapical rim, the quadrangular lateral side and the dorsal (or ventral) triangular side (negative 259). Cyst 9453/31. 2, 3. Lateral view, note the small antapical depression (negative 79). 2. Apical view (negative 77). 3. 4,8. Cyst 9453/2. 4. Lateral-apical view (negative 4). Lateral-apical view (negative 9). 8. Cyst 9453/32; detail of cyst wall (negative 82), x 5000. 5. 6,7, 10. Cyst 9453/13; holotype. Apical view, the small hole on the apical side is considered to be an effect of preservation (negative 33). 6. 7. Dorsal (or ventral) view (negative 35). 10. Lateral view (negative 36). 9,11. Cyst 9453/37. Cross-section of cyst wall (negative 125), x 5000. 9. Apical view, cyst without operculum (negative 91). 11.

Sphaerodinella albatrosiana (Kamptner, 1963) Keupp et Versteegh, 1989, var. spinulosa var. nov. 12, 13, 16. Cyst 9453/38; holotype. 12. Detail cyst surface (negative 99), x 5000. 13. Cross-section of cyst wall (negative 98), x 5000. 16. Cyst, apical-dorsal view (negative 97), x 1500. 14, 15. Cyst 9451/24. 14. Cyst, apical-lateral view (negative 18), x 1500. 15. Detail cyst surface (negative 19), x 5000.

366

PLATE V

G.J.M. VERSTEEGH

367

NEW PIOCENE AND PLEISTOCENE CALCAREOUS DINOFLAGELLATE CYSTS FROM SOUTHERN ITALY AND CRETE

ing the connection between the two antapical plates present on peridinioid dinoflagellates. In that case the rectangular faces on either side of the "antapical rim" have a lateral orientation whereas the triangular faces in between are orientated dorsally and ventrally. The distance between both lateral faces sometimes shows an abrupt decrease directly above the antapical rim. The cyst surface shows a tapetum of differently orientated calcareous crystal rods.

Holotype of type species: Follisdinellum splendidum

Dimensions: Holotype: length 40 Ixm, apical face

Diagnosis: Obliquipithonelloid calcareous dino-

32 ktm x 33 lam. Other specimens: length 30-34 ~tm, apical face 30-38 lxm x 23-30 ~tm. n = 6.

flagellate cysts. The precingular and antapical plates, and sometimes the 1' plate, are reflected by intratabular lobes. The archaeopyle has an apical orientation and is probably formed by the release of plate 3'.

Occurrence: Frequent in sample 9453 of the Singa section, upper Pliocene. U n c o m m o n in the Subrecent sediments of the Yucatan Basin (Wall and Dale, 1968a). Comparison: C. perfectum resembles Calcicarpinum bivalvum sp. nov. but differs in having five rather than three faces and in having a central protrusion on the operculum. C. perfectum differs from Calcicarpinum tetraedricum, Deflandre, 1948, in having five rather than four faces and consequently having an antapical rim rather than an antapical point.

Follisdinellum gen.

(Plate V, 6, 8-11).

Derivation of name: Latin, follis, balloon. With reference to the shape of the cyst.

Repository: Palynology, Netherlands.

Laboratory of Palaeobotany University of Utrecht,

and The

Comparison: Follisdinellum most closely resembles the genus Calciperidinium, but differs by lacking a peridinioid shape. Thus far, the type species is the only species within this genus. Follisdinellum splendidum sp.

nov. (Plate V, 1-11)

1968a "Clavate-type" Wall and Dale, p. 1406, text-fig.3, figs. 7-9, pl. 172, figs. 16-18. 1968b "Lobate-cyst" Wall and Dale, pl. 4, fig. 25.

nov.

Type species: Follisdinellum splendidum sp. nov.

Derivation of name: Latin, splendidus, splendid. With reference to the splendid shape of the cyst.

PLATE V

Follisdinellumsplendidumgen. nov., sp. nov. (All photographs x I000, except where otherwise stated.) 1,4.

1. 4. 2,5. 2. 5. 3. 6,8-11. 6. 8. 9. 10. 11. 7.

Cyst T20-S1/80-81/2. Lateral view, apical side to the right (negative 9). Apical view, note the eight lobes and archaeopyle (negative 7). Cyst T20-S1/80-81/I0. Apical view, note the eight lobes and archaeopyle (negative 35). Lateral view (negative 34). Cyst T20-S1/80-81/66; apical view, operculum only partly removed (negative 201). Cyst T20-SI/80-81/5; holotype. Lateral view (negative 19). Cross-section of cyst, note the subspherical endocoel (negative 231). Ventral view (negative 16). Cross-section of antapical cyst wall (negative 232), x 5000. Detail of operculum (negative 18), x 10,000. Cyst T20-Sl/80-81/27; paratype, antapical view, note the two antapical and seven precingular lobes as well as a smaller eighth lobe reflecting plate 1' (negative 90).

368

PLATE VI

G.J,M. VERSTEEGH

NEWPIOCENEANDPLEISTOCENECALCAREOUSDINOFLAGELLATE CYSTSFROMSOUTHERNITALYANDCRETE

Holotype: Cyst no. T20-S1/80-81/5, SEM micrographs 17-20, 231,232 (Plate V, 6, 8-11).

Paratype: Cyst no. T20-S1/80-81/27, SEM micrograph 90 (Plate V, 7). Type locality: Piston core T87-2-20G, southern continental margin of Crete, 34°.58 ', 4 N, 23°.44 ', 8E. Type stratum: Upper Pleistocene, sapropel $5, + 1 2 x 104a.

Repository: Palynology, Netherlands.

Laboratory of Palaeobotany and University of Utrecht, The

Diagnosis: A species of Follisdinellum with in general nine intratabular lobes: seven precingular and two antapical. An assessory tenth lobe may be positioned between the precingular lobes. The conical cylinder between the antapical face and the ring of precingular lobes displays no tabulation. The diameter of this cylinder decreases towards the antapex.

Description: Calcareous dinoflagellate cysts having a subspherical endocoel, a roughly conical shape and a compact wall, composed of numerous randomly orientated calcite needles. The apical face is convex and subcircular in outline. Centered at the apical face a subcircular field with different wall structure, interpreted as the operculum, may be present. The operculum may be completely absent. Sometimes, however, a hole is present in

369

the operculum through which the organism may have left the cyst. Seven intratabular lobes, arranged in a transversal ring, reflect the precingular plates. Additionally, an eighth lobe, mostly of smaller size may be interpolated. From this seven or eight lobed ring towards the antapical area the cyst tapers and no tabulation is reflected. The antapical area is two-lobed. Both lobes are considered to be homologous with plates 1~" and 2". The position of the usually smaller, additional "eighth" lobe on the multi-lobed ring relative to that of the antapical lobes, suggests this "eighth" lobe to be homologous with plates 1' or 4". For calcareous dinoflagellate cysts with clear tabulation the 1' plate is much smaller than the precingular plates whereas the 4" plate is just normal in size. Hence, the often smaller, additional "eighth" lobe is considered to be homologous with the 1' plate.

Dimensions: Holotype: length 42 ~tm, width 45 ~tm, width endocoel 30 lam. Other specimens: length 37-41 ~tm, width 40-46 ~tm, n = 8. Occurrence: Frequent in core T87-2-20G, southern continental margin of Crete, late Pleistocene. Also known from Recent sediments of the Mediterranean and near Bermuda (Wall and Dale, 1968a, b). Comparison: The combination of a seven or eight lobed ring and a two lobed antapical face readily separates this taxon from all other calcareous dinoflagellate cysts.

PLATE VI

Melodomunculaberlinensisgen. nov., sp. nov. (All photographs x 1500, except where otherwisestated.) 1-3, 7. 1. 2. 3. 7. 4. 5, 6. 5. 6.

CystT20-S1/80-81/109; holotype. Apical view (negative 310). Antapicalview (negative 308). Lateral view (negative307). Detail of antapical side, margins of the antapical paraplates are indicated by sutures (negative315), x 5000. Cyst T20-S1/80-81/86; apical view, the margin of archaeopyleis weaklyindicated by a suture. Cyst T20-S1/80-81/16. Antapicalview,margins of the antapical paraplates are indicatedby sutures, right corresponds with dorsal (or ventral) cyst side (negative57). Detail of cyst surface (negative 56), x 5000.

370

PLATE VII

G JM. VERSTEEGH

NEW PIOCENE AND PLEISTOCENE CALCAREOUS DINOFLAGELLATE CYSTS FROM SOUTHERN ITALY AND CRETE

Remark: Forms with a nine lobed equatorial ring as described by Wall and Dale (1968a) have not been observed in the material from Crete. Genus Melodomuncula gen. nov.

Type species: Melodomuncula berlinensis sp. nov. Holotype of type species: Melodomuncula berlinensis (Plate VI, 1-3, 7). Derivation of name: Latin, melis, honey; Latin, domuncula, little house. With reference to the shape, which resembles the chamber of a honey bee. Repository: Laboratory of Palaeobotany and Palynology, University of Utrecht, The Netherlands. Diagnosis: Obliquipithonelloid dinoflagellate cysts, characterised by a hexagonal apical and antapical face. From the margins of the apical face six quadrangular faces are developed which are bordering the corresponding margins of the also hex-

371

agonal antapical face. The apical face displays a centered archaeopyle.

Remarks: Melodomuncula differs from Praecalcigonellum Keupp et Versteegh, 1989 by having a more reduced tabulation and an obliquipithonelloid rather than an orthopithonelloid crystal orientation in the outer wall of the cyst. Gonellum Keupp, 1987 differs by having only five large prismatic fields corresponding with the fused equivalences of pre- and postcingular plate cycles. Both cysts may be the result of a convergent evolutionary development. Thus far, the type species is the only representative of the genus. Melodomuncula berlinensis sp. nov. (Plate VI, 1-7,

Fig. 7)

Derivation of name: Latin, berlinensis, from Berlin. Holotype: Cyst no. T20-S1/80-81/109, SEM micrographs 307-310, 315 (Plate VI, 1-3, 7). Type locality: Piston core T87-2-20G, southern continental margin of Crete, 34°.58 ', 4 N, 23°.44 ', 8E.

PLATE VII Obliquipithonella spp. Keupp et Mutterlose, 1984. From the top of this plate downwards the distal overgrowth of the surface crystals decreases (All photographs x 1000, except where otherwise stated.) O. patriciagreeleyae (Bolli, 1974) Keupp et Mutterlose, 1984. 1, 2. Cyst 9652/4; the complete cyst wall is covered by large plates formed by groups of distally fused crystal ends. 1. Apical view (negative 3). 2. Detail of cyst surface (negative 4), x 5000. O. thayeri (Bolli, 1974) Keupp et Mutterlose, 1984. 3, 4. Cyst 9453/23; single slender distally not extended surface crystals alternate with large plates of fused crystal ends 3. Apical view (negative 59). 4. Detail of cyst surface (negative 60), x 5000. O. muhistrata (Pflaumann et Krasheninnikov, 1978) Keupp et Mutterlose, 1984. 5, 6. Cyst 9453/27; poor fusion and distal extension of the surface crystals 5. Apical view (negative 68). 6. Detail of cyst surface (negative 70), x 5000. O. carteri (Bolli, 1974) Keupp et Mutterlose, 1984. According to Keupp (1981, p. 38) this cyst type represents the inner wall of Orthopithonella cysts 7, 8. Cyst 9651/53; due to erosion of the outer wall of this specimen of O. multistrata the innerwall, with typical O. carteri characteristics, is visible. 7. Latero-apical view (negative 60). 8. Detail of cyst surface (negative 61), x 5000.

372

G.J.M. VERSTEEGH

A

B

C

D

Fig. 7. Schematicdiagramof Melisdomunculaberlinensisgen. nov., sp. nov. (A) Apicalview.(B) Antapicalview.(C) Dorsal/ventral view. (D) Lateralview.

Type stratum: Upper Pleistocene, sapropel $5, + 1 2 x 104a.

25 lam x 19 lam. Other specimens: length 15-18 lam, width 20-26 rtm. n = 4.

Repository: Laboratory of Palaeobotany and Palynology, University of Utrecht, The Netherlands.

Occurrence: Core T87-2-20G, southern continental margin of Crete, upper Pleistocene.

Diagnosis: A species of Melodomuncula characterised by apical and antapical faces of the same size which are parallel. The corresponding corners of these faces are connected by straight ridges, each pair of ridges enclosing a rectangular face. All rectangular faces are about the same size. At the center of the apical face a subspherical archaeopyle may be present.

Comparison: The combination of obliquely orientated calcite crystals in the outer wall and hexagonal apical and antapical faces readily separates M. berlinensis from all other calcareous dinoflagellate cyst species.

Description: Calcareous dinoflagellate cyst having a subspherical endocoel, a polygonal shape and a wall composed of numerous randomly orientated calcite needles. The apical and antapical faces are hexagonal and have the same size and all margins are of the same length. Both hexagonals are flat and have a parallel orientation. On polar view, they have not, or only slightly, been twisted relative to each other. Each corner o n the apical face is connected with the corner directly below (on the antapical face) by a straight ridge. Between each pair of ridges a flat rectangular face is developed. The apical face displays a centered subspherical archaeopyle. On the central part of the antapical face both antapical plates may be indicated by weakly developed sutures. Five of the six margins of the antapical face may correspond with the upper margins of the postcingular plates while the sixth one is located in the sulcal area. Dimensions: Holotype: length 19 lam, hexagonal faces 20 lam x 27 ~tm, quadrangular faces

Genus Obliquipithonella Keupp et Mutteriose, 1984 Type species: O. multistrata (Pflaumann et Krasheninnikov, 1978) Keupp et Mutterlose, 1984. All encountered Obliquipithonella species belong to the Obliquipithonella patriciagreeleyae/ multistrata group sensu Keupp, 1981. Differences between the species of this group are considered to be dependant on environmental conditions during cyst-formation (Keupp, 1981; Bandel and Keupp, 1985). As a result, the species concerned are often not clearly separated from each other. Obliquipithonella carteri (BoUi, 1974) Lentin et Williams, 1985 (Plate VII, 7, 8) Remarks: This cyst type is considered to represent the isolated inner wall of representatives of the Obliquipithonella patriciagreeleyae/multistrata group (Keupp, 1981). In Eocene sediments this species is equivalent to specimens of Bicarinellum castaninum Deflandre, 1948 without tabulation (Keupp, 1984). O. carteri had not been reported from sediments younger than Eocene in age

NEWPIOCENEAND PLEISTOCENECALCAREOUSDINOFLAGELLATECYSTSFROMSOUTHERNITALYAND CRETE

(Keupp, 1984). In the present study it was found in samples of late Pliocene age.

Pseudopithonella gen.

373

nov.

Type species: Pseudopithonella striatula sp. nov. Obliquipithonella multistrata

(Pflaumann et Krasheninnikov, 1978) Lentin et Williams, 1985 (Plate VII, 5, 6)

Hoiotype of type species: Pseudopithonella striatula

Remarks: This form had not been reported from

Derivation of name: Greek, pseudo, false. With

sediments younger than Cretaceous in age, but has now been found in samples of late Pliocene and late Pleistocene age.

reference to the orientation of the surface crystals.

Obliquipithonella patriciagreeleyae (Bolli, 1974)

(Plate IX, 3, 4).

Repository: Laboratory of Palaeobotany and Palynology, Netherlands.

University

of

Utrecht,

The

Lentin et Williams, 1985 (Plate VII, 1, 2)

Diagnosis: Spherical to elongate obliquipithonelRemarks: This form had not been reported from

Remarks: This form had not been reported from

loid dinoflagellate cysts. Except for the archaeopyle no tabulation is indicated. The surface layer of the cyst shows a pattern of areas (called "unidirectional areas") in which all calcite crystals have the same orientation. The crystal orientation is different for each area. Below the cyst surface the cyst wall consists of randomly arranged calcite crystals.

sediments younger than Cretaceous in age. In the present study it was found in samples of late Pliocene age.

Comparison: In contrast to the Cretaceous Pithonelloideae, which display only one unidirec-

sediments younger than Cretaceous in age. Here it is reported from samples of late Pliocene age.

Obliquipithonella thayeri (Bolli, 1974) Lentin et Williams, 1985 (Plate VII, 3, 4)

PLATE VIII (see p. 374)

Praecalcigonellum schizosaeptum sp. nov. (All photographs x 1000, except where otherwise stated.) 1-5,7,8. 1. 2. 3. 4. 5. 7. 8. 6.

Cyst 9453/52; holotype, positioned on the second left lateral side. Pentagonal, apical side, showing archaeopyle suture (negative 253). Pentagonal right lateral side (negative 260). Trigonal right lateral side (negative 285). Pentagonal dorsal side (negative 257). Trigonal left lateral side (negative 258). Antapical view, showing the trigonal and pentagonal right lateral sides and the pentagonal dorsal side (negative 286). Detail of apical cyst surface (negative 255), x 5000. Cyst 9452/UI; apical view (negative 3).

PLATE IX (see p. 375)

Pseudopithonella striatula gen. nov., sp. nov. (All photographs × 1000, except where otherwise stated.) 1, 6. 1. 6. 2, 5. 2. 5. 3, 4. 3. 4.

Cyst T20-$2/87-88/8. Apical view (negative 10). Detail of cyst surface, note the parallel rows of rod like crystals (negative 2), × 5000. Cyst T20-S1/80-81/59. Cross-section of cyst wall, note the platy crystals (negative 240). Detail of cyst surface, note the rod like appearance of crystals on cyst surface (negative 179), x 10,000. Cyst T20-$2/87-88/1; holotype. Lateral view (negative 5). Detail of cyst surface, note the areas with different crystal orientation (negative 6), x 5000.

374 PLATE VIII

(for description see p. 373)

G.J.M. V E R S T E E G H

NEW PIOCENE AND PLEISTOCENE CALCAREOUS DINOFLAGELLATE CYSTS FROM SOUTHERN ITALY AND CRETE

PLATE IX

(for description see p. 373)

375

376

GJ.M. VERSTEEGH

tional area (the complete cyst), the cyst surface of this new genus shows at least two of these areas. It differs from other Obliquipithonelloid genera in having the crystals on the cyst surface arranged in areas with identical orientation rather than a irregular or random crystal arrangement.

"rods" have a parallel orientation. Along the borders of the unidirectional areas these rows may be broken down and a random crystal orientation may be observed. On cross-section the single layered cyst wall shows a dense structure of randomly orientated, often plate-like, crystals.

Pseudopithonella striatula sp. nov. (Plate IX, 1-6)

Dimensions: Holotype: width 59 Ixm, thickness wall 9 gm. Other specimens: width 57-52 p.m, thickness wall 6.4 gm. n = 2. Length-width ratio of surface crystals 3-6.

Derivation of name: Latin, striatus, striation. With reference to the striate appearance of the cyst surface.

Holotype: Cyst no. T20-$2/87-88/1, SEM micro-

Occurrence: Common in core T87-2-20G, southern continental margin of Crete, upper Pleistocene.

graphs 5, 6, 13 (Plate IX, 3, 4).

Type locality: Piston core T87-2-20G, southern continental margin of Crete, 34°.58 ', 4 N, 23°.44 ', 8E.

Type stratum: Upper Pleistocene, sapropel $3, _+7.6 × 104 a.

Repository: Laboratory of Palaeobotany and Palynology, Netherlands.

University

of

Utrecht,

The

Comparison: This form differs from all other described calcareous dinoflagellate cysts by its pseudopithonelloid surface structure.

Subfamily ORTHOPITHONELLOIDEAEKeupp, 1987 Tribus ORTHOPITHONELLEAEKeupp et Versteegh, 1989

Genus Praecalcigonellum Keupp et Versteegh, 1989

Type

species:

Praecalcigonellum polymorphum

Diagnosis: A spherical to subspherical cyst of Pseudopithonella. The crystals of the unidirectional

(Keupp, 1980b) Keupp et Versteegh, 1989.

areas are arranged in parallel rows. The direction of these rows is different for each area.

Praecalcigonellum schizosaeptum sp.

nov.

(Plate VIII, 1-8; Fig. 8)

Description: Orthopithonelloid calcareous dinoflagellate cysts having a subspherical endocoel and shape. The crystals within the "unidirectional areas" (areas in which all crystals have the same orientation) are arranged in parallel rows. On the cyst surface, these crystals may have a rod like appearance. In each row the long sides of these

A

s

B

s

C

1968a "Tetrahedral-type"Wall and Dale, pp. 1403-1405, textfig. 3, figs. 1-3, pl. 172, fig. 13. 1989 Praecalcigonellumsp. 1. Keupp and Versteegh, p. 21 l, pl. 2, fig. 12.

Derivation of name: Greek, schizos, to divide; Latin, saeptum, septum.

D

E

Fig. 8. Schematicdiagram of Praecalcigonellumschizosaeptumgen. nov., sp. nov. (A) Apical view. (B) Antapical view. (C) Ventral view. (D) Dorsal view. (E) Left lateral view.

377

NEW PIOCENE AND PLEISTOCENE CALCAREOUS DINOFLAGELLATE CYSTS FROM SOUTHERN ITALY AND CRETE

Holotype: Cyst no. 9453/52, SEM micrographs 253-258, 260, 261,280-287 (Plate VIII, 1-5, 7, 8).

Type locality: Sample Calabria, south Italy.

9453,

Singa

section,

Occurrence: Rare in the upper Pliocene sediments of the Singa section, and core T87-2-20G, southern continental margin of Crete, upper Pleistocene. Also reported from Recent sediments of Bermuda and plankton samples from Woods Hole (Wall and Dale, 1968a).

Type stratum: Upper Pliocene, + 20 x 105 a. Repository: Laboratory of Palaeobotany and Palynology, Netherlands.

University

of

Utrecht,

The

Diagnosis: A species of Praecalcigonellum characterised by a flattened apical face on the widest part of the cyst and a very reduced sulcal area. Except for the transversal ridge that separates the apical face from the rest of the cyst, all ridges have a longitudinal orientation.

Comparison: P. schizosaeptum differs from other species of Praecalcigonellum by its approximately wedge-formed, poorly tabulated antapex which is not distinctly separated from the middle part of the cyst. This new species resembles Calcicarpinum tetraYdicum Deflandre, 1948, but P. schizosaeptum has a orthopithonelloid, rather than an obliquipithonelloid structure of the outer wall.

Tribus CALCIODINELLEAEKeupp et Versteegh, 1989

Description: Orthopithonelloid dinoflagellate cysts with a subspherical endocoel and a flattened five sided apical field containing a circular central archaeopyle. This apical field is separated from the rest of the cyst by a pentagonal "transversal" ridge. From this ridge towards the antapical pole the diameter of the cyst decreases. The lowermost 1/5 of the cyst shows a stronger decrease in diameter than above. From the antapical pole three ridges develop in opposite directions. Two of these ridges bifurcate in the lower 1/5 of the cyst, each forming a pair of branches. Between the branches of each pair and the transversal ridge two flattened triangular faces are enclosed. Between the transversal ridge, the facing branches of the bifurcating ridges and the lower unbranched part of these ridges, a flattened pentagonal face is present. The four branches enclosing the triangular faces end at the transversal ridge, each on a different corner. The third ridge bifurcates near the transversal ridge and both corresponding branches stay close together, they contact the transversal ridge at the remaining corner. Two other pentagonal faces are present at either side of this third, apically branching, ridge.

Genus

Sphaerodinella

Keupp et Versteegh, 1989

Type species: S. albatrosiana (Kamptner, 1963) Keupp et Versteegh, 1989.

Sphaerodinella albatrosiana

var.

spinulosa var.

nov.

(Plate IV, 12-16) 1977a Thoracosphaerasp. 1. Fiitterer, p. 717, pl. 7, figs. 1-7.

Derivation of name: Latin, spinula, little spine. Holotype: Cyst no. 9453/38, SEM micrographs 18, 19 (Plate IV, 12, 13, 16).

Type locality: Sample Calabria, south Italy.

9451,

Singa

section,

Type stratum: Upper Pliocene, _ 20 × 105 a. Repository: Laboratory of Palaeobotany and Palynology, Netherlands.

University

of

Utrecht,

The

Dimensions: Holotype: length 42 ~tm, width 40 lxm.

Diagnosis: A variety of Sphaerodinella albatrosiana

Other specimens: 34-42 ~tm.

with a small capitate spine on each connection between muri on the cyst wall.

length

35-43 I~m, width

378

G.J.M. VERSTEEGI-I

Occurrence: Rare in the upper Pliocene sediments of the Singa section. Also rare in uppermost Miocene and Pliocene sediments from the Serra Leone Rise (Fiitterer, 1977a). Dimensions: Holotype: width 30 ~tm, length of spines ~<1 lam. Other specimens: width 18-30 p.m length of spines ~<1 I.tm. Remarks: Except for the small capitate spines on the

cyst

wall,

this

type

is

identical

to

S. albatrosiana. Although Fiitterer (1977a) considered the presence of spines to be of fundamental specific rank, separation on species level on the basis of this small difference alone seems not appropriate.

Discussion None of the encountered obliquipithonelloid calcareous dinoflagellate cysts were hitherto known to occur in Pliocene or Pleistocene sediments. The species Obliquipithonella multistrata (Pflaumann et Krasheninnikov, 1978) Lentin et Williams, 1985; O. patriciagreeleyae (Bolli, 1974) Lentin et Williams, 1985, and O. thayeri (Bolli, 1974) Lentin et Williams, 1985, were not even known from Tertiary sediments. The presence of these Jurassic and Cretaceous forms, especially in the samples from the Singa section, may be due to reworking. This may be confirmed by the presence of reworked Cretaceous elements in organic-walled dinoflagellate cyst assemblages from the Singa section (Zevenboom and Versteegh, in prep.). On the other hand, there still is a general lack of taxonomic information on Tertiary calcareous dinoflagellate cysts. Thus the possibility that the forms represent long ranging species should not be completely ruled out. Furthermore, the encountered Obliquipithionella species are all representatives of the O. patriciagreeleyae-multistrata group (Keupp, 1981). Morphological variation within this group may have an environmental rather than a genetic origin (Keupp, 1981), whereas living representatives have been reported from Recent sediments (Akselman and Keupp, 1990; Keupp, in press). Six of the newly described taxa (Bicarinellum tricarinelloides sp. nov., Calciperidinium asymmet-

ricum gen. nov, sp. nov., Calcicarpinum bivalvum sp. nov., C. perfectum sp. nov., Follisdinellum splendidum gen. nov., sp. nov., and Sphaerodinella albatrosiana var. spinosa var. nov.) have been mentioned in the literature before. They are only reported from Pliocene or younger sediments. The present occurrences are therefore considered to be autochthonous. Both species which have not been recorded before (Pseudopithonella striatula gen. nov, sp. nov. and Melodomuncula berlinensis gen. nov., sp. nov.), may be either reworked from Cretaceous sediments or autochthonous. However, because of the fact that these forms have never been encountered in the relatively well known Cretaceous assemblages, it is here considered that the two species represent Pleistocene elements. Conclusions Rich assemblages of calcareous dinoflagellate cysts and other calcareous cysts have been encountered in sediments from southern Italy (Pliocene-early Pleistocene) and the southern continental margin of Crete (Pleistocene). Despite the occurrence of forms that might represent reworked Cretaceous elements, the assemblages appeared to yield valuable information on species diversity among Pliocene and Pleistocene calcareous dinoflagellate cysts. Evidently the Mediterranean realm is an excellent study area for a further reconstruction of the still incompletely known Late Neogene and Quaternary history of these forms, and subsequent assessment of their potential in stratigraphical and paleo-environmental analyses.

Acknowledgements H. Keupp is kindly thanked for the opportunity to work at the Institut f/Jr Palfiontologie, Fachbereich Geologie, FU Berlin and for his advise and inspiring enthusiasm during this study. Constructive criticism of H. Brinkhuis and H. Visscher (Univ. Utrecht) greatly improved the manuscript. U. Heier (FU Berlin), P. Hoen, H. Elsendoorn and E. Rohling (Univ. Utrecht) are thanked for their technical assistance. Financial support from the Utrechts Universiteits Fonds and

NEWPIOCENEAND PLEISTOCENECALCAREOUSDINOFLAGELLATECYSTSFROMSOUTHERNITALYAND CRETE

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