new late triassic calcareous algae from hydra,...

ACTA PALAEONTOLOGICA ROMANIAE V. 2 (1999), P. 139-156

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NEW LATE TRIASSIC CALCAREOUS ALGAE FROM HYDRA, GREECE

OVIDIU DRAGASTAN1, BÄRBEL KUBE2 & DETLEV K. RICHTER2

Abstract. The Late Triassic sequence of Kap Kastello studied by one of the authors (B. K.) provided an association of calcareous algae dominated by the families Pseodoudoteaceae and Halimedaceae. An analysis of the characters of the cortical system indicates the presence within the Family Halimedaceae of two evolutionary – lines: one having cylindriform cortical siphon , starting from the Ordovician with the genera Dimorphosiphon – Palaeoporella; and a second one having vesiculiferous cortical siphons starting in the Permian with the genera Saxonia - Tauridium and Halimeda. The fallowing new taxa are introduced: Halimeda helladica n.sp., H. discreta n.sp., Hydracara kubeae n.g., n.sp., Felixporidium triasicum n. sp., Metasolenopora semiconcentrica n. sp. and Parachaetetes rhaeticus n. sp.

Keywords: Late Triassic, calcareous algae.

1 University of Bucharest. Faculty of Geology and Geophysics, Laboratory of Paleontology, Bd. N. Bălcescu 1, 70111 Bucureşti 2 Ruhr – Universität, Bochum, Institut für Geolgie, Universitätsrasse 150, D-44801 Bochum, Germany

INTRODUCTION

The Tethyan carbonate platforms of Middle to Late Triassic age are important for their widespread reefal environments with high algal diversity (e.g. FLÜGEL, 1982, FLÜGEL & FLÜGEL-KAHLER, 1992, BECK et al., 1996). In view of faunal and floral provinces, algae are of special interest because of their position within the Tethyan ocean (KUBE et al., 1998a, 1998b).

A Middle to Late Triassic reef basin complex dominates the geology of Hydra island, consisting of

deep water carbonates on the southern thrust sheet, while especially the northeastern part of the island is characterized by shallow water limestones (RÖMERMANN, 1968, RICHTER, 1999). SCHÄFER & SENOWBARI-DARYAN (1982) described a vertical as well as horizontal facies differentiation, caused by the progradation of the carbonate platform towards the south. In the area of Kap Kastello, a brecciated level of uppermost Triassic to Early Jurassic age overlies lagoonal carbonates of Norian / Rhaetian age, which cover a Carnian prograding reef - complex (Fig.1, 2).

Figure 1 - Sketch map of Hydra island including the area studied at Kap Kastello (for detail see fig. 2a).

Upper left: position of Hydra within the Hellenic zones after Mountrakis (1986).

O. DRAGASTAN, B. KUBE & D.K. RICHTER

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In 1994, TURNSEK & SENOWBARI-DARYAN investigated the reef limestones east of Hydra Chora and described Carnian/Lowermost Norian coral associations. SCHÄFER & SENOWBARI-DARYAN (1983) for the first time studied the microbiota of the Norian/Rhaetian Loferites. DRAGASTAN et al. (1997) described new Norian and Rhaetian taxa from the lagoonal facies of the Kap Kastello section. Polyphase tectonic activity, associated with the formation of breccias at the Triassic/Jurassic boundary, caused the

rapid subsidence of the carbonate platform. The lithology and genesis of these breccias are discussed in RICHTER & FÜCHTBAUER (1981), FÜCHTBAUER & RICHTER (1983) and RICHTER (1994). KUBE et al. (1998a) for the first time discribed the facies-development of the Kap Kastello section in detail.

Now, a revision of microfacies investigation in connection with more samples has resulted in recognition of several new algal taxa, which is the aim of this presentation.

Figure 2 - 2a. Geological map of the Kap Kastello area showing the location of the investigated section (thick black line). 2b. Schematic columnar section of Late Triassic / Jurassic facies evolution (from Kube et al., 1998). Late Triassic shallow water limestones (backreef limestones, loferites) are overlain by Rhaetian / Liassic breccias (Breccia succession). Basinal deposits are represented by Middle / Late Jurassic radiolarites.


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GEOLOGICAL SETTING Within the Hellenic geotectonic zonation, the

sedimentary series of Hydra are located in the Subpelagonian zone (Fig.1) - the southwestern margin of the Pelagonian Platform transitional to the Pindos zone (JACOBSHAGEN, 1986). The Lower Permian to Late Jurassic succession from a lithostratigraphic series with a thickness of more than 1500 - 2200 m (RICHTER,1999) - distributed in two EW-striking thrust sheets, dislocated by vertical faults (Fig.1). The sequence documents the build-up, fracture and subsidence of three carbonate platforms (RICHTER & FÜCHTBAUER, 1981). RICHTER (1994) considered the multiphase formation, with internal breccias and mass flows, to be associated with strike-slip faults, activated during the Hellenide rifting phase.

Permian shallow-marine deposits, the first carbonate platform, are overlain by Scythian gravity flows and deep water limestones of Lowermost Anisian age. Reef limestones rich in Shamovella characterize the formation of the second carbonate platform of Middle to Late Anisian in age. Following Illyrian mass-flows, tuffites and ammonoid-bearing red limestones demonstrate massive tectonic activity. This event caused strong facies differentiation, with basinal deposits on the southern and shallow-water carbonates on the northern thrust sheet of Hydra. During the evolution of the Late Triassic third carbonate platform, deposition of laminated cherty limestones indicate deep-water sedimentation in the south, while reef limestones document the growth of a carbonate platform on the northern thrust sheet. The transitional area of this reef-basin complex is represented by well-bedded fore-slope limestones containing gravity flows with shallow-water bioclasts. The progradation of the platform towards the southwest resulted in horizontal as well as vertical facies zonation (SCHÄFER & SENOWBARI-DARYAN, 1982). The final lagoon dominated phase of the third carbonate platform - investigated in microfacial detail by KUBE et al. (1998)- contains our sequence with the new algal taxa presented in this paper:

In the Kap Kastello region, backreef limestones, which are overlain by Norian/Rhaetian Loferites, cover reef-limestones up to 1000m thick (Fig. 2). A lateral facies differentiation of discrete reef and debris-rich interreef deposits is developed at the top of the coarsely bedded Carnian Reef limestones. The dominance of algal rich sediments (especially encrusting cyano-phyceans), as well as the overlying deposition of lagoonal carbonates, indicate patch reefs growing in a backreef area. Late Triassic lagoonal sedimentation started with deposition of 80m thick Carnian backreef-limestones. Ostracod-mudstones of a low energy environment represent the background sedimentation. Grainstone intercalations with high species-diversity (algae, bivalves, echinoderms, sponges, corals, etc.) as well as the occurrence of ooids, indicate the temporary existance of a higher energetic realm. The upper part of the backreef facies is characterized by zebra limestones. The lower part of the 450 m thick sub- to supratidal Norian/Rhaetian loferites is dominated by peloidal loferites subsequently replaced by algal mat loferites, formed by stromatolites. The succession consists of an alternation of loferitic layers and subtidal micritic limestones, mainly composed of ostracod-mudstones and characterized by the occurrence of megalodontids. Loferites layers are the characterstic facies, composed of dolomicritic mats with embedded calcified fenestrae as well as irregular dolomitic peloids.

Calcarenitic intercalations of algae, foraminifers, ostracods and ooids occur sporadically. At the Triassic / Jurassic transition, tectonic activity caused rapid subsidence of the carbonate platform, accompanied by internal breccias and mass flows. The 80 m thick Breccia succession of internal breccias and mass flows is overlying by the loferitic limestones in the Kap Kastello area. Middle to Late Jurassic pelagic cherts (Radiolarites), covering Liassic red limestones (Ammonitico Rosso), indicate the final deepening of the hydriotic depositional environment.

The Jurassic drifting of the Pindos ocean effected the development of a convergent plate margin. Subduction towards the northeast (Pelagonian platform) took place until Tertiary time, and the hydriotic succession became part of an accretionary wedge (e.g. RICHTER, 1999).

STRATIGRAPHY The Late Triassic of the Kap-Kastello profile

provided a Carnian, Norian, Rhaetian sequence and a Rhaeto-Lias transitional breccia facies.

Carnian - The biota of reef limestones is dominated by a coral-sponge assemblage of Voltzeia - Stuorseia and Pantokratoria fasciculata.

The foraminifers which occur in the reef - environment are represented by Spiriamphorella sp., Auloconus permodiscoides (Oberhauser), Diplotremina sp., D. cf. alta (Kristan - Tollmann), “Tubiphytes” gracilis SchÄfer & Senowbari-Daryan, “T.” multisiphonatus SchÄfer & Senowbari-Daryan, and Reophax sp.

Cyanophyceans are not so widespread in the reef environment. They occur sporadically, having small thalli, such as Rivularia lemaitrae (Dragastan, 1989) and Ortonella myrae Racz (DRAGASTAN, 1993, DRAGASTAN et al., 1998).

The dasycladaleans Oligoporella minutuloidea (Herak, 1967) and O. hydrae Dragastan have robust, small, well calcified thalli. This assemblage developed on the reef in the small pools and cavities.

Ladinella porata Ott very often occurs in the Carnian sequence. In our opinion this is a microsponge which preferred to live in reef cavities.

Norian - The major characteristic of the Norian biota is the presence of different microorganisms of the lagoon communities, clearly dominated by various groups of foraminifers: Ataxophramiidae (Agglutisolena), Involutinacea - Aulotortidae (Aulotortus), and by green algae: Bryopsidales - Pseudoudoteaceae, Dasycladales (Gyroporella, Pseudogyroporella, Salpingoporella). Cyanophyceans are represented by Rivularia lemaitrae (Dragastan) and Polytrichella fructicosa Schäfer & Senowbari-Daryan, 1983.

Crustose, red algae (Parachaetetes - Solenopora) occur in surrounding of the patch reef communities in the lagoon. The involutinid species, mainly the aulo-tortids represented by Aulotortus sinuosus Weynschenk, A. gaschei (Koehn - Zaninetti & Brönnimann), A. communis (Kristan), and A. tenuis (Kristan) are mostly found in the lower part of the Norian.

The algae were well developed in lagoons in the Norian. The dasycladalean and bryopsidalean green algae became dominant, in comparison with cyanophyceans and red algae.

Medium sized, well calcified Gyroporella vesiculifera Gümbel and Pseudogyroporella norica Dragastan, P. cryptica Dragastan, with small size thalli, together with the Aciculella sp., a calcareous sporangiophore, indicate a Norian age.


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Figure 3 - Stratigraphical ranges of algae, foraminifers and microproblematica in the Upper Triassic Kap Kastello section, Hydra


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The flora of green siphoneous algae, Bryopsidales – Pseudoudoteaceae, is dominant in the Norian. The taxa, with large or small thalli, are strongly calcified, such as Pseudoudotea magna Dragastan et al., 1997, P. littlerorum Dragastan et al., 1997, and Hydraea calcarata Dragastan et al., 1997. Garwoodia maxima Dragastan, the ancestral pseudoudoteacean, has the same Norian sratigraphic range.

Rhaetian - The patch-reef lagoon and the tidal flat environments also contain Retiophyllia, a hexacoral, and diverse foraminifers (involutinids - Aulotortidae, Triasininae), and algae (dasyclads, halimedaceans, udoteaceans, rhodophytes, cyanophyceans and microproblematicae).

The presence of Triasina hantkeni Majzon, which is considered Rhaetian in age throughout the whole Triassic Tethyan (De Castro, 1990), is a very good marker for the Norian - Rhaetian boundary.

In the lagoon, the dasycladalean with large, cylindrical well calcified thalli, such as Griphoporella curvata (Gümbel), Heteroporella zankli (Ott) and Macroporella retica Zanin Buri, are common. Griphoporella curvata and Macroporella retica also very often occur as broken thalli. The above mentioned species could be considered as index species for the Rhaetian.

The presence of green siphoneous algae, such as halimedaceans (Halimeda helladica n. sp., H. discreta n. sp., Felixporidium triasicum n. sp., Hydracara kubeae n. g. n. sp.) and udoteaceans (Pseudopenicillus aegaeicus Dragastan et al., 1997), which have segmented thalli, represent a characteristic Rhaetian algal microflora (Fig. 3)

The red algae Metasolenopora semiconcentrica n. sp. and Parachaetetes rhaeticus n. sp. have a mainly Rhaetian stratigraphic range.

PALEOALGOLOGICAL DESCRIPTION (The algae are introduced and described by Ovidiu Dragastan)

Phylum CYANOPHYTA Family RIVULARIACEAE

Genus Rivularia C. Agardh ex. Bornet & Flahault, 1886 (see remarks of P. C. Silva, in Taxon 29, 1, 1980)

Rivularia lemaitrae (Dragastan, 1989 non 1969) Dragastan, 1985

Pl. 1, Figs. 5 – 6 1969 Cayeuxia lemaitrae n. sp. – Dragastan in Diaconu & Dragastan, pl. I , fig. 8 1983 C. lemaitrae Schäfer & Senowbari - Daryan, Taf. 1, Fig. 6 1985 Rivularia lemaitrae (Dragastan), p. 110, pl. I, Figs. 4 - 10

Lectotypification in 1989, Rev. Roum. Géol., Géophys., Géogr., Géologie, t. 33, Bucharest

Paratypes: Carnian (Sample Lo30), Norian (Sample Lo2), Hydra, Greece, Coll. L. P. B. V, No. 1051

Description: Thalli medium size (width 2. 10 - 2. 40 mm and high 1. 80 - 2. 0 mm) built by superposed hemispherical bushes, sometimes with growth zones (Pl. 1, Fig. 5). The thalli are crossed by pseudobifurcate tubes with diameters of 70 - 90 ηm and a reduced angle of divergence (50).

Remarks: Carnian thalli show growth zones and in the Norian occuras bushes; sometimes broken in hemispherical thalli debris.

Genus Polytrichella Schäfer & Senowbari - Daryan,

1983

Polytrichella fructicosa Schäfer & Senowbari - Daryan, 1983

Pl. 1, Figs. 1 - 4 1983 Polytrichella fructicosa n. g., n. sp. Schäfer & Senowbari - Daryan, p. 106 - 107, Taf. 2, Fig. 1, 2 and 8

Paratype: Norian (Sample Lo12), Hydra, Greece, Coll L. P. B. V, No. 1052

Description: Thallus bush - like (Pl. 1, Fig. 3), crossed by fine V - shaped dichotomic, tubes disposed in bundles with microsparite sheaths difficult to observe. Width of thallus 2.50 - 2.75 mm, height 1. 50 - 1. 80 mm. Tube – bundles are 60 - 120 µm in width and comprise 5 - 6 tiny tubes with diameters of 10 - 20 µm (Pl. 1, Fig. 4). The thallus is crossed by “voids” which correspond to boring organisms.

Phylum CHLOROPHYTA

Class BRYOPSIDOPHYCEAE Round, 1963 Order BRYOPSIDALES Schaffner, 1922

Soborder BRYOPSIDINEAE Hillis - Colinvaux, 1984 Family PSEUDOUDOTEACEAE Dragastan et. al., 1997

Ancestral - ecorticatae Genus Hedstroemia Rothpletz, 1913 Hedstroemia klausi Dragastan, 1989

Pl. 2, Fig. 1 1989 Hedstroemia klausi n. sp. Dragastan, p. 424, pl. 6, fig. 1-2

Paratype: Rhaetian (Sample Lo29/3), Hydra, Greece, Coll. L. P. B. V. No. 1053

Description: Thallus fan - like, small; 2. 5 mm wide and 1. 20 mm high. Thallus crossed by dichotomic siphons with a proximal diameters of 75 - 80 µm that an large distally to 100 - 110 µm. Siphon walls, smooth, nonperforate.

Genus Garwoodia Wood, 1941

Garwoodia polytomica Dragastan, 1989 Pl. 1, Fig. 7

1989 Garwoodia polytomica n. sp. Dragastan, p. 436, pl. 10, fig. 1-2

Paratype: Norian (Sample Lo2), Hydra, Greece, Coll. L. P. B. V. No1054

Description: Thallus hemispherical, width 6. 0 mm, height 3. 0 mm. Thallus consists of polytomic (4) branched siphons disposed at an angle of 900. Siphons 72 - 90 µm in diameter.

Garwoodia maxima Dragastan, 1989

Pl. 2, Figs. 8 - 10 1989 Garwoodia maxima n.sp. Dragastan, p. 443, pl 12, fig. 1-2

Paratypes: Carnian (Sample Lo20), Norian (Sample Y2), Hydra, Greece, Coll. L. P. B. V. No. 1055, No. 1056

Description: Fan - like thallus, 5. 0 mm width and 4. 0 mm high, consisting of 3 - 6 branched siphons. Siphons branch at an angle of 900.

ECORTICATAE

Genus Pseudoudotea Dragastan et. al, 1997 Pseudoudotea magna Dragastan et. al, 1997

Pl. 2, Figs. 2 - 6 1997 Pseudoudotea magna n. g., n. sp. Dragastan et. al., p. 108, Text - Fig. 7b, Pl. 8, Figs. 1- 10

Paratypes: Norian (Sample Lo20), Hydra, Greece, Coll. L. P. B. V, No. 1057 (exception Pl. 2, Fig. 4 which represents Isotype figured in 1997)

Description: Thallus hemispherical, width 6. 0 - 6. 6 mm, height of 3. 0 mm, composed of 1 or 3 fan - like blades. The blades show diffuse growth - zones and are crossed by tubular, dichotomic-branched siphons with a thick calcareous sheath crossed by pores. The siphons have a proximal diameter of 30 - 36 µm and a distal


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diameter of 84 - 90 µm. Sheath thickness is 12 - 24 µm. Remarks: The species is a marker for the Early

Norian occuring frequently in this interval.

Genus Hydraea Dragastan et al., 1997 Hydraea calcarata Dragastan et al., 1997

Pl. 2, Fig. 7 1997 Hydraea calcarata n. g., n. sp. Dragastan et al., p. 110, Pl. 9, Fig. 6 - 10

Paratype: Norian (Sample Lo20), Hydra, Greece, Coll. L. P. B. V, No. 1058

Description: Broken blade-like thallus with width of 1.50 mm and height of 0. 90 mm, having small marginal lobes margins. The blade is crossed by small cylindrical siphons with diameters of 36 - 48 µm. The sheath - wall is well-calcified with very fine pores.

Suborder HALIMEDINEAE Hillis - Colinvaux, 1984

Family HALIMEDACEAE Link, 1832 Genus Halimeda Lamouroux, 1812

Halimeda helladica n. sp. Pl. 3, Figs. 1 - 2

Derivatio nominis: “helladica” from Greece Holotype: Pl. 3, Fig. 1, Rhaetian (Sample Lo29),

Hydra, Greece, Coll. L. P. B. V, No. 1059 Isotype: Pl. 3, Fig. 2, Rhaetian (Sample Lo29/2),

Hydra, Greece, Coll. L. P. B. V, No. 1060 Diagnosis: Thallus composed of cylindrical

segments crossed by strongly calcified medullary siphons, dichotomically branched; short and angularly disposed. The cortex system is formed by 3 series of utricle - siphons: primary simple, short vesiculiferous - bulbous; second long, dichotomic cylindrical – curvate; and third short, cylindrical dichotomic. The medulla clearly is differentiated from the cortex.

Description: Thallus built by cylindrical segments. The contact or node area between segments is strongly

calcified. The segments have an uneven cylindrical shape, crossed by a large medulla which shows cylindrical, dichotomic siphons, short and inflated at the points of branching, and angularly disposed between them. The transition from medulla to cortex is marked clearly by microcrystalline calcite.

The cortex is also well-calcified and is crossed by 3 series of utricle - siphons. The primary siphons are simple, vesiculiferous bulbous, followed by second, long cylindrical - curvate dichotomic siphons, and third short, cylindrical siphons which are also dichotomic.

The calcification shows differential preservation: the medulla in microcrystalline calcite and the cortex strongly calcified (Pl. 3, Fig. 2).

Dimensions in mm: Length of segment (L) = 3. 6mm; Diameter of segments (D) = 1.50 - 2.0mm; diameter of medulla (dm) = 0. 50 - 0. 60; diameter of the medullary siphons (dms) = 0. 050 - 0. 12; thickness of the cortex (thc) = 0. 50 - 0. 70; diameter of cortical siphons: primary (dcs1) = 0. 050 - 0. 060; secondary (dcs2) = 0. 020 - 0. 040; tertiary (dcs3) = 0. 020; length of the cortical siphons: primary (lcs1) = 0. 10 - 0. 12; secondary (lcs2) = 0. 16 - 0. 17; tertiary (lcs3) = 0. 045 - 0. 060.

Remarks: The new species can be compared with Halimeda marondei (Flügel, 1988) from the Late Triassic of Thailand. The species had two kind of segments, large and small with different modes of preservation of the medulla and the cortex, which in our opinion belong to two different taxa. The utricle - siphons with 3 series have a different shape and disposition, and belongs to the vesiculiferous - bulbous second evolutionary - line of the Halimedaceae (Fig. 5).

The Thailand species do not belong to the genus Boueina Toula which has cylindriform primary utricle -siphons and corresponds to the first evolutionary - line of the Halimedaceae, which evolved in the Ordovician (Fig. 4).

Figure 4 - Characteristic features of the cortical system within the first evolutionary line of the Halimedaceae (1-2. Dimorphosiphon – Aphroditicodium after Elliott, 1982; 3. Halimeda goreaui, H. copiosa; 4. H. opuntia f. trilobata, after Littler & Littler, 1997 and Littler & Littler, in press).


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Figure 5 - Characteristic features of the cortical system within the second evolutionary line of the Halimedaceae

during the Permian - Cretaceous.


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Generally speaking the cortex system is essential for species recognition. Without information concerning the dimensions, length and diameter of the siphons no effective comparison is possible.

The new species belongs to the second evolutionary - line of the Halimedaceae, but in comparison with the Permian taxa Saxonia pygmaea Gebhard & Schneider, 1993, Tauridium and Halimeda soltanensis Poncet, 1989, the cortical system is more advanced. The Permian taxa aslo have 3 series of utricular-siphons, with the exception of Tauridium which has additional series (?).

The Middle Jurassic Leckhamptonella llewellyae Elliott, 1982 (Elliott, 1982) shows another cortical system having 3 or 4 series of utricular siphons (2 vesiculiferous and 2 cylindrical).

The Cretaceous species of Halimeda (H. camenitzae Dragastan & Bucur, 1979; H.fluegeli Bucur, 1994; Banatea bucurii n. sp.; H. paucimedullaris Schlagintweit & Ebli, 1998 and H. agharkari Badve & Nayak, 1983) have a transitional pattern of primary utricular - siphons, from vesiculiferous to subconical - elongate, cylindrical and additional series (3, 4, 5 series).

Halimeda discreta n. sp.

Pl. 3, Figs. 3 - 6 Derivatio nominis: “discreta” from the delicate inner

structure of the thallus. Holotype: Pl. 3, Fig. 3, Rhaetian (Sample Lo29/1),

Hydra, Greece, Coll. L. P. B. V, No. 1061. Isotypes: Pl. 3, Figs. 4 - 6, Rhaetian (Sample

Lo29/1), Hydra, Greece, Coll. L. P. B. V, No. 1062, No. 1063, No. 1064.

Diagnosis: Thallus small, delicate, discrete, built by tiny cyilindrical segments. The segments are crossed axially by a narrow medullary area with fine, cylindrical dichotomically branched siphons disposed vertically, parallel and linear in the axial - medulary zone. Cortex very thin, more calcified in comparison of the medulla.

The cortex is composed by 3 series of utricular - siphons: primary (simple, vesiculiferous) and secondary and tertiary (fine, cylindrical and dichotomically branched) (Fig. 6).

Description: Thallus very small, composed of tiny slender cylindrical, segments, ranging in length from 1. 2 - 1. 5 mm. In axial - longitudinal sections (Pl. 3, Figs. 3 - 5), segments have a different cylindrical shape, with “constrictions” crossed by a narrow medullar area. The medullary siphons are long, very thin, and dichotomically branched, with a parallel - vertical- linear disposition.

The cortex is more heavily calcified, and is crossed by very fine siphons with 3 series of utricles. The primary cortical siphon are simple, small, and vesiculiferous – bulbous, followed by a secondary long, cylindrical dichotomically branched siphons and tertiary short, cylindrical and also dichotomic siphons.

The cortical utricular - siphons have a very small diameter. The cross section (Pl. 3, Fig. 6) is circular, having a medulla preserved in microcrystalline calcite with dispersed medullary - siphons. The cortex correspond to the 3 series of utricular - siphons very fine and small in diameter.

Dimensions in mm: L = 1. 2 - 1. 5; D = 0. 36 - 0. 66; dm = 0. 12 - 0. 30; dms = 0. 012 - 0. 015; thc = 0. 12 - 0. 14; diameter of cortical siphons: dcs1 = 0. 020 - 0. 024; dcs2 = 0. 010; dcs3 = 0. 006 - 0. 008; length of cortical siphons: lcs1 = 0. 030 - 0. 036; lcs2 = 0. 070 - 0. 072; lcs3 = 0. 020 - 0. 024.

Remarks: Halimeda discreta n. sp. from the Rhaetian belongs to the second evolutionary line of the Halimedaceae with simple vesiculiferous primary cortical siphons. The new species can be compared with H. nana Pia, 1932, but differs in the medullary and cortical utricular - siphons. Comparison with other Cenozoic species of Halimeda is very difficult due to lack of clear descriptions of the cortical systems and shape and dimension details (see H. eocaenica, H. praeopuntia, etc). Consequently, all Cenozoic species need revision for correct identifications and comparisons to be made.

Genus Felixporidium Dragastan, 1999

Felixporidium triasicum n. sp. Pl. 4, Figs. 1 - 3

Derivatio nominis: “triasicum” from the Triassic. Holotype: Pl. 4, Fig. 1, Rhaetian (Sample Y2), Hydra,

Greece, Coll. L. P. B. V, No. 1065 Isotypes: Pl. 4, Figs. 2 - 3, Rhaetian (Sample Y1 and

Y2), Hydra, Greece, Coll. L.P.B.V, No. 1066, No. 1067. Diagnosis: Thallus hemispherical, built by ovoidal

blades. The thallus is crossed by a narrow round medullary axial - area which continued into the blades. The blades have a cortex system with primary simple, subvesiculiferous siphons, followed by dichotomically branched secondary cylindrical siphons.

The tertiary and fourth series of siphons, are also dichotomic branched (Fig. 7) and have a cylindrical shape.

Description: Thallus built by ovoid - blades (segments) disposed bilaterally along the main medullary siphonal area (Pl. 4, Fig. 1). Each blade has cylindrical connective, medullary siphons from which the cortical series of utricular - siphons continued. Sometimes the blades are found disarticulated and isolated from the main thallus. The strongly calcified cortex is crossed by primary, simple subvesiculiferous siphons. The secondary, tertiary and fourth series of utricular siphons are cylindrical, dichotomically branched and long or short.

In cross section the thallus has more or less a circular outline with small lobes or protuberances; the axial part is crossed by a narrow, medullary area and has a thick cortex with 4 series of utricular - siphons.

Dimesions in mm: Width of thallus (W) = 3. 0 - 3. 2; height of thallus (H) = 4. 5 - 4. 8; width of the blade -

Figure 6 - Reconstruction of the cortical system of Halimeda discreta n. sp.. (1, 2 ,3 = cortical system)


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segment (w), basal part = 0. 36 - 0. 40; height of the blade segment (h) = 1. 0 - 1. 20; dm = 0. 060 - 0. 080; cortex system: dcs1 = 0. 040 - -. 080; dcs2 = 0.024 - 0.040; dcs3 = 0.012 - 0. 20; dcs4 = 0.010 - 0.015; length of cortical siphons: lcs1 = 0. 20 - 0. 30; lcs2 = 0. 10 - 0. 15; lcs3 = 0. 080 - 0. 10; lcs4 = 0. 040 - 0. 060.

Remarks: The new species is different from Felixporidium atanasiuii Dragastan, 1999 and F. alpidicum Dragastan, 1999 from the Late Jurassic - Early Cretaceous by the structure of cortex system with 4 series of utricular - siphons, of which the primary series is subvesiculiferous and the other three are cylindrical, dichotomic, and long or short.

Hydracara n. g.

Derivatio nominis: from Hydra island and the latin “cara” which means a plant.

Diagnosis: Thallus built by long cylindrical, segments crossed axially by a narrow medullary area with few medullary siphons, that are dichotomic branched.

The cortex is well-calcified and has simple, ampuliform siphons (Fig. 8), which in the distal part may be dichotomic, but this feature difficult to observe.

Type genus: Hydracara kubeae n. g., n. sp. Remarks: The thallus segments, crossed by

dichotomically branched medullary siphons and with a simple cortex, resemble the Recent genus Codium Stockhouse. From this point of view the systematic position in the Family Halimedaceae remains questionable.

Comparing with genus Egericodium Flügel, Velleditis, Senowbari - Daryan & Riedel, 1991/1992 from the Ladinian - Carnian of Hungary, the only resemblance is the presence of a narrow medullary area. In genus Egericodium the cortex is very thick, but unfortunately a real description of the cortical siphons was not presented.

The genus Collareocodium Brandner & Resch, 1980 from the Ladinian Wettestein Limestone north of Innsbruck shows a cylindrical, dichotomically branched thallus with irregular segmentation and constrictions, sometime connected with collar - like concentric rings. The thallus is crossed by a narrow medullary zone and has a well-described cylindriform cortical system. This genus belongs to the first evolutionary - line of the Halimedaceae and is distinct from the new taxon.

The genus Nipponophycus Yabe & Toyama, 1928 from the Late Jurassic also has a branched cylindrical thallus and crossed by a narrow medullary zone. The cortex has only 2 or 3 series of utricles: the primary siphons are vesiculiferous in shape.

Hydracara kubeae n. sp.

Pl. 3, Figs. 7 - 8 Derivatio nominis: Species dedicated to Bärbel Kube

from Ruhr University Bochum, who studied many geological sections of Hydra Island.

Holotype: Pl. 3, Fig. 7, Rhaetian (Sample Lo29), Hydra, Greece, Coll. L. P. B. V, No. 1068.

Isotypes: Pl. 3, Fig. 8, Rhaetian (Sample Lo29/2), Hydra, Greece, Coll. L. P. B. V, No. 1069.

Diagnosis: Thallus composed of long, cylindrical segments crossed by a narrow medullar zone with few, cylindrical dichotomically branched siphons. The strongly calcified cortex has a simple utricular system. The cortical siphons are ampuliform, simple and probably with an incision at the distal end.

Description: Thallus formed by long, cylindrical segments having some irregularities, such as small constrictions. In vertical section (Pl. 3, Fig. 7) the segment show a narrow medullary zone crossed by cylindrical dichotomically branched siphons, that are more or less parallel and few as number. The thick cortex is pierced by simple, primary ampuliform siphons, possible with an incision at the distal end (Fig. 8). In cross section the thallus segments are round and in some areas show the ampuliform siphons, that are small in diameter at the distal end (Pl. 3, Fig. 8).

Dimensions in mm: L = 2. 70 - 3. 30; D = 0. 55 - 0. 90; dm = 0. 25 - 0. 30; dms = 0. 020 - 0. 070; thc = 0. 20 - 0. 30; dcs1 - proximal = 0. 10 - 0. 12, distal = 0. 070 - 0. 085; lcs1 = 0. 20.

Remarks: The new species differs in the inner morphology of the medullary and cortical zones from Collareocodium oenipontanum Brandner & Resch, 1980 (Ladinian), Egericodium hungaricum Flügel, Velleditis, Senowbari - Daryan & Riedel, 1991/1992 (Ladinian - Carnian reefs) and Nipponophycus ramosus Yabe & Toyama, 1928 (Late Jurassic).

The new taxon is close to the Recent genus Codium in its simple utricular system.

Figure 7 - Reconstruction of the cortical system of Felixporidium triasicum n. sp.. (ma = medullar siphon; 1, 2, 3 ,4 = cortical system)

Figure 8 - Reconstruction of the cortical system of Hydracara kubeae n. g., n. sp.


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Class CHLOROPHYCEAE Order DASYCLADALES

Family TRIPLOPORELLACEAE Pia, 1920 Tribe SALPINGOPORELLEAE

Subtribe SALPINGOPORELLINAE (Systematics after Berger & Kaever, 1992)

Genus Uragiella Pia, 1922 Uragiella minima n. sp.

Pl. 4, Figs. 4 - 5 Derivatio nominis: “minima” from the small size of

the thallus. Holotype: Pl. 4, Fig. 4, Rhaetian (Sample Y2/2),

Hydra, Greece, Coll. L. P. B. V, No. 1070 Isotype: Pl. 4, Fig. 5, Rhaetian (Sample Y2), Hydra,

Greece, Coll. L. P. B. V, No. 1071 Diagnosis: Thallus small, cylindrical crossed by large

axial hollow. The ramifications are only primary, euspondylous, uragielliform being narrow in both the proximal and distal parts. In the middle, they widen sharply. The shape of the ramifications is very close to the acrophorous type.

Description: Thallus cylindrical having a large axial hollow. In axial longitudinal sections the thallus shows euspondyle verticils, with primary ramifications uragielliform. The ramifications have a distinctive shape: narrowing to the proximal part after widening sharply, and continuing to became narrow to the distal part.

Dimensions in mm: Length of thallus (L) = 2. 80; diameter of thallus (D) = 1. 0 - 1. 20, diameter of the axial siphons (d) = 0. 56 - 0. 60; diameter of the ramification: proximal = 0. 10 - 0. 18, distal = 0. 10 - 0. 15; in the middle = 0. 050 - 0. 070, length of ramification = 0. 20 - 0. 22.

Remarks: The new Late Triassic species is very small and differs from Uragiella supratriassica Bystricky, 1967, U. liasica Lebouche & Lemoine, 1963 and U. suprajurassica (Gümbel, 1891) Pia, in the shape of its ramifications and thallus dimensions. Uragiella ragusina Sokac, 1992 from the Dogger also has a small thallus, but differs from the new species by “pronounced species - specific characteristic, the variable shape of the ramifications which can be globular, sack or bag shaped or similar to the pyriform type”(Sokac, 1992).

Phylum RHODOPHYTA

Class RHODOPHYCEAE Family SOLENOPORACEAE

Genus Metasolenopora Pia, 1930 Metasolenopora semiconcentrica n. sp.

Pl. 4, Figs. 6 - 8 Derivatio nominis: “semiconcentrica” from the

partially concentric disposition of tube - cells in the thallus.

Holotype: Pl. 4, Fig. 6, Rhaetian (Sample Lo29 b), Hydra, Greece, Coll. L. P. B. V, No. 1072

Isotype: Pl. 4, Fig. 7 - 8, Rhaetian (Sample Lo29 b, Y2/2), Hydra, Greece, Coll. L. P. B. V, No. 1073, No. 1074

Diagnosis: Thallus more or less nodular - hemispherical in shape composed of tube - cells with few horizontal partitions (2 - 3) at different level along the tubes. In cross - section the tube - cells have a semiconcentric disposition.

Description: Thallus hemispherical – nodular, formed by tube - cells straight in a compact structure. The tube - cells have few horizontal partitions (2 - 3) along the tube. The semiconcentric disposition is a characteristic feature of the species that is only observed in cross - sections (Pl. 4, Figs. 6 - 7).

Dimensions in mm: W = 3. 0 - 4. 0; H = 2. 20 - 2. 50; diameter of the tube - cells = 0. 024 - 0. 050, number of horizontal partitions along tube - cells = 2 - 3.

Remarks: The genus Metasolenopora was introduced by Pia (1930) for solenoporacean algae which have few horizontal partititions and tube - cells in concentric rows, being reevaluated by Maslov (1961), Poignant (1991). The species Solenopora endoi Flügel, 1975, S. styriaca Flügel, 1960, S. simionescui Dragastan, 1989 from the Triassic may or not have horizontal partitions, and do not have a concentric arrangement of tube - cells.

Parachaetetes Deninger, 1906 Parachaetetes rhaeticus n. sp.

Pl. 4, Figs. 9 - 11 Holotype: Pl. 4, Fig. 11, Rhaetian (Lo29 a), Hydra,

Greece, Coll. L. P. B. V. No. 1075 Isotypes: Pl. 4, Fig. 9 - 10, Rhaetian (Lo29 b), Hydra,

Greece, Coll. L. P. B. V. No. 1076, No. 1077 Diagnosis: Thallus small, flabelliform, formed by very

small tube - cells having many horizontal partitions disposed more or less at the same level.

Description: Thallus flabelliform, small, sometimes flaring toward the margin. Tube - cells very small, and present many horizontal partitions more or less at the same level (Pl. 4, Fig. 11)

Dimensions in mm: W = 0. 50 - 0. 60; H = 0. 26 - 0. 30; diameter of the tube - cells = 0. 012 - 0. 020; distance between partitions = 0. 024 - 0. 040.

Remarks: The new species is distinguished by its small thallus, horizontal partitions and semiconcentric disposition of tube – cells from Parachaetetes maslovi Flügel, P. triasinus (Vinassa de Regny) and P. mariae (Moisceev) from the Middle and Late Triassic, which have more partitions and lack concentric disposition of the tube - cells.

The evolutionary - lines of halimedaceans during the Phanerozoic

In spite of the difficulties of thallus preservation, the record fossil halimedacean evolution can be traced from the Ordovician, and possibly from the Cambrian, to the Pleistocene. In the Recent there are over 30 species of Halimeda.

The thallus of Halimeda is composed by two parts: the holdfast and the erect - segmented part of the thallus. Monographic studies of Recent Halimeda have been made by HILLIS - COLINVAUX (1959, 1980, 1986, 1991), LITTLER & LITTLER (1997) and LITTLER & LITTLER (in press).

A few macroscopic characters are helpful in taxonomic determination of species: “the appearance of the plant as a whole of the holdfast system and the shape of the majority of the segments… Microscopic taxonomic characters which are most used are: pattern of medullary siphons at the node and pattern and extent of cortex. All these characters vary somewhat with age and the position of the segment on the thallus. The different pattern of nodal siphons found in different groups of Halimeda species are useful in delimiting groups of species and are considered to reflect fundamental phyletic divergence” (HILLIS - COLINVAUX, 1980).

In a synthesis of “Udoteaceae”, BASSOULLET et. al. (1983) commented on the above characteristics in delimiting the species of Halimeda as follows: “It is difficult to delimit fossil species by external appearance of segments alone because the characters, as can be


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observed on living specimens, are extremely variable, even in one individual. Internal characteristics are important: mainly the aspect, arrangement and size of the medullary and cortical filaments”.

In the most of the cases from Ordovician to Palaeocene halimedacean thalli are composed of cylindrical segments, that are simple or very rarely branched, and sometimes a little flattened.

During the Eocene thalli - segments appeared that are cylindrical, compressed or flattened, cuneate, or reniform with the top margin entire or lobate.

The thalli - segments are crossed by a medullary zone with many or few(2 - 4; 4 - 6, over 6) siphon - series, that are vertically or axially disposed. The dichotomically branched siphons have a parallel arrangement between series of medullary siphons.

The medullary zone is surrounded by a cortex of variable thickness which is crossed by 2 to 7 series of utricular - siphons.

In our opinion the most important character for fossil identification, at species level remains the morphology of the cortical system. The cortical system is crossed by siphons of variable shape and number of series. These are very important for delimiting the species.

The concept of the utricular system as a main character for species recognition was introduced recently by LITTLER & LITTLER, 1997 and LITTLER & LITTLER, in press.

Many authors over the years have introduced new taxa of fossil Halimedaceae. Some of these taxa lack a complete description that includes, diameter and length measurements of siphons, and number of siphon - utricular series. This has diminished the utility of these taxa, making correct identification or comparison difficult.

Taking into account characteristic of halimedacean taxa described from the Palaeozoic - Mesozoic and part of the Cenozoic, two evolutionary lines can be distinguished.

The first evolutionary line includes all taxa having cylindriform primary utricular siphons in the cortex system, and evolved from Ordovician until Recent. The line begins with multibranched siphons (1 - 4) of the genera Dimorphosiphon - Palaeoporella (Ordovician) and continued until the Permian with the genus Aphroditicodium (Fig. 4).

During the Mesozoic, the presence of cylindriform primary siphons in the cortex was observed in the genera Collareocodium (Middle Triassic), Boueina (Late Triassic - Cretaceous) and Arabicodium (Jurassic - Cretaceous). Some species of the genus Arabicodium which do not preserve the medullary siphons must be transfered to the genus Pseudopenicillus (paper in preparation). Within this line during the Cretaceous – Cenozoic, species of Halimeda appeared that maintained the same character of cylindriform primary siphons such as H. robusta, H. corneola, H. densituba, and H. triradiata described by BADVE & KUNDAL.

Species of Halimeda described from the Turonian - Eocene (KUSS & HERBIG, 1993) to Pleistocene have not been included in this scheme, because many of the taxa lack a clear description of the cortical system, e.g. H. elliotti, H. praeopuntia and H. praemonilis. Many of the above mentioned species must be revised.

During the Pliocene - Pleistocene - Recent interval this line is continued with the same character

(cylindriform primary siphons) in the taxa Halimeda goreaui, H. copiosa, H. opuntia f. triloba, and H. opuntia f. opuntia (Fig. 4).

The second evolutionary line corresponds to the taxa having vesiculiferous - bulbous, primary cortical siphons. This line begins in the Permian with Saxonia, Tauridium and Halimeda soltanensis. The cortical system is formed by simple, vesiculiferous primary siphons continued by secondary and tertiary dichotomic branched siphons, or having more utricle siphons series as in Tauridium and Halimeda soltanensis (Fig. 5). Saxonia pygmaea shows a combination between primary vesiculiferous siphons and secondary and tertiary cylindrical siphons. Unfortunately this taxon is invalid because the authors designated two holotypes for the species.

During the Mesozoic this combination of vesiculiferous and cylindrical siphons in the cortical system is observed in Halimeda marondei, H. helladica n. sp. and H. discreta n. sp., all species from the Late Triassic.

The Middle Jurassic genus Leckhamptonella maintains these characters, but with the difference of increasing number of utricular - siphons (4 series) in the cortex.

During the Early Cretaceous, cortical systems preserve the same characteristics. In H. camenitzae the primary siphons became a little conical in shape and have 3 series of utricles; also in Banatea bucurii n. g., n. sp. The species H. fluegeli shows 4 series of utricular - siphons, 2 vesiculiferous and 2 cylindrical.

The multibranched cortex system (1 – 5) is shown by H. paucimedullaris in the Late Cretaceous and by H. agarkari near the Cretaceous / Cenozoic boundary. At this boundary some species of Halimeda have a tendency to change the shape of the primary siphons to conical - elongate, as in H. agarkari and H. chiplonkari. The latter species only has 1 - 3 series of utricular - siphons. Due to incomplete descriptions the Cenozoic taxa of Halimedaceae were not introduced in this scheme.

The transitional pattern from vesiculiferous to conical - elongate primary siphons is observed also in Recent species, such as Halimeda incrassata, H. scabra, H. simulans, H. tuna f. platydiscus.

In conclusion, analysis of the cortical system indicates the presence of two evolutionary lines within the Family Halimedaceae:

• an older one (Ordovician - Recent) represented by cylindriform primary siphons

• a younger one (Permian - Recent) having vesiculiferous - bulbous, primary siphons. These different morphologies of cortical siphons

during the Phanerozoic, developing from simple series of utricles (1 - 2) to multibranched series of utricles (1-3; 1-4; 1-5; 1-7) reflect functional strategies for obtainning nutrients.

Acknowledgements Thanks are due to Prof. dr. Jan Veizer, Head of the

Geological Department, Ruhr - Universität Bochum for financial support. For technical assistance we thank Mr. Ress (Ruhr Univ. Bochum) and Mr. Dumitrescu (University of Bucharest).


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PLATES Plate I - Kap Kastello, Hydra, Greece. Figs. 1-4. Polytrichella fructicosa SchÄfer & Senowbari, 1983. 1, entire thallus; 2-4, part of thallus crosed by

microborings (v) filled by coarsely crystalline calcite, Norian. Figs. 5-6. Rivularia lemaitrae (Dragastan, 1989 non 1969), vertical section, Carnian–Norian. Fig. 7. Garwoodia polytomica Dragastan, 1989, oblique vertical section, Norian. Fig. 8. Hydracara kubeae n. g. n. sp. and Triasina hantkeni, Rhaetian. Plate II - Kap Kastello, Hydra, Greece Fig. 1. Hedstroemia klausi Dragastan, 1989, vertical axial section; dichotomic siphons expanding distally, Rhaetian. Figs. 2-6. Pseudoudotea magna Dragastan et al., 1997; 2, entire thallus; 3-6, varionsly orientated section, siphons

from original, 1997, Early Norian. Fig. 7. Hydraea calcarata Dragastan et al., 1997, axial vertical section, small cylindrical dichotomic siphons, Norian. Figs. 8-10. Garwoodia maxima Dragastan, 1989, axial vertical and oblique cross-section, 3 to 6 siphons branch at an

angle of 900, Norian. Plate III - Kap Kastello, Hydra, Greece Figs. 1-2. Halimeda helladica n. sp. 1, holotype, axial vertical section, Rhaetian, Coll. L.P.B.V No.1059; 2, isotype,

Rhaetian, Coll. L.P.B.V No. 1060. Figs. 3-6. Halimeda discreta n. sp. 3, holotype, axial vertical section, Rhaetian, Coll. L.P.B.V No.1061; 4-6, isotypes, Rhaetian, Coll. L.P.B.V No. 1062, 1063, 1064.

Figs. 7-8. Hydracara kubeae n. g. n. sp. 7, holotype, axial vertical section, Rhaetian, Coll. L.P.B.V No.1068; 8, isotype, cross section, Rhaetian, Coll. L.P.B.V No. 1069.

Plate IV - Kap Kastello, Hydra, Greece Figs. 1-3. Felixporidium triasicum n. sp. 1, holotype, vertical oblique section, Rhaetian, Coll. L.P.B.V No. 1065; 2-3,

Isotypes, vertical and cross section of blade-segments, Rhaetian, Coll. L.P.B.V No. 1066 and No. 1067. Figs. 4-5. Uragiella minima n. sp. 4, holotype, vertical oblique section, Rhaetian, Coll. L.P.B.V No. 1070; 5, isotype,

Rhaetian, Coll. L.P.B.V No. 1071. Figs. 6-8. Metasolenopora semiconcentrica n. sp. 6, holotype, vertical section, Rhaetian, Coll. L.P.B.V No. 1072; 7-8,

isotypes, Rhaetian, Coll. L.P.B.V No. 1073 and No. 1074. Figs. 9-11. Parachaetetes rhaeticus n. sp. 11, holotype, vertical section, Rhaetian, Coll. L.P.B.V No. 1075; 9-10,

isotypes, Rhaetian, Coll. L.P.B.V No. 1076 and No. 1077. .

new late triassic calcareous algae from hydra,...

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