Dasycladales Algae from the Norian-Rhaetian Reef Carbonates of Argolis Peninsula, Greece

ACTA PALAEONTOLOGICA ROMANIAE (2014) V. 10 (1-2), P. 39-46

DASYCLADALES ALGAE FROM THE NORIAN-RHAETIAN REEF CARBONATES OF ARGOLIS PENINSULA, GREECE

Baba Senowbari-Daryan

Received: 10 March 2014 / Accepted: 30 July 2014 / Published online: 29 September 2014

Abstract Three new dasycladales algae, including Probolocuspis sarmeikensis nov. sp. Probolocuspis? tenuipora nov. sp. and Diplopora obliguspora nov. sp. are described from Mavrovouni Mountains (Argolis Peninsula, Greece). The specimens were found embedded in micritic sediments between the reef builders in the Norian-Rhaetian reef carbonates near the town of Sarmeika. The genus Probolocuspis was known from the Ladinian-Carnian of Iran and the Alps. Argolis Peninsula is the third locality, where this genus was found. The stratigraphic range of the genus should be extended from Ladinian to Norian (perhaps to Rhaetian?).

Keywords: Dasycladales, Probolocuspis, Diplopora, Norian, Rhaetian reef, Argolis, Greece.

INTRODUCTION

Norian-Rhaetian reef structures, so called „Dachsteinkalk-Reefs“ in the Alps, are known from several localities in tethyan realm (Alps, Sicily, Turkey, Iran, Pamir Mountains) and outside of the Tethys (USA, Canada). Such “Dachsteinkalk-Reefs“ also occur in Mavrovouni Mountains (Argolis Peninsula, Greece). Available information about the occurrence of Norian- Rhaetian reefs within the so called „Pantokrator- limestones“ in Greece are limited to few publications of e. g. Renz (1908), Bachmann & Risch (1979) and Flügel (1983). Detailed investigations about the facies and palaeontological content of these reefs are not carried out. An example for Norian-Rhaetian reef carbonates within the „Pantokrator-limestones“ are the reef structures and shallow-water carbonates in the Mavrovouni Mountains. These structures reach a size of about two Kilometers and are exposed near the town of Sarmeika, southeast of the town of Adami (Argolis Peninsula, compare Fig. 1). Matarangas et al. (1995) were the first authors, who reported the occurrence of a relatively large reef near Sarmeika. The age of organisms in this reef, especially of the corals was dated into Carnian by these authors. Senowbari-Daryan et al. (1996) and Senowbari-Daryan et al. (2002) described some typically Norian-Rhaetian reef organisms from the same reef complex near Sarmeika. Further investigations on reef communities confirmed the Norian-Rhaetian age of these reef limestones. Hypercalcified sponges, including chambered sphinctozoans, non-chambered inozoans, disjectoporids and chaetetids are the most abundant reef builders, followed by scleractinian corals and other reef organisms. The Upper Triassic to Lower Jurassic shallow-water carbonates of “Pantokrator-limestones” (Partsch, 1887; Fig. 1 Geographic position of the Norian-Rhaetian reef locality Renz, 1955; Flügel, 1983) are widely distributed in the north of the town of Sarmeika, Peloponnes, Greece. Adriatic-Ionic region and in the Argolis Peninsula, northeastern Peloponnes in Greece (Fig. 1). The main algal mats, known as “Loferitic facies”. This facies type facies of the „Pantokrator- limestones“ is represented by is known from several localities in northern and southern bedded limestones and dolomites indicating their parts of the Tethys (Alps: e. g. Fischer 1964; Enos & depositional environment in lagoon and in tidal flats with Samankassou 1998; Satterley 1996; Haas 1982; Sicily:

______Geozentrum Nordbayern, FG Palaeoumwelt, Loewenichstr. 28, 39 91054 Erlangen, Germany. [email protected] Baba Senowbari-Daryan

Catalano et al. 1974), and own observations from Oman. Upper Triassic “Loferitic facies” from the Argolis Peninsula were described by Vartis-Matarangas & Matarangas (1991), Pomoni-Papaioannou (2008), and Photiades et al. (2010).

Depository: The holo- and paratypes of described algae are deposited in the Geozentrum Nordbayern, FG Palaeoumwelt (Material: Senowbari-Daryan, Trias, Argolis).

SYSTEMATIC PALAEONTOLOGY

Devision Chlorophycophyta Papenfuss, 1955 Class Chlorophyceae Kützing, 1843 Order Dasycladales Pascher, 1931 Family Triploporellaceae (Pia, 1920) Genus Probolocupsis Brönnimann, Zaninetti, Moshtaghian & Huber, 1974 Type species: Probolocupsis espahkensis Brönnimann, Zaninetti, Moshtaghian & Huber, 1974 Further species: P. aculeata Nittel, 2006; P. sarmeikensis nov. sp.

Remarks: Brönnimann et al. (1974) described individual and disintegrated laterals of the thallus as “tooth-like”, “bifide” or “trifide” elements, if two or three laterals are still preserved together. The elements were compared with conodonts, but because of the calcareous skeleton they were interpreted as “tooth-like microfossil” (“incertae sedis”) by these authors. Senowbari-Daryan & Majidifard (2003) revealed the nature of these elements documenting them as disintegrated parts of the dasycladalean thallus of Probolocupsis. They gave following diagnosis to the thallus of Probolocupsis: “Cylindrical thallus with trichophorous laterals of first order only, with euspondyl(?) arrangement” (Senowbari-Daryan & Majidifard 2003: p. 108). In the meantime a second species of the genus with a clearly euspondyl arrangement of the laterals is described as Probolocupsis aculeata by Nittel (2006) from the Ladinian Wetterstein limestones of the Northern Calcareous Alps, Austria.

Probolocupsis sarmeikensis nov. sp. (Fig. 2; Fig. 4a-n) Derivatio nominis: Named from the town Sarmeika near the locality. Holotype: Fig. 4l (magnification in Fig. 2). Paratypes: Fig. 4a-k, m-n. Locus typicus: Norian-Rhaetian reef carbonates, north of the town of Sarmeika in the Mavrovouni Mountains (Fig. 1). Stratum typicum: Norian-Rhaetian reef carbonates. Diagnosis: Cylindrical thallus with a narrow axial cavity. Trichophorous laterals of the first order are usually amalgamated at the base appearing as a thick wall around Fig. 2 Probolocupsis sarmeikensis nov. sp. (Holotype). The the axial cavity. Euspondyl arrangement of the laterals. magnification from Fig. 4l shows the spine-like extended individual laterals (some of the internal pores: see arrows). Laterals are perpendicular to the thallus axis.

Dasycladales algae from the Norian-Rhaetian reef carbonates of Argolis Peninsula, Greece

Differential diagnosis: See comparison after the description of the species. Material: Numerous specimens in several thin sections.

Description: Almost all specimens of this cylindrical alga are cut in cross sections. Two specimens illustrated in Fig. 4c and the holotype Fig. 4l (H) exhibit the most characteristics of the alga in longitudinal section. The thallus of the holotype reaches a length of almost 10 mm with a diameter of 2.2 mm. The axial cavity of the holotype has a diameter of about 0.4 mm. Both longitudinal sections show that the laterals are arranged perpendicular to the axial cavity. The alga is characterized by a narrow axial cavity of 0.25-0.5 mm and a thick thallus wall. Ratio of axial cavity/thallus diameter varies between 11 % and 25% (see Tab. 1). Individual laterals appear spine-like on the thallus outside. Marginal sections of the laterals may appear as points or small white circles (Fig. 4e, g, k). Because of the recrystallization of the majority of specimens the base of laterals is not well recognizable. The well preserved specimen illustrated in Fig. 4j exhibits that the laterals are arisen as single from the axial cavity, but others show two or three laterals arisen from the common base (Fig. 4h). Each spine-like lateral contains a very narrow tube (Fig. 2; Fig. 4m: arrows). Length of laterals is usually 1 mm, their diameter about 0.08-0.15 mm, their number in each is 22-40. In the holotype as well as in the majority of specimens the recrystallization amalgamated the base of laterals, appearing as a thick wall around the axial cavity. Some biometrical data of Probolocupsis sarmeikensis nov. sp. are listed in Table 1.

Table 1. Some biometrical data of the thallus of Probolocupsis sarmeikensis nov. sp. D) diameter of the thallus, d) diameter of the axial cavity. “Specimen” indicates the number of the measured specimens of corresponding Plates and Figures. All measurements are in mm.

Specimen D d d/D Thin section Pl. 1, Fig. 3 2.0 0.4 20 % G05C Pl. 1, Fig. 8 2.4 0.36 15 % G05C Pl. 1, Fig. 8 2.5 0.38 15 % G05C Pl. 1, Fig. 12, 2.2 0.4 18 % G05C H Pl. 1, Fig. 12 2.2 0.4 18 % G05b Pl. 1, Fig. 12 2.4 0.4 17 % G05b Pl. 1, Fig. 5 2.6 0.32 12 % G05b Pl. 1, Fig. 1 3.2 0.34 11 % G05b Pl. 1, Fig. 2 2.4 0.45 19 % G05b Pl. 1, Fig. 2 2.0 0.4 20 % G05b Pl. 1, Fig. 7 2.0 0.5 25 % G05b Pl. 2, Fig. 4 1.0 0.25 25 % G12

Fig. 3 Diplopora obliqusipora nov. sp. (Holotype). The Comparison: The distinctly narrow axial cavity of magnification from Fig. 5c shows the relics of metaspondyle Probolocuspis sarmeikensis nov. sp. differs (without arrangement of the tufts (large arrows, also the cross sections of regard to the stratigraphic age) this species from the type the laterals at the base of the thallus indicate the metaspondyle species – Probolocupsis espahkensis Brönnimann et al. – arrangement of the laterals). Small arrows indicate the tufts and from P. aculeata Nittel clearly. Differences of the arising from the common base. The thin wall around the axial cavity is clearly recognizable.

Baba Senowbari-Daryan skeletal elements are additional criteria for distinguishing Table 2. Some biometrical data of the thallus of Probolocuspis? this species from the known species. tenuiparia nov. sp. All measurements are in mm. For Schlagintweit et al. (2013) illustrated in Fig. 3a some abbreviations see Table 1. dasycladalean specimens (maybe recrystallized) from the “Lower Gosau Subgroup of Austria” as Milanovicella Specimen D D d/D Thin hammudai (Radoičić 1975), which are very similar to section recrystallized specimens of P. sarmeikensis nov. sp. Pl. 2, Fig. 7 1.0 0.25 25 % G08a According to the diagnosis of Milanovicella, given by Pl. 2, Fig. 10 0.45 0.075 17 % G08a Granier & Berthou (1994) and compared with Pl. 2, Fig. 6 0.3 0.05. 17 % G08a illustrations of the type material of Radoičić (1975), Pl. 2, Fig. 9 0.4 0.07 18 % G08a described as Likanella hammudai and revised by Pl. 2, Fig. 9 0.4 0.12 30 % G08a Schlagintweit (1990) the species of Sarmeika can not be Pl. 2, Fig. 9 0.8 0.10 13 % G08a attributed to this genus. Pl. 2, Fig. 8 0.3 0.05 17 % G08a There are some morphological similarities between the Pl. 2, Fig. 5 0.4 0.05 13 % G08b species Neogyroporella? gawliki described from the Pl. 2, Fig. 5 0.5 0.07 14 % G08b Upper Jurassic-Lower Cretaceous of the Northern Pl. 2, Fig. 5 0.45 0.07 16 % G08b Calcareous Alps by Schlagintweit (2005, compare also Pl. 1, Fig. 5 0.36 0.08 22 % G08b Schlagintweit 2011) and P. sarmeikensis. The general appearance of P. sarmeikensis is similar to Family Diploporellaceae (Pia, 1920), Deloffre 1988 Teutloporella echinata, described by Ott (1975) from the Genus Diplopora Schafhäutl, 1863 Ladinian of the Lombardian Alps, northern Italy. The Type species: Diplopora annulata (Schafhäutl, 1853) individual laterals in P. sarmeikensis are arranged 1863 perpendicular to the axial cavity, but they are orientated Diplopora obliquspora nov. sp. oblique in T. echinata. Laterals are risen as single from (Fig. 3; Fig. 5a-d) the axial cavity (later they can be branched: Fig. 4d, h) in ? 1979 Clypeina sp.- Flügel, pl. 2, fig. 9. P. sarmeikensis, but they may be doubled? in T. echinata. ? 1980 Clypeina sp.- Senowbari-Daryan, p. 67, pl. 14, fig. In addition the shape of laterals in P. sarmeikensis is 5-6. always spine-like and their cross sections are always Derivatio nominis: Obliquus (latin) = oblique, porus circular or oval, but the shapes of laterals (see Ott 1975: (latin) = pore. Named for the oblique running laterals. fig. 3) and their cross sections in T. echinata are different. Holotype: Specimen illustrated in Fig. 5c (magnification Advanced interpretations of these features deserve a in Fig. 3, thin section G75). comparing study of the material from Alps and Argolis. Locus typicus: Norian-Rhaetian reef carbonates near the town Sarmeika (Fig. 1). Probolocuspis? tenuiparia nov. sp. Stratum typicum: Norian-Rhaetian reef carbonates. (Fig. 5e-l) Diagnosis: Cylindrical thallus with a relatively wide axial Derivatio nominis: Tenuis (latin) = thin, and paries (latin) cavity. Tufts of trichophorous laterals are originated from = wall. Named from the thin thallus wall of the species. the common base. Oblique orientation of the laterals. A Holotype: Fig. 5h. thin wall at the base of the thallus wall surrounds the Locus typicus: Norian-Rhaetian reef carbonates north of axial cavity. Metaspondyl arrangement of the laterals. the nearby town Sarmeika (Fig. 1). Material: Four specimens. Stratum typicum: Norian-Rhaetian reef carbonates. Diagnosis: Cylindrical thallus with thin wall and wide Description: The thallus of this cylindrical and tiny alga axial cavity. Laterals, corresponding to the thin thallus is composed of a relatively thick thallus wall. The length wall, are short. Individual laterals are visible only as of the holotype (Fig. 3) is about 8 mm with thallus small spines on the outer surface of the thallus. diameter (D) of 1.4 mm. Diameter of axial cavity (d) is Material: Numerous species in several thin sections. 0.6 mm (about 43%). Characteristic of the alga is the metaspondyl arrangement of the laterals as indicated by Description: Similar to the preceding described species the relict appearance of the tufts through the thallus wall most specimens of this alga are cut in cross sections. The (Fig. 3: large arrows). Each tuft is composed of about five thallus walls of all specimens are recrystallized, the laterals (in longitudinal section three are recognizable, nature of laterals is visible only as small spines on the Fig. 3: small arrows). outer surface of the thallus. The arrangement of the Laterals are oriented oblique to the axial cavity with laterals is not clear and therefore the attribution to the about 30-75° off-axis angle). genus Probolocupsis is uncertain. Around the axial cavity a thin wall of only 0.06 mm The specimen designated as holotype (Fig. 5h) is an thickness is pierced by the base of laterals. Diameter of oblique to longitudinal section exhibiting clearly the tip individual laterals is about 0.05 mm. Individual laterals of the needle- or spine-like individual laterals on one are extended spine-like over the thallus surface appearing side. Some specimens (Fig. 5e-f, j-k), cut in cross as circles in cross section (see at the base of the thallus). sections clearly show this character of the thallus. For the Discussion: The attribution of this species to the genus biometrical data and dimensions of the thallus see Table Diplopora is based on the metaspondyle arrangement of 2.

Dasycladales algae from the Norian-Rhaetian reef carbonates of Argolis Peninsula, Greece

Fig. 4 a to n Probolocupsis sarmeikensis nov. sp. from the reef carbonates near the town of Sarmeika, Peloponnes, Greece. a Cross section exhibiting the narrow axial cavity and the thick thallus wall. Laterals appear as white points in section through them. G058, x10. b Cross section through two poorly preserved specimens. G058, x8. c Oblique longitudinal section through an incomplete? specimen exhibits the laterals, which are oriented perpendicular to the axial cavity. G05C, x16. d Cross section of a specimen exhibiting the partly recrystallized laterals of the thallus. G05A, x16. e Cross section through an apparently poorly preserved specimen. Laterals appear as white points in section through them. G058, x16. f Cross section through an incomplete specimen. G050, x8. g Sections through a complete and an incomplete specimen. Laterals appear as white points around the incomplete specimen. G05B, x16. h Cross sections through two specimens. G05C, x12. i Cross section through a relatively well preserved specimen. G05C, x16. j Oblique section similar to Fig. i. G05C, x10. k Cross section exhibiting the pores (arrows) in the laterals. G05B, x16. l Longitudinal section through the holotype (for magnification see Fig. 2) and cross sections through two paratypes. G05C, x6. m Cross section through a recrystallized specimen. Arrows indicate the pores within the laterals. G05B, x16. n Cross section through a strongly recrystallized specimen causing the disappearance of the axial cavity. G025, x16. 43

Baba Senowbari-Daryan

Fig. 5 Diplopora obliguspora nov. sp. (a to d) and Probolocuspis tenuiparia nov. sp. (e to l) from the Norian-Rhaetian reef carbonates of Sarmeika, Peloponnes, Greece. a Specimen exhibits the thin wall around the axial cavity and the spine-like extended laterals. Thin section G1, x32. b Similar section as Fig. a showing the common base of oblique orientation of individual laterals. Thin section G89, x12. c (Holotype). Longitudinal section exhibiting the metaspondyle arrangement of the laterals, tuft of laterals and the thin wall around the axial cavity (for more information see Fig. 3). Thin section G75, x8. d Two cross sections exhibiting the spine-like extended individual laterals. G12, x20. e Cross sections through several specimens with thin thallus wall. Thin section G012, x20. f Sections through three specimens with relatively thick thallus wall but the individual laterals are recognizable. Thin section G08a, x25. g Oblique to longitudinal section. Thin section G08a, x20. h Holotype. The oblique section shows the individual laterals at the base. Thin section G08a, x20. i Sections through four specimens showing the laterals in part. Thin section G08a, x20. j Sections similar to Fig. i. Thin section G08a, x20. k Cross sections clearly showing the spine-like extended laterals. Thin section G08a, x40. l Similar to k. Thin section G40/1, x12.

Dasycladales algae from the Norian-Rhaetian reef carbonates of Argolis Peninsula, Greece the laterals and the tufts arising from the common base. des Centres de Recherches Exploration-Production Cysts within the axial cavity were not found in the Elf-Aqutaine, 3(2): 569-587. Sarmeika species. Such spine-like extended individual Flügel, E., 1983. Microfazies der Pantokrator-Kalke laterals of the new species – like in Probolocupsis (Lias) von Korfu, Griechenland. Facies, 8: 263-300. sarmeikensis described above – are not known from the Photiades, A., Pomoni-Papaioannou, F. & Kostopoulou, other species of the genus Diplopora. V., 2010. Correlation of the Late Triassic and Early Diplopora obliquspora – similar sections were described Jurassic Lofer-type carbonates from the Peleponnesus from the Norian-Rhaetian reef carbonates of the Northern Penninsula, Greece. Bulletin of the Geological Calcareous Alps as Clypeina sp. by Flügel (1979) and Society of Greece, 43(2): 726-736. Senowbari-Daryan (1980). Both specimens from the Granier, B. & Berthou, P-Y., 1994. Description de Alps, including their dimensions of the thallus are similar Milanovicella mamciliana n. gen., n. sp., algue to the Sarmeika species illustrated in Fig. 5a. The dasycladale du Portlandien de LÁlgarve central classification of these specimens as Diplopora (Portugal), et Validation de quelques taxons affins. obliqusipora is uncertain. Revue de Micropaléontologie, 37(2): 113-121. Diplopra obliquspora differs from the known species of Granier, B. R. C. & Grgasović, T., 2000. Les Algues the genus by the inclination of the laterals, very thin Dasycladales du Permien et du Trias – Nouvelle laterals and the outer surface of the thallus. Dimensions tentative dìnventaire bibliographique, géographique are given in table 3. et stratigraphique. Geologia Croatica, 53(1): 1-197. Haas, J., 1982. Facies analysis of the cyclic Dachstein Table 3. Some biometrical data of the thallus of Diplopora Limestone Formation (Upper Triassic) in the Bakony obliquspora nov. sp. (H= holotype). All measurements are in Mts., Hungary. Facies, 6: 75-84. mm. For abbreviations see Table 1. Kützing, F. T., 1843. Phycologia, Generalis oder Anatomie, Physiologie und Systematik der Tange. FA Specimen D D d/D Thin Brockhaus, Leipzig: 311-313. section Matarangas, D., Marcopoulou-Diacantoni, A. & Vartis- Pl. 2, Fig. 3 1.4 0.6 43 G75 Matarangas, M., 1995. Carnian reef facies from (H) Pantokrator Limestones of Argolis Peninsula (NE Pl. 2, Fig. 1 1.5 0.5 33 G1? Mavrovouni), Peleponnese. Geological Society of Pl. 2, Fig. 2 0.8 0.3 38 G89 Greece, Special Publication 4(1): 218-225. Pl. 2, Fig. 4 1.0 0.3 30 G12 Nittel, P., 2006. Beiträge zur Stratigraphie und Mikropaläontologie der Mitteltrias der Innsbrucker ACKNOWLEDGMENTS Nordkette (Nördliche Kalkalpen, Austria). Geo-Alp, 3(5): 93-145. M. Vartis Matarangas and D. Matarangas (both Athen) Ott, E. 1975. 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