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Revista Espaola de Micropaleontologa, 37(1), 2005, pp.

95-103 Instituto Geolgico y Minero de Espaa ISSN: 0556-655X

TRIASSIC DASYCLADALEAN ALGAE FROM THE DOLOMITES (NORTHERN ITALY): STRATIGRAPHIC ASSESSMENT
D. BASSI AND A. FUGAGNOLI
Dipartimento delle Risorse Naturali e Culturali, C.so Ercole I d'Este 32, I-44100 Ferrara, Italy E-mails: bsd@unife.it; fggnna@unife.it

Abstract The use of the calcareous green algal group Dasycladales for biostratigraphic correlation in the Triassic of the Dolomite area (northern Italy) has, up till now, suffered from the poor state of knowledge concerning their taxonomic inventory and stratigraphic distribution. In this study, the Triassic Dasycladales from the Dolomite area are listed together with the assessment of their stratigraphic ranges. At least 34 dasycladalean species (14 genera) are known from the Dolomite area. The greatest species richness occurs during the Pelsonian-Illiryan (26 species from 10 genera) and corresponds to the late Anisian carbonate platform development well known in the literature. At the AnisianLadinian boundary, most of the species disappear. Abundant and diverse dasycladaleans have not been identified during the development of the extensive Ladinian to Rhaetian carbonate platforms as might be expected. The biostratigraphic record of the Dasycladales is highly biased by provincialism and facies dependence.
Key words: Triassic, green algae, Dasycladales, biostratigraphy, Dolomites, northern Italy.

Resumen El uso de las algas verdes calcreas perteneciente al grupo Dasycladales para la correlacin bioestratigrfica en el Trisico del rea de las Dolomitas (norte de Italia) ha sufrido hasta ahora del escaso conocimiento sobre su taxonoma y su distribucin estratigrfica. En este trabajo se presenta un listado de especies de Dasycladales trisicas de las Dolomitas y se valora el rango estratigrfico de cada una de ellas. Se conocen al menos 34 especies, distribuidas en 14 gneros, del rea de las Dolomitas. La mayor riqueza especfica tiene lugar durante el Pelsoniense/Illiriense (26 especies pertenecientes a 10 gneros) y se corresponde con el desarrollo de las plataformas carbonatadas durante el Anisiense superior, bien conocidas en la literatura. La mayora de las especies desaparece en el lmite Anisiense-Ladiniense. A pesar de lo que cabra esperar, no se han encontrado abundantes ni diversas asociaciones de Dasycladales en las amplias plataformas carbonatadas en el intervalo Ladiniense-Rhetiense. El registro bioestratigrfico de Dasycladales est altamente sesgado por el provincialismo y la dependencia de facies. Palabras clave: Trisico, algas verdes, Dasycladales, bioestratigrafa, Dolomitas, norte de Italia.

died since the second half of the 19th century, are represented by blue-green algae (e. g., cyanobacteria), green Marine benthic calcareous algae are important bio- algae (udoteaceans and dasycladaleans), as well as by genic components of Triassic shallow water carbonate solenoporacean and gymnocodiacean red algae. In the successions. The Triassic calcareous algal floras, stu- Triassic, about 50 genera and 200 species of calcare-

INTRODUCTION

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ous algae have been described (Flgel, 1991; Barattolo, 1991). The extinction rate at the PermianTriassic boundary is 80% for dasycladaleans genera (Granier & Grgasovic, 2000) and 53% for all algal genera (Flgel, 1991). In the Early Triassic calcareous algae (except for spongiostromates) are absent at a global scale (Flgel, 1991; Kiessling et al., 2003). The Permian-Triassic biotic extinction led to a profound decline in reef production, but platform growth was less affected even if the facies changed considerably. It cannot yet be tested whether latitudinal variations in the distribution of Lower Triassic photozoan-type platforms are correlated with climatic fluctuations (Kiessling et al., 2003). A climate feedback from the coal-forest extinction has been proposed for explaining the Lower Triassic intensified warmth; such a climate feedback contributed to delayed biotic recovery that generally did not begin until mid-Triassic, but appears to have resumed first at high latitudes late in the Early Triassic (Kidder & Worsley, 2004). The first Triassic skeletal algae have been found in the lowermost Middle Anisian of the western Tethys. Starting from the Anisian, carbonate platform settings are characterised by highly diverse dasycladalean floras (93 species; Granier & Grgasovic, 2000) which indicate a rapid post-Permian diversification. Dasycladalean diversity decreases from the Anisian to the Late Triassic with a high taxonomic turnover during the Middle and Late Triassic and reaches a low at the Ladinian/Carnian boundary (Ott, 1972a; Flgel, 1991; Barattolo, 1991). The Triassic sedimentary successions of the Dolomite area (DA, Southern Alps, northern Italy; Fig. 1) show such a high species diversity of Dasycladaleans. To elucidate the history of this important calcareous algal group in the DA, we have collected all the literature data concerning Triassic dasycladalean records allowing the dasycladalean species diversity to be resolved at the stage level for the first time. This analysis reveals previously unrecognised patterns which may be useful for the biostratigraphy of the DA stratigraphic successions. No systematic revision was made herein on the quoted species; many taxonomic determinations have been made using different and outdated diagnostic criteria and, therefore, need to be revised. Nonetheless, the stratigraphic position of many species have to be checked according to the updated DA stratigraphy as well as to the stratigraphic correlations with other fossil groups. The aims of this study are, therefore, (a) to provide a bibliographic, geographic and stratigraphic inventory of the Triassic dasycladalean taxa recorded so far in the DA, (b) to update the taxonomic list according to

the most recent studies (e. g., Granier & Grgasovic, 2000; Bucur & Enos, 2001), and (c) to assess the stratigraphic level to which these species have been identified. The Dolomites are located in the eastern Southern Alps. They are bounded by the Neogene Insubric Lineament to the North and by the Valsugana-Pieve di Cadore Thrust to the South (Fig. 1). During the Triassic, the DA underwent extensional conditions due to strike-slip tectonics which is also responsible for local compression features (Doglioni, 1984, 1987). The Dolomites consist of separate blocks controlled by roughly E-W and N-S structural trends. These show different subsidence rates resulting in varying thicknesses of the same lithostratigraphic units within the Triassic sedimentary cover (De Zanche et al., 1993). Triassic deposits consist of continental, lagoonal, carbonate platform and basinal units which are variously stacked and interfingered. These successions are characterised by a great variety of facies (e. g., Leonardi,

FIGURE 1Geographic location of the Dolomites (Southern Alps, northern Italy).

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1967a; Assereto et al., 1977; Broglio Loriga et al., 1990; De Zanche, 1990; Bosellini, 1984, 1991; Masetti et al., 1991; Russo et al., 1991; Gaetani et al., 1992; Brack & Rieber, 1993; De Zanche et al., 1993; Wignall & Hallam, 1993). Particularly noteworthy are the Anisian to Carnian carbonate platforms, recording an evolution from regional muddy banks to isolated highrelief buildups (e. g., Bosellini et al., 2003). Stratigraphic synthesis and sequence stratigraphy of the whole Triassic interval in the Southern Alps have been proposed by De Zanche et al. (1993) and by Gianolla et al. (1998). Although none of these studies considered dasycladalean algae in the biostratigraphy of the Triassic successions of the DA, Pia (1930a, Fig. 3) in his biostratigraphic division of Middle Triassic of Southern Alps established several dasycladalean zones and defined the Anisian-Ladinian boundary on the base of the first appearance of Diplopora annulata Schafhutl. Pias pioneer work has been often quoted and a Ladinian age for D. annulata-bearing strata has been accepted by most of the authors (see references in Granier & Grgasovic, 2000). Among these Ott (1972a, b) recognised four different algal floras in the Triassic of Alps (DA included) that may have biostratigraphic value.

STRATIGRAPHIC ASSESSMENT
In the present paper, we only refer to Triassic dasycladalean species identified in the DA if they have been described and/or illustrated in the cited studies. We also include the records of Triassic dasycladaleans from the Brenta Dolomites (Giudicarie Alps) which represent the transitional zone between the Lombardy Basin and the DA (e. g., Venzo & Fuganti, 1965; Gaetani, 1969; Peloso & Vercese, 1982). The stratigraphic distributions of the taxa are considered as given by the cited authors. For those species revised in later studies, the valid names following Granier & Grgasovic (2000), Bucur & Enos (2001), and Granier & Hofmann (2002) are used. Species are listed alphabetically. Aciculella bacillum Pia, 1930b: Senowbari-Daryan et al., 1993 (uncertain assignment). Clypeina besici Pantic in Granier & Deloffre, 1994, non 1965: Flajs, 1977 (uncertain assignment). Diplopora annnulata (Schafhutl, 1853) 1863: Stoppani, 1858; Gmbel, 1872; Salomon, 1895; Tornquist, 1899; Steinmann, 1907; Pia, 1920; Bubnoff, 1921; Ogilvie Gordon, 1927; Epting et al., 1976;

Gaetani et al., 1981; Senowbari-Daryan et al., 1993; Rinaldo & Jadoul, 1994 (uncertain assignment); Pugliese, 1997. Diplopora cadorica Ogilvie-Gordon, 1935: Ogilvie Gordon, 1935. Diplopora cellulata Hurka, 1967: Hurka, 1967; Farabegoli & Levanti, 1982 (uncertain assignment). Diplopora comelicana Fois, 1979: Fois, 1979. Diplopora monregalensis (Baretti, 1919) Granier & Hofmann, 2002: Ogilvie-Gordon, 1927; Rossi, 1967; Bechstdt & Brandner, 1970; Gaetani et al., 1981; Senowbari-Daryan et al., 1993; Pugliese, 1997; Granier & Hofmann, 2002. Diplopora nodosa Schafhutl, 1863, emend. De Castro, 1979: Gmbel, 1872; Salomon, 1895; Pia, 1920; Farabegoli & Levanti, 1982. Euteutloporella triasina (Schauroth, 1859) De Castro, 1993: Schauroth, 1855; Gmbel, 1872; Tornquist, 1900; Pia, 1912. Griphoporella curvata (Gmbel, 1872) Pia, 1915 emend. Barattolo et al., 1993: Pugliese, 1997. Griphoporella? guembeli (Salomon, 1895) Pia, 1920: Salomon, 1895; Pia, 1920. Gyroporella vesiculifera Gmbel, 1872, emend. Benecke 1876: Pia, 1920; Pugliese, 1997. Kantia debilis (Gmbel, 1872) Baretti, 1919, emend. Gven 1979: Gmbel, 1872. Kantia dolomitica Pia, 1912 emend. Gven, 1979: Pia, 1912. Kantia praecursor (Pia, 1920) Gven, 1979: Hurka & Schmid, 1971. Macroporella alpina Pia, 1912: Farabegoli & Levanti, 1982; Senowbari-Daryan et al., 1993. Macroporella perforatissima Pia, 1920: Casati et al., 1982. Oligoporella pilosa var. varicans Pia, 1935: Senowbari-Daryan et al., 1993. Oligoporella serripora Pia, 1912: Pia, 1912. Physoporella croatica Herak in Granier & Deloffre, 1994 non 1958: Hurka, 1969. Physoporella intusannulata Hurka, 1967: Hurka, 1967. Physoporella leptotheca Kochanski-Devid, 1967: Fois, 1979. Physoporella lotharingica (Benecke, 1898) Pia, 1931: Pugliese, 1997. Physoporella pauciforata var. gemerica Bystricky, 1962: Dieni & Spagnulo, 1964; Casati et al., 1982. Physoporella pauciforata var. pauciforata Pia ex Bystricky, 1964: Steinmann, 1907; Pia, 1920; Ogilvie Gordon, 1927; Hurka, 1969; Farabegoli & Guasti, 1980; Casati et al., 1982.

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Physoporella pauciforata var. sulcata Bystricky, 1962: Dieni & Spagnulo, 1964; Gaetani et al., 1981; Casati et al., 1982. Poncetella helvetica (Pia, 1912) Gven, 1979: Hurka, 1967. Poncetella hexaster (Pia, 1912) Gven, 1979: Bubnoff, 1921 (uncertain assignment); SenowbariDaryan et al., 1993. Scinderella scopuliformis Grgasovic & Sokac, 2002: Senowbari-Daryan et al., 1993; Grgasovic et al., 2002. Terquemella brentai Pugliese, 1997: Pugliese, 1997. Teutloporella herculea (Stoppani, 1857) Pia, 1912: Salomon, 1895; Ogilvie-Gordon, 1927. Teutloporella peniculiformis Ott in Granier & Deloffre, 1994 non 1963: Ogilvie-Gordon, 1935 (uncertain assignment); Epting et al., 1976; Casati et al., 1982; Senowbari-Daryan et al., 1993. Teutloporella vicentina var. nana Pia, 1912: Tornquist, 1899; Pia, 1912; Accordi, 1955; Leonardi, 1967b; Epting et al., 1976; Scheuber, 1990. Teutloporella vicentina var. vicentina Pia, 1912: Pia, 1912.

daleans have been recorded and illustrated in the DA sensu stricto; in the Brenta Dolomites only Griphoporella curvata (Gmbel) Pia, Gyroporella vesiculifera Gmbel and Terquemella brentai Pugliese have been identified. Oligoporella serripora Pia, Diplopora cadorica Ogilvie Gordon, Diplopora cellulata Hurka, Diplopora annulata (Schafhutl), and Diplopora monregalensis (Baretti) show the longest stratigraphic ranges.

In the Triassic formations of the Dolomites, 34 species belonging to 14 genera were identified ranging from the Anisian to the Rhaetian (Fig. 2). In the Aegean, dasycladaleans are very rare being represented only by Poncetella helvetica (Pia) Guven and Oligoporella serripora Pia; the latter is, however, of uncertain stratigraphic position and taxonomic assignment. During the Bithynian, only 5 species (belonging to 5 genera) are recorded. The diversity reaches a peak during the PelsonianIlliryan with 26 species (from 10 genera). Most of the species belong to the genera Physoporella Steinmann and Kantia Pia. The Pelsonian-Illiryan boundary seems to be marked by the appearance of the Kantia species such as K. praecursor (Pia) Guven, K. dolomitica Pia, as well as Diplopora comelicana Fois, D. monregalensis (Baretti), Euteutloporella triasina (Schauroth) De Castro, Physoporella leptotheca Kochanski-Devid, Scinderella scopuliformis Grgasovic & Sokac, Teutloporella vicentina var. nana Pia, and T. var. vicentina Pia. Most of the species disappear at the Anisian-Ladinian boundary. Twelve species (from six genera) have been recog- F IGURE 2Dasycladalean algae in the Triassic of the Dolomites and their stratigraphic distribution. Grey bars nised in the Ladinian and three species (all of uncertain after Granier & Deloffre (1994); dashed lines point to taxonomic assignment, from two genera) in the lower uncertain taxonomic assignments; *Brenta Dolomites. The Carnian. No dasycladalean species occur in the Upper stratigraphic distribution of Scinderella scopuliformis is Carnian. During the Norian and Rhaetian, no dasyclaafter Grgasovic et al. (2002).

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The peak of dasyclad diversity reached in the Pelsonian-Illiryan corresponds to the wide distribution of Anisian carbonate platforms in the DA. Although a dasycladalean abundance has been recorded during the Ladinian to Rhaetian (e. g. Granier & Deloffre, 1994; Granier & Grgasovic, 2000), no abundant and diversi fied dasycladaleans have been identified in the DA as would be expected in the carbonate platform successions of this time span. Further studies are needed to verify if this lack of data are due to the real absence of dasycladaleans or to the incompleteness of the palaeontological data.

BIOSTRATIGRAPHIC REMARKS
Early Triassic Tethyan carbonate platforms were formed without the contribution of reefal debris. The Anisian and Ladinian shelf platforms are similar to those of the Early Triassic with regard to the importance of micrite and microbial carbonates, but differ by the increasing quantitative importance of dasycladalean green algae, the increasing frequency of crinoids and molluscs, and the scarcity of ooids. Late Triassic platform carbonates resemble those of the Middle Triassic, but have a higher contribution of foraminifera, molluscs and oolites, and fewer dasycladaleans (e.g., Maurer, 2000; Keim & Schlager, 2001; Kiessling et al., 2003). This general trend is followed by the dasycladaleans in the DA which show the highest diversity in the Anisian-Ladinian. The decrease in diversity toward the Norian and Rhaetian (Fig. 2) is coupled with the higher contribution of foraminifera, molluscs and oolites. While Anisian and Ladinian platforms carbonates originated within or near the core tropics, Norian and Rhaetian platforms are absent or rare in the equatorial zone (Kiessling et al., 2003). The comparison between the stratigraphic distribution of dasycladalean species provided by Granier & Deloffre (1994) and that presented herein show some differences (Fig. 2). Poncetella helvetica (Pia) occurs in the Aegean-Bithynian of the DA while Granier & Deloffre (1994) indicated a younger stratigraphic distribution limited to the Ladinian. Physoporella leptotheca Kochanski-Devid appears in the Illiryan of the DA, while Granier & Deloffre (1994) indicate Norian age for this species. Traditionally, assemblage zones (coenozones) have been applied to shallow water Triassic successions in Europe. The application of Triassic calcareous algae for biostratigraphy is restricted to dasycladaleans and is strongly affected by environmentally controlled dis-

tribution patterns. Algal assemblages, rather than the stratigraphic range of single taxa, enable the recognition of time intervals which might correspond to several ammonite or conodont zones in the Triassic (Flgel, 1991). In the Triassic of the Apuseni Mountains (Romania), Dragastan (1981) defined six coenozones based on benthic foraminifera and dasycladaleans. Most of these coenozones can be also recognised in the DA except for the Diplopora annulata-D. nodosa association (Ladinian in Dragastan, 1981) which, in the DA, has been found spanning the Pelsonian-Ladinian. Grgasovic & Sokac (2003) recog nised several fossil dasycladalean assemblages in the Triassic carbonate platform successions of Croatia. The strong analogy of these assemblages with those of the DA is represented by the diversity peak reached in the Pelsonian-Illiryan. The Anisian-Ladinian boundary in Croatia is marked by the first appearance of Kantia dolomitica Pia and Diplopora annulata Schafhutl which in the DA are already present since the Illiryan and Pelsonian respectively. Although the record of Clipeina besici Pantic in the DA is questionable (i. e. Flajs, 1977), it is restricted to the Carnian as in Croatia. Gyroporella vesiculifera Gmbel and Gryphoporella curvata (Gmbel) characterise the Norian and the Rhaetian in both areas. According to Grgasovic & Sokac (2003), correla tions between the Triassic stratigraphic distributions of dasycladaleans and other fossil groups (e. g. ammonites, benthic foraminifera, conodonts, etc.) has not yet been sufficiently established and still needs to be elaborated. An interesting statistical approach to the palaeobiogeography of dasycladalean distribution has been performed by Rasser & Fenninger (2002) for JurassicCretaceous taxa of the Northern Calcareous Alps. Such an analysis could also be developed for the Triassic forms in order to verify possible analogies between similar localities as DA and Croatia. Furthermore, taphonomic bias represent a main problem. The preservation of the aragonitic dasycladalean extraskeleton as a whole and of the taxonomic features in particular requires a micritic matrix, encrustation and/or micritisation. Although studies on skeletal minerology exist (e. g. Flajs, 1977), taphonomic consideration on dasycladalean have been restricted to only a few taxa (e. g. De Castro, 1993). The biostratigraphic record of the Dasycladales in the DA is highly biased due to provincialism and facies dependence. The potential use of this group for biostratigraphic interpretations is also hampered by the poor knowledge concerning their stratigraphic distributions and taxonomic inventory. We suggest that the definition of for-

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Geognostisch-palontologische Beitrge, Mnchen, 2, mal biozones is an appropriate first order approach for 296-317. a stratigraphic correlation using dasycladaleans. As . 1898. Diplopora und einige andere Versteinerungen im suggested by Rasser & Fenninger (2002), dasycladaleelsass-lothringischen Muschelkalk. Mitteilungen der an biozones should, however, only be defined for disgeologischen Landesanstalt von Elsass-Lothringen, 4, tinct bioprovinces.

ACKNOWLEDGEMENTS
Grateful thanks are due to I. I. Bucur (Cluj-Napoca) and T. Grgasovic (Zagreb) for discussing biostrati graphic problems of the Triassic dasycladaleans and to R. Rettori (Perugia) for the useful comments on an early draft of the manuscript. We thank T. Grgasovic and an anonymous referee for their thoughtful review of the manuscript. We acknowledge the help of S. Ros Franch in translating the Spanish abstract and of J. H. Nebelsick in correcting the English text. The research was funded by the local research fund 2002 of the University of Ferrara.

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MANUSCRITO RECIBIDO: 23 enero, 2004 MANUSCRITO ACEPTADO: 10 mayo, 2004