Mesozoic cyanobacteria and calcareous ? Algae of the Apennine platform (Latium and Abruzzi, Italy)

MESOZOIC CYANOBACTERIA AND CALCAREOUS ? AIJGAE OF THE A P E N N I N E PLATFORM (LATIUM AND ABRUZZI, ITALY) ANNA MANCINELLI, DIEGO F E R R A N D E S Mancinelli, A., Ferrandes, D., 2001. Mesozoic cyanobacteria and calcareous ? Algae of the Apennine platform (Latium and Abruzzi, Italy). [Cyanobact6ries et ?algues calcaires du Mesozoique de l'Apennin (Latium et Abruzzes, Italie)]. [Cyanobatteri e ? algae calcaree del Mesozoico dell'Appennino (Lazio e Abruzzi, Italia]. Geobios 34 (5), 533546. Villeurbanne, le 30.10.2001. Manuscrit d4pos6 le 16.05.2000; accept6 d6finitivement le 16.02.2001 ABSTRACT - Eighteen species of cyanobacteria and calcareous ?algae belonging to 'Rivularia', Ortonella, Hedstroemia and Garwoodia are described and illustrated. They come from Mesozoic carbonate sediments (Jurassic and Cretaceous) cropping out in several stratigraphic sections located in southern Latium and Abruzzi (central Italy). The abundance of these microfossils at different intervals of time allowed us to establish their stratigraphic range in the Apennine platform and, in addition, to estimate their paleoecologic significance. © 2001 Editions scientifiques et m6dicales Elsevier SAS KEYWORDS: CYANOBACTERIA, CALCAREOUS ?ALGAE, SYSTEMATICS, BIOSTRATIGRAPHY, PALEOECOLOGY, MESOZOIC, ITALY. R]~SUMt~ - Les s6diments de plate-forme carbonat6e du Mesozoique qui affieurent dans le Latium m6ridional et dans les Abruzzes ont mis en 6vidence une riche association d'algues calcaires longtemps assign6es par divers auteurs aux Codiaceae ou r6unies sous le nom g6n6ral de Cayeuxia; elles sont caract6ris6es par la pr6sence de thalles calcaires noduleux constitu6s de filaments tubulaires plus ou moins rapproch6s et diversement bifurqu6s, assign6es par Pia (1927) dans le groupe informel 'Porostromata'. De r6centes 6tudes pour la plupart taxonomiques (Riding 1991; Dragastan 1993; Dragastan et al. 1997) ont mises en 6vidence qu'il s'agit de taxons diff6rents, quelques uns avec une importante valeur stratigraphique et utile pour la pal6o6cologie. Les 6tudes d6taill6es de ces algues nous ont permis de d6crire et d'illustrer 18 esp6ces qui appartiennent aux genres 'Rivularia', Ortonella, Hedstroemia et Garwoodia; de fournir la r6partition stratigraphique de ces esp6ces dans les localit6s consid4r6es; de formuler quelques hypoth6ses sur leur signification pal6o6cologique. Beaoucoup de ces esp6ces, jamais observ6es jusqu'~ pr6sent en Italie, ont 6t6 signal6es ici pour la premi6re lois. Leur d6couverte a permis d'effectuer d'utiles corr61ations avec les s6diments contemporains du bassin de la Tethys. © 2001 ]~ditions scientifiques et m4dicales Elsevier SAS MOTS-CL]~S: CYANOBACTt~RIES, ?ALGUES CALCAIRES,SYST]~MATIQUE, BIOSTRATIGRAPHIE,PALI~OI~COLOGIE, MESOZOIQUE, ITALIE.

INTRODUCTION The Mesozoic carbonate sediments of the Apennine platform (sensu Merlini & Mostardini 1986) contain very rich communities of calcareous ?algae and cyanobacteria till now generically called 'Codiaceae' or with the generic name Cayeuxia. They belong to many genera characterized by calcareous nodular thalli composed of tubular filaments, more or less packed and branched, which m an y authors have assembled in the 'Porostromata' group (sensu Pia 1927). For a long time, little importance has been attached to these species and their systematic position has hardly been discussed; this taxonomic uncertainty means th at these genera and species are not as useful as they might be for solving stratigraphic and paleoenvironmental problems. The present article is concerned with the genera 'Rivularia' (ROTH, 1802)AGARDH, 1824, Ortonella

GARWOOD, 1914, Hedstroemia ROTHPLETZ,1913 and Garwoodia WOOD, 1941 and their different species, contained in the Mesozoic carbonate sediments of the Latium-Abruzzi area (central Italy). The particularly rich fossil associations allowed us to establish the stratigraphic range of the analyzed species in the Apennines, as well as to propose some hypotheses on their paleoecological significance. The analyzed samples come from several stratigraphic sections previously investigated and described in detail (Chiocchini & Mancinelli 1977, 1978; Chiocchini et al. 1994); in Figure 1 these sections are grouped according to the main facies: 'paleopiattaforma' (sensu Chiocchini & Mancinelli 1978), inner shelf facies (M. Aurunci) back-reef included (M. Cairo and M. d'Ocre), and shelf-edge facies (Gran Sasso d'Italia).

534

Stratigraphical sections "paleopiatteforma"

(upper Tries-lower Uas) 1 Montedella Selva 2 Pizzo Cefalone shelf-edge

3 Colle Matzo 4 Cirna d'Anzano 5 Costa di Vat]elunga 6 Costa dell'Orsa 7 Cesta di Piecipopoli

FIGURE 1 - Distribution of the Jurassic and Cretaceous carbonate facies in the central-southern Apennines (after Carbone 1978; Chiocchini et al. 1986, modified) and location map of the analysed stratigraphic sections. Distribution des facies carbonatds au Mesozo~que dans l'Apennin centre-mdridional (d'apr~s Carbone 1978; Chiocchini et al. 1986, modifid) et localisation des successions dtudides.

back-reef

8 Monte Cefalone 9 Colle S. Lucia-La Querce 10 Villa S, Lucia 11 Colle S, Lucia-La Cesa 12 Omo di Cairo 13 Costalunga inner shelf

14 M. Moneta-Grotte dei Serpenti

15 Monte Sorgenza

sh~fedQefac~ea

~ 1 "paleopiattaforma" ,ql,lq~;~ (Upper Trlas-Lower Use)

~i]

~

~l! ITIll stepeandpata~cfaeces

inner shelf facies

PALEONTOLOGY

Because of the classification of the recent Rivularia is based on details of the soft organic part, not preserved in the fossils, at present we believe more correct to call 'Rivularia' the fossil calcified skeletons to distinguish them from the recent specimens; further studies are needed to clarify the precise affinities among the fossil and the recent Rivularia. Moreover, some uncertainties exist about the systematic position of Hedstroemia and Garwoodia. In fact, according to Dragastan (1991) and Dragastan et al. (1997), these genera, showing politomic branches and a rather complex inner structure, belong to the calcareous Algae (Phylum Siphonophyta Margiulius, 1974). On the contrary, Riding (1991) believes that Hedstroemia is a cyanobacterium, while the larger tubes and the bigger size of Garwoodia make it difficult to consider it a cyanobacterium, so '...it seems best to place Garwoodia in a separate group to await clarification of its systematic position' (op. cit., pag. 71). At the moment the systematic position of Garwoodia and Hedstroemia and the phylogenetic relationships among the fossil and the recent Rivulariacea are not fully understood; however, we believe that it may be useful to illustrate the analyzed fossil associations sampled in the Apennines and to establish their stratigraphic range, in order to assess possible correlations among Mesozoic carbonate sediments in the Mediterranean area. The following species are described and illustrated, according to the taxonomic criteria proposed by Dragastan (1993) and Dragastan et al. (1997): 'Rivularia' carpathica DRAGASTAN,1985 'Rivularia' dianae (DRAGASTAN• BUCUR, 1978) 'Rivularia' fruticulosa (JOHNSON & KASKA, 1965)

back reef facies

~ ' ,~,~ volcanites (Pile-Pleistocene)

'Rivularia' kurdistanensis (ELLIOTT, 1957) 'Rivularia' lissaviensis (BORNEMANN,1887) ' R i v u l a r i a ' p i a e (FROLLO, 1938) 'Rivularia' theodori DRAGASTAN,1985 Ortonella lemoineae DRAGASTAN,1971 Ortonella perconigi DRAGASTAN,1972 Garwoodia bardosi DRAGASTAN, 1985 Garwoodia fissa DRAGASTAN,1989 Garwoodia fluegeli DRAGASTAN,1985 Garwoodia maxima DRAGASTAN,1989 Garwoodia polytomica DRAGASTAN,1989 Garwoodia toomeyi DRAGASTAN,1989 Hedstroemia moldavica DRAGASTAN,1985 Hedstroemia racasiensis DRAGASTAN,1989 Hedstroemia villosa DRAGASTAN,1989' CYANOBACTERIA Phylum CYANOPHYTA(Sachs, 1874) Order NOSTOCALESGeitler, 1925 Family RIVULARIACEAERabenhorst, 1865 Genus R i v u l a r i a (RoTH, 1802) AGARDH,1824 Type-species

-

Rivularia dura

ROTH, 1 8 0 2 .

Remarks - Because of the classificationof the recentRivularia is based on many features not preserved in the fossils,we believe more correctto call 'Rivularia' the calcifiedfossilskeletons.

'Rivularia' carpathica DRAGASTAN,1985 Fig. 2.3; Fig. 3.2, 3 1985 Rivularia carpathica nov. sp. - Dragastan, pl. 15, fig. 7; p]. 16, figs. 1-2 1998 Rivularia carpathica DRAGASTAN- Dragastan et al., p]. 3, fig. 5 (not fig. 4). D e s c r i p t i o n - Hemispherical thallus, 4.5 m m high, 3.0 mm wide, characterized by many growth

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4

6 FIGURE 2 - 1. Hedstroemia racasiensis DRAGASTAN(X 30); Colle S. Lucia stratigraphic section (M. Cairo); lower Aptian. 2. Garwoodia fluegeli DRAGASTAN(X 30); Monte della Selva stratigraphic section (Gran Sasso d'Italia); Hettangian. 3. 'Rivularia' carpathica DRAGASTAN(× 30); Coste di Vallelunga stratigraphic section (Gran Sasso d'Italia); Berriasian. 4. 'Rivularia' theodori DRAGASTAN(X 30); Villa S. Lucia stratigraphic section (M. Cairo); Barremian. 5. Garwoodia maxima DRAGASTAN(X 30); Villa S. Lucia stratigraphic section (M. Cairo); Barremian. 6. Hedstroemia viUosa DRAGASTAN(X 30); Coste di Vallelunga stratigraphic section (Gran Sasso d'Italia); Berriasian. 1. Hedstroemia racasiensis DRAGASTAN(X 30); Colle S. Lucia (M. Cairo); Aptien infdrieur. 2. Garwoodia fluegeli DRAGASTAN(× 30); Monte della Selva (Gran Sasso d'Italia); Hettangien. 3. ~Rivularia' carpathica DRAGASTAN(× 30); Coste di Vallelunga (Gran Sasso d'Italia); Berriasien. 4. ~Rivularia' theodori DRAGASTAN(X 30); Villa S. Lucia (M. Cairo); Barremien. 5. Garwoodia maxima DRAGASTAN(× 30); Villa S. Lucia (M. Cairo); Barremien. 6. Hedstroemia villosa DRAGASTAN(× 30); Coste di Vallelunga (Gran Sasso d'Italia); Berriasien.

zones and a compact inner structure. The tube filam e n t s are dichotomically b r a n c h e d u n d e r a r a t h e r small angle of divergence (less t h a n 3°); their diam e t e r is about 0.010-0.020 mm.

Stratigraphic range - 'Rivularia' carpathica, so far known only in the Tithonian of Romania (Dragastan 1985; Dragastan et al. 1998), has been found for the first time in the Apennines (Berriasian of the Gran Sasso d'Italia).

536

5

2

3

FIGURE 3 - 1. 'Rivularia' dianae (DRAGASTAN& BUCUR)(× 30); Coste di Va]lelunga stratigraphic section (Gran Sasso d'Italia); Berriasian. 2. 'Rivularia' carpathica DBAGASTAN(X 20); Coste dl Vallelunga stratigraphic section (Gran Sasso d'Italia); Berriasian. 3. 'Rivularia' carpathica DRAGASTAN(X 30); Coste di Vallelunga stratigraphic section (Gran Sasso d'Italia); Berriasian. 4. 'Rivularia' kurdistanensis (ELLIOTT)(X 30); Monte della Selva stratigraphic section (Gran Sasso d'Italia); Hettangian. 5. Garwoodia fissa DRAGASTAN(X 25); Colle S. Lucia stratigraphic section (M. Cairo); Upper Barremian. 1. 'Rivularia' dianae (DRAGASTAN& BUCUR) (X 30); Coste di Vallelunga (Gran Sasso d'Italia); Berriasien. 2. 'Rivularia' carpathica DRAGASTAN(X 20); Coste di Vallelunga (Gran Sasso d'Italia); Berriasien. 3. 'Rivularia' carpathica DRAGASTAN(X 30); Coste di Vallelunga (Gran Sasso d'Italia); BeT~'iasien. 4. 'Rivularia' kurdistanensis (ELLIOTT) (X 30); Monte della Selva (Gran Sasso d'Italia); Hettangien. 5. Garwoodia fissa DRAOASTAN(X 25); Colle S. Lucia (M. Cairo); Barremien supdrieur.

'Rivularia' dianae (DRAGASTAN & BUCUR, 1978)

1985

Fig. 3.1 1978

Diversocallis dianae nov. sp. - Dragastan & Bucur, pl. 2. figs 1-2; pl. 3, figs 1-2.

1998

Rivularia dianae (DRAGASTAN• BUCUR)- Dragastan, pl. 15, figs 2-3. Rivularia dianae (DRAGASTAN& BUCUR) - Dragastan et al., pl. 3, fig. 4.

537

2 3

4

6

5 FIGURE4 - 1. Ortonella lemoineae DRAGASTAN(X 30); Colle S. Lucia stratigraphic section (M. Cairo); upper Barremian. 2. Hedstroemia moldavica DRAGASTAN(X 30); Colle S. Lucia stratigraphic section (M. Cairo); upper Barremian. 3. Garwoodia toomeyi DRAGASTAN(X 30); Colle S. Lucia stratigraphic section (M. Cairo); Barremian. 4. Ortonella perconigi DRAGASTAN(X 30). Coste di Vallegunda stratigraphic section (Gran Sasso d'Italia); upper Barremian. 5. Garwoodia bardosi DRAGASTAN(× 30); Costa Lunga stratigraphic section (Gran Sasso d'Italia); upper Barremian. 6. 'Rivularia' kurdistanensis (ELLIOTT)(X 30); Costa dell'Orsa stratigraphic section (Gran Sasso d'Italia); Valanginian. 1. Ortonella lemoineae DRAGASTAN(X 30); Colle S. Lucia (M. Cairo); Barrdrnien supdrieur. 2. Hedstroemia moldavica DRAGASTAN(X 30); Colle S. Lucia (M. Cairo); Barrgmien supdrieur. 3. Garwoodia toomeyi DRAGASTAN(X 30); Colle S. Lucia (M. Cairo); Barr~mien. 4. Ortone]la perconigi DRAGASTAN(X 30); Coste di Vallelunga (Gran Sasso d'Italia); Barrdmien supdrieur. 5. Garwoodia bardosi DRAGASTAN (× 30); Costa Lunga (Gran Sasso d'Italia); Barr~mien supgrieur. 6. 'Rivularia' kurdistanensis (ELLrOTT) (× 30); Costa dell'Orsa (Gran Sasso d'Italia); Valanginien. - Fanlike thallus, 3.2-6.0 m m wide and 2.1-5.0 m m high, Almost parallel filaments with rare dichotomic branching; the filaments Description

diverge under a very small angle and their diameter is 0.015-0.020 m m . Frequent voids (a typical f e a t u r e of the recent m a r i n e species of

538

•

FIGURE 5 - Main occurrences of 'Rivularia' kurdistanensis. The numbers are related to the following references. 1. Elliott 1957 (Aptian). 2. Dufaure 1958 (Oxfordian-Tithonian). 3. Johnson 1968 (BerriasianoAptian). 4. Bouroullec & Deloffre 1968, 1970 (Sinemurian-Aptian). 5. Masse 1976 (Berriasian-Aptian). 6. Flugel 1979 (Oxfordian-Tithonian). 7. Canerot 1979 (Tithonian-Berriasian). 8. Dragastan 1985; Dragastan et al. 1996 (upper Oxfordian). The asterisk indicates the present paper. Principaux gisements de 'Rivularia' kurdistanensis d'apr~s: 1. Elliott 1957 (Aptien); 2. Dufaure 1958 (Oxfordien-Tithonique); 3. Johnson 1968 (Berriasien-Aptien); 4. Bouroullec & Deloffre 1968, 1970 (SindmurienAptien); 5. Masse 1976 (BerriasienAptien); 6. Flugel 1979 (OxfordienTithonique); 7. Canerot 1979 (Tithonique-Berriasien); 8. Dragastan 1985, Dragastan et al. 1996 (Oxfordien supgrieur). Astdrisque: gisments dtudigs.

l r 8

'Rivularia') are present in the central part of the thallus. Stratigraphie r a n g e - 'Rivularia' dianae is known in the Kimmeridgian-Berriasian of Romania (Dragastan & Bucur 1978; Dragastan 1985; Dragastan et al. 1998); it has been found for the first time in the Apennines (Berriasian of the Gran Sasso d'Italia).

'Rivularia' fruticulosa (JOHNSON • KASKA, 1965) Fig. 6.2 1965

Cayeuxia fruticulosa nov. sp. - Johnson & Kaska, pl. 12. figs 1-2; pl. 13, figs 1-3. 1985 Rivularia fruticulosa (JOHNSON • KASKA) - Dragastan, pl. 10, figs 3-7, pl. 11, figs 1-5 (and references therein). 1990 Rivularia fruticulosa (JOHNSON& KASKA)- Kuss, pl. 19, figs 3, 10, 11. 1992 Rivularia fruticulosa (JOHNSON • KASKA) - Dragastan, figs 5, 6; pl. 3, figs 1-7, pl. 11, fig. 4. - Hemispherical, nodular, subspherical thallus, or ellipsoidal and irregular, sometimes very flaring, about 3.0 mm high and 5.0-7.0 mm wide. The inner structure is 'lax' and shows, between the filaments, wide zones filled with micrite. The fine tube filaments diverge under an angle of 5°-10 °" Description

Stratigraphie r a n g e - 'Rivularia' fruticulosa, observed for the first time in the Maastrichtian of Guatemala (Johnson & Kaska 1965), is known in the following localities: Honduras, Lower Cretaceous (Johnson 1969); Romania, Tithonian-lower Aptian (Dragastan 1985, 1992); Sinai, lower Bajocian (Kuss 1990). This species has been found for the first time in the Apennines (Kimmeridgian of the Gran Sasso d'Italia).

'Rivularia' kurdistanensis (ELLIOTT, 1957) Fig. 3.4; Fig. 4.6 1957 Cayeuxia kurdistanensis nov. sp.- Elliott, pl. 25. figs 6-7. 1985 Rivularia kurdistanensis (ELLIOTT) - Dragastan, pl. 14, figs 8-14 (and references therein). 1996 Rivularia kurdistanensis (ELLIOTT) - Dragastan et al., pl. 7, fig. 3. - Small nodular, fanlike thallus, 1.52.3 mm wide and 1.0-1.5 mm high, crossed by

Description

tubular filaments 'V' dichotomically branched, which diverge under an angle of 3°-5 °. The relatively small diameter of the tubular filaments (0.015-0.035 mm) and their regular arrangement cause the rather compact inner structure of the thallus.

Stratigraphic r a n g e - 'Rivularia' kurdistanensis, a rather widespread species observed for the first time in the Aptian of Iraq (Elliott 1957), is known in many localities (see Fig. 5). In the Apennines, 'R.' kurdistanensis has been found in all the examined areas (M. Aurunci, M. Cairo and Gran Sasso D'Italia); it ranges from Hettangian to lower Aptian.

'Rivularia' lissaviensis (BORNEMANN, 1887) Fig. 6.1 1887 Zonotrichites lissaviensis nov. sp. - Bornemann, pl. 5. figs 1-2; pl. 6, figs 1-2. 1985 Rivularia lissaviensis (BORNEMANN)- Dragastan, figs 1-2; pl. 1, figs 1-3; pl. 3, figs 1-2; pl. 4, figs 1-6; pl. 5, figsl-5; pl. 6, figs 1-2; pl. 7, figs 1-4; pl. 81, figs 1-5 (references the rein). 1986 Cayeuxia andamanica - Badve & Kundal, fig. 2. 1992 Rivularia lissaviensis (BORNEMANN) - Dragastan, pl. 1, figs 1-2; pl. 2, fig. 1. 1994 Rivularia lissaviensis (BORNEMANN)- Bucur~ pl. 21, figs 9, 11. - Subspherical, hemispherical, fanlike or columnar erect thalli, varying in size (up to 60 mm high), crossed by tube filaments whose diame~ ter varies between 0.020 and 0.060 mm; they are about parallel and ~V' dichotomically branched under a rather variable angle of divergence (2 ° to 20°). The columnar erect thalli show many growth zones and, sometimes, a progressive growth by little tufts (see Fig. 6.1). Description

Stratigraphic r a n g e - 'Rivularia' lissaviensis is a rather widespread species; observed for the first time in the Upper Rhaetian of Poland-Silezia (Bornemann !887), it is known in many localities (see Fig. 7). In the Apennines 'Rivularia' lissaviensis has been found in all the examined areas (M. Aurunci, M. Cairo and Gran Sasso D'Italia); it ranges from Hettangian to Valanginian.

539

2 7

3

4

5

FIGURE6 - 1. 'Rivularia' lissaviensis (BoRNEMANN)(X 15); Monte della Selva stratigraphic section (Gran Sasso d'Italia); Hettangian. 2. 'Rivularia'fruticulosa (JOHNSON• KASKA)(X 15); Colle Marzo stratigraphic section; Kimmeridgian. 3. Hedstroemia moldavica DRAGASTAN (× 30); Villa S. Lucia stratigraphic section (M. Cairo); Barremian. 4. Garwoodia maxima DRAGASTAN(× 30); Villa S. Lucia stratigraphic section (M. Cairo); Barremian. 5. 'Rivularia' cf. theodori DRAGASTAN(× 30); Colle S. Lucia stratigraphic section (M. Cairo); upper Barremian. 1. 'Rivularia' lissaviensis (BORNEMANN)(X 15); Monte della Selva (Gran Sasso d'Italia); Hettangien. 2. 'Rivularia' fruticulosa (JOHNSON~z KASKA)(X 15); Colle Marzo; Kimmdridgien. 3. Hedstroemia moldavica DRAGASTAN(X 30); Villa S. Lucia (M. Cairo);Barremien. 4. Garwoodia maxima DRAGASTAN(X 30); Villa S. Lucia (M. Cairo); Barremien. 5. 'Rivularia' cf. theodori DRAGASTAN(X 30); Colle S. Lucia (M. Cairo); Barremien supgrieur.

'Rivularia'piae (FROLLO,1938) Fig. 8.1 1938

Cayeuxia piae nov. sp. - Frollo, pl. 18.

1985 Rivularia piae (FROLLO) - Dragastan, pl. 3, figs 5-6; pl. 9, figs 1-5; pl. 10, figs 1-2 (and references therein). 1988 Rivularia piae (FROLLO) - Dragastan, pl. 2, fig. 1. 1990 Rivularia piae (FROLLO) - Kuss, pl. 19, figs 1-2.

540 7 - Main occurrences of 'Rivularia' lissaviensis. The numbers are related to the following references. 1. Bornemann 1887 (upper Rethian). 2. Kamptner,1951 (Tithonian). 3. Derin & Reiss 1966 (Bajocian). 4. Praturlon 1966 (Hettangian-Toarcian). 5. Bouroullec & Deloffre 1968, 1970 (Oxfordian-Aptian). 6. Deloffre 1962 (Sinemurian). 7. Catalano et al. 1974 (Tithonian). 8. Duzbastilar 1978 (Oxfordian-Tihonian). 9. Bucur 1980-81, 1994 (Berriasian-lower Aptian), Dragastan 1985 (upper Valanginianlower Aptian). 10. Badve & Kundal 1986 (Cretaceous). The asterisk indicates the present paper. Principaux gisements de 'Rivularia' lissaviensis d'apr~s: 1. Bornemann 1887 (Rhdtien supdrieur); 2. Kamptner 1951 (Tithonique); 3. Derin & Reiss 1966 (Bajocien); 4. Praturlon 1966 (HettangienToarcien); 5. BourouUec & Deloffre 1968, 1970 (Oxfordien-Aptien); 6. Deloffre 1962 (Singmurien); 7. Catalano et al. 1974 (Tithonique); 8. Duzbastilar 1978 (Oxfordien-Tihonique); 9. Bucur 1980-81, 1994 (Berriasien-Aptien infgrieur), Dragastan 1985 (Valanginien supgrieur-Aptien infgrieur); 10. Badve & Kundal 1986 (Crgtacg). Astgrisque: gisments gtudigs. FIGURE

1991 Rivularia piae (FROLLO)- Diizbastilar, pl. 1, fig. 3-4. 1993 Rivularia piae ( F R O L L O ) - Bucur et al., pl. 2, fig. 6. 1993 Rivularia piae ( F R O L L O ) - Dragastan, pl. 2, figs 2-3; pl. 3, fig. 2. 1993 Rivularia piae ( F R O L L O ) - Dragastan et al., pl. 2, fig. 4. 1994 Rivularia piae ( F R O L L O ) - B u c u r , pl. 21, fig. 8. 1996 Rivularia piae ( F R O L L O ) - Dragastan et al., pl. 4, fig. 4; pl. 5, figs 2-3; pl. 6, fig. 3-4; pl. 7, fig. 2.

- Hemispherical, subspherical or e l l i p s o i d a l t h a l l u s , 3.0-4.5 m m w i d e a n d 1.5-3.0 m m h i g h , c r o s s e d b y t u b u l a r f i l a m e n t s , r a d i a l l y to c h a o t i c a l l y a r r a n g e d , 'V' d i c h o t o m i c a l l y b r a n c h e d , w h i c h d i v e r g e u n d e r a n a n g l e o f 5°-10 ° . T h e t h a l l u s shows a 'semi-lax' inner structure, the filaments a r e s u b p a r a l l e l a n d t h e i r i n n e r d i a m e t e r is a b o u t 0.065-0.090 m m .

Stratigraphic range - Observed for the first time in the Oxfordian-Aptian of Romania (Dragastan, 1985), 'Rivularia' theodori is known in the following localities: Kimmeridgian of Aquitaine, France (Dufaure 1958); Valanginian of Greece (Richter et al. 1992); Barremian of Romania (Bucur 1993); early Cretaceous of Greece (Dragastan et al. 1998). This species has been found for the first time in the Apennines (Barremian of Gran Sasso d'Italia).

Description

F a m i l y SCITONEMATACEAE K u t z i n g , 1883 G e n u s O r t o n e l l a GARWOOD, 1914 Type-species - Ortonella furcata GARWOOD,1914 Ortonella lemoineae DRAGASTAN, 1971 Fig. 4.1

Stratigraphic range - 'Rivularia' piae is a very widespread species; observed for the first time in the Tihonian of Romania (Frollo 1938), it is known in many localities (see Fig.9). In the Apennines 'Rivularia'piae has been found in all the examined areas (M. Aurunci, M. Cairo and Gran Sasso D'Italia); it ranges from Hettangian to Albian. ' R i v u l a r i a ' theodori DRAGASTAN, 1985 Fig. 2.4; Fig. 3.3

1958 Cayeuxia sp. - Dufaure, pl. 14, fig. 16. 1985 Rivularia theodori nov. sp. - Dragastan, pl. 3, figs 3-4; pl. 14, figs 1-7. 1992 Rivularia theodori DRAGASTAN- Richter et al., pl. 4, fig. 4. 1993 Rivularia theodori DRAGASTAN- B u c u r , pl. 2, fig. 9. 1993 Rivularia theodori DRAGASTAN- Dragastan, pl. 2, fig. 5. 1998 Rivularia theodori DRAGASTAN- Dragastan et al., pl. 2, fig. 5.

D e s c r i p t i o n - S m a l l f a n l i k e t h a l l u s , 1.0-1.6 m m w i d e a n d 0.8-1.2 m m h i g h , c r o s s e d b y t u b e filam e n t s 'V' d i c h o t o m i c a l l y b r a n c h e d w h i c h d i v e r g e u n d e r a n a n g l e o f 3°-10°; t h e y a p p e a r r a t h e r r e g u l a r l y a r r a n g e d , b r a n c h i n g o n l y once, a t a b o u t t h e s a m e h e i g h t . T h e i n n e r d i a m e t e r o f t h e f i l a m e n t s is 0.060 - 0.075 m m .

1971 1985 1992 1998

Ortonella lemoineae nov. sp. - Dragastan, pl. 10, figs 5-8. Ortonella lemoineae DRAGASTAN- Dragastan, pl. 16, fig. 3. Ortonella lemoineae DRAGASTAN- Richter et al., pl. 4, fig. 5. Ortonella lemoineae DRAGASTAN- Dragastan et al., pl. 3, fig. 3.

Description

- H e m i s p h e r i c a l , n o d u l a r t h a l l u s , 1.11.3 m m w i d e a n d 0.9-1.1 m m h i g h , c r o s s e d b y r a d i a l t u b e f i l a m e n t s ~V" d i c h o t o m i c a l l y b r a n c h e d , w h i c h d i v e r g e u n d e r a v e r y w i d e a n g l e (40°-50°). T h e t u b e a r e long, s t r a i g h t o r s l i g h t w a v y , w i t h a d i a m e t e r o f 0.035-0.055 m m . T h e t h a l l u s s h o w s a 'lax' inner structure.

Stratigraphic range - Observed for the first time in the upper Neocomian of Romania (Dragastan 1971), this species is known in the following localities: Valanginian of Greece (Richter et al. 1992), late Berriasian-Valanginian of Romania (Dragastan et al. 1998). Ortonella lemoineae has been observed for the first time in the Apennines; it has been found in the upper Barremian of Gran Sasso d'Italia. Ortonella perconigi DRAGASTAN, 1992 Fig. 4.4

1992 Ortonella perconigi nov. sp. - Dragastan, pl. 4, figs 4-6.

541

2

I

3

FIGURE 8 - 1. 'Rivularia' piae (FROLLO)(X 30); M. Sorgenza stratigraphic section (M. Aurunci); Aalenian. 2. Hedstroemia moldavica DRAGASTAN(X 30); Costalunga stratigraphic section (M. Cairo); Callovian. 3. 'Rivularia' theodori DRAGASTAN(× 30); Villa S. Lucia stratigraphic section (M. Cairo); Barremian. 4. Garwoodia polytomicaDRAGASTAN(X 15); Colle S. Lucia stratigraphic section (M. Cairo); upper Barremian. 1. 'Rivularia' piae (FROLLO) (X 30); M. Sorgenza (M. Aurunci); Aalenien. 2. Hedstroemia moldavica DRAGASTAN(X 30); Costalunga (M. Cairo); Callovien. 3. 'Rivularia' theodori DRAGASTAN(X 30); Villa S. Lucia (M. Cairo); Barremien. 4. Garwoodia polytomica DRAGASTAN(X 15); Colle S. Lucia (M. Cairo);Barremien supgrieur.

D e s c r i p t i o n - Subspherical, fanlike thallus about 1.0 m m wide and 1.1 m m high, which show a 'lax' i n n e r structure. The radially a r r a n g e d tube fila-

ments are dichotomic, isotomic ~Y' branched; t h e y diverge u n d e r an angle of about 50 ° . The main tube filament, before branching, shows a funnel shape

542 and its diameter is about 0.035 mm (proximal part) and 0.080 mm. (distal part, just under the branching). The isotomic tube filaments are short and their diameter is 0.050-0.055 ram. The distance between two successive dichotomic branchings is about 0.030 mm. Stratigraphic range - Known only in the Berriasian of Romania (Dragastan 1992), Ortonella perconigi h a s been observed for the first time outside the type-locality. In the Apennines it has been found in t h e Berriasian of G r a n Sasso d'Italia.

CALCAREOUS ? ALGAE Phylum ? CLOROPHYTAPascher, 1914 Class ? BRIOPSIDOPHYCEAERound, 1963 Order ? BRYOPSIDALESSchaffner, 1922 Family ? PSEUDOUDOTACEAEDragastan, 1997

1997

- Small hemispherical thallus, slightly zonated, 2.6-5.7 mm wide and 2.9-5.7 mm high, crossed by moniliform siphons, dichotomically branched under a right angle. The siphons are straight and parallel; their diameter is 0.030-0.060 ram, in the proximal part and near the top, respectively. The sheath-walls have a thickness of about 0.030 mm. In cross section the particular arrangement of the siphons forms a honeycomb characteristic feature; the space between the sheath-walls is filled with microcrystalline calcite. Description

Stratigraphic range - Garwoodia bardosi is known in t h e lower A p t i a n of Romania ( D r a g a s t a n 1985) a n d in t h e Valanginian of Greece (Richter et al. 1992); this species h a s b e e n found for the first time in t h e A p e n n i n e s (upper B a r r e m i a n of Monte Cairo).

Genus Garwoodia WOOD, 1941 Type-species - Garwoodia gregaria

(NICHOLSON,

although possibly a n alga, is best regarded as Microproblematicum.

Garwoodia bardosi DRAGASTAN,1985 Fig. 4.5 1985 1992

Garwoodia fissa DRAGASTAN,1989 Fig. 3.5

1888).

Remarks - According to Riding, 1991 the genus Garwoodia,

Garwoodia bardosi nov. sp. - Dragastan, pl. 20, figs 15. Garwoodia bardosi DRAGASTAN- Richter et al., pl. 4, figs 6-7.

Garwoodia bardosi DRAGASTAN- D r a g a s t a n et al., pl. 2, figs 2-5; pl. 4, fig. 1.

1989 1997 1998

Garwoodia fissa nov. sp. - Dragastan, pl. 9, figs 1-4. Garwoodia ]~ssa D R A G A S T A N - D r a g a s t a n et al. pl. 4, fig. 2. Garwoodia fi88a D R A G A S T A N - D r a g a s t a n et al., pl. 11, fig. 1-3.

Description - Hemispherical thallus, 1.8-5.0 mm wide and 1.5-4.0 mm high, composed of dichotomic cylindro-conical siphons and rarely by trichotomic ones, branched under a right angle. The siphons show a diameter of 0.060 up to 0.200 mm. The shea-

FmURE 9 - Main occurrences of 'Rivularia' piae. The numbers are related to the following references. 1. Frollo 1938 (Tithonian). 2. Pfender 1939 (Hettangian-Toarcian). 3. E1]iott 1957 (Aptian). 4. Johnson 1964 (Oxfordian-Albian). 5. Johnson & Kaska 1965 (OxfordianBarremian). 6. Roda 1965 (Aalenian-Tithonian). 7. Praturlon 1966 (Sinemurian-lower Albian). 8. Fenninger & Holtz 1967 (Kimmeridgian-Tithonian). 9. Bouroullec & Deloffre 1970 (Sinemurian, upper Oxfordian-Tithonian). 10. Basson & Edgel i971 (Bajocian-Tithonian, Aptian). 11. Luperto-Sinni 1976 (Coniacian-Maastrichtian). 12. Duzbastilar 1978, 1991 (Hettangian-Toarcian). 13. Chiocchini & Mancinelli 1978 (Hettangian-Toarcian). 14. Canerot 1979 (Berriasian-lower Hauterivian). 15. Dragastan 1985 (Tithonian). 16. Kuss 1990 (lower Bajocian). 17. Bucur 1993, 1994 (Barremian-lower Aptian). 18. Dragastan et al. 1993.19. Dragastan eta]. 1996 (Tithonian-Hauterivian). The asterisk indicates to the present paper. Principaux gisements de 'Rivularia' piae d'apr~s: 1. Frollo 1938 (Tithonien); 2. Pfender 1939 (Hettangien-Toarcien); 3. Elliott 1957 (Aptien); 4. Johnson 1964 (Oxfordien-Albien); 5. Johnson & Kaska 1965 (Oxfordien-Barremien); 6. Roda 1965 (Aalenien-Tithonien); 7. Praturlon 1966 (Sindmurien-Albien infgrieur); 8. Fenninger & Holtz 1967 (Kimmdridgien-Tithonien); 9. Bouroullec & Deloffre 1970 (Sindmurien, Oxfordien supdrieur-Tithonien); 10. Basson & Edgel 1971 (Bajocien-Tithonien, Aptien); 11. Luperto-Sinni 1976 (Coniacien-Maastrichtien); 12. Duzbastilar 1978, 1991 (Hettangien-Toarcien); 13. Chiocchini & Mancinelli 1978 (Hettangien-Toarcien); 14. Canerot 1979 (Berriasien-Hauterivien infdrieur); 15. Dragastan 1985 (Tithonien); 16. Kuss 1990 (Bajocien infdrieur); 17. Bucur 1993, 1994 (Barremien-Aptian infdrieur); 18. Dragastan et al. 1993; 19. Dragastan et al. 1996 (Tithonien-Hauterivien). Astdrisque: gisements dtudi~s.

543 th-walls of the siphons are very thin (0.010 mm), so they are rarely preserved. Stratigraphie range - Garwoodia fissa is known only in t h e T i t h o n i a n - l o w e r A p t i a n of R o m a n i a ( D r a g a s t a n 1989; D r a g a s t a n et al. 1997; D r a g a s t a n et al. 1998). In the Apennines it has been found in the upper Barremian-Albian of Monte Cairo.

Garwoodia fluegeli DRAGASTAN,1985 Fig. 2.2 1985 Garwoodia fluegeli nov. sp. - Dragastan, fig. 3; pl. 19, figs 1-8. 1989 Garwoodia fluegeli DRAGASTAN Dragastan, pl. 11, fig. 3. 1993 Garwoodia fluegeli DRAGASTAN Dragastan, pl. 5, fig. 4; pl. 7, fig. 4. 1997 Garwoodia fluegeli DRAGASTAN Dragastan et al., pl.3, fig. 2.

the siphons appear separated by large spaces filled with microcrystalline calcite. In the cross sections, the pattern distribution of the siphons resembles a petaloid bunch. Stratigraphic range - Observed for the first time in the Tithonian of Romania ( D r a g a s t a n 1989) Garwoodia polytomica is also known in the Kimmeridgian-Berriasian of the same region (Dragastan 1997). In the Apennines this species h a s been found in the Aalenian of Colle Marzo, in the Berriasian of Gran Sasso d'Italia, and in the B a r r e m i a n of M. Cairo.

Garwoodia toomeyi DRAGASTAN,1989 Fig. 4.3

-

-

-

Hemispherical thallus, 2.70-2.80 mm wide and 1.7 mm high, crossed by dichotomic or trichotomic siphons, laterally branched under a right angle; their diameter is about 0.040-0.050 mm. The siphons, polygonal in cross section, show a typical honeycomb feature. Description

G a r w o o d i a m a x i m a DRAGASTAN, 1 9 8 9 Fig. 2.5; Fig. 6.4

Garwoodiamaxima nov. sp. - Dragastan, pl. 12, figs 1-2. Garwoodia maxima DRAGASTAN Dragastan, pl. 8, figs 1-4. Garwoodia maxima DRAGASTAN- Dragastan et al., pl. 1, -

Garwoodiatoomeyi nov. sp. - Dragastan, pl. 10, figs 3-8. Garwoodiatoomeyi DRAGASTAN Dragastan, pl. 6, fig. 3. Garwoodia toomeyi DRAGASTAN Dragastan et al., pl. 1, -

-

fig. 4.

-

Stratigraphie range - Garwoodia fluegeli is known in t h e B a r r e m i a n - A p t i a n of Romania (Dragastan 1937). In the Apennines this species is very widespread, r a n g i n g from Hettangian to Barremian.

1989 1993 1997

1989 1993 1997

Description - Hemispherical thallus 1.75-2.90 mm wide and 3.90-4.0 mm high, crossed by dichotomic siphons, branched under a right angle. Before branching, the main siphon has a rather small diameter (0.045 mm), compared with the following bigger diameter after branching (0.070-0.090 ram). The siphon sheaths, non-perforated, are preserved in microcrystalline calcite; their thickness is about 0.010-0.015 mm.

Stratigraphic range - Known only in the KimmeridgianTithonian of Romania (Dragastan 1989, 1993; D r a g a s t a n et al. 1997), Garwoodia toomeyi h a s been observed for the first time outside the type-locality. In t h e Apennines it has been found in the B a r r e m i a n of M. Cairo.

figs 1-3.

Fanlike thallus, 6.0 mm wide and 4.0 mm high. The main axial siphon, slightly moniliform is dicho- or trichotomic branched, under a right angle. The siphons, whose diameter is about 0.09 mm, have a non-perforated sheath, preserved in microcrystalline calcite or in granular coarse crystalline calcite the thickness of which is 0.0180.030 mm. The sheath-walls of the siphons are separated and the space among them is filled with microcrystalline calcite and looks like a 'double wall'. Description

Stratigraphie range - Observed for the first time in the Tithonian of Romania (Dragastan 1989) Garwoodia maxima is known also in the Upper Trias of Romania (Dragastan 1997) and of Greece (Dragastan et al. 1997). In the Apennines this species h a s been found only in the G r a n Sasso d'Italia area, where it ranges from B a r r e m i a n to Aptian.

Garwoodia polytomica DRAGASTAN,1989 Fig. 3.4 1989 1997

Genus Hedstroemia ROTHPLEZ,1913

-

Garwoodiapolytomica nov. sp. - Dragastan, pl. 10, figs 12. Garwoodiapolytomica DRAGASTAN Dragastan et al., pl. 1, -

figs 5-6; pl. 2, fig. 7. Description - Fanlike, flabelliform thallus, slightly zonated, 4.0-5.5 mm wide and 1.80-3.0 mm high. The siphons are polytomically branched (4-6 or 7) under a right angle. The main siphon has a diameter of 0.090-0.120 mm, the branched siphons of 0.040-0.060 mm, until 0.10 mm in their distal part. The sheath-wall of the siphons, rather irregular in shape, is generally preserved in microcrystalline calcite; its thickness is 0.010-0.020 mm. Frequently

Type-species - Hedstroemia halimedoidea

R O T H P L E Z ,

1913.

Remarks - According to Riding, 1991 the genus Hedstroemia belongs to cyanobacteria.

Hedstroemia moldavica DRAGASTAN,1985 Fig. 3.2; Fig. 6.3 1958 1964 1966 1985

Cayeuxia sp. - Dufaure, pl. 4, figs 3-4. Cayeuxia sp. - Colacicchi, fig. 17. Cayeuxia sp. - Derin & Reiss, fig. 56. Hedstroemia moldavica nov. sp. - Dragastan, pl. 21, figs 19; pl. 25, fig. 1.

- Hemispherical, fanlike thallus, 1.13.2 mm wide and 1.0-2.4 mm high, with a 'semilax' inner structure. The siphons are trichotomic, diverge under an angle of about 40-5 ° and their diameter is 0.45-0.60 mm; they show a characteristic fork shape and the diameter growing distally. Description

Stratigraphie range - Observed for the first time in the Bajocian-Callovian of France (Dufaure 1958), Hedstroemia moldavica is known in the following localities: Cenomanian of the Apennines (Colacicchi 1964); Bajocian of Israel (Derin & Reiss 1966); Tithonian-Aptian of Romania (Dragastan 1985); B a r r e m i a n of the same region (Bucur 1993). In the Apennines this species h a s been found in all the investigated areas and it extends from Bajocian to Cenomanian.

Hedstroemia racasiensis DRAGASTAN,1989 Fig. 2.1 1989

Hedstroemia racasiensis nov. sp. - Dragastan, pl. 3, fig. 1; pl. 4, fig. 1.

544

J L

U A

I

R

A S D 0

S

S

I

C

G G E R M

HETTANGIA~ $tNEMUR~A~ ......

A }

CRETACEOUS L

M |

LOWER CRETACEOUS

................................................................................

"Rivulada" piae "Rivularia" lissaviensis "Rivulada" kurdistanensis ,,,,

Garwoodia fluegeli

~

,

Garwoodia polytomica Hedstroemia moldavica "Rivularia" fruticulosa Hedstroemia villosa Hedstroemia racasiensis "Rivularia" carpathica "Rivularia" dianae ~..

Ortonella perconigi

!'

Garwoodia maxima "Rivutaria" theodori Garwoodia toorneyi Garwoodia fissa Ortonella lemoineae

I

Garwoodia bardosi

"paleopiattaforma"-- . . . .

Ik:¸ inner shetf facies . . . . .

back reef facies ~

t

shelfedge facies . . . . . . . .

FIGURE 10 - Stratigraphic range of the analyzed species and their distribution in the different facies of the Apennine platform.

Distribution stratigraphique des esp~cies dtudides dans les facies de la plate-forme Appenninica.

Description - Hemispherical, fanlike thallus, 2.53.5 mm wide and 1.4-2.5 mm high, composed of long waved siphons, radially arranged, with a rather variable diameter, 0.030 at the base up to 0.080 mm in their distal part. The branchings are dichotomic or, more frequently, trichotomic; the siphons diverge under an angle of 5°-10 °. Stratigraphic range - Known only in the KimmeridgianTithonian of Romania (Dragastan 1989), Hedstroemia racasiensis has been observed for the first time outside the typelocality. In the Apennines it has been found in the Berriasian of Gran Sasso d'Italia and in the lower Aptian of M. Cairo.

Hedstroemia villosa

DRAGASTAN, 1989 Fig. 2.6

1989 1992

Hedstroemia villosa nov. sp. - Dragastan, pl. 6, figs 3-4. Hedstroemia villosa DRAGASTAN- Richter et al., pl. 4, figs 3.

Description - Columnar erect thallus, 6.0 mm high and 2.0-4.0 mm wide, composed of thin long and rather parallel or slightly divergent siphons; their diameter is 0.010-0.015 mm (about 0.030 mm in their distal part). The branchings are dichotomic and trichotomic, and diverge under a very small angle (2°-3°).

Stratigraphic range - Observed for the first time in the Berriasian-Hauterivian of Romania (Dragastan 1989) Hedstroemia villosa is also known in the Valanginian of Greece (Richter 1992). In the Apennines this species has been found in the Berriasian-lower Aptian of Gran Sasso d'Italia and M. Cairo.

CONCLUSIONS The rich fossil association that we have recognized in our material, sampled in the Jurassic and Lower Cretaceous carbonate sediments cropping out in the Latium-Abruzzi area, shows that: - all the species, except 'Rivularia' piae and Hedstroemia moldavica have been observed for the first time in Italy; - many species have a very short stratigraphic range, as shown in the Figure 10. They are: 'Rivu-

laria' fruticulosa, 'Rivularia' dianae, 'Rivularia' carpathica, Ortonella perconigi, Garwoodia toomeyi, 'Rivularia' theodori, Garwoodia bardosi and Ortonella lemoineae; - the same figure shows that the recognized species seems to be particularly frequent and abundant during some intervals of time, i.e., middle Jurassic (Bajocian-Callovian), early Cretaceous (BerriasianValanginian and Barremian-lower Aptian).

545 The abundance of these species in some stratigraphic sections of the Apennine carbonate platform, deposited near the shelf edge, confirm that these algae are conditioned by particular water factors, controlled by the hydrodynamism; they seem to prefer the back-reef environment, characterized by shallow water and a more or less high hydrodynamism and, subordinately, the shelf edge environment. On the contrary, their aboundance in the innermost shelf-lagoon seems to be restricted to some intervals of time (i.e. Lias-lower Dogger), as shown in Figure 10. Acknowledgements - This research was supported by the grant of the MURST (Fondo di Ricerca d'Ateneo 1998, 1999), A. Mancinelli responsable. The authors thank the two referees O. Dragastan and R. Riding for their suggestions.

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