Bollettino della Società Paleontologica Italiana, 52 (2), 2013, 123-138. Modena

The Mt Perda Liana section (Middle , central-eastern ): revised stratigraphy and faunas

Iginio Dieni, Francesco Massari & Vladan Radulović

I. Dieni, Dipartimento di Geoscienze, Università degli Studi di Padova, Via Gradenigo 6, I-35131 Padova, ; [email protected] F. Massari, Dipartimento di Geoscienze, Università degli Studi di Padova, Via Gradenigo 6, I-35131 Padova, Italy; [email protected] V. Radulović, Department of Palaeontology, Faculty of Mining and Geology, Kamenička st. 6, P. O. Box 162, RS-11000 Belgrade, Serbia; [email protected]

KEY WORDS - Middle Jurassic, Sardinia, Lithostratigraphy, Biostratigraphy, .

ABSTRACT - The Jurassic in age stratigraphic section of Mt Perda Liana, the classic locality of the “Tacchi” area (central-eastern Sardinia), has been studied from a stratigraphic and palaeontologic viewpoint. A rich fauna is contained in a limestone interval comprised between the siliciclastic deposits of the Genna Selole Formation (Bajocian-lowermost Bathonian) and the typical dolostones of the Formation. This limestone interval, also present in other localities of the “Tacchi” area, is here formally distinguished as member of the Dorgali Fm. (Perda Liana Member) due to its peculiar lithology and faunal content. It mainly consists of shelf-lagoonal micritic limestones, particularly rich in infaunal bivalves. The presence of the brachiopods Kallirhynchia oranensis (Flamand, 1911) and Holcothyris angulata Buckman, 1918, allows the attribution of the Perda Liana Member to the Lower Bathonian.

RIASSUNTO - [La sezione di Monte Perda Liana (Giurassico Medio, Sardegna centro-orientale): revisione stratigrafica e fauna a brachiopodi] - Viene illustrata la sezione stratigrafica della classica località di Perda Liana, ubicata nella regione dei “Tacchi”. Una ricca fauna, in parte già nota, è contenuta nell’intervallo calcareo compreso tra i sedimenti silicoclastici della Formazione di Genna Selole, attribuita al Bajociano e al Batoniano basale, e le tipiche dolomie della Formazione di Dorgali. Questo intervallo calcareo, identificato in alcune altre località dei “Tacchi”, viene qui distinto per i suoi caratteri peculiari di fauna e litologia come membro della Formazione di Dorgali e formalizzato come nuova unità litostratigrafica (Membro di Perda Liana). Si tratta prevalentemente di calcari micritici indicativi di un ambiente di bassa energia identificabile come laguna aperta, il cui fondo ospitava una ricca fauna di bivalvi fossatori. Particolarmente significativi dal punto di vista biostratigrafico si sono rivelati i brachiopodiKallirhynchia oranensis (Flamand, 1911) e Holcothyris angulata Buckman, 1918, indicativi del Batoniano Inferiore. Terebratula lamarmorae Meneghini, 1857, viene considerata come nomen oblitum a favore di Holcothyris angulata Buckman, 1918, che diviene pertanto nomen protectum.

INTRODUCTION The continental to marginal-marine sedimentation gave way to marine carbonate ramp sedimentation, more The lower part of the Jurassic succession of eastern completely preserved in the Ogliastra, Supramonte and Sardinia is generally made up of dolostones which failed Baronie areas of eastern Sardinia - as exemplified by generally to give information useful for a biostratigraphic the Dorgali Formation, Mt Tului Limestone, locally attribution, due to the scarcity or absence of fossils. grading basinwards into S’Adde Limestone, and Mt However, in places the succession includes calcareous Bardia Limestone (Amadesi et al., 1961; Casellato et al., lenses to layer bundles of limited lateral persistence, 2012, and references therein). The deposits share several containing rich fossil assemblages, mostly bivalves and characteristics of the Jurassic carbonate platforms of the less abundant brachiopods, allowing to establish some south-European margin (Stampfli et al., 2002; Costamagna firm chronostratigraphic points. One of the richest section, & Barca, 2004), for instance in Provence (Fourcade et useful to this purpose, is that of Mt Perda Liana in the al., 1977) and in the Corsica autochthonous succession “Tacchi” area of central-eastern Sardinia, which is taken (Peybernès et al., 2001). Similar record of Middle Jurassic into account in this paper, by illustrating stratigraphic and transgressive carbonates bounded at the base by a marked palaeontologic data. unconformity is known in the Briançonnais (Bourbon et The first unit of the local Jurassic cover is represented al., 1973), and part of the Swiss and French “Préalpes by continental to marginal-marine siliciclastic deposits médianes” (“Domain intermédiaire”; Septfontaine, 1983). of the Genna Selole Formation (defined by Dieni et al., 1983), transgressive on the Palaeozoic and locally substrate and bearing palynomorphs and plant macrorests GEOLOGIC AND STRATIGRAPHIC SETTING indicating a Bajocian-Bathonian age (Dieni et al., 1983; (I. Dieni & F. Massari) Del Rio, 1984). The basal, coarse-grained conglomerates record alluvial fan sedimentation accounting for denudation Geologic setting of a rugged relief, with deep erosion of the substrate. The Monte Perda Liana (sometimes written as This episode is conceivably related to the rifting which Perdaliana or Perda ’e Liana; hereafter simply called affected the south-European margin in the Middle Jurassic, Perda Liana) is an isolated, tower-shaped mount (1293 m generating a swell-and-basin topography, concomitantly a.s.l.), located in central-eastern Sardinia, in the territory with the initial Tethys opening (Lemoine et al., 1978). of Gairo (boundary area between the Barbagia of Seulo

ISSN 0375-7633 doi:10.4435/BSPI.2013.21 124 Bollettino della Società Paleontologica Italiana, 52 (2), 2013

remains of a much wider carbonate platform covering most part of Sardinia. Their morphology somewhat recalls that of the mesas of the Monument Valley (U.S.A.). As outlined above, the Perda Liana section, like most part of the “Tacchi”, is the record of the lower part of the Jurassic succession of eastern Sardinia, more completely developed in the Ogliastra, Supramonte and Baronie areas. An appreciable synthesis on the Middle Jurassic succession of the “Tacchi” area, highlighting the state of the art on these deposits, especially from the viewpoints of the palaeogeography and regional geology, was presented by Costamagna & Barca (2004). They distinguished three units within the Genna Selole Formation on lithostratigraphic grounds: the conglomeratic “Laconi- Lithofacies”, the sandy-clayey “Nurri-Escalaplano Lithofacies” and the mixed siliciclastic-carbonate “Ussassai- Lithofacies”, grading upwards into the Dorgali Formation. They argued that the Genna Selole type-section, located in the Ogliastra region, is Fig. 1 - Distribution of Jurassic formations in central-eastern Sardinia (from Dieni et al., 1985, modified). poorly representative and inadequate to illustrate the full range of lithotypes in the Tacchi area. For this reason they proposed a formal redefinition of the formation, which would include the three above mentioned lithofacies, and Ogliastra). It is one of the so called “Tacchi” (Fig. 1) introducing two parastratotypes in the areas of Escalaplano of central-eastern Sardinia, hills generally rising from the and Perdasdefogu, characterised by good exposures Palaeozoic basement (Fig. 2) and consisting of carbonate and continuity. The threefold subdivision proposed by mesas. They are true natural monuments of characteristic Costamagna & Barca (2004) was accepted by Scanu et al. aspect, resembling castles, towers and ruins (Barrocu (2012) in a preliminary study of the Jurassic plant fossils & Gentileschi, 1966; Fig. 3a), and represent isolated based on the revision of the Lovisato Collection.

Fig. 2 - Geologic sketch-map of the Mt Perda Liana area (from Atzeni, 1967). 1: limestone and dolostone (Dorgali Formation, Middle Jurassic); 2: quartz-conglomerate, arenite and carbonaceous claystone (Genna Selole Formation, lower Middle Jurassic); 3: quartz-porphyry in dykes and subvolcanic bodies (Variscan cycle); 4: Variscan granite; 5: graptolite-bearing slate (Wenlock, ); 6: phyllite (Llandovery, Silurian); 7: Orthis-bearing slate and quartz-phyllite (); 8: fault. I. Dieni et alii - Bathonian of Mt Perda Liana and its brachiopods 125

Fig. 3 - View of the Perda Liana tower with trace (dashed line) of the measured section of Fig. 4 (a) and the first stratigraphic description (b) due to La Marmora (1857, fig. 29 at p. 157).

The stratigraphic succession of Perda Liana commonly marly, with conchoidal fracture, in places The stratigraphic succession of Perda Liana is containing a small fine-grained quartz fraction. A lower palaeontologically well known as it yielded abundant and plane-parallel-bedded interval 19 m thick shows muddy varied macrofaunas, formerly studied by La Marmora interbeds, plant remains, molluscs (among which very (1857; Fig. 3b), Meneghini (1857), Fucini (1894, 1899), abundant infaunal bivalves, usually in life position), Pampaloni (1900) and Dainelli (1903). A preliminary study common brachiopods and characteristic bioclastic infilling of the section was performed by De Biasi (1985). of burrows. It grades into an interval 20.5 m thick, The lower siliciclastic part of the succession (of which where compaction on thorough bioturbation resulted in a only the uppermost portion is represented in the log of Fig. predominantly flaser to nodular bedding (Fig. 4). 4) may be attributed to the Genna Selole Formation (sensu 4. Unit IV, 8.5 m thick, is a succession of three, Dieni et al., 1983), and may be correlated to the lithofacies normally graded layers, individually comprising a lower recognized in the “Tacchi” area by Costamagna & Barca grainstone interval with hummocky cross bedding and (2004) respectively as “Laconi-Gadoni Lithofacies” and ripples in the upper part, and an upper bioturbated, flaser- “Nurri-Escalaplano Lithofacies”. Conversely, we do bedded wackestone/packstone interval similar to the not share the opinion of these authors concerning the lithology of the underlying unit. The macrobenthos is inclusion of their “Ussassai-Perdasdefogu Lithofacies”, represented by molluscs and rare brachiopods. mostly consisting of limestone, in the Genna Selole 5. Unit V, about 5 m thick, consists of thin- to medium- Formation. Considering its palaeontological content bedded grainstone layers, generally graded and rippled/ and environmental setting we believe that this unit can bioturbated in the upper part. The nerineid Cossmannea be formally distinguished as a member of the Dorgali cf. eudesii (Morris & Lycett, 1851) locally occurs in this Formation (Perda Liana Member, see below), due to its interval together with other sparse molluscs and plant essentially carbonate lithology. remains. The interval is bounded at the base by an erosional Six units may be distinguished in the sedimentary surface draped by a coarse layer of quartz pebbles. succession: 6. Unit VI corresponds to the typical, thick-bedded dolostone facies of the Dorgali Formation, of which only 1. Unit I (not illustrated in the log) is represented by the initial part has been measured. This facies forms the about 15 m of poorly outcropping, scarcely cemented vertical walls of the Perda Liana tower, with an estimated whitish, clast-supported quartz conglomerate, with clasts height of about 70 m (Fig. 3). of pluricentimetric dimensions, clearly derived from veins of the underlying crystalline schists. The interval including units III to V has been chosen 2. Unit II, a few metres thick, consists of an association as type-section of the Perda Liana Member of the Dorgali of massive fine quatzarenite and mudstone, with common Formation. plant remains and lignite seams. The top layer is a tightly cemented arenite with sub-spherical ferruginous concretions (formerly iron sulphides) commonly bearing MATERIAL AND METHODS plant remains in their core. 3. Unit III consists of grey-beige, very fine-grained, After the necessary field survey, the Perda Liana heavily bioturbated, limestone (wackestone and mudstone), section has been logged and about 50 samples of the 126 Bollettino della Società Paleontologica Italiana, 52 (2), 2013

Fig. 4 - Stratigraphic log of the type-section of the Perda Liana Member. I. Dieni et alii - Bathonian of Mt Perda Liana and its brachiopods 127 various lithofacies were collected. In addition, a number associated with plant remains, such as Viviparus scoticus, of thin sections were prepared. The numerous macrofossils probably transported from nearby areas. collected are housed in the Museo di Geologia e Paleontologia of the Università di Padova except for the brachiopods of the Meneghini Collection which are FAUNAL CONTENT housed in the Museo di Storia Naturale e del Territorio of the Università di Pisa. The Perda Liana fossil content is certainly the richest and taxonomically most various assemblage within the Middle Jurassic successions of Sardinia. Most fossils RESULTS studied by former authors (Meneghini, 1857, who determined the macrofaunas collected by La Marmora; Microfacies description Pampaloni, 1900, who presented a brief list of fossils, Unit III mostly consists of a wackestone, rarely most of them subsequently studied in detail by Dainelli, packstone, with sparse angular quartz grains of variable 1903) were mostly obtained from the scree deposits at the size, sometimes sorted out and concentrated into the foot of the tower-like hill, given their abundance in the burrows. Bioturbation is diffuse and framboidal clusters of limestone debris fallen from the cliffs. Here we present a iron sulphide are quite common. Remains of bivalves and preliminary list of most characteristic taxa, while a more gastropods, sometimes micritized, are present with variable complete palaeontologic and biostratigraphic study of abundance and commonly show marginal micro-borings. the Middle Jurassic macrofaunas of eastern Sardinia will They are preserved as calcite moulds after aragonite, be the subject of another paper in progress. Of the fossils with lacking evidence of shell collapse indicating that here reported, several specimens come from the detritus; calcite precipitation into dissolution voids occurred after however those presented alongside the stratigraphic log lithification of the sediment. Other minor constituents of Fig. 4 were collected in situ. observable in thin sections are foraminifers (such as In addition to the following list, the presence of rare nodosariids, textulariids, miliolids, “siphovalvulinas” specimens of large discoidal solitary corals and teeth of and encrusting forms), echinoderm fragments, ostracods, hybodontid sharks (Fig. 5) is worth mentioning. sparse ooids and peloids. The thin sections 3 and especially 4 (Fig. 4) show a peculiar composition, with abundant Brachiopods sponge spicules and radiolaria. Brachiopod remains are Kallirhynchia oranensis (Flamand, 1911) particularly abundant in the upper part of the third unit. Holcothyris angulata Buckman, 1918 The three layers of unit IV consist of a grainstone in the lower part grading into a packstone and wackestone Bivalves upwards. Worth noting is the presence in this unit, among Grammatodon (Cosmetodon) sp. the foraminifers, of Bosniella croatica (Gušić, 1969) and Modiolus (M.) imbricatus J. Sowerby, 1818 Amijiella amiji (Henson, 1948). Unit V mostly consists Inoperna plicata (J. Sowerby, 1819) of an ooidal grainstone. Pinna subcancellata (Lissajous, 1923) Trichites cf. thurmanni Choffat, 1888 Environmental interpretation Pteria plana (Morris & Lycett, 1854) Units I and II were deposited in a continental setting Pteroperna costatula (Deslongchamps, 1824 (alluvial fan to braided stream), grading into brackish, (=Pteroperna fucinii Dainelli, 1903) and finally marine-marginal (marshy coastal plain) Costigervillia crassicosta (Morris & Lycett, 1853) environment. Unit III may be attributed to a low-energy Eligmus perdalianae (Meneghini, 1857) open lagoon (shelf lagoon) environment. The local abundance of siliceous organisms suggests blooms related to episodes of high productivity. Unit IV records a high- energy, storm-related back-barrier setting, alternating with a low-energy lagoonal setting. The erosional surface at the base of unit V is thought to represent a “ravinement” surface marking the transition to storm-dominated, inner- ramp, marine deposits. The interval comprising units III to V corresponds to the calcareous variety (b) of the “Ussassai-Perdasdefogu lithofacies” of Costamagna & Barca (2004), typical of the north-eastern “Tacchi” area. In a study of salinity-controlled Bathonian benthic faunas of the Causses (southern ) Fürsich et al. (1995) attribute the Pholadomya lirata and Ceratomya striata associations to a normal marine environment. In the Perda Liana unit III bivalves of the above associations are well represented and, together with the evidence of low-energy environment, suggest that the assumed shelf- lagoon environment was in a wide connection with the open sea. Only the lowermost part of the third unit contains Fig. 5 - Tooth of hybodontid shark (a: upper view; b: lateral view) some adapted to a lower-salinity environment (MGP 31180). 128 Bollettino della Società Paleontologica Italiana, 52 (2), 2013

Entolium (E.) corneolum (Young & Bird, 1828) succession of eastern and central Sardinia, has in general Camptonectes (C.) laminatus (J. Sowerby, 1818) quite uniform dolomitic lithology and is usually barren Plicatula sp. of fossils, so that its age is poorly constrained. However, Placunopsis socialis Morris & Lycett, 1853 (= important biostratigraphic data are yielded by some locally Placunopsis pampalonii Dainelli, 1903) occurring lenses of even thin packages of limestone beds, Plagiostoma hellicum (d’Orbigny, 1850) in places partly dolomitized, generally containing abundant Trigonia (T.) cf. hemisphaerica Lycett, 1850 fossils. These fossil-bearing limestones occur at the base, Mesomiltha bellona (d’Orbigny, 1850) as in the case of Perda Liana, or at various levels within Protocardia (P.) lycetti Rollier, 1912 the Dorgali Formation, such as the lenses of limestone to Pachymya (Arcomya) meneghinii (Dainelli, 1903) silty limestone identified in the Monte Albo area (Janna Pholadomya (Bucardiomya) lirata (J. de C. Sowerby, Portellitos and Punta Casteddu sections) containing 1818) brachiopods indicating the Bremeri and Retrocostatum Homomya gibbosa (J. Sowerby, 1813) zones, such as Burmirhynchia turgida Buckman and Gresslya peregrina (Phillips, 1829) Arceythyris diptyca (Oppel), which testify the Upper Ceratomya striata (J. de C. Sowerby, 1812) Bathonian (Alméras et al., 1997). Actinostreon gregareum (J. Sowerby, 1815) Here we reserve however the name of Perda Liana Member only to the lower calcareous package of the Gastropods Dorgali Formation, which should correspond to the Viviparus scoticus (Tate, 1873) (= “Viviparus variety “b” of the “Ussassai-Perdasdefogu lithofacies” aurelianus Benn.” reported by Dorn, 1940, p. 326, in of Costamagna & Barca (2004), characterised by the “Tacchi” area) predominantly calcareous lithology and confined in the Globularia ranvillensis (d’Orbigny, 1850) (= Natica north-eastern part of the “Tacchi” area. In our opinion, parthenica Meneghini, 1857) even their mostly dolomitic variety “a” should not be Harpagodes wrightii (Morris & Lycett, 1851) distinguished as a separated unit from the essentially dolomitic facies of the Dorgali Formation. There is Echinoids no doubt that the first two lithofacies distinguished by Acrosalenia hemicidaroides Wright, 1851 Costamagna & Barca (2004), i.e. the “Laconi-Gadoni and Nurri-Escalaplano lithofacies”, characterised by terrigenous lithology, are to be included in the Genna BIO- AND CHRONOSTRATIGRAPHY Selole Formation. On the other hand, considering the lithostratigraphic and environmental relationships of the Among the fossils occurring in the studied section, limestone package with the dolostones of the Dorgali some forms are long ranging, either from the Bajocian Formation, in our opinion this carbonate unit should be to the Kimmeridgian (Modiolus imbricatus, Inoperna regarded as a member of this formation, instead of part plicata, Entolium corneolum, Actinostreon gregareum, of the Selole Formation as claimed by Costamagna & Gresslya peregrina, etc.), or from the Bajocian to the Barca (2004). Bathonian or the Callovian (Bosniella croatica, Amijiella The fossil content of the Perda Liana Member, amiji; Pteroperna costatula, Camptonectes laminatus, dominated by bivalves, is generally quite uniform and Pholadomya lirata, Homomya gibbosa, Ceratomya striata, generally indicates an Early Bathonian age (see above). etc.). Other taxa, such as Pinna subcancellata, Placunopsis As type-section of the Perda Liana Member of the Dorgali socialis, Plagiostoma hellicum, Pachymya meneghinii, Formation, we selected the Perda Liana section, whose Viviparus scoticus, Globularia ranvillensis, Harpagodes characteristic features (lithology, thickness, sedimentary wrightii, Acrosalenia hemicidaroides, etc., have a more structures, fossils, etc.) are presented in the stratigraphic restricted stratigraphic distribution, being exclusive of the log of Fig. 4. The lower boundary of the member is with Bathonian. The Bathonian age of the assemblage is thus the siliciclastic facies of the Genna Selole Formation, the confirmed, as formerly established by Pampaloni (1900) upper boundary is marked by the first dolostone beds of and Dainelli (1903). A further age precision is provided the Dorgali Formation. As supplementary section we may by the two species of brachiopods Kallirhynchia oranensis mention the one exposed NW to the village of (Fig. and Holcothyris angulata. K. oranensis occurs in one level 1) (locality Gedili, near Bruncu Pranedda) for the lower (6: Fig. 4) while H. angulata was collected from two levels part of the member, already studied by Dorn (1940), and (7 and 9: Fig. 4). They document the Oranensis Zone of the one located SW of the same village (locality Matta the brachiopod biostratigraphic zonation, which in turn can Prana, western slope of Bruncu Matzeu), for the upper part. be correlated to the Lower Bathonian Zigzag Zone of the ammonite chronostratigraphic zonation (Alméras et al., 1997). Therefore, the Perda Liana calcareous interval, here SYSTEMATIC PALAEONTOLOGY erected as lower member of the Dorgali Formation, can be dated, at least in its type-section, to the Early Bathonian. (I. Dieni & V. Radulović) Repositories - The studied specimens are housed in FORMAL INSTITUTION the following institutions: OF THE PERDA LIANA MEMBER - Museo di Geologia e Paleontologia, Università di Padova, Dieni Collection (MGP); The Dorgali Formation, which is widespread and - Museo di Storia Naturale e del Territorio, Università typical of the lower part of the Jurassic carbonate di Pisa, Meneghini Collection (MUP); I. Dieni et alii - Bathonian of Mt Perda Liana and its brachiopods 129

- Museo di Storia Naturale, Sezione di Geologia e 1994 Kallirhynchia oranensis (Flamand) sous-espèce A (Rousselle) Paleontologia, Università di Firenze, Dainelli Collection - Alméras et al., pp. 221, 225-228, 231, pl. 2, fig. 5. (IGF). 1994 Kallirhynchia oranensis (Flamand) sous-espèce quefaitensis (Rousselle) - Alméras et al., p. 231, pl. 2, fig. 6. 1995 Kallirhynchia oranensis (Flamand) - Ouali-Mehadji, p. 81, pl. 2, figs 5-6. rachiopoda Phylum B Duméril, 1806 1996 Kallirhynchia oranensis (Flamand) - Alméras et al., pp. 606, Order Kuhn, 1949 616, figs 10b-c. Family Tetrarhynchiidae Ager, 1965 2008 Kallirhynchia oranensis (Flamand) - Alméras & Fauré, p. 592, pl. 1, figs 3-10; tabs 1-2; text figs 3-6, figs 1-5 (cum Genus Kallirhynchia Buckman, 1918 syn.).

Kallirhynchia oranensis (Flamand, 1911) Material - Twenty-one specimens from the Dieni Figs 6, 7a-e, 10a-d Collection (MGP 31346-31351, 31357-31359, 31368- 31379, mostly collected in situ), four specimens from v 1857 Rhynchonella subobsoleta Davids - Meneghini, p. 273, pl. E, the Meneghini Collection (MUP I 13899, 13892, 13882a, figs 20a-c. 13882b) and two specimens from the Dainelli Collection v 1857 Rhynchonella sp. ind. - Meneghini, p. 276. (IGF 3487F, 3488E), all from Perda Liana, except for the v 1903 Rhynchonella cfr. subobsoleta Davidson - Dainelli, p. 270. specimen 3488E, which comes from the Tacco of , + 1911 Rhynchonella royeri d’Orbigny var. oranensis nov. var. - central Sardinia. Therefore, a total sum of 27 specimens Flamand, p. 911, pl. 11, figs 13-16. is available. The exact stratigraphic position of the 1911 Rhynchonella sublacunosa (non Szajnocha) - Flamand, p. 907, pl. 7, fig. 5. specimens coming from the Meneghini and Dainelli 1911 Rhynchonella dumortieri (non Szajnocha) - Flamand, p. 908, collections is unknown. pl. 7, fig. 6. ? 1935 Rhynchonella (Cymatorhynchia) obsoleta Sowerby - Description - Oosterbaan, p. 91. External characters (Figs 7a-e, 10a-d): Shell medium in 1964 Rhynchonella royeri d’Orb. var. oranensis Flam. - Rousselle, size (Tab. 1), rounded sub-pentagonal outline, subglobose p. 320. in lateral profile, lenticular in anterior view. Dorsi- 1965 Rhynchonella aff. royeriana oranensis Flamand - Rousselle & Demarez, p. 36, pl. 4a, figs 1-2 (forme A) and 3-4 (forme biconvex, dorsal valve twice more convex than the ventral B); text-pl. on p. 37, tab. 1, text-figs 1-2. valve. Maximum width at/or slightly anterior to mid- 1965 Kutchirhynchia oranensis (Flamand)? - Rousselle, p. 53, pl. length; maximum thickness lies slightly posterior of mid- 4, figs 5-14. length. Lateral commissure straight; anterior commissure 1965 Kutchirhynchia oranensis (Flamand)? sous-espèce A - moderately and roundly uniplicate or slightly asymmetric. Rousselle, p. 54, pl. 4, figs 5, 6, 8-11; text-figs 23-24. Fold not well developed, low and narrow, only slightly 1965 Kutchirhynchia? oranensis (Flamand) - Rousselle, p. 56, pl. raised above the slopes at anterior one-third. Sulcus narrow 4, figs 1-4. and shallow, occurring at the posterior half. Beak strong, 1965 Kutchirhynchia oranensis (Flamand)? quefaïtensis nov. moderate in height, sub-erect, foramen large, circular, subsp. - Rousselle, p. 56, pl. 4, figs 7, 12-14; text-fig. 25. 1965 Rhynchonella sublacunosa (non Szajnocha) Flamand - hypothyrid. Beak ridges sharp, short. Interareas small but Rousselle, p. 58, pl. 4, figs 15-16. well defined, slightly concave. Ribs coarse, subangular to 1986 Kutchirhynchia oranensis (Flamand) - Alméras & Gupta, p. sharp, on each valve numbering 16-21, with four-six on 425. fold and three-five in sulcus. Apical angle from 86° to 105°.

Specimen L W T W/L T/L T/W NRDV Commissure Remarks MGP 31346 16.0 16.5 7.6 1.03 0.47 0.46 18 asymmetric MGP 31368 16.1 16.4 10.9 1.02 0.68 0.66 18 symmetric MGP 31347 18.6 21.0 13.4 1.13 0.72 0.64 21 symmetric Fig. 7a MGP 31348 19.4 25.3 16.1 1.30 0.83 0.64 18 symmetric Fig. 7b MGP 31349 19.7 21.0 13.0 1.07 0.66 0.62 18 ?asymmetric Fig. 7c MGP 31350 21.2 23.7 17.3 1.12 0.82 0.73 20 symmetric sectioned, Fig. 7d MGP 31369 21.3 21.3 13.7 1.00 0.64 0.64 18 symmetric MGP 31370 23.8 26.1 16.6 1.10 0.70 0.64 19 symmetric MGP 31351 26.6 26.5 20.3 0.99 0.76 0.77 18 asymmetric Fig. 7e IGF 3488E 15.8 16.7 9.0 1.06 0.57 0.54 18 symmetric MUP I 13899 17.5 17.7 11.2 1.01 0.64 0.63 21 symmetric Fig. 10a MUP I 13892 19.7 20.0 12.4 1.01 0.63 0.62 16 symmetric Fig. 10b MUP I 13882a 21.6 20.8 14.2 0.96 0.66 0.68 21 asymmetric Fig. 10c MUP I 13882b 23.5 22.2 14.4 0. 94 0.61 0.65 21 symmetric Fig. 10d

Tab. 1 - Biometric parameters of the Kallirhynchia oranensis (Flamand, 1911) studied specimens. L = length; W = width; T = thickness; NRDV = number of ribs on the dorsal valve. 130 Bollettino della Società Paleontologica Italiana, 52 (2), 2013

Fig. 6 - Transverse serial sections of Kallirhynchia oranensis (Flamand, 1911) through the specimen MGP 31350 (the same as Fig. 7d), Lower Bathonian of Perda Liana. Numbers indicate the distance in mm from the tip of the ventral umbo. I. Dieni et alii - Bathonian of Mt Perda Liana and its brachiopods 131

Fig. 7 - Brachiopods from the Lower Bathonian of Perda Liana. a-e: Kallirhynchia oranensis (Flamand, 1911): a = MGP 31347; b = MGP 31348; c = MGP 31349; d = MGP 31350, used for transverse serial sections of Fig. 6; e = MGP 31351. f-i: Holcothyris angulata Buckman, 1918: f = MGP 31352; g = MGP 31353; h = MGP 31355; i = MGP 31356, weakly crushed, used for transverse serial sections of Fig. 9. All the specimens, coated with ammonium chloride, are in dorsal (1), ventral (2), lateral (3) and anterior (4) views.

Internal characters (Fig. 6): Dental plates slender Crural bases crescent-like, concave laterally, slightly and straight, ventrally divergent, totally disappearing projected ventrally and dorsally from the hinge plates. at the level of full teeth development. Delthyrial cavity Crura curving ventrally, with widened and thickened distal trapezoidal; lateral umbonal cavities triangularly rounded. ends (similar in appearance to a diabolo in transverse Deltidial plates inwardly curved. Hinge teeth globular and sections), calcariform, terminating at a distance of 0.26 crenulated, inserted in deep dental sockets with lateral dorsal valve length from the dorsal umbo. denticula. Outer hinge plates initially ventrally deflected, anteriorly becoming horizontal, relatively wide, and Remarks - Meneghini (1857) described and figured straight. Septalium not present. Septalial plates short, from the Jurassic of Perda Liana (the same locality of pendant. Euseptoideum reduced to a short and low ridge. the studied specimens) as Rhynchonella subobsoleta 132 Bollettino della Società Paleontologica Italiana, 52 (2), 2013

Davidson some specimens (MUP I 13899, 13892, 13882a, v 1857 Terebratula Lamarmorae nov. sp. - Meneghini, p. 271, pl. E, 13882b) which have the same external morphology as figs 19a-b [Nomen oblitum]. the specimens collected by us. He distinguished three 1894 Terebratula Lamarmorae Mgh. - Fucini, p. 152. morphotypes, which, according to him, are very close to 1899 Terebratula Lamarmorae Mgh. - Fucini, p. 121. each other. Their external morphology fits well with our + 1918 Holcothyris angulata nom. nov. - Buckman, p. 192, pl. 10, figs 1-4 [Nomen protectum]. specimens in general shape, dimensions, in tendency to 1940 Holcothyris angulata Buckman - Sahni, p. 10, text-fig. 7. asymmetry, type and number of ribs on the slopes and on 1940 Holcothyris angulata Buckman var. depressa nov. - Sahni, the fold and in the sulcus. Dainelli (1903) described one p. 11, pl. 1, figs 17-18. external mould from the Bathonian of the same locality and 1940 Holcothyris angulata Buckman var. areolata nov. - Sahni, p. one small coeval specimen from the Tacco of Seui (central 11, text-fig. 8. Sardinia). Due to their poor state of preservation, following 1960 Holcothyris angulata Buckman - Makridin, p. 298. Meneghini he identified both forms as Rhynchonella cfr. 1963 Holcothyris angulata Buckm. - Ovčarenko, p. 33. subobsoleta. Having carefully examined this material, we 1965 Holcothyris angulata Buckman - Muir-Wood, p. 780, fig. arrived to the conclusion that both specimens are to be 642,2. referred to K. oranensis. 1967 Holcothyris angulata Buckman - Ovčarenko, p. 27. 1971 Holcothyris angulata Buckman - Alméras, p. 334, pl. 58, figs The internal morphology of the sectioned specimen 7-12; pl. 59A-B. (Fig. 6) entirely coincides with that observed in Rousselle 1979 Holcothyris angulata Buckman - Jin (Ching) et al., p. 198. & Demarez (1965, serial sections of six specimens in 1981 Holcothyris angulata Buckman - Doescher, p. 98. text-plate of p. 37), Alméras & Fauré (2008, figs 1-5), and 1983 Holcothyris angulata Buckman - Ovčarenko, p. 111, pl. 10, especially with the reconstructions of crura by Rousselle figs 5-10; pl. 11, figs 1-6; pl. 12, figs 1-4; pl. 13, figs 1-4; (1965, fig. 24), which are calcariform with widened and text-fig. 26. thickened distal ends (similar in appearance to a diabolo). 1983 Holcothyris angulata Buckman - Cooper, p. 91. This supports the inclusion of this species within the genus 1986 Holcothyris angulata Buckman - Alméras & Gupta, pp. Kallirhynchia Buckman, 1918. 422-425, 427, pl. 1, figs 1-8; pl. 2, figs 1-4; text-figs 1, 3. Kallirhynchia oranensis differs externally from the 1986 Holcothyris angulata Buckman morphe depressa - Alméras & Gupta, p. 424, pl. 1, fig. 9; text-fig. 2. Upper Aalenian (Ludwigia murchisonae-Graphoceras 1987 Holcothyris angulata Buckman - Yang & Shi, p. 36, pl. 1, concavum zones) Globirhynchia subobsoleta (Davidson, figs 1-3; text-figs 4A-B. 1852, p. 91, pl. 17, fig. 14) of England and France by its 1990 Holcothyris angulata Buckman - Sun, p. 241, pl. 5, figs 5-8; larger size and by the fewer number of ribs. In addition, text-fig. 13. the two taxa can be clearly distinguished by their 1990 Holcothyris angulata Buckman - Ovčarenko, p. 50. internal morphology, especially by the development of 1991 Holcothyris angulata Buckman - Alméras et al., p. 18. two different types of crura: with widened distal ends, 1991 Holcothyris angulata Buckman - Alméras et al., p. 447, pl. calcariform (a variant of raduliform) in Kallirhynchia 21, figs 8-11; text-figs 9-11. and canaliform in Globirhynchia (Savage et al., 2002, fig. 1991 Holcothyris angulata Buckman - Georgescu, p. 241. 901, 3e-3m, reproduced from Shi & Grant, 1993, fig. 69), 1993 Holcothyris angulata Buckman - Georgescu, p. 43. 1995 Holcothyris angulata Buckman - Radulović, p. 375. which places them in two different families. Interior of the 1995 Holcothyris angulata Buckman - Ouali-Mehadji, p. 124. Sardinian specimens and especially distal ends of the crura 1996 Holcothyris angulata Buckman - Alméras et al., pp. 606, (a diabolo in transverse sections), are identical with those 616, text-fig. 10a. in the type species of the genus, Kallirhynchia yaxleyensis 1997 Holcothyris angulata Buckman - Alméras et al., p. 177. (Davidson, 1878), from the Upper Bathonian of Lorraine, 1998 Holcothyris angulata Buckman - Sun & Zhang, p. 256, pl. France (Savage et al., 2002, fig. 921, 2d-2j, reproduced 8, figs 5-8; pl. 10, figs 8-11; text-fig. 36. from Laurin, 1984, fig. 182). However, it differs externally 2006 Holcothyris angulata Buckman - Lee et al., p. 2113, fig. 1401, from oranensis in having a globose shape, larger number 7a-m. of ribs (25-31), and always symmetrical uniplication with 2008 Holcothyris angulata Buckman - Wang et al., p. 99. high rectangular linguiform extension. 2008 Holcothyris angulata Buckman - Alméras & Fauré, p. 660, text-fig. 33. 2009 Holcothyris angulata Buckman - Löwner, p. 99. Stratigraphic and geographic range - According to Alméras & Fauré (2008) K. oranensis occurs in the uppermost Bajocian (Parkinsonia parkinsoni Zone), Lower Nomenclatural note - In 1857 Meneghini established Bathonian - Zigzagiceras zigzag-Procerites (Siemiradzkia) the new species Terebratula lamarmorae from the “Great aurigerus zones - and Middle Bathonian (Cadomites Oolite” (Bathonian; Dainelli, 1903) of Perda Liana, central bremeri Zone) of Morocco, western Algeria, and Nepal. Sardinia. The name lamarmorae was given in honour Our finding in Sardinia represents therefore the first record of Albert de La Marmora, ending with an “ae”, though, of this taxon on the northern margin of the Tethys. etymologically, it should have been written with an “ai” (I.C.Z.N., Art. 31.1.2, 2000). Nevertheless, the correction of the name would be an unjustified emendation, and Order Terebratulida Waagen, 1883 the original name (although it has an incorrect genitive Family Postepthyrididae Tchorszhevsky, 1974 ending) should be retained (I.C.Z.N., Art. 31.1.3.). In 1918 Buckman, apparently unaware of the paper by Genus Holcothyris Buckman, 1918 Meneghini (1857), erected Holcothyris angulata, the type species of his genus Holcothyris Buckman, 1918, from Holcothyris angulata Buckman, 1918 the Namyau Beds (Bathonian) of Burma. H. angulata Figs 7f-i, 8-9, 10e-f has been generally accepted and is in widespread use in I. Dieni et alii - Bathonian of Mt Perda Liana and its brachiopods 133

Fig. 8 - Transverse serial sections of Holcothyris angulata Buckman, 1918, through the specimen MUP I 13880a, syntype of Terebratula lamarmorae Meneghini, 1857 (figured by its author in pl. E, fig. 19b and refigured here in Fig. 10e), Lower Bathonian of Perda Liana. Numbers indicate the distance in mm from the tip of the ventral umbo. 134 Bollettino della Società Paleontologica Italiana, 52 (2), 2013

Fig. 9 - Transverse serial sections of Holcothyris angulata Buckman, 1918, through the specimen MGP 31356 (the same as Fig. 7i), Lower Bathonian of Perda Liana. Numbers indicate the distance in mm from the tip of the ventral umbo. I. Dieni et alii - Bathonian of Mt Perda Liana and its brachiopods 135 the palaeontologic and stratigraphic literature, although Cardinal process well developed, nearly flat with it is a junior synonym of Terebratula lamarmorae distinct myophore. Dorsal umbonal cavity present. Meneghini, 1857. The senior name T. lamarmorae has Crural bases high and slender, ventrally tapering, project not been used as a valid name since 1899, when it was above hinge plates. Hinge plates narrow, sub-horizontal. quoted for the last time by Fucini (1899) (I.C.Z.N., Art. Euseptoideum short and low, bordered laterally by two 23.9.1.1.). Conversely, the name H. angulata Buckman small ridges. Crural processes high, ventrally convergent has been used in more than 25 papers by more than 10 and tapering. Transverse band relatively wide, high and authors in the period from 1918 to 2009 (see synonymy flat. Loop thin, as long as 0.42-0.45 of dorsal valve length, list) (I.C.Z.N., Art. 23.9.1.2.). Therefore, both conditions with very long terminal points. necessary for reversal of precedence under I.C.Z.N., Art. 23.9 are met, and following Art. 23.9.2 we cite the two Remarks - Referring to the external characters, such as names together and explicitly state that the younger name rounded sub-pentagonal outline, the shape of the anterior angulata Buckman is valid and must be maintained as commissure, and the finely capillate ornamentation, nomen protectum, while the senior name lamarmorae where exfoliated, the studied specimens fit very well Meneghini is invalid, being nomen oblitum. the Lower Bathonian (Zigzagiceras zigzag Zone) Holcothyris angulata Buckman, 1918, of the Tethyan Material - Six specimens from the Dieni Collection Realm. The only difference is the sporadical presence of (MGP 31352 - 31356, 31360, collected in situ) and the umbonal dorsal sulcus in the latter, which is variable two specimens from the Meneghini Collection (MUP I in character (see below). By their internal characteristics, 13880a, 13880b), all from Perda Liana, central Sardinia, the sectioned specimens from Perda Liana are very close for a total sum of eight specimens available. The exact to the specimens of angulata described by Alméras stratigraphic position of the specimens coming from the (1971), Ovčarenko (1983), Alméras & Gupta (1986), and Meneghini Collection is unknown. Alméras et al. (1991). Mainly, the aspects of the hinge plates, the size of the crural bases, the shape and the size Description - of transverse band and especially long terminal points External characters (Figs 7f-i, 10e-f): Medium size are very similar. (Tab. 2), rounded sub-pentagonal outline, lateral margins Ovčarenko (1983, p. 112), based on a very large slightly rounded, always longer than wide, moderately number of specimens (more than 3000), of H. angulata ventri-biconvex. Maximum width and thickness at mid- from many Bathonian localities of Pamir, stated that length. Beak massive, wide, erect, in close contact with “in the dorsal umbonal part a small furrow is usually the dorsal umbo totally obscuring symphytium, with large developed, which sometimes reaches the anterior end” circular mesothyrid to permesothyrid foramen. Beak whereas “in adults in the relief of the posterior part it ridges short and rounded. Lateral commissure gently may be absent”. Moreover, Alméras & Gupta (1986) and ventrally arched in the posterior two-thirds or nearly Alméras et al. (1991) figured specimens of this species of straight, and sharply curved ventrally at the anterior; Buckman from the Lower Bathonian of India and Algeria anterior commissure slightly bisulcate. A shallow sulcus devoid of sulcus in the dorsal umbo. Yang & Shi (1987, p. developed on the anterior one-third of the dorsal valve, 47), studying external and internal characters on material deepens anteriorly, bordered by rounded and parallel from the Lower Bathonian of China belonging to the folds. Ventral valve with median rounded fold originating genus Holcothyris, stated in the emended diagnosis of on anterior one-third, separated by shorter and narrower the genus that dorsal valve is “variably furrowed”. As in sulci. Fine growth lines, much more numerous and all the specimens of H. angulata from Burma figured by crowded anteriorly, and ornament of fine radiating capilli Buckman (1918) and Sahni (1940) and those from China (about two-three capilli per mm near the anterior margin) studied by Sun (1990), the sulcus of the dorsal umbo is where exfoliated. present. From the above facts it can be concluded that the dorsal umbonal sulcus is not always present. It is absent Internal characters (Figs 8-9): Two specimens, both in all newly collected specimens from Perda Liana the smaller (Fig. 8, L = 22.3 mm) from Meneghini’s and in the two specimens figured by Meneghini (1857). Collection and the larger (Fig. 9, L >27.8 mm) from The two specimens figured by Meneghini (1857, as the Dieni Collection, have been serially sectioned for Terebratula lamarmorae) have a slightly higher anterior studying the internal morphology. commissure when compared with the specimens collected

Specimen L W T W/L T/L T/W Remarks MGP 31352 16.5 14.5 8.0 0.88 0.48 0.55 Fig. 7f MGP 31353 20.8 18.8 12.2 0.90 0.59 0.65 Fig. 7g MGP 31354 21.8 17.0 >9.3 0.78 -- -- MGP 31355 26.8 22.5 14.8 0.84 0.55 0.66 Fig. 7h MGP 31356 >27.8 23.6 18.0 -- -- 0.76 sectioned, Fig. 7i MUP I 13880a 22.3 20.4 14.4 0.91 0.65 0.70 sectioned, Fig. 10e MUP I 13880b 29.2 23.5 15.7 0.80 0.54 0.67 Fig. 10f

Tab. 2 - Biometric parameters of the studied specimens of Holcothyris angulata Buckman, 1918. L = length; W = width; T = thickness. 136 Bollettino della Società Paleontologica Italiana, 52 (2), 2013

Fig. 10 - Brachiopods from the Lower Bathonian of Perda Liana described and figured by Meneghini (1857). The specimens are housed in the Museo di Storia Naturale e del Territorio of the Università di Pisa (MUP). a-d: Kallirhynchia oranensis (Flamand, 1911): a = MUP I 13899, described but not figured by Meneghini (1857, p. 276) as Rhynchonella“ sp. ind.”; b = MUP I 13892, specimen determined by Meneghini (1857, p. 273, pl. E, fig. 20c) as Rhynchonella“ subobsoleta Davids”; c = MUP I 13882a, specimen figured by Meneghini (1857, pl. E, fig. 20b) as “Rhynchonella subobsoleta Davids”; d = MUP I 13882b, specimen figured by Meneghini (1857, pl. E, fig. 20a) asRhynchonella “ subobsoleta Davids”. e-f: Holcothyris angulata Buckman, 1918 (nomen protectum): e = MUP I 13880a, syntype of Terebratula lamarmorae figured by Meneghini (1857, p. 271, pl. E, fig. 19b; nomen oblitum), used for transverse serial sections of Fig. 8; f = MUP I 13880b, syntype of Terebratula lamarmorae figured by Meneghini (1857, pl. E, fig. 19a). All the specimens, coated with ammonium chloride, are in dorsal (1), ventral (2), lateral (3) and anterior (4) views. by us at Perda Liana, the same locality from which the Sinai, eastern Africa, northern Africa, and Maghreb on Meneghini types come. The serial transverse sections of the southern margin of Tethys. one type of Meneghini Collection (MUP I 13880a; Fig. 8) and of a specimen collected by us (MGP 31356; Fig. 9) have revealed the same internal morphology. ACKNOWLEDGEMENTS Holcothyris rotundata Sahni (1940), co-occurring in We are indebted to L. Chiappini (Museo di Storia Naturale Burma with H. angulata and Kallirhynchia oranensis e del Territorio, Università di Pisa) and E. Cioppi (Museo di (Flamand, 1911), differs from the Sardinian specimens Storia Naturale, Sezione di Geologia e Paleontologia, Università of angulata by almost spherical shell, and in having di Firenze) for the kind loan of the specimens of Meneghini and a shorter biostratigraphic range (lower part of the respectively Dainelli collections in their care. Svetlana Nikolaeva Zigzagiceras zigzag Zone; Alméras, 2008). (Natural History Museum, London) and Alessandro Minelli (Dipartimento di Biologia, Università di Padova) helped with Stratigraphic and geographic range - The species is nomenclatural questions. We are grateful to M. Murino (Tortolì) widespread in the Lower Bathonian (Zigzagiceras zigzag for his generous present of fossils from Perda Liana. The reviewers F. Jadoul and A. Vörös are thanked for their constructive and very Zone, sometimes post zigzag) of the Tethyan realm: helpful comments. F.M. Dalla Vecchia is acknowledged for his China, Burma, Pamir, Turkmenia, Crimea, Northern suggestion concerning the fish teeth. N. Michelon and S. Castelli Caucasus, Romania, France, Sardinia, and ?England (Università di Padova) are thanked for their technical contribution. on the northern margin of Tethys; India, , The research was partially supported by the Ministry of Education I. Dieni et alii - Bathonian of Mt Perda Liana and its brachiopods 137 and Science of the Republic of Serbia, Project No. 176015 (grant Cooper G.A. (1983). The Terebratulacea (Brachiopods), Triassic to VR). to Recent: a study of the brachidia (loops). Smithsonian Contributions to Paleobiology, 50. ıx + 445 pp. Costamagna C. & Barca S. (2004). Stratigrafia, analisi di facies, REFERENCES paleogeografia ed inquadramento regionale della successione giurassica dell’area dei Tacchi (Sardegna orientale). Bollettino Ager D.V. (1965). Mesozoic and Cenozoic Rhynchonellacea. 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