Carnets Geol. 20 (13)

E-ISSN 1634-0744 DOI 10.2110/carnets.2020.2013

Stratigraphic and taxonomic considerations on the Late Cretaceous rudist fauna of Aksai Chin (Western Tibet, China) from the DE FILIPPI Collection

Jingeng SHA1

Simone FABBI2*

Riccardo CESTARI3

Lorenzo CONSORTI4

Abstract: The rudist fauna collected in western Tibet in the Aksai Chin area by the DE FILIPPI expedi- tion in 1914 has been reprised and redescribed. This fauna is composed of Radiolites cf. lusitanicus, Radiolites sp., Gorjanovicia cf. endrissi, ? Sauvagesia sp., Sphaerulites sp., Durania sp., and Gyropleu- ra sp. The rudist-bearing beds belong to the Tielongtan Group of the Tianshuihai terrane. The Turo- nian-? Coniacian Xiloqzung Formation (Fm.) bear the older rudists (Radiolites cf. lusitanicus, Radiolites sp., Sphaerulites sp., Durania sp.), whereas younger ages have been determined through microfossil analysis which, compared with the western Neotethys records, suggests an early-mid Campanian age. This allowed to ascribe the younger rudists of the collection (Gorjanovicia cf. endrissi, ? Sauvagesia sp., Gyropleura sp., Radiolites sp.) to the Dongloqzung Formation. Our data confirm that rudist-bea- ring facies in the Tielongtan Group extend at least up to the middle Campanian. The Aksai Chin rudist assemblage should belong to the Southwestern Asian assemblage of the Eastern Mediterranean Sub- province. Key-words: • Hippuritida; • Tethysian biostratigraphy; • Upper Cretaceous of Tibet; • paleobiogeography; • museum collection

Citation: SHA J., FABBI S., CESTARI R. & CONSORTI L. (2020).- Stratigraphic and taxonomic considera- tions on the Late Cretaceous rudist fauna of Aksai Chin (Western Tibet, China) from the DE FILIPPI Col- lection.- Carnets Geol., Madrid, vol. 20, no. 13, p. 249-272.

1 State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, 39 East Beijing Road, Nanjing 210008 (China) 2 * Corresponding author ISPRA - Geological Survey of Italy, via V. Brancati 48, 00144 Rome (Italy) [email protected] 3 Strata GeoResearch Srl Spin Off INGEO Dept. of Engineering and Geology, University D'Annunzio Chieti-Pescara Via Dei Vestini, 31 66100 Chieti (Italy) 4 University of Trieste, Dept. of Mathematics and Geosciences, via Weiss 2, 34128 Trieste (Italy)

Published online in final form (pdf) on September 21, 2020 [Editor: Robert W. SCOTT; technical editor: Bruno GRANIER]

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Résumé : Considérations stratigraphiques et taxonomiques sur la faune de rudistes du Cré- tacé supérieur d'Aksai Chin (Tibet occidental, Chine) appartenant à la Collection DE FILIPPI .- La faune de rudistes récoltée dans la région d'Aksai Chin dans l'ouest du Tibet par l'expédition DE FILIP - PI en 1914 est réexaminée et redécrite. Cette faune est composée de Radiolites cf. lusitanicus , Radioli- tes sp., Gorjanovicia cf. endrissi , ? Sauvagesia sp., Sphaerulites sp., Durania sp. et Gyropleura sp. Les couches à rudistes appartiennent au Groupe Tielongtan du terrane de Tianshuihai. La Formation Xiloq- zung du Turonien - ? Coniacien recèle les rudistes les plus anciens ( Radiolites cf. lusitanicus , Radiolites sp., Sphaerulites sp., Durania sp.), tandis que l'analyse du contenu microfossilifère par comparaison avec celui de la Néotéthys occidentale suggère un âge Campanien inférieur à moyen pour les rudistes les plus jeunes. Ceci permet d'attribuer les rudistes les plus jeunes appartenant à la collection ( Gorja- novicia cf . endrissi , ? Sauvagesia sp., Gyropleura sp. et Radiolites sp.) à la Formation de Dongloqzung. Nos données confirment que les faciès à rudistes du Groupe Tielongtan sont présents au moins jus- qu'au Campanien moyen. L'association de rudistes de l'Aksai Chin devrait être rattachée à l'association du sud-ouest asiatique au sein de la Sous-Province méditerranéenne orientale. Mots-clefs : • Hippuritida ; • biostratigraphie de la Téthys ; • Crétacé supérieur du Tibet ; • paléobiogéographie ; • collection de musée Introduction (i.e. , Loqzung Mts./Luokezongshan subregion – Chinese Academy of Sciences, 2000), the first The Aksai Chin is a remote region at the scientific data known to date are those collected western termination of the Tibetan Plateau (Chi- by the Italian DE FILIPPI Expedition made in 1913- na), extending at its westernmost point to the 1914 (DE FILIPPI , 1915; DAINELLI , 1934; reviewed Karakorum Pass (Fig. 1.a). It is essentially a wide by GAETANI , 2011). This expedition crossed the system of plateaus, partly occupied by salt lakes Western Loqzung Mts., through the Kizil Pass and (e.g. , Aksai Chin Lake), the average elevation of reached the Thaldàt Basin and the Aksai Chin La- which is greater than 4800 m. Northward, it is ke (Fig. 1.c). Later data was collected by the bordered by and separated from the Tarim Basin "Comprehensive Scientific Expedition to the Tibe- by the Kunlun Range, a mountain chain that ex- tan Plateau" (Chinese Academy of Sciences, ceeds 7000 m asl. The Upper Karakash River, 1998, 2000). During the DE FILIPPI expedition, which is one of the main tributaries of the Tarim which crossed the Himalayan Range from Kash- River, flows in the western part of the Aksai Chin, mir to Baltistan and Xinjiang and lasted for about along a deeply incised valley. In the central sec- two years, almost one month was spent in the tor of the region a high mountain ridge exceeding Aksai Chin Region (DAINELLI , 1934). The ~1000 6400 m asl, is known as Western Loqzung Moun- rock and fossil samples collected during the expe- tains/Luokezongshan (Fig. 1.b). dition are stored at the Natural History Museum Aksai Chin is almost uninhabited and very dif- of Florence University (Italy), and represent still ficult for researchers to investigate because the today one of the most complete geological archi- average altitude is more than 4800 m asl, with a ves of the area (GAETANI , 2011). frigid climate and reduced oxygen levels, which The first goal of this paper is to re-evaluate in result in a wide desert land. Furthermore, travel- modern terms the taxonomy of this remarkable ling is extremely difficult across the region with a rudist collection from the DE FILIPPI expedition in single, wide, long road (G219) (Fig. 1.c) built at th order to refine the Upper Cretaceous stratigra- the end of 20 century. phy. The second goal is to place this area into a Because of such issues, the geology of Aksai wider paleobiogeographic framework by compa- Chin was sporadically investigated through time ring it with more recently published research and, with few exceptions, most of the significant (WEN , 1998; WEN et al. , 1998; Chinese Academy expeditions in the area were made in the first half of Sciences, 2000). of 20th century (HEDIN , 1909; DE TERRA , 1932; NORIN , 1946). After such pioneering explorations Geological setting this remote area was almost forgotten by geolo- The geological setting of the study area is gists until the '80s of the 20th century, when it complex because of the superimposition of multi- was investigated by the "Comprehensive Scienti- ple orogenic phases ( e.g. , MOLNAR & TAPPONNIER , fic Expedition to the Tibetan Plateau" in the years 1975; SENGOR et al. , 1988; DECELLES et al. , 2002; 1987-1990 (Chinese Academy of Sciences, 1998, YIN & HARRISON , 2000; MATTERN & SCHNEIDER , 2000). Also it was only marginally the object of 2000). The subsequent accretion of crustal blocks studies of the huge Sino-English "geotraverse of at the southern margin of the Siberian Craton or Tibet" (CHANG et al. , 1986, 1989; Sino-UK Geolo- paleo-Asia (Fig. 2) persisted from Paleozoic to gical Expedition Team, 1990), and subsequently Cenozoic times (DEWEY et al. , 1988; WINDLEY , of the Sino-French expedition, which followed the 1988; CHANG et al. , 1989; GAETANI , 1997; ZANCHI route of the G219 Highway (MATTE et al. , 1996; & GAETANI , 2011; ANGIOLINI et al. , 2013; FAISAL et GAETANI , 2011). Regarding the area east of the al. , 2014). The northernmost of these crustal Karakorum Pass, and in particular between the blocks is represented by the western Kunlun Ran- Upper Karakash Valley and the Thaldàt Basin ge, a Paleozoic orogen rejuvenated during the

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Figure 1: (a) and (b) Location and geographical setting of the study area (basemap retrieved from Google Earth). (c) close-up of the study area, the dashed line is the route of the DE FILIPPI expedition through the W-Loqzung Mts., yellow stars indicate sampling localities. early Cenozoic collisional stage of the Himalaya- A complex system of terranes south of the Karakorum-Tibet orogenic system (TAPPONNIER & Kunlun block, drifted northward after the frag- MOLNAR, 1977; CHANG et al., 1986, 1989; YONGUN mentation of the northern margin of Gondwana & HSU, 1994; MATTE et al., 1996; Chinese Acade- (SENGOR et al., 1988). It was accreted during the my of Sciences, 2000; XIAO et al., 2005; ZHANG et late Middle-Late Permian to the Eocene (Fig. 2), al., 2019). The southern boundary of the western and includes the Songpan-Ganzi-Hoxhil terrane, Kunlun range is represented by the regional scale the Qiantang composite terrane, the Tianshuihai Altyn Tagh – Karakash strike-slip fault or Kunlun- terrane, the Lhasa terrane, and the Himalaya ter- Maqe Fault (Fig. 2 ; MATTE et al., 1996; SHA & rane (MOLNAR & TAPPONNIER, 1975; HUANG & CHEN, FÜRSICH, 1999; XIAO et al., 2005; SHA et al., 1987; DEWEY et al., 1988; WINDLEY, 1988; CHANG 2004; ZANCHI & GAETANI, 2011). et al., 1989; GAETANI et al., 1990; MATTE et al.,

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Figure 2: Tectonic framework of the study area. The yellow star indicates the study area. TST=Tianshuihai terrane, S-G-H=Songpan-Ganzi-Hoxhil terrane; Modified after YIN & HARRISON, 2000 (basemap retrieved from Google Earth).

1996; SHA & FÜRSICH, 1999; SHA, 1998; YIN & and 171 Ma ages of the plutonic rocks (mostly HARRISON, 2000; SHA et al., 2004; ZANCHI & granitoid) were determined using various radiome- 40 39 GAETANI, 2011; SHA, 2018). tric methods ( Ar/ Ar, K/Ar, U/Pb) (MATTE et al., Most of the Aksai Chin Region belongs to the 1996; MATTERN & SCHNEIDER, 2000). This magmatic Tianshuihai terrane (PAN et al., 1992; MATTE et activity ceased in the Middle Jurassic (MATTERN & al., 1996; Chinese Academy of Sciences, 2000; SCHNEIDER, 2000). WITTLINGER et al., 2004; XIAO et al., 2005) or Unconformably covering the Paleozoic-Triassic Kara-Kunlun and Uygur terranes (MATTERN et al., deformed rocks is a marine sedimentary succes- 1996; MATTERN & SCHNEIDER, 2000). The Tianshui- sion consisting of a few hundred (?) meters of hai terrane is a small crustal block interposed Jurassic black shales, sandstones and limestones between the larger Kunlun, Pamir, Songpan-Gan- (Fig. 3). This is, in turn, covered by a thick Creta- zi-Hoxhil, and Qiantang terranes (Fig. 2). It was ceous stratigraphic succession made of conglo- interpreted as an accretionary wedge produced merates, sandstones and limestones, containing by the northward subduction of the Paleotethys macrofossils such as ammonites, echinoids, bival- oceanic crust beneath the Kunlun block (MATTERN ves (including rudists), corals among others (PA- & SCHNEIDER, 2000; XIAO et al., 2005). The gene- RONA, 1928; STEFANINI, 1928; DAINELLI, 1933- ral stratigraphy of the Tianshuihai terrane (Fig. 3) 1934; WEN et al., 1998, 2000a, 2000b). The Ju- is characterized by a thick (>> 6000 m) rassic-Cretaceous sedimentary successions of the ? Carboniferous to Permian succession consisting Aksai Chin are poorly investigated, but since the of fusulinid limestones (marbles), green schists end of the last century, a general stratigraphic and brachiopod-bearing marbles that overly poor- subdivision was established by WEN et al. (1998, ly known early Paleozoic rocks (MATTE et al., 2000a, 2000b) based on paleontology and super- 1996). These are overlain by Permo-Triassic dark position. In this paper we contribute to the know- grey slates and barely metamorphic flyschoid ledge of the Cretaceous stratigraphy of the area sandstones (CHANG et al., 1989; MATTE et al., through the analysis of rudists collected in the 1996; MATTERN et al., 1996). The stratigraphic surroundings of the Kizil Pass by the Italian succession was strongly deformed during the scientists Giotto DAINELLI and Olinto MARINELLI in Triassic - ? Early Jurassic (Cimmerian) orogeny 1914. These fossils are from the Tielongtan (MATTERN & SCHNEIDER, 2000; SENGOR et al., Group (see below), which crops out, strongly de- 1988). Arc magmatism affected the Tianshuihai formed, in the Western Loqzung Mts. (DAINELLI, accretionary wedge, with batholiths intruding the 1933-1934; MATTE et al., 1996; WEN et al., sedimentary succession trending parallel to the 2000a, 2000b). orogen (MATTERN & SCHNEIDER, 2000). The 215

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Figure 3: Sketch of the general stratigraphy of the W-Loqzung Mts. (modified after MATTE et al., 1996; WEN et al., 2000a, 2000b).

The Cretaceous of Aksai Chin the range of Tielongtan Group to be early Ceno- manian to late Maastrichtian (e.g., XI et al., Stratigraphy 2019), although they do not provide any good A discontinuous stratigraphic succession crops evidence. out in the Aksai Chin, characterizing the Loqzung Palaeontology Mts./Luokezongshan stratigraphic subregion (WEN The fossil assemblages that are the object of et al., 1998, 2000a, 2000b), previously studied this study, led PARONA (1917, 1928) to subdivide by pioneers such as HEDIN (1909), PARONA (1917; the Upper Cretaceous succession in two parts: 1. 1928), STEFANINI (1917), DOUVILLÉ (1926) and A Cenomanian - ? "Senonian" limestone and mar- DAINELLI (1933-1934). The Lower Cretaceous in- ly limestone with abundant macrofossils such as cludes both continental and marine units: the ammonoids in the lowermost part, with rudist bi- Aptian-? Albian Yapaqin Group (Fig. 3) is made of valves, echinoids and miliolids (PARONA, 1917, interbedded marine (limestone) and non-marine 1928; DAINELLI, 1933-1934); 2. An ? "upper Se- facies (SUN & ZHANG, 1983; WEN et al., 2000a); nonian" limestone with bivalves (including ru- this unit likely corresponds to the possibly Albian dists), bryozoans, brachiopods, and possibly orbi- crinoid red limestone and underlying conglomera- toids (PARONA, 1928; DAINELLI, 1933-1934). This tes, described by PARONA (1928) and DAINELLI subdivision was obviously only tentative, and (1933-1934), and to the Albian limestones infer- they did not reconstruct a reliable stratigraphy, red by MATTE et al. (1996). The occurrence of but the stratigraphic succession seems roughly to Barremian limestone with Choffatella sp. and or- correspond to the Tielongtan Gp., which is com- bitolinids described by DOUVILLÉ (1916, 1926) in posed of interbedded brown calcareous sandsto- samples collected by HEDIN (1909) is considered nes, marls, red conglomerates, gray massive bio- not reliable. clastic limestones, and brown sandstones, also The Lower Cretaceous is disconformably cove- bearing bioclastic limestones in the upper part red by marls and limestones belonging to the Tie- (YANG et al., 2014). This group yields, in fact, longtan Group (Fig. 3), which marks a regional abundant marine bivalves, in particular, rudists, transgression considered it to be Cenomanian which are abundant along the eastern slopes of (SUN & ZHANG, 1983; XI et al., 2019) or Turonian Western Loqzung Mts. (WEN et al., 2000a, (WEN et al., 2000a, 2000b). Elsewhere it directly 2000b). According to WEN et al. (2000b) the Xi- overlies the Upper Jurassic (e.g., SUN & ZHANG, loqzung/Xiluokezong Formation yields a rich ru- 1983). The Tielongtan Gp. is a very thick (up to dist assemblage with Rhedensia, Vaccinites, >3300 m) stratigraphic unit. It is subdivided into Bournonia, Medeella, Biradiolites, Praeradiolites, the Xiloqzung/Xiluokezong (269-970 m thick), Li- Sauvagesia, and Durania. The Litian Formation tian (200-652 m thick) and Donglokzung/Don- yields Lapeirousia, Distefanella, Hippurites, and gluokezong (584-1719 m thick) formations. Its Gorjanovicia. The Dongloqzung/Dongluokezong age was suggested as being late Turonian to Formation yields Lapeirousinae, Praeradiolites, Campanian (even to early Maastrichtian) based Bournonia, Gorjanovicia, and Hippuritidae (Table on rudists and gastropods (WEN et al., 1998, 1). Some of these determinations should be re- 2000a, 2000b). Recently, some authors inferred viewed and are not considered here.

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Table 1: Stratigraphic chart of rudist taxa of the Aksai Chin region cited in the literature and notes.

Material and methods Unfortunately, most of the nearly 100 speci- mens of the collection are broken isolated frag- This is a study of rudist-bearing samples sto- ments or fragmental in whole rock. The list of red in the Natural History Museum of the Univer- specimens in the collection, with register num- sity of Florence (Table 2), from the Aksai Chin bers (Table 2), includes several specimens that area (Fig. 1). Sampling was carried out by uncon- are grouped under the same register number. ventional methods, and it is very difficult to refer Ten samples have been cut to check the shell each specimen to a precise locality. PARONA structure and for general microfacies analysis, re- (1917, 1928) and DAINELLI (1934) erroneously sulting in 20 thin sections. Due to limitations of said that the samples were from the "Lingzi- the preservation of fossil material and that only a Thang area", which is actually the Western Loq- few specimens were sectioned, only gross zung Mts. The actual collection area is marked by descriptions of the main features of the rudists well-developed rudist banks, with specimens col- have been perfomed. Shell shape and structure lected in the surroundings of Kizil Pass, "along description follows DECHASEAUX et al. (1969), PONS the valley descending from the pass toward NE in & VICENS (2008), PONS et al. (2012), and SKELTON the left riverside" and "beyond Kizil II camp site" (2013). (PARONA, 1917). In addition, we have tried our best to refer each specimen to a formation, fol- Abbreviations: AB=Anterior Band; PB=Poste- lowing the stratigraphic nomenclature established rior Band; L= Ligament ridge; CA= Cardinal ap- paratus; IGF = Register number. by WEN et al. (1998, 2000a, 2000b).

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Table 1: Stratigraphic chart of rudist taxa of the Aksai Chin region cited in the literature and notes.

Paleontological note Family Radiolitidae ORBIGNY, 1847

The rudist bivalves (Order HIPPURITIDA NE- Original determination (PARONA, 1928): WELL, 1965) of the DE FILIPPI fossil collection were Radiolites cf. peroni (CHOFFAT) originally determined by PARONA (1917, 1928), who recognized three (or possibly more) species Radiolites cf. lusitanicus (BAYLE, 1857) of radiolitids and one of monopleurids: Radiolites (Fig. 4.a-g) indicus (STOLICZKA) (Radiolites radiosus ORB. Material: 5 quite well-preserved right valves group); Radiolites cf. peroni (CHOFFAT); Radiolites and 7 fragments (IGF 103258, 103260, 103261, sp.; Durania f. ind. (cf. D. arnaudi (CHOFF.)), then identified as Durania mutabilis and Gyropleura 103262, 103263). cenomanensis (ORB.). Locality: beyond Kizil 2 (Fig. 1) Inferred formation: Xiloqzung Fm. Age: ? Turonian-? Coniacian

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Table 2: Aksai Chin Rudist collection stored at the Natural History Museum of Florence University (Italy). A catalo- gue number can include one or more specimens.

IGF Description N. Order Family Genus Species Locality Nation Epoch Age Year Collectors

Fragments Gyropleura Upper DAINELLI & 103252 with rudists 7 Hippuritoida Monopleuridae - Kizil 2 CHINA Turonian 1914 cenomanensis Cretaceous MARINELLI and matrix Fragments beyond Upper DAINELLI & 103253 with rudists 3 Hippuritoida Radiolitidae Radiolites Radiolites sp. CHINA Turonian 1914 Kizil 2 Cretaceous MARINELLI and matrix Fragments beyond Upper DAINELLI & 103254 with rudists 11 Hippuritoida Radiolitidae Radiolites Radiolites sp. CHINA Turonian 1914 Kizil 2 Cretaceous MARINELLI and matrix Fragments beyond Upper DAINELLI & 103255 with rudists 3 Hippuritoida Radiolitidae Radiolites Radiolites sp. CHINA Turonian 1914 Kizil 2 Cretaceous MARINELLI and matrix Fragments beyond Upper DAINELLI & 103256 with rudists 3 Hippuritoida Radiolitidae Radiolites Radiolites sp. CHINA Turonian 1914 Kizil 2 Cretaceous MARINELLI and matrix Fragments beyond Upper DAINELLI & 103257 with rudists 8 Hippuritoida Radiolitidae Radiolites Radiolites sp. CHINA Turonian 1914 Kizil 2 Cretaceous MARINELLI and matrix

Single Radiolites cf. beyond Upper DAINELLI & 103258 1 Hippuritoida Radiolitidae Radiolites CHINA Turonian 1914 specimen peroni Kizil 2 Cretaceous MARINELLI

Broken Aksai Upper DAINELLI & 103259 7 Hippuritoida Radiolitidae Radiolites Radiolites sp. CHINA Turonian 1914 specimens Chin Cretaceous MARINELLI

Radiolites cf. beyond Upper DAINELLI & 103260 Fragments 2 Hippuritoida Radiolitidae Radiolites CHINA Turonian 1914 peroni Kizil 2 Cretaceous MARINELLI

Radiolites cf. beyond Upper DAINELLI & 103261 Fragments 3 Hippuritoida Radiolitidae Radiolites CHINA Turonian 1914 peroni Kizil 2 Cretaceous MARINELLI

Radiolites cf. beyond Upper DAINELLI & 103262 Fragments 2 Hippuritoida Radiolitidae Radiolites CHINA Turonian 1914 peroni Kizil 2 Cretaceous MARINELLI

Radiolites cf. beyond Upper DAINELLI & 103263 Fragments 3 Hippuritoida Radiolitidae Radiolites CHINA Turonian 1914 peroni Kizil 2 Cretaceous MARINELLI

beyond Upper DAINELLI & 103264 Fragments 4 Hippuritoida Radiolitidae Radiolites Radiolites sp. CHINA Turonian 1914 Kizil 2 Cretaceous MARINELLI

beyond Upper DAINELLI & 103265 Fragments 7 Hippuritoida Radiolitidae Radiolites Radiolites sp. CHINA Turonian 1914 Kizil 2 Cretaceous MARINELLI

beyond Upper DAINELLI & 103266 Fragments 3 Hippuritoida Radiolitidae Radiolites Radiolites sp. CHINA Turonian 1914 Kizil 2 Cretaceous MARINELLI

Aksai Upper DAINELLI & 103267 Fragments 15 Hippuritoida Radiolitidae Radiolites Radiolites sp. CHINA Turonian 1914 Chin Cretaceous MARINELLI

Single Radiolites beyond Upper DAINELLI & 103268 1 Hippuritoida Radiolitidae Radiolites CHINA Turonian 1914 specimen indicus Kizil 2 Cretaceous MARINELLI

beyond Upper DAINELLI & 103269 Fragments 15 Hippuritoida Radiolitidae Radiolites Radiolites sp. CHINA Turonian 1914 Kizil 2 Cretaceous MARINELLI

Two Durania beyond Upper DAINELLI & 103270 2 Hippuritoida Radiolitidae - CHINA Turonian 1914 specimens mutabilis Kizil 2 Cretaceous MARINELLI Fragments Gyropleura Upper DAINELLI & 103942 with rudists 1 Hippuritoida Monopleuridae Gyropleura Kizil 2 CHINA Turonian 1914 cenomanensis Cretaceous MARINELLI and matrix Fragments Gyropleura Upper DAINELLI & 103943 with rudists 1 Hippuritoida Monopleuridae Gyropleura Kizil 2 CHINA Turonian 1914 cenomanensis Cretaceous MARINELLI and matrix Fragments beyond Upper DAINELLI & 103944 with rudists 1 Hippuritoida Radiolitidae Radiolites Radiolites sp. CHINA Turonian 1914 Kizil 2 Cretaceous MARINELLI and matrix Fragments beyond Upper DAINELLI & 103945 with rudists 1 Hippuritoida Radiolitidae Radiolites Radiolites sp. CHINA Turonian 1914 Kizil 2 Cretaceous MARINELLI and matrix Fragments beyond Upper DAINELLI & 103946 with rudists 1 Hippuritoida Radiolitidae Radiolites Radiolites sp. CHINA Turonian 1914 Kizil 2 Cretaceous MARINELLI and matrix Fragments beyond Upper DAINELLI & 103947 with rudists 1 Hippuritoida Radiolitidae Radiolites Radiolites sp. CHINA Turonian 1914 Kizil 2 Cretaceous MARINELLI and matrix

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Description : Elongate shell, subcylindrical, up Original determination (PARONA , 1928): to >90 mm high showing slightly inclined growth Radiolites sp. lamellae sets and dominant normal cellular shell structure (Fig. 4.a, g, g'). In transverse sections Several specimens, mainly fragments, in the the cardinal apparatus is well developed (Fig. collection were described by PARONA (1928) as 4.f). On the external side, well impressed growth Radiolites sp. (Table 2). Most of the material cycles are present with robust non-protruding shows no diagnostic characters, and its determi- costae (Fig. 4.a-b, d). Growth cycles are well im- nation is thus left as Radiolites sp. sensu PARONA pressed also interiorly, in the inner shell of elon- (1928). But some distinction is made between gated specimens, from which the outer shell layer two groups of specimens that show different cha- has spalled away (Fig. 4.e). The radial bands are racters. Note that register numbers can be repea- on indented folds producing smooth sinuses (Fig. ted because different taxa can be found in the sa- 4.a-b, g). The ligament ridge is well marked and me polished slab. projected towards the ventral side (Fig. 4.e-g). Radiolites sp. 1 Discussion : This fossil material was described (Fig. 5.a-e) as Radiolites cf. peroni by PARONA (1917, 1928). Radiolites peroni was considered as a distinct Material: Several poorly preserved right valves species but only differs from Radiolites lusitanicus and fragments (IGF 103253, 103254, 103255, by the number of folds in the interband, which is 103256, 103257, 103259, 103264, 103265, variable and not a distinctive character (STEUBER , 103266, 103267, 103269, 103944, 103946, 1999). For this reason and the common occurren- 103947). ce of both species in the same stratigraphic inter- Locality: Kizil 2; beyond Kizil 2. val (PEJOVIC , 1957; BERTHOU , 1973; STEUBER , Inferred formation: Xiloqzung/Litian/Dongloq- 1999) led STEUBER (1999) to consider R. peroni a zung Fm. junior synonym of R. lusitanicus. More fossil ma- terial close to the type locality in Portugal has Age: ? Turonian to ? Campanian been recently described, with a shell structure Description : Only right valves, commonly in marked by well differentiated lamellar sets inte- the form of inner shell fragments, from which the riorly inclined with regular, well developed, small outer shell layer has spalled away, have been costae, externally barely visible radial bands and studied. Shape is conical to low cylindrical, with normal cells passing locally to very elongate cells well-developed growth cycles (Fig. 5.a-b). Shell is from the inner towards the external part of the marked by growth lamellar sets with both outer shell layer (TROYA -GARCÍA , 2016). compact and normal cellular structure (Fig. 5.c- Due to the poorly preserved morphostructural d), with this last more developed on the dorsal features of the studied material, a definite attri- side. bution of this fossil material to this species is not Discussion: This material differs from other possible due to: (a) the notable presence of com- specimens by its elongate cylindrical shape and pact structure together with the normal cellular the dominant compact structure (Fig. 5.e), which structure of growth lamellae sets (Fig. 4, inset contrasts to the cellular structure of Radiolites g'), (b) the ligament ridge that is more projected sp. 2. toward the inner cavity in our material with Radiolites sp. 2 respect to the material described by TROYA -GARCÍA (2016), and (c) the unclear organization of radial (Fig. 5.f-g) bands. Despite this, we have no sufficient data to Material: Few poorly preserved right valves refute the original determination by PARONA , thus and fragments (IGF 103253, 103254, 103254B, we stand on its attribution, updated to modern 103267). taxonomy ( e.g. , STEUBER , 1999). Locality: Kizil 2; beyond Kizil 2. R. lusitanicus has a broad geographical Inferred formation: Litian/Dongloqzung Fm. distribution, being reported from both the southern and northern margins of Neotethys, and Age: ? Campanian from Portugal to the Mediterranean area, Middle- Description : Only right valves have been stu- East, Afghanistan and western Tibet ( e.g. , CHOF - died; shell form is conical to low cylindrical, FAT , 1886; DOUVILLÉ , 1910; PARONA , 1917, 1928; structure of growth lamellar sets is normal cellu- STEUBER , 1999; STEUBER & LÖSER , 2000; EL HEDENY , lar with large cells more developed in the thicker 2007). Originally described as a Turonian species, dorsal side with short ligament ridge (Fig. 5, inset its stratigraphic range is between the late Ceno- f'), whereas it is compact in the thinner antero- manian and the Coniacian (STEUBER , 1999; EL ventral side, where the radial structures are mar- HEDENY , 2007; ZAKHERA , 2011). ked by slightly developed folds (as in 103267, Fig. 5.f-g).

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Figure 4: Radiolites aff. lusitanicus (ex cf. peroni in PARONA, 1928). (a) IGF103260 (figured as R. cf. peroni by PARO- NA, 1928, Pl. 16, fig. 2), two joined specimens in lateral view. (b, c) IGF 103261, lateral dorsal view and longitudinal section (showing slightly inclined growth lamellae sets) of right valve, respectively. (d) IGF103260, lateral dorsal view of right valve. (e) IGF 103263, dorsal view of right valve, only a thin layer of inner shell preserved, with well impressed growth cycles and invaginated ligament ridge. (f) IGF 103261, transverse section of right valve. (g, g') IGF 103258, transverse section of two right valves with detail of the outer shell layer with normal cellular (left) and compact (right) structure in inset f'. Scale bars = 1 cm. Discussion: This material differs from the Description: Two transverse sections of right other group by its cylindro-conical shape, the do- valves are characterized by a thick compact outer minant normal cellular structure (Fig. 5.f-g) and shell layer; radial bands are marked by slightly radial band organization. developed folds with a slight external central in- Besides these two groups, other material dentation of the AB, which is separated from the originally described as Radiolites sp. has PB by a narrow interband; a thin triangular liga- mentary infolding is also present (Fig. 6.a). been tentatively assigned to different taxa. Discussion: The radial bands organization, Gorjanovicia cf. endrissi (BOEHM, 1927) the thick compact outer shell and the ligamentary (Fig. 6.a) infolding suggest a similarity of our specimens to Gorjanovicia endrissi as described by FENERCI- Material: IGF 103945, 103947 MASSE et al. (2011). The lack of any well-preser- Locality: beyond Kizil 2 ved material only allows an uncertain attribution Inferred formation: Dongloqzung Fm. to Gorjanovicia cf. endrissi. Age: Campanian

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Figure 5: Radiolites sp. (a, b) IGF103265, ventral and dorsal views, respectively of elongate specimens, only a very thin layer of the inner shell is preserved, with well impressed growth cycles and invaginated ligament ridge. (c, d) IGF103267 and IGF 103269 respectively, thin sections of the outer shell showing growth lamellar sets with both compact and normal cellular structures. (e) IGF103267, transverse section of the right valve showing dominant com- pact structure. (f, g) IGF103267, transverse sections of right valves showing normal cellular with well-developed cells in the dorsal side and short ligament ridge (inset f'). Scale bars = 1 cm.

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Figure 6: (a) IGF103267, Transverse section of the right valve of Gorjanovicia cf. endrissi. (b) IGF 103947, trans- verse section of the right valve, juvenile stage, of ? Sauvagesia sp. Scale bars = 1 cm.

? Sauvagesia sp. Description: This single specimen of the col- lection consists of a large right valve (up to 200 (Fig. 6.b) mm in diameter) partly deformed and broken on Material: IGF 103947 the ventral side (Fig. 7.a). The shell is cylindro- Locality: beyond Kizil 2 conical, at least 35 cm high, the right (lower) val- ve is very thick in the antero-dorsal side (up to Inferred formation: Dongloqzung Fm. 80 mm) with a wide radial, down-and-outward Age: Campanian fold separating two badly preserved radial struc- Description: Another taxon previously as- tures (Fig. 7.a). Shell is composed of growth la- signed to Radiolites sp. is a single specimen cut mellar sets with normal cellular structure and in a polished section from a rock slab (IGF cells commonly are elongate in the outer part, 103947). The transverse section of the lower where lamellar sets are downfolded (Fig. 7.a, c-d) (right) valve, juvenile stage, shows a sub-roun- giving rise to a foliaceous aspect. Growth cones ded external perimeter and thick outer shell layer cannot be observed in the outer surface as it is composed of growth lamellar sets with normal eroded and badly preserved. Ligament ridge is cellular structure. Ligament ridge is present and robust at the base (Fig. 7.b), presumably with a delimiting an undeveloped cardinal apparatus, at long pedicle. Left valve is absent. the edge of the dorsal side (Fig. 6.b). Discussion: The specimen was assigned to Discussion: Although determined by a well- Radiolites indicus by PARONA (1917), but the few preserved single specimen, the polished slab re- preserved features suggest a different assign- veals a shape and structure that could be possi- ment to the genus Sphaerulites. The shell struc- bly assigned to Sauvagesia sp. by the characte- ture is marked by growth lamellar sets with nor- ristic small cells of the cellular shell structure. mal cellular structure composed of elongate cells. The radial bands, although not clearly visible, are Original determination (PARONA, 1917): separated by a typical main fold in between Radiolites indicus (STOLICZKA) them. The cellular structure differs slightly from the radial, essentially discontinuous ridges of the Sphaerulites sp. cellular structure typical of Sphaerulites (e.g., (Fig. 7.a-d) TROYA-GARCÍA, 2016). The poorly preserved fea- Material: 1 right valve (IGF 103268) tures and the scarcity of material do not allow a specific assignment, but we consider invalid the Locality: beyond Kizil 2 generic emendation by PARONA, due to the analy- Inferred formation: Xiloqzung Fm. zed characters, which led us to assign this speci- Age: ? Turonian-? Coniacian (for its occurren- men to Sphaerulites, rather than to Radiolites. ce in association with R. cf. lusitanicus).

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Figure 7: Sphaerulites sp. (a) IGF103268, conical shaped specimen seen from above showing thick ligament ridge; in the ventral side the presumably position of the radial bands is separated by an upward fold. (b) IGF103268, inter- nal view of growth cycles and robust ligament ridge. (c) IGF103268, growth lamellar sets with normal cellular struc- ture characterized by elongated cells. (d) IGF103268, side view of sub-horizontal and downfolded lamellae sets pro- jecting laterally in grain-supported sediment. Sphaerulites indicus is a species not commonly rial, S. indicus would be possibly an invalid taxon. reported in the scientific literature. It was esta- In addition, the hypothetical geographical distri- blished by STOLIZCKA (1871) from specimens in bution of the species should oddly be limited only Upper Cretaceous brownish marly limestone at to southeastern India and, eventually, the Aksai Odium (NW from Ottakkovil) south-east India. Chin area. These issues led us to consider, that if DOUVILLÉ (1910) suggested that S. indicus was si- the species is valid, its actual occurrence in the milar to R. peroni, but he did not emend the Himalayan area is very dubious, and thus we as- taxon, and he noted that the figures by STOLIZCKA sign our specimen to Sphaerulites sp. and the quality of the material was poor. S. in- Original determination (PARONA, 1928): dicus was then described as Radiolites indicus by Durania mutabilis (STOL.) PARONA (1917) and listed in KÜHN (1932) and SAN- CHEZ (1981). Our unique specimen and the sectio- Durania sp. ning limitations do not allow a better taxonomic (Fig. 8.a-b) redescription that could result in a definite speci- Material: 1 almost complete right valve, 1 bro- fic determination. Only few species are currently ken and weathered right valve, 1 fragment of assigned to genus Sphaerulites, and S. indicus is right valve (IGF 103256, 103270). not included (as in TROYA-GARCÍA, 2016). Due to the scarce reports and poor quality of the mate- Locality: beyond Kizil 2

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Figure 8: (a, b) Durania sp., IGF103270, (a) transverse section of right valve showing well developed folds and ra- dial bands ; (b) ventral view of a right valve of Durania sp. showing ribs corresponding to the radial bands (figured by PARONA , 1917, p. 135). IGF103270. (c-g) Gyropleura sp., (c) IGF 103943, ventral view of a single spirogyrate val- ve; (d) IGF 103942, fragments of made of compact calcite structure, embedded in grain-supported limestone; (e) IGF 103252, bouquet of several spirogyrate valves of Gyropleura sp.; (f) IGF103942, thin section of a shell frag- ment: outer shell layer (OL) is made of compact calcite lamellar sets, a very thin middle shell layer (ML) separate it from the inner shell layer (IL) made of white neomorphic calcite substituting the original aragonite. Scale bars 1 cm. (g.) IGF103942, same shell fragment shown in (f) with crossed polars, showing calcite crystals oriented perpendicu- lar to the growth lines, typical of rudists. Inferred formation: Xiloqzung Description . Although the material is not well Age: ? Turonian-? Coniacian (for its occurren- preserved, the few specimens have an elongate ce in association with R. cf. lusitanicus ) cylindrical shape (Fig. 8.b) and well-developed

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growth laminar sets with normal cellular structure Age and stratigraphic considerations (Fig. 8.a). The radial bands have growth laminar sets that are concave in radial section and they Rudists are marked externally by a dozen thin costellae Comparing the original sedimentological de- (Fig. 8.b) separated by a protruding main inter- scription of the succession, which yielded the stu- band fold. Well-developed folds form external died material (as said, collected more than one costae; no ligament ridge has been detected. century ago in an inaccessible area, DAINELLI , Discussion . The studied material was first 1933-1934), with the recently assessed stratigra- described as Durania f. ind. (cf. D. arnaudi phic setting of the area (WEN et al. , 2000a, (CHOFF .)) by PARONA (1917) and then as Durania 2000b), we infer that the rudist beds represented mutabilis (STOL .) by PARONA (1928), who erro- in the collection belong to the Tielongtan Group. neously placed Radiolites mutabilis STOLICZKA in The lower part of the Tielongtan Gp. (Xiloq- "Durania mutabilis ". However later R. mutabilis zung Formation) is Turonian-Coniacian p.p. ac- was reassigned to Bournonia mutabilis by DOU - cording to WEN et al. (2000a, 2000b). The older VILLÉ (1910), which is reported in the Maastrich- rudists determined at generic rank by WEN et al. tian Ottakkovil Fm. of southern India (MOHANTI & (1998, 2000a, 2000b) in the Xiloqzung Fm. were PONS , 1999; DIHMAN et al. , 2019). Except for the considered as Turonian, by the occurrence of papers by PARONA , " D. mutabilis " was no longer Vaccinites , Bournonia and other taxa (Table 1), cited in the palaeontological literature (with the which ruled out the hypothesis of a Cenomanian only exception of a paper by MORRIS & SKELTON age. (1995), who considered D. mutabilis as a senior Cenomanian and Turonian beds bearing Eora- synonym of D. gaensis ), and is considered the diolites hedini , Sauvagesia garianica and Hippuri- result of an erroneous taxonomic attribution by tes aff. requieni were noted by DOUVILLÉ (1916) PARONA . The described specimens can be, therefo- and KLINGHARDT (1934), who studied samples col- re, determined only at the generic rank, as Dura- lected by S. HEDIN and H. DE TERRA (Table 1), nia sp. respectively. Nevertheless, the stratigraphic ran- Family Monopleuridae ge of E. hedini and Sauvagesia garianica being (MUNIER -CHALMAS , 1873) not well constrained, and H. requeni definitely a Turonian taxon, these occurrences have to be Gyropleura sp. (Fig. 8.c-g) considered as most likely Turonian . Material: 4 rock samples with several speci- As we have already discussed, age determina- mens embedded in limestone matrix, some thin tions by PARONA (1917, 1928), who ascribed the section has been performed (IGF 103252, studied rudist fauna to the upper Cenomanian- 103942, 103943) Turonian, are not reliable. The age of the older Locality: Kizil 2 rudist assemblage of our material ( R. cf. lusitani- cus , Durania sp., Sphaerulites sp., Radiolites sp. Inferred formation: Dongloqzung Fm. 1) is not well-constrained as well, but the occur- Age: Campanian rence of Radiolites cf. lusitanicus , which is a Ce- Description : Small (< 20 mm) spirogyrate nomanian-Coniacian taxon (STEUBER , 1999), is valves (Fig. 8.c, e), difficult to study in terms of significant, whereas other taxa only determined shell structures and morphostructural analysis. A at generic rank do not provide any additional in- thin section performed on the whole rock has formation, being Durania and Radiolites Albian- shown shell fragments made of plano-parallel Maastrichtian genera, and Sphaerulites a Ceno- growth laminar sets with compact calcite in the manian-Campanian genus (STEUBER et al. , 2016). outer shell layer and the inner layer made of neo- The Turonian-Coniacian p.p. age of the Xiloq- morphic calcite, in some case showing the middle zung Fm. is fully consistent with the stratigraphic layer, aragonitic in origin (Fig. 8.f-g). distribution of Radiolites lusitanicus and is thus Discussion : Gyropleura is a monopleurid ru- inferred for the rudist-bearing beds of the succes- dist represented by few samples in the studied sion characterized by its occurrence, which collection. The specimens described as Gyropleu- should belong to the Xiloqzung Fm. ra cenomanensis by PARONA (1917) are difficult to Ages younger than the Coniacian were infer- determine at specific rank, being small and em- red in the literature and confirmed by analyzing bedded in the rock (Fig. 8.d-e) and not showing the studied material. The rudist assemblages of any distinctive character. For this reason, nothing the Litian Formation, composed of Lapeirousia , can be added in a paleontological point of view. It Hippurites , etc. determined by WEN et al. (1998, is worth noting that when morphostructural ana- 2000a), were considered as being approximately lyses can be performed only with difficulty, speci- Santonian but the lack of specific determinations fic attribution is easily erroneous. Similar spirogy- does not allow a greater precision of stratigraphic rate rudists occur beginning in the Aptian and up attribution. Our material provided no additional to the end of the Cretaceous (STEUBER et al. , data regarding this unit. 2016). The studied specimens are associated with a Campanian microfossil assemblage and are considered as Gyropleura sp.

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Figure 9: Chronostratigraphically important benthic Foraminifera and stromatoporoids recovered from the matrix of the rudist specimens. (a) IGF 103944, Sarmentofascis zamparelliae SCHLAGINTWEIT et al.. Scale bar 2 mm. (b) IGF 103269, Nezzazatinella sp. Scale bar 0.5 mm. (c) IGF 103264, Navarella sp. CIRY & RAT. Scale bar 0.5 mm. (d, f) IGF 103942 and 103946, Calveziconus lecalvezae CAUS, basal and longitudinal sections respectively. Scale bar 0.5 mm. (e, g) IGF 103264 and 103946, Orbitolinidae gen. ind. Scale bar 0.5 mm. (h, i) IGF 103269, Murgeina apula (LUPERTO SINNI). Scale bar 0.1 mm. (j, k) IGF 103942, Cibicidoides sp. Scale bar 0.1 mm. (l) IGF 103942, Dicyclina schlumbergeri MUNIER-CHALMAS. Scale bar 0.5 mm. The upper part of the Tielongtan Gp. is repre- Some of the studied rudists have been refer- sented by the the Dongloqzung Fm.; the occur- red to this unit (Gorjanovicia cf. endrissi, ? Sauva- rence of Lapeirousinae, which did not appear until gesia sp., Radiolites sp., Gyropleura sp.) with the Santonian, and Praeradiolites, Bournonia, etc. only exception of G. cf. endrissi, which is essential- (Table 1), which appeared slightly earlier but ly a Campanian taxon (FENERCI-MASSE et al., 2011), extended into Maastrichtian, led WEN et al. (1998, other taxa do not provide any additional infor- 2000a, 2000b) to infer a mainly Campanian or mation, but their age has been constrained based possibly Maastrichtian age. on micropaleontological analysis (see below).

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Benthic Foraminifera particular, the identification of Calveziconus lecal- Along with the study of rudist assemblages, vezae is supported by its subepidermal network micropaleontological analyses performed through formed by two orders of beams and rafters, and thin sections on the rock matrix embedding the by radial undulated septula. Orbitolinidae gen. rudist specimens revealed the occurrence of so- ind. (Figs. 9.e, g, 10.b) lacks the second order of me stratigraphically relevant foraminifers, which beams. Their occurrence in such levels is justified provide important information on the temporal by LUPERTO SINNI & RICCHETTI (1978) as well, who position of the samples. found some strictly comparable forms in the Up- per Cretaceous carbonates of southern Italy, and The carbonate matrix embedding Radiolites in co-assemblage with Calveziconus . Their identi- sp. is represented by a bioclastic packstone- fication should be accompanied by a close taxo- wackestone and subordinate floatstone (Figs. 9.a, nomical analysis, likely resulting in the descrip- 10.k), with Dicyclina schlumbergeri MUNIER -CHAL - tion of a new taxon, which falls beyond the scope MAS (Fig. 9.l), possible Murgeina apula (LUPERTO of the present contribution. This foraminiferal as- SINNI ) (Fig. 9.h-i), possible Navarella CIRY & RAT semblage, if correlated with the European re- (Fig. 9.c), a possible Nezzazatinella sp. (Fig. 9.b), cords, suggests an age ranging between the and a probable specimen structurally close to a lower and the middle Campanian. This age of the macrospheric ? Keramosphaerina STACHE (Fig. European assemblage is firmly constrained by the 10.i). We have observed three features suppor- Strontium Isotope Stratigraphy studies made ting the identification of Navarella (Fig. 9.c): 1) through the levels bearing Calveziconus lecalve- the microgranular shell texture; 2) the accumula- zae (STEUBER et al. , 2005; FRIJIA et al. , 2015; VIL - tion of coarse carbonate grains over the outer LALONGA et al. , 2019) and Sarmentofascis zampa- shell surface, and 3) the streptospiral arrange- relliae (SCHLAGINTWEIT et al. , 2016). Cibicidoides ment typical of Navarella . The geometry of cham- sp. is commonly recorded in the Maastrichtian bers over the central area of the cross section (CONSORTI & RASHIDI , 2019) and the lower Paleo- suggests a degree of involution rather than a per- cene (see Gavelinellidae indet. in SERRA -KIEL et fectly symmetrical disposition of lumens that, in a al. , 2020), but its stratigraphic range would planispiral geometry, should appear on both si- widen into older or younger levels, thus limiting des. Regarding ? Keramosphaerina sp. (Fig. 10.i), its biostratigraphic potential. Besides giving criti- another possible identification may be a micro- cal and more detailed stratigraphic information spheric Hellenalveolina (HOTTINGER et al. , 1989). on the Tibetan rudists, this assemblage allows us That species, however, has more distinctly strep- to definitely consider that the rudist-bearing fa- tospiral chambers than our specimen. Furthermo- cies of the Tielongtan Gp. extends up to, at least, re, the proloculus size is substantially smaller the middle Campanian. than that of a megalospheric Hellenalveolina . Mo- reover, Hellenalveolina seems geographically Comparison of the Aksai Chin fauna restricted into the Upper Cretaceous of Europe. with other Neotethys faunas Having a look at the initial chambers of Keramo- sphaerina in DEVOTO (1964) we have found a clo- Rudist faunas from the Himalaya-Karakorum- se match. Anyway, due to the scarcity of material Tibet range and the Tarim basin are known from at our disposal we prefer keeping this identifica- the late Aptian and are well-developed into the tion under open nomenclature. The identification Maastrichtian (LAN & WEI , 1995; WEN et al. , 1998; of Murgeina apula (see CHIOCCHINI et al. , 2012) is SCOTT et al. , 2010; RAO et al. , 2015, 2017, 2020; supported by the thickly calcified umbos on both SHA & CESTARI , 2016; RAO , 2019). sides of the test. Nezzazatinella sp. can be linked SCOTT et al. (2010) reported 36 genera and 44 to the Santonian taxon figured by CHIOCCHINI et species occurring in the region in the Albian- al. (2012) as N. picardi , but a more specific taxo- Maastrichtian interval. Most of the taxa were nomic designation is hampered by the lack of considered endemic and only 10 species were diagnostic sections. considered as being cosmopolitan. This western Tibetan foraminiferal assemblage In particular, the uppermost Lower Cretaceous is also known in the western Neotethys where it Langshan Formation and the Upper Cretaceous ranges from upper Santonian to at least middle Tielongtan Group of Lhasa, Qiantang and Tian- Campanian (RADOIČIĆ et al. , 2010; FRIJIA et al. , shuiai terranes and the Tarim basin, yielded rich 2015; CONSORTI et al. , 2017); this age is attribu- rudist faunas (WEN , 1998; WEN et al. , 1998; ted to the Litian and Donglozung formations. SCOTT et al. , 2010). The genera determined from The packstone matrix embedding Gyropleura the Langshan Fm. include Monopleura , Auroradio- sp. contains, along with other abundant undeter- lites , Eoradiolites , Requienia , Magallanesia , Sel- mined microfossils, the stromatoporoid Sarmen- laea , and Toucasia (SCOTT et al. , 2010; RAO et al. , tofascis zamparelliae SCHLAGINTWEIT , FRIJIA & PA- 2015, 2017). Differently, the Upper Cretaceous RENTE (Figs. 9.a, 10.k), Calveziconus lecalvezae Tielongtan Group yielded a fauna dominated by CAUS (Figs. 9.d, f, 10.a, d), a small and high-coni- warm, shallow-water Tethysian rudists, such as cal agglutinated foraminifer closely resembling Hippurites , Vaccinites , Praeradiolites , Biradiolites , the Orbitolinidae gen. ind. in LUPERTO SINNI & RIC - Durania , Sauvagesia , Distefanella , and Lapeirou- CHETTI (1978: Pl. 43, figs. 3, 7), Cuneolina sp., sia , among others. They are more abundant and (Fig. 10.e) and Cibicidoides sp. (Fig. 9.j-k). In diverse than rudists of the Himalaya, Shiqunhe-

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Figure 10: Benthic Foraminifera and facies of the matrix of rudist specimens. (a, d) IGF 103269; 103943, Calveziconus lecalvezae CAUS. Scale bar 0.5 mm. (b) IGF 103269, Orbitolinidae gen. ind. Scale bar 0.5 mm. (c) IGF 103264, Thick-shelled undetermined miliolid. Scale bar 0.5 mm. (e) IGF 103269, Cuneolina sp. Scale bar 0.5 mm. (f, g) IGF 103942; 103946, Nezzazatinella sp. Scale bar 0.5 mm. (h) IGF 103942, Nezzazatinella sp. scale bar 0.25 mm. (i) IGF103254, megalospheric ? Keramosphaerina sp. Scale bar 1 mm. (j) IGF 103254, Packstone facies rich in miliolid foraminifers. Scale bar 2 mm. (k) IGF 103944, floatstone with Sarmetofascis zamparelliae. Scale bar 2 mm. Xainza (Southeastern Tethys) and southwestern bed by LAN & WEI (1995) such as the Campanian margin of the Tarim Basin (Northeastern Tethys) age of Gyropleura beds. (WEN, 1999; SCOTT et al., 2010). In the Kunlun The taxa described in this paper should be terrane and Tarim Basin, Late Cretaceous rudists considered cosmopolitan; in particular Radiolites are Biradiolites, Gyropleura, Lapeirousella, Oscu- lusitanicus and Gorjanovicia endrissi are known ligera, and Sauvagesia (LAN & WEI, 1995; WEN, from most of the Tethysian realm (Fig. 11). 1999; RAO, 2019). Radiolites indicus (sensu PARONA), as said, is The rudist fauna of the Aksai Chin in the Tie- considered an invalid taxon and is useless for pa- longtan Group is not rich and has low variability leogeographic considerations. Gyropleura sp., compared with published data. In addition, it is Praeradiolites sp., Sphaerulites sp., Sauvagesia similar in terms of age with the Western Tarim sp., Durania sp., and Radiolites sp. provide no fauna (Yigeziya and Wuyitake formations) descri- further information on their distribution pattern,

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Figure 11: Geographical distribution of the rudist species from the Western Loqzung Mountains discussed in this pa- per. not being determined at specific rank. Along with Neotethys including the central and southern the described rudists, also the Turonian Sauvage- Apennines (Italy: FRIJIA et al., 2015; SCHLAGINT- sia garianica and Hippurites aff. requieni descri- WEIT et al., 2016; CONSORTI et al., 2017), Dinari- bed by KLINGHARDT (1934) are considered as cos- des (Croatia: STEUBER et al., 2005), Apulian plat- mopolitan taxa, the first having been found in the form (Italy: LUPERTO SINNI & RICCHETTI, 1978) and Aksai Chin, Lybia and Tunisia (PARONA, 1921; BUS- southeastern Pyrenees (northeast Spain: VILLA- SON, 1967), and the second being a common LONGA et al., 2019, among others). It is thus affir- taxon of both the northern and southern margins med that during the Campanian a biogeographic of Neotethys (e.g., CESTARI & SARTORIO, 1995; connection, represented by foraminifers and De- STEUBER & LÖSER, 2000). mospongiae, was established between the Tibe- In contrast, the Albian - Cenomanian rudists tan and Mediterranean areas. The confidence in Auroradiolites gilgitensis (= Eoradiolites gilgiten- the age assignment is provided by a direct corre- sis) and Eoradiolites hedini (= Praeradiolites lation with the European fauna, which stratigra- hedini) described by MATTE et al. (1996) and DOU- phic age is calibrated by means of Strontium Iso- VILLÉ (1916) are considered as endemic taxa (RAO tope (e.g., STEUBER et al., 2005; FRIJIA et al., et al., 2015), being only known from the north- 2015; VILLALONGA et al., 2019). However, we can- eastern margin of the Neotethys (Fig. 11). not exclude the presence of a biogeographic ef- fect, likely due to faunal dispersal, which may No further paleobiogeographic consideration slightly affect such age-correlation. Like other can be done regarding the rudist assemblages shallow-water taxa, large Upper Cretaceous ag- described by WEN et al. (1998, 2000a, 2000b), glutinating and porcelaneous foraminifera were being only generic determinations of cosmopoli- possibly subject to provincialism and periodic po- tan genera, except for Rhedensia, which is not a pulations diversification (e.g., HOTTINGER et al., valid genus according to STEUBER et al. (2016) 1989; GOLDBECK & LANGER, 2009). This is the rea- and Distefanella, which is a radiolitid with an son why Navarella from Tibet would not exactly extremely elongate shell whose occurrence is li- match with the European forms. Even in Europe, mited to the central Mediterranean Tethys the specific diversity (or the occurrence of diffe- (CESTARI et al., 2018). rent varieties) may be higher than the expected, The Aksai Chin rudist assemblage should be- as MAYNC (1954) demonstrates. In the Tibetan long to the Southwestern Asian assemblage of samples, in fact, there are certain morphotypes the Eastern Mediterranean subprovince, as defi- such as Nezzazatinella sp. and ? Keramosphaeri- ned by KAUFFMAN (1973) and SCOTT et al. (2010). na sp., or the Orbitolinidae gen. ind., that cannot Concerning benthic foraminifers and Demo- be assigned to any species so far known. These spongiae, the specimens identified in the studied are likely new taxa, but their detailed systematic microfacies show strong affinities with that known placing is hampered by the lack of additional ma- in the shallow-water deposits of central-western terial.

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Conclusions Bibliographic references The rudist fauna of the DE FILIPPI collection ANGIOLINI L., ZANCHI A., ZANCHETTA S., NICORA A. & from western Tibet has been reprised and rede- VEZZOLI G. (2013).- The Cimmerian geopuzzle: scribed. The previous taxonomic attribution of New data from South Pamir.- Terra Nova , vol. this fauna has been mostly considered inaccura- 25, no. 5, p. 352-360. te, with the only exception of R. cf. lusitanicus , BAYLE E. (1857).- Nouvelles observations sur which has been renamed after the recent emen- quelques espèces de rudistes.- Bulletin de la dation of the taxon R. peroni . Regarding the Société Géologique de France , vol. 2, no. 14, other specimens no definitely specific attributions p. 647-719. have been possible due to the poor quality of the BERTHOU P.-Y. (1973).- Le Cénomanien de l'Estré- material. 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