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1 Zootaxa 0000 (0): Copyright 2014 Magnolia Press Article ISSN (print edition) ZOOTAXA ISSN (online edition) Volutidae (Mollusca: Gastropoda) of the Lakhra Formation (Earliest Eocene, Sindh, Pakistan): systematics, biostratigraphy and paleobiogeography DIDIER MERLE 1 5, JEAN-MICHEL PACAUD 1, GRÉGOIRE MÉTAIS 1, ANNACHIARA BARTOLINI 1, RAFIQ A. LASHARI 2, IMDAD A. BROHI 2, SARFRAZ H. SOLANGI 2, LAURENT MARIVAUX 3 & JEAN-LOUP WELCOMME 4 1 Sorbonne Universités - CR2P - MNHN, CNRS, UPMC-Paris6. Muséum national d Histoire naturelle, Département Histoire de la Terre. Case postale 38, 8 rue Buffon, F Paris cedex 05 (France) 2 Centre for Pure and Applied Geology, University of Sindh, Allama I.I. Kazi Campus, Jamshoro 76080, Pakistan 3 57, chemin de la Mort-Aux-Ânes, Villeneuve-lès-Maguelone, France 4 Laboratoire de Paléontologie, Institut des Sciences de l Évolution de Montpellier (ISE-M, UMR-CNRS 5554), c.c. 064, Université Montpellier 2, place Eugène Bataillon, F Montpellier Cedex 05, France 5 Corresponding author. dmerle@mnhn.fr Table of contents Abstract Introduction Material and methods Systematics (D. Merle & J.-M. Pacaud) Family Volutidae Rafinesque, Subfamily Volutinae Rafinesque, Tribe Lyriini Pilsbry & Olsson, Genus Lyria Gray, Lyria sp Genus Mitreola Swainson, Mitreola brohii sp. nov Lyriini incertae sedis Genus Lyriopsis gen. nov Lyriopsis cossmanni (Vredenburg, 1923) comb. nov Volutinae incertae sedis Genus Lyrischapa Aldrich, Lyrischapa haimei (d Archiac & Haime, 1853) Lyrischapa sismondai (d Archiac & Haime, 1853) Lyrischapa vredenburgi sp. nov Lyrischapa blanfordi (Vredenburg, 1923) comb. nov Lyrischapa brevispira sp. nov Subfamily Athletinae Pilsbry & Olsson, Genus Athleta Conrad, Subgenus Volutospina Newton, Athleta (Volutospina) noetlingi (Cossmann & Pissarro, 1909) Subgenus Volutopupa Dall, Athleta (Volutopupa) intercrenatus (Cossmann & Pissarro, 1909) comb. nov Athleta (Volutopupa) citharopsis sp. nov Subgenus Volutocorbis Dall, Athleta (Volutocorbis) eugeniae Vredenburg, Athleta (Volutocorbis) burtoni Vredenburg, Athleta (Volutocorbis) cf. wynnei Cox, Athleta (Volutocorbis) lasharii sp. nov Subfamily Volutilithinae Pilsbry & Olsson, Genus Volutilithes Swainson, Volutilithes welcommei sp. nov Accepted by M. demaintenon: 23 May 2014; published:?? Month

2 Volutilithes sindhiensis sp. nov Genus Pseudaulicina Chavan in Furon & Kouriatchy, Pseudaulicina coxi sp. nov Genus Sindhiluta gen. nov Sindhiluta lakhraensis sp. nov Incertae sedis Subfamily Volutodermatinae Pilsbry and Olsson, Genus Pakiluta gen. nov Pakiluta solangii sp. nov The Early Cenozoic diversification of volutids in Pakistan Conclusion Acknowledgments References Abstract The paleobiodiversity of the Volutidae (Mollusca: Gastropoda) of the Ranikot Group (Sindh, Pakistan) and particularly of the Lakhra Formation (SBZ 5 biozone, Earliest Eocene), is reconsidered on the basis of new material collected during recent field trips. Ten new species are described (Mitreola brohii sp. nov., Lyrischapa vredenburgi sp. nov., L. brevispira sp. nov., Athleta (Volutopupa) citharopsis sp. nov., A. (Volutocorbis) lasharii sp. nov., Volutilithes welcommei sp. nov., V. sindhiensis sp. nov., Pseudaulicina coxi sp. nov., Sindhiluta lakhraensis sp. nov. and Pakiluta solangii sp. nov.) and one species is in open nomenclature (Lyria sp.). Three new genera are described: Lyriopsis gen. nov. [Volutinae,?Lyriini, type species: Lyriopsis cossmanni (Vredenburg, 1923)], Sindhiluta gen. nov. [Volutilithinae, type species: Sindhiluta lakhraensis n. sp.] and Pakiluta gen. nov. [?Volutodermatinae, type species: Pakiluta solangii n. sp.]. Two new combinations are proposed: Lyriopsis cossmanni (Vredenburg, 1923) comb. nov. and Athleta (Volutopupa) intercrenatus (Cossmann & Pissarro, 1909) comb. nov. Lectotypes are designated for Lyria cossmanni Vredenburg, 1923, L. feddeni Vredenburg, 1923, Volutospina noetlingi Cossmann & Pissarro, 1909, V. intercrenata Cossmann & Pissarro, 1909 and Athleta (Volutocorbis) victoriae Vredenburg, With 21 species, this volutid fauna is the most diverse recorded from the Tethys Ocean during Earliest Eocene time. The assemblage is characterized by a strong turnover marked by regional speciation and the appearance of many western Tethyan invaders. Although at the species level, the assemblage documents a strong provincialism, at the genus level, the high number of shared genera between Eastern Tethyan and Old World Tethyan realms begins a phase of long-term homogeneity of volutid assemblages from the Tethyan paleobiogeographic province. Key words: Neogastropoda, Muricoidea, Early Paleogene, Eastern Tethys, systematics, new taxa Introduction The Indus Basin (Pakistan) has been known since the 19 th century for its rich invertebrate faunas from the Early Paleogene of the Eastern Tethys. They allow documentation of the biotic impacts of two global events, the Cretaceous/Paleogene crisis and the Paleocene-Eocene Thermal Optimum (= PETM) in the context of the India- Asia collision (Jablonski 1998, 2008). The molluscan faunas have been the subject of several monographs: d Archiac & Haime (1853), Cossmann & Pissarro (1909), Vredenburg (1923, 1924, 1928) and Douvillé (1929) for the Ranikot Group in Sindh, Eames (1951a, 1952) for the Sulaiman Range and Cox (1930) from the Samana Range, but little research has been carried out since the 1950s. The molluscs of the Ranikot Group are typical of this lack of subsequent interest because, although the monographs by Cossmann & Pissarro (1909) and Vredenburg (1923, 1924, 1928) demonstrated that this fauna is one of the richest found in the Early Paleogene of the Eastern Tethys, no recent research has been undertaken. Taking account of the potential for further work, field trips in the Jhirak area and in the Lakhra Dome have been undertaken in 2010, 2011 and During these field trips we collected new material in order to better document different aspects of the paleobiodiversity of the Ranikot Group and particularly, of the Lakhra Formation - reevaluation of the species diversity, evolution of the fauna through time and comparisons with other parts of the Tethyan Ocean. The present paper is devoted to the volutids, which are abundant in the Lakhra Formation, and will be followed by further papers about other gastropod families. The volutids belong to the clade Neogastropoda. They are carnivorous burrowing molluscs and are usually considered to be thermophilic gastropods, although the majority of 2 Zootaxa 0000 (0) 2014 Magnolia Press MERLE ET AL.

3 living species occurs in warm temperate to temperate waters and some even in cold waters and in abyssal depths (Darragh 1988). Members of this family are common in the Paleogene and because of their large size, many previous authors described them in regional monographs. For Pakistan, previous works describing volutids were by d Archiac & Haime (1853), Cossmann & Pissarro (1909), Vredenburg (1923), and Cox (1930). Givens (1991) studied the genus Lyrischapa. Abbreviations Localities: Stn 1 (N25 40'35.5"; E68 11'17.6") Lakhra Dome, Old Indus Coal pit, uppermost Bara Formation. Stn 2 (N25 42'8.3"; E68 12'12.8") Lakhra Dome, Sarwar Coal pit, uppermost Bara Formation. Stn 3 (N 25 25'15.56"; E68 11'16.60") Jamshoro area, Khnu Brohi coal pit, uppermost Bara Formation. Stn 4 (N25 42'58.0"; E68 10'34.1") Lakhra Dome, East of Lakhra village section, base of the Lakhra Formation. Stn 5 (N25 40'19.6"; E68 11'23.1") Lakhra Dome, Old Indus section, uppermost Lakhra Formation. Stn 6 (N25 02'57.1"; E68 15'11.2") Jhirak area, Jhirak section, middle part of the Lakhra Formation. Repository: NHMUK PI: Natural History Museum (Paleontology Invertebrates), London. CPAG: Centre for Pure and Applied Geology, University of Sindh, Jamshoro, Pakistan. GSI: Geological Survey of India, Calcutta. MNHN.F: Muséum National d Histoire Naturelle (collection de Paléontologie), Paris. Other abbreviations: RAN: Ranikot I: Invertebrate Characters: ssc: subsutural cord shc: shoulder cord Dimensions: H: height D: greatest diameter Geological setting. The Cenozoic sedimentary deposits of Pakistan can be divided into two distinct basins: the Indus Basin and the Balochistan Basin. The Sulaiman and Kirthar Range form the Upper and Lower Indus basins respectively, and they are classically separated by the Jacobabad re-entrant (Bender & Raza 1995). In the Lower Indus Basin, the early Paleogene deposits (Paleocene to Early Eocene) are designated as the Ranikot Group. They are well exposed in the Karachi Arc (Sindh Province, southern Pakistan), an invergent fold-thrust belt along the western margin of the Indo-Pakistan Plate that protrudes eastwards into the Lower Indus Basin (Schelling 1999) and particularly in the Lakhra Dome and in the Jhirak-Jimpir area (Fig. 1). The earliest systematic geological investigation in the Hyderabad region was that of Blanford (1879), who produced a geological map showing the Lakhra Dome. The Jhirak-Jimpir area is also historically well-known thanks to the geological description by Vredenburg (1906, 1909) and to the paleontological descriptions by Cossmann & Pissarro (1909) and Vredenburg (1923, 1924, 1928). The Ranikot Group consists in ascending order of three distinct lithostratigraphic units: the Khadro, Bara, and Lakhra Formations (Cheema et al. 1977), but recently some authors included the Sohnari Formation in this group (see Frederiksen 1994). The Khadro Formation (Cardita beaumonti beds of early authors) and the Sohnari Formation are not dealt with in this study. The Bara Formation This formation includes non-marine, brackish water and inner shelf marine deposits. It ranges in thickness from more than 1,000 m in the Karachi Trough 50 km southwest of the Lakhra Dome (Quadri and Shuaib 1986) to less than 60 m in the northern part of the Sulaiman Range (Williams 1959). At Lakhra, the coal-bearing Bara Formation consists of shales, sandstones, marls and coal bands on the western side. In addition, EARLY EOCENE VOLUTIDS FROM PAKISTAN Zootaxa 0000 (0) 2014 Magnolia Press 3

4 the presence of marls and lenticular argillaceous limestone bands at Lakhra in the west reflect marine water conditions (Baqri 1997). Following Frederiksen (1994) who studied angiosperm pollen, the age of the Bara Formation is not younger than early Selandian, but according to Wakefield & Monteil (2002), the top of the Bara Formation could be late Thanetian. FIGURE 1. Location of the fossil localities (stn 1 to 6) in a simplified geological map showing the Ranikot Group in the Jimpir-Jhirak Area (JJA) and the Lakhra Dome (LD). From the Bara Formation, we sampled volutids in three localities, two at Lakhra (Old Indus Coal pit (stn 1) and Sarwar Coal pit (stn 2)) and one at Jamshoro (Khnu Brohi coal pit (stn 3)) near Hyderabad. All the material was collected on waste heaps resulting from coal extraction in the upper part of this formation. At Lakhra, the assemblage is dominated by Calyptraphorus indicus Cossmann & Pissarro, 1909 and turritellids for the gastropods and carditids, corbulids and lucinids among the bivalves. At Jamshoro, the assemblage is quite different, consisting of a Chama bed containing Calyptraphorus indicus, turritellids, rare Vetigastropoda, carditids, nuculids, venerids, corals, shark teeth and crabs (Charbonnier et al. 2013). Although the assemblages from Lakhra and Jamshoro are slightly different, they both indicate deposition in shallow water marine environments. 4 Zootaxa 0000 (0) 2014 Magnolia Press MERLE ET AL.

5 The Lakhra Formation The Lakhra Formation conformably overlies the Bara Formation and consists of evenly bedded impure limestone alternating with bioclastic sandstone and calcareous shale. The most fossiliferous beds correspond to bioclastic sandstones containing a rich molluscan fauna described by Cossmann & Pissarro (1909). The most common gastropods are Calyptraphorus indicus and Turritella ranikoti (Vredenburg, 1928). The age of the Lakhra Formation is quite controversial in the literature. In the classical book Stratigraphy of Pakistan, Shah (1977, 2009) indicated that the Lakhra Formation is entirely Late Paleocene in age, whereas Wakefield & Monteil (2002) suggested that it straddles the Paleocene/Eocene boundary. In the Jhirak area, located about 30 km SSE of the Lakhra Dome, the transition between Bara and Lakhra Formations is particularly well exposed on the southern bank outfall drainage channel. Here, in the first calcareous sandstone horizon at the base of the Lakhra Formation, we found a quite well-preserved assemblage of larger foraminifers, characterized by the co-occurrence of Alveolina vredenburgi Davies, 1937 (formerly A. cucumiformis Hottinger, 1960), Miscellanea miscella (d Archiac & Haime, 1853) and Ranikothalia nuttali (Davies, 1927), which allows correlation of this horizon with shallow-water benthic biozone SBZ 5 (Serra-Kiel et al. 1998; Hottinger 2009). According to criteria used to characterize the base of Eocene of the GSSP section (Dababiya, Egypt: Aubry et al. 2007), the Paleocene/Eocene (P/E) boundary is defined by the onset of the negative Carbon Isotope Excursion (CIE), which has been correlated directly with the base of the P5b and NP9b planktonic zones. Regarding the standard shallow benthic zonation (Serra-Kiel et al. 1998; Hottinger 2009), correlation with the CIE is not obvious, because the shallow-water sediments are generally more affected by diagenesis than deep marine sediments (Zhang et al. 2013) and moreover their record is often discontinuous. Following the recalibration of the shallow benthic zonation of Scheibner and Speijer (2009) from Egyptian sections, the P/E boundary should be located between SBZ4 and SBZ5. However, recent investigations on a relatively continuous shallow-water limestone succession in Tibet (Zhang et al. 2013) suggested that the onset of the CIE (i.e., the P/E boundary) is located in the upper part of SBZ5, characterized by the First Occurrence (FO) of Alveolina vredenburgi. Thus the base of the Lakhra Formation is probably close to the P/E boundary, at least in the Jhirak area. Further studies are required to test the synchroneity or the diachroneity of the base of the Lakhra Formation in other outcrops, including the area of the type section (Lakhra Dome). However, preliminary stratigraphic results indicate that the Calyptraphorus indicus facies, which marks the base of the Lakhra Formation in the region of Jhirak (Vredenburg 1909), is also present at the base of the Lakhra village section (stn 4) in the Lakhra Dome. From the Lakhra Formation, we sampled volutids at three localities, two at Lakhra (Lakhra village section (stn 4) and Old Indus section (stn 5) and one at Jhirak (stn 6)). The Lakhra village section displays the base of the Lakra Formation, whereas in the Old Indus section the uppermost Lakhra Formation is well exposed. In a bioclastic sandstone located at the base of the Lakhra village section, the fossil assemblage contains a diversified fauna containing molluscs (Calyptraphorus indicus facies), corals, echinids, crabs, large foraminifers, ray teeth and turtle bones. The main part of the volutid material described here comes from this bed, in which we identified about 150 species of gastropods. In the uppermost part of the Old Indus section, we found another bed of bioclastic sandstone containing a rich assemblage of invertebrates. Both assemblages are comparable to the fauna from Jhirak described by Cossmann & Pissarro (1909). At Jhirak, we collected material from the middle part of the Lakhra Formation, but the assemblage sampled was far less diverse than those found in the Lakhra Dome. Material and methods In the Jhirak area and in the Lakhra Dome, the fossilized molluscs found in the Lakhra Formation are preserved as internal molds or as recrystallized shells. For the gastropods, the internal molds are usually useless for identification at low taxonomic ranks, but the recrystallization of the shells in calcite neomorphs replicated the original aragonite shells perfectly, and their delicate details, such as spiral threads, spines or growth lines, can be observed and used for identification. However, the shells are often partially embedded in their sedimentary matrix (a bioclastic sandstone) that masks interesting details (Fig. 2) and as such they need preparation in order to extract them from the matrix. Thus, the material presented here has been extracted as much as possible with a pneumatic engraving pen. The type and figured material is housed at the Centre for Pure and Applied Geology of the University of Sindh (Jamshoro, Pakistan) and a copy (plastotype) of this material is housed at the Muséum national d Histoire naturelle, Paris (collection de Paléontologie). EARLY EOCENE VOLUTIDS FROM PAKISTAN Zootaxa 0000 (0) 2014 Magnolia Press 5

6 FIGURE 2. Volutids embedded in the bioclastic sandstone of the Lakhra Formation (stn 4, sample 2014). A. A specimen of Lyrischapa vredenburgi sp. nov. showing only a part of its first whorls. The white arrow indicates the location of the protoconch. B. A specimen of Lyrischapa blanfordi (Vredenburg, 1923) partially (base and columellar folds) embedded in the sedimentary matrix. Scale bars: 50 mm. FIGURE 3. Stratigraphic ranges of Pakistani volutid species in the Kadhro Formation (Danian), Hangu and Bara Formations (Selandian-Thanetian) and Lakhra Formation (Earliest Eocene, Ypresian). Blue lines: probable autochthonous species descending from the Indo-Pakistan stock; black line, species persisting from formation to another one; red lines: probable invaders. Note that the low species richness found in the Uppermost Bara Formation results to the poor preservation of the shell material (decalcified shells) and thus corresponds to taphonomic bias. 6 Zootaxa 0000 (0) 2014 Magnolia Press MERLE ET AL.

7 Systematics (D. Merle & J.-M. Pacaud) Family Volutidae Rafinesque, 1815 Subfamily Volutinae Rafinesque, 1815 Tribe Lyriini Pilsbry & Olsson, 1954 Genus Lyria Gray, 1847 Type species. Voluta nucleus Lamarck, 1811 (Recent, Eastern Australia) by original designation [= Voluta pattersonia Perry, 1811]. Lyria sp. (Fig. 4A) Description. Fragment of a narrow spire, H 14.8, D 9.0 mm. Protoconch eroded, probably paucispiral. Teleoconch of four whorls, the last broken. Whorls high, weakly convex, delineated by a deep, linear suture. Axial sculpture of strong but narrowly rounded costae, almost corresponding from whorl to whorl. No spiral sculpture. Costae opisthocline, rather straight. Columella with at least three folds. Material. 1 spm (stn 5: CPAG.RAN.I.1, cast MNHN.F.A50339). Comments. Although the aperture of this specimen is lacking, the shape of its teleoconch whorls is similar to those found in other members of Lyria, as for example in Lyria subturgidula (d Orbigny, 1850) from the Middle Eocene of the Paris Basin. By its high whorls and its straight costae, this specimen is easily distinguishable from Lyria cossmanni Vredenburg, 1923 (=?L. feddeni Vredenburg, 1923) from the Lakhra Formation and from Lyria samanaensis Cox, 1930 from the Hangu Formation (Samana Range, North Pakistan). Both species belong to another genus, Lyriopsis nov. gen., described below. Stratigraphic range. Lakhra Formation: Lakhra Dome. Genus Mitreola Swainson, 1833 Type species. Mitra monodonta Lamarck, 1803 (Middle Eocene, Paris Basin, France) by subsequent designation (Herrmannsen 1847). Mitreola brohii sp. nov. (Fig. 4B H) Etymology. Dedicated to Professor Imdad A. Brohi, Centre for Pure and Applied Geology, University of Sindh, Jamshoro, Pakistan. Type locality. Stn 4: Lakhra Dome, Lakhra village section, base of the Lakhra Formation. Type material. Holotype (stn 4, CPAG.RAN.I.2; cast MNHN.F.A50340), paratype 1 (stn 4, CPAG.RAN.I.3; cast MNHN.F.A50341), paratype 2 (stn 4, CPAG.RAN.I.4; cast MNHN.F.A50342), paratype 3 (stn 4, CPAG.RAN.I.5; cast MNHN.F.A50343). Other material. 23 spm (stn 4, MNHN). Description. Shell ovately fusiform, H 30 33, D mm (holotype H 31.5 not complete, D 13.9 mm). Protoconch not preserved. Teleoconch of 7 8 whorls. Spire high, occupying 36% of total shell height. Spiral whorls slightly convex. Last whorl rather narrow. Suture linear. Axial sculpture of coarse costae, tending to disappear on penultimate whorl. Costae straight, orthocline, reaching from suture to suture on spire. On first whorl: 12 costae; from second to sixth whorls: 10 to 11 costae; last whorl: no costae. Spiral sculpture of ca fine threads, present only on four earliest whorls. On last whorl, very fine threads on base of aperture. Aperture EARLY EOCENE VOLUTIDS FROM PAKISTAN Zootaxa 0000 (0) 2014 Magnolia Press 7

8 lenticular, elongate, occupying 63% of total height, 36% of diameter. No posterior notch. Outer lip thin, not thickened externally. Inner lip bearing a callus developed more anteriorly than posteriorly. Four strong columellar folds, the anteriormost oblique. Siphonal canal short, slightly curved dorsally. Siphonal notch shallow, siphonal fasciole poorly developed. Comparisons. According to Darragh (1985, 1988), Mitreola is a Tethyan genus during the Middle Eocene and its range extended from Australia to the Eastern Atlantic (Europe). However, no species of Mitreola has been recorded between Australia and the western Tethys and, therefore, this record of a species of Mitreola from the Earliest Eocene of the Eastern Tethys fills this gap. M. brohii displays the usual characters of Mitreola as well as atypical characters. An elongate, lenticular aperture with an inner lip bearing four columellar folds, the most basal being oblique, are shared with many species of Mitreola. Coarse costae (e.g. M. raricosta (Lamarck, 1803) and M. maxwelli Le Renard, 1994 from the Lutetian of the Paris Basin) or a lack of axial sculpture (e.g. M. labratula (Lamarck, 1803) from the Lutetian of the Paris Basin) are observed in many species of the genus, but the disappearance of the axial sculpture on the last whorls is less frequent. This character of M. brohii is shared only with M. salaputium Darragh, 1988 from the Late Eocene of South West Australia. M. brohi differs from all other Mitreola species in lacking an externally thickened outer lip, which often bears a variably developed labral nodule (Cernohorsky 1970; Cate 1972). These authors and Darragh (1988) regarded the presence of a labral nodule as a diagnostic character of the genus, but some species of Mitreola (e.g. M. labratula (Lamarck, 1803), M. mutica (Lamarck, 1803) and M. subplicata (Deshayes, 1835) from the Lutetian of the Paris Basin, France) lack a nodule. Stratigraphic range. Lakhra Formation: Lakhra Dome. Lyriini incertae sedis Genus Lyriopsis gen. nov. Type species. Lyria cossmanni Vredenburg, 1923 by present designation. Early Eocene from Lakhra Formation (Sindh, Pakistan). Etymology. Combination of Lyria with the suffix opsis (resembling in appearance). Gender feminine. Diagnosis. A ventricose Lyria shape with costae slightly opisthocline, rather flexuous in their posterior part, not corresponding from whorl to whorl. Whorls rather depressed, slightly convex, terraced. Four oblique, strong columellar folds. Protoconch bulbous. Included species. Lyria cossmanni Vredenburg, 1923 (Lakhra Formation, Sindh Province, Pakistan) and Lyria samanaensis Cox, 1930 (Hangu Formation, Paleocene, Khyber Pakhtunkhwa, Pakistan). Discussion. Cossmann & Pissarro (1909), Vredenburg (1923) and Cox (1930) attributed these two species to the genus Lyria and compared them to three Eocene species from the Paris Basin: Lyria (Lyria) harpula (Lamarck, 1803), L. (L.) subturgidula (d Orbigny, 1850) (= Voluta turgidula Deshayes, 1835, not of Brocchi, 1814) and L. (Pseudolyria) coroni (Morlet, 1888). According to these authors, strong but narrowly rounded costae and a biconic shape are shared with the Eocene Lyria species of the Paris Basin and even with many other Cenozoic species of Lyria. Thus, Lyriopsis is placed in the tribe Lyriini, but four characters distinguish members of Lyria from Lyriopsis: the morphology of the costae, the construction of the columellar folds, the shape of the whorl, and the protoconch. In L. (Lyria), the posterior ends of the costae are straight, whereas they are obviously sinuous in Lyriopsis. Regarding the subgenus Pseudolyria Martin, 1931 [type species Pseudolyria ventricola Martin, 1931 by monotypy], the costae are more sinuous than in L. (Lyria), but the sinuosity is more anteriorly placed than in Lyriopsis. Moreover, the outer lip of L. (Pseudolyria) is crenulated, whereas it is smooth in Lyriopsis. In L. (Lyria), the inner lip of most species displays two or three strong basal folds and several weaker folds adapically, whereas only four strong folds have been observed in the species referred to Lyriopsis. The shape of the whorl is rather depressed, slightly convex and terraced in Lyriopsis species, whereas the whorls are strongly convex in most species of Lyria (Lyria). For these reasons, the placement of Lyriopsis in the Lyriini is provisional and uncertain. 8 Zootaxa 0000 (0) 2014 Magnolia Press MERLE ET AL.

9 FIGURE 4. A. Lyria sp., CPAG.RAN.I.1, stn 5; B H. Mitreola brohii sp. nov., B C (holotype, CPAG.RAN.I.2, ventral (B) and dorsal (C) views, stn 4), D (paratype 1, CPAG RAN.I.3, ventral view, stn 4), E F (paratype 2, CPAG.RAN.I.4, ventral (E) and dorsal (F) views, stn 4), G H (paratype 3, CPAG.RAN.I.5, juvenile, ventral view (G) and enlarged view of the spire (H), stn 4); I K. Lyriopsis cossmanni (Vredenburg, 1923), I J (CPAG.RAN.I.6, ventral view (I) and enlarged view of the spire (J), stn 4), K (CPAG.RAN.I.7, ventral view, stn 4); L. Lyrischapa haimei (d Archiac & Haime, 1853), CPAG.RAN.I.8, ventral view, stn 6 ; M N. Lyrischapa sismondai (d Archiac & Haime, 1853), M (CPAG.RAN.I.10, ventral view, stn 6), N (CPAG.RAN.I.9, dorsal view, stn 3). Scale bars: B G, I, K N = 10 mm; A, H, J = 5 mm. EARLY EOCENE VOLUTIDS FROM PAKISTAN Zootaxa 0000 (0) 2014 Magnolia Press 9

10 Lyriopsis cossmanni (Vredenburg, 1923) comb. nov. (Fig. 4I K) Lyria cossmanni Vredenburg, 1923: 264 (new name for Lyria sihesurensis sensu Cossmann & Pissarro, 1909, pl. 2, fig. 26 and pl. 3, figs 25 26, incorrect identification as Voluta sihesurensis d Archiac & Haime, 1853). Lyria sihesurensis (non Voluta sihesurensis d Archiac & Haime, 1853). Cossmann & Pissarro, 1909: 29, pl. 2, fig. 26, 31 and pl. 3, figs [new combination]. Lyria feddeni Vredenburg, 1923: 265 (new name for Lyria sihesurensis sensu Cossmann & Pissarro, 1909, pl. 2, fig. 31, incorrect identification as Voluta sihesurensis d Archiac & Haime, 1853); Cotter in Vredenburg 1928: 37. Lyria cossmanni. Cotter in Vredenburg 1928: 37. Description. Shell biconic, H 30 32, D mm. Protoconch smooth, rounded, bulbous, of 2¼ whorls, strongly differentiated from teleoconch. Teleoconch of 4 to 4 ¼ whorls. Spire low, occupying 23% of total shell height. Spiral whorls rather depressed, weakly convex, terraced. Last whorl ventricose. Suture slightly channelled. Axial sculpture of strong but narrowly rounded costae, not corresponding from whorl to whorl. Costae slightly opisthocline, extending from suture to suture on spire, posterior part rather flexuous, extending to base. First whorl: 18 costae, spire: costae, last whorl: 9 10 costae. Spiral sculpture of ca. 30 fine threads present only from base to anterior half of last whorl. No spiral sculpture on spire. Aperture narrowly ovate, occupying 60% of total height, 28% of diameter. Posterior notch shallow. Outer lip thickened externally. Inner lip almost straight anteriorly, slightly curved posteriorly. Parietal callus weakly developed. Four strong columellar folds. Base of siphonal canal not preserved. Material. 2 spm (stn 4: CPAG.RAN.I.6 7, cast MNHN.F.A ). Comments. According to Vredenburg (1923), Cossmann & Pissarro (1909) made a mistake in identifying the Ranikot material as Lyria sihesurensis, which represents another species from the Middle Eocene of the Salt Range (Pakistan). From the monograph of Cossmann & Pissarro (1909), Vredenburg (1923) distinguished two species: Lyria cossmanni (in reference to Lyria sihesurensis sensu Cossmann & Pissarro, 1909, pl. 2, fig. 26 and pl. 3, figs 25 26) and L. feddeni (in reference to Lyria sihesurensis sensu Cossmann & Pissarro, 1909 pl. 2, 31 only). The figured material of L. cossmanni corresponds to three specimens from Jhirak (two well-preserved juveniles and a less well-preserved adult). The specimen illustrated on plate 3, fig. 26 is a well-preserved juvenile and is designated here as the lectotype of L. cossmanni. The material collected in the Lakhra Dome is similar to the specimens of L. cossmanni from Jhirak and shares fine threads present only from the base to anterior part of the last whorl and opisthocline costae flexuous over the posterior part of the whorls. The only illustrated specimen of L. feddeni is designated here as the lectotype of the species. Unfortunately, it is poorly preserved and it is difficult to establish differences between L. cossmanni and L. feddeni, although they were published in the same work (respectively Vredenburg 1923, pp. 264 and 265). In respect to article of the ICZN (Determination of precedence of names or acts by the First Reviser) we act as first revisers and give precedence to the name Lyria cossmanni by those who consider L. cossmanni and L. feddeni to be conspecific; thus the valid name for the species is Lyriopsis cossmanni (Vredenburg, 1923). L. cossmanni can be compared to L. samanaensis Cox, 1930 from the Paleocene (Köthe 1988; Wardlaw et al. 2007; Afzal et al. 2009) of the Hangu Formation (Samana Range, North Pakistan). Cox (1930) compared L. samanensis to L. feddeni, but as seen above, the illustrated specimen of L. feddeni does not allow accurate identification. Lakhra specimens are very similar to the holotype of L. samanaensis. They share the same number of costae on the spire and on the last whorl, a flexuous morphology of these costae, spiral threads on the base of the last whorl, four oblique and strong columellar folds on the inner lip, and a pyriform outline. These numerous similarities suggest strongly that Lyriopsis cossmanni and Lyriopsis samanaensis belong to the same species. However, the protoconch Lyriopsis samanaensis is missing and this character is needed before we can regard L. samanensis as a junior synonym of L. cossmanni. Kachhara et al. (2011) reported the occurrence of L. feddeni in the Paleocene from Southwestern Kachchh, Gujarat (India), but this identification is based only on an internal cast lacking specific characters. Stratigraphic range. Lakhra Formation: Jhirak (Cossmann & Pissarro 1909; Vredenburg 1923) and Lakhra Dome (this paper). 10 Zootaxa 0000 (0) 2014 Magnolia Press MERLE ET AL.

11 Volutinae incertae sedis Genus Lyrischapa Aldrich, 1911 Type species. Lyrischapa harrisi Aldrich, 1911 (Middle Eocene, Mississippi, USA) by original designation. Discussion. Wenz (1943) assigned Diconomorpha Wenz, 1943 [type species Diploconus elegans Douvillé, 1929 = Diploconus Douvillé, 1929, non Candèze (1860) [Insecta] by original designation] to the Pholidotominae Cossmann, 1896, but Volutocristata Gardner and Bowles, 1934 [type species: V. chiapasensis Gardner, 1934 by original designation] and Lyrischapa to the Volutinae, in spite of close similarities between these three genera. Givens (1979) demonstrated that Volutocristata is a junior synonym of Lyrischapa and, following Pilsbry & Olsson (1954), he assigned Lyrischapa to the Fulgorariinae, because of the deviated protoconch, numerous columellar folds, a shallow siphonal notch and a weak siphonal fasciole. Later, Givens (1991) suggested that Diconomorpha might be a junior synonym of Lyrischapa. In addition, he considered that the assignment of Lyrischapa to the Fulgorariinae is incorrect, because members of this subfamily are distinguished from Lyrischapa by quite different shapes (fusiform to ovate), absence of an anal sulcus on the outer lip, and generally fewer columellar folds. Species of Lyrischapa have a biconic shape, a spiral row of variably developed shoulder spines, weak spiral sculpture on the last whorl, and five to six strong columellar folds. Several Athletinae and particularly the genus Athleta share with Lyrischapa a biconic shape and a row of shoulder spines. For this reason, Vredenburg (1923) compared his new species A. blanfordi with Athleta (Athleta). However, the early whorls of Athleta (Athleta) species display a subsutural and two abapical rows of nodules, which are missing in Lyrischapa. This character is widespread in juveniles of Athleta species and is even present in adults of A. (Volutocorbis) species. Thus, the similarities between the shells of Athleta and Lyrischapa are superficial. The shells of Lyrischapa more closely resemble species of the Amoriinae or the Volutinae. In the Amoriinae, they can be compared with Nannamoria Iredale, 1929 [type species: N. amicula Iredale, 1929]. Several species (e.g. N. ralphi (Finlay, 1930) from the Middle Miocene of Australia or N. paraboloides Darragh, 1988 from the Late Neogene of Australia) have a biconic Athleta shape, developed shoulder spines and weak spiral sculpture on the last whorl, but differ from Lyrischapa by the presence of only four strong columellar folds (instead of five or six to eight). With respect to the Volutinae, the shells of Lyrischapa can be compared with Voluta ebraea Linnaeus, 1758 or V. musica Linnaeus, 1758 (Recent, Brazil), which exhibit five to six strong columellar folds and several weaker folds adapically, and some specimens have a biconic shape with developed shoulder spines. However, a morphological type close to Lyrischapa is uncommon in the Amoriinae and Volutinae and could be the result of convergent and repeated evolution. Thus, like Givens (1991), we conclude that the subfamilial position of this genus is uncertain, but we believe that an assignment to the Fulgorariinae or Athletinae is incorrect. Lyrischapa haimei (d Archiac & Haime, 1853) (Fig. 4L) Voluta haimei d Archiac & Haime, 1853: 325, p1. 31, figs. 26, 27. Voluta haimei d Archiac, 1850: 298 (nomen nudum). Aulicina haimei. Cossmann & Pissarro 1909: 26, pl. 2, fig. 29 only [new combination]. Volutoconus funiculifer Cossmann & Pissarro, 1909: 27, pl. 3, figs Aulica (Aulicina) haimei. Vredenburg 1923: 269; Cotter in Vredenburg 1928: 38 [new combination]. Eovasum haimei. Cox 1930: 186 in part (not pl. 21, figs 1 4), see (Givens 1991) [new combination]; Davies 1934: 297. Lyrischapa haimei. Givens 1991: 664, figs [new combination]. Material. 1 spm (stn 6: CPAG.RAN.I.8, cast MNHN.F.A50346). Comments. In his paper on Lyrischapa, Givens (1991) gave a revised description of L. haimei. He recognized two differential characters distinguishing it from L. sismondai (d Archiac & Haime, 1853), with which it has often been confounded in the past: 1) a shallower shoulder sinus, and 2) the spire is covered with crowded spiral threads alternating in two sizes (a character already cited by Vredenburg 1923). In addition, Vredenburg (1923) noted that the protoconch of L. sismondai is larger than that of L. haimei. From the middle part of Lakhra Formation at Jhirak, we collected a juvenile specimen of H 22 mm (Fig. 4L). Although it is a little crushed, it displays these three EARLY EOCENE VOLUTIDS FROM PAKISTAN Zootaxa 0000 (0) 2014 Magnolia Press 11

12 distinguishing characters of L. haimei. Stratigraphic range. Lakhra Formation: Jhirak (Cossmann & Pissarro 1909; Vredenburg 1923; this paper). Lyrischapa sismondai (d Archiac & Haime, 1853) (Fig. 4M N) Voluta sismondai d Archiac & Haime, 1853: 326, p1. 31, figs. 25. Voluta sismondai d Archiac, 1850: 298 (nomen nudum). Aulicina pusiola Cossmann & Pissarro, 1909: 26, pl. 2, fig. 34, pl. 3, figs. 4, 5. Volutoconus corrugatus Cossmann & Pissarro, 1909: 28, pl. 3, figs. 6, 7. Aulica sismondai. Vredenburg 1923: 267 [new combination]. Aulica (Aulicina) sismondai. Cotter in Vredenburg 1928: 38. Eovasum sismondai. Cox 1930: 186 in part (pl. 21, figs 1 4), see Givens (1991) [new combination]. Eovasum haimei (misidentification as Voluta haimei d Archiac & Haime, 1853). Iqbal 1972: 64, pl. 17, fig. 8. Lyrischapa sismondai. Givens 1991: 666, figs , 4.1, 4.2, 4.5 (non figs 4.3, 4.6) [new combination]. Material. 1 spm (stn 3: CPAG.RAN.I.9, cast MNHN.F.A50347); 1 spm (stn 6: CPAG.RAN.I.10, cast MNHN.F.A50348). Comments. In order to avoid confusion with Lyrischapa haimei, Givens (1991) gave a supplementary description for L. sismondai. According to this author, L. sismondai exhibits a larger protoconch, a deeper shoulder sinus and its spiral sculpture differs by having threads of uniform size and spacing on the spire, rather than two sizes of threads alternating. The two specimens collected at Jhirak in the Lakhra Formation and at Jamshoro in the Bara Formation are no more than mm in height and have a high spire. They are rather similar to specimens from the Hangu Formation (NHMUK PI G49443, G49444, G49445) and from the Lakhra Formation (NHMUK PI G50319) illustrated in the paper by Givens (1991), but they differ from the largest specimen (NHMUK PI G92737) from the Ypresian (cf. Wardlaw et al. 2007) of the Nammal Formation, Salt Range, Pakistan) illustrated by this author (Givens 1991, figs 4.3, 4.6). Numerous specimens collected in the Lakhra Dome provide a good ontogenetic series linking juveniles with the largest specimens, such as those figured by Givens (1991) and Cossmann & Pissarro (1909, pl. 2, fig. 30, pl. 8, fig. 2). Although they share a large protoconch, these juveniles display many differences from L. sismondai. At the same size (H 40 mm), they exhibit a lower spire and a wider shape, their spiral sculpture is obsolete on the spire and missing on the last whorl, except on the base which bears some fine threads, and their shoulder spines are acute and better developed. Specimens of a larger size and very large specimens of around H 80 mm have the same characters on their early teleoconch whorls, showing that they belong to the same species, which is described here below as Lyrischapa vredenburgi sp. nov. However, they also demonstrate that the specimens attributed to L. sismondai do not correspond to the early stages of growth of the largest specimens often attributed to this species. Thus, the supplementary description by Givens (1991) is still a composite description and the character 9 12 prominent spines on the shoulder should be replaced by: 9 10 short, rounded spines on the shoulder. By its biconical shape, by the configuration of the shoulder sinus and by its short, rounded shoulder spines, L sismondai appears closely related to Diconomorpha elegans (Douvillé, 1929) [type species of Diconomorpha Wenz, 1943 = Diploconus Douvillé, 1929, non Candèze (1860) by original designation] from the Danian of Sindh (Kadhro Formation). Although these Danian specimens do not display the superficial sculpture observed in L. sismondai, because they are poorly preserved, they are so similar that, according to Givens (1991), generic distinction seems unjustified. Stratigraphic range. Hangu Formation: Hangu Shales (Cox 1930); Upper Bara Formation: Jamshoro (this paper); Lakhra Formation: Jhirak (Cossmann & Pissarro 1909; this paper). Lyrischapa vredenburgi sp. nov. Fig. 5A H Aulicina haimei (non Voluta haimei d Archiac & Haime, 1853). Cossmann & Pissarro 1909: 25, pl. 2, fig. 30, pl. 8, fig. 2. Athleta blanfordi Vredenburg, 1923 pars: 255, proposed for Aulicina haimei d Archiac & Haime, 1853 sensu Cossmann & Pissarro, 1909, pl. 8, fig. 2 only. Lyrischapa sismondai (non Voluta haimei d Archiac & Haime, 1853). Givens 1991, 666, figs 4.3, Zootaxa 0000 (0) 2014 Magnolia Press MERLE ET AL.

13 FIGURE 5. A H. Lyrischapa vredenburgi sp. nov., A B (paratype 1, CPAG.RAN.I.12, protoconch and first teleoconch whorl in ventral (A) and dorsal (B) views, stn 4), C E (holotype, CPAG.RAN.I.11, ventral (C), dorsal (D) and apical (E) views, stn 4), F (paratype 2, CPAG.RAN.I.13, ventral view, stn 4), G (paratype 3, CPAG.RAN.I.14, ventral view, stn 4), H (paratype MNHN.F.J12751, Cossmann coll., Jhirak). Scale bars: A H = 10 mm. EARLY EOCENE VOLUTIDS FROM PAKISTAN Zootaxa 0000 (0) 2014 Magnolia Press 13

14 Etymology. Dedicated to E. W. Vredenburg for his work on the geology and paleontology of the Ranikot Group. Type locality. Stn 4: Lakhra Dome, Lakhra village section, base of the Lakhra Formation. Type material. Holotype (stn 4: CPAG.RAN.I.11, cast MNHN.F.A50349), paratype 1 (stn 4: CPAG.RAN.I.12, cast MNHN.F.A50350), paratype 2 (stn 4: CPAG.RAN.I.13, cast MNHN.F.A50351), paratype 3 (stn 4: CPAG.RAN.I.14, cast MNHN.F.A50352), three paratypes (Jhirak, Cossmann coll. MNHN.F.J12751). Other material. 16 spm (stn 4: MNHN); 3 spm (stn 5: MNHN). Description. Shell biconical, wide, H 77 85, D mm (holotype (subadult) H 38.3, D 31.1 mm). Protoconch smooth, globose, slightly deviated, of two whorls; H 16, D 11 mm when complete (Fig. 5A B). Transition from protoconch to teleoconch not defined. Teleoconch of 3 to 3.5 whorls. Spire low, occupying 10% of total shell height. Spiral whorls almost flat, separated by shallow suture undulating between shoulder spines of preceding whorl. Last whorl rather triangular. Spiral sculpture on spire of obsolete threads, visible only on first teleoconch whorl. On juvenile specimens of one whorl, last whorl sculptured with very fine spiral threads covering three abapical quarters of shell, threads disappearing on succeeding whorls. Axial sculpture of spiny shoulder costae only. First whorl: 11 costae, becoming more prominent on second half of whorl. Second whorl: 9 10 costae; third whorl: 9 12 costae. No shoulder sinus. Aperture narrow, occupying 87% of total height, 28% of diameter. Inner lip straight, columella with five strong folds succeeded posteriorly by one weaker fold; inner lip callus narrow. Outer lip thin, not thickened externally. Siphonal canal short; siphonal notch weak; siphonal fasciole low, weak, but delineated by carina spreading dorsally on gerontic specimens. Comparisons. From the Lakhra Formation, this species can be compared with Lyrischapa blanfordi (Vredenburg, 1923). Both species share obsolete threads on the first teleoconch whorl and the disappearance of the spiral sculpture on the last few whorls. As in L. vredenburgi, L. blanfordi exhibits shoulder spines and lacks a shoulder sinus. L. blanfordi differs in its less numerous costae on the last whorl (8 9 instead of 9 12), its shoulder spines oriented adapically and its narrower shape. The number of columellar folds varies from 5 to 6, whereas those of L. blanfordi vary from 6 to 8. L. vredenburgi also has a strong similarity to L. crassa (Douvillé, 1929) described from the Danian of the Kadhro Formation, at Jakhmari and at Ranikot. The two species share a wide biconic shape, a low spire, 12 costae on the last whorl and the presence of shoulder spines. The type specimen of L. crassa is poorly preserved, but it apparently differs from A. vredenburgi by having only four columellar folds and more rounded shoulder spines. L. crassa was originally assigned to Diconomorpha, but the close similarities between A. crassa and A. vredenburgi constitute a further argument to consider Diconomorpha as a junior synonym of Lyrischapa. Stratigraphic range. Lakhra Formation at Lakhra and Jhirak; Nammal Formation, Salt Range (Givens 1991). Lyrischapa blanfordi (Vredenburg, 1923) comb. nov. (Fig. 6A F) Athleta blanfordi Vredenburg, 1923: 255. Aulicina haimei (non Voluta haimei d Archiac & Haime, 1853). Cossmann & Pissarro 1909: 25, pl. 2, fig. 27 only. Athleta blanfordi. Cotter in Vredenburg 1928: 36. Description. Shell biconic, elongate, H 68 85, D mm. Protoconch smooth, globose and slightly deviated, of two whorls, H 6, D 10 mm when complete. Protoconch/teleoconch transition not obvious. Teleoconch of 3 whorls. Spire low, occupying 6% of total shell height. Spire whorls almost flat, separated by shallow suture undulating between shoulder spines of preceding whorl. Last whorl elongate, rather conical. Spiral sculpture on spire of obsolete threads, visible only on first teleoconch whorl. On juvenile specimens of one whorl, last whorl sculptured by very fine threads, covering abapical three quarters of shell, disappearing on succeeding whorls. Axial sculpture of only spiny shoulder costae. Shoulder spines oriented adapically on last whorl. First whorl: 11 costae, becoming more prominent on second half of whorl. Second whorl: 8 9 costae; third whorl: 8 9 costae. No anal sulcus. Aperture narrow, occupying 89% of total height, 18% of diameter. Inner lip straight, columella with six strong folds succeeded posteriorly by two weaker folds; inner lip callus narrow. Outer lip thin, not thickened externally. Siphonal canal short, slightly curved to right towards aperture; siphonal notch weak; siphonal fasciole low, weak, but delineated by carina spreading dorsally on gerontic specimens. Material. 28 spm (stn 4: 3 spm CPAG.RAN.I.15 17, cast MNHN.F.A and 25 spm MNHN). 14 Zootaxa 0000 (0) 2014 Magnolia Press MERLE ET AL.

15 FIGURE 6. A F. Lyrischapa blanfordi (Vredenburg, 1923), A C (CPAG.RAN.I.17, ventral (A), dorsal (B) and apical (C) views, stn 4), D E (CPAG.RAN.I.16, ventral (D) and dorsal (E) views, stn 4), F (CPAG.RAN.I.15, ventral view, stn 4); G I. Lyrischapa brevispira sp. nov., holotype, CPAG.RAN.I.18, ventral (G), dorsal (H) and apical (I) views, stn 4. Scale bars: A H = 10 mm. EARLY EOCENE VOLUTIDS FROM PAKISTAN Zootaxa 0000 (0) 2014 Magnolia Press 15

16 Comments. Although Cossmann & Pissarro (1909) confounded specimens of L. haimei with L. blanfordi, L. blanfordi was originally described in the genus Athleta and later Givens (1991) did not discuss it in his revision of the genus Lyrischapa. Vredenburg (1923) compared L. blanfordi with two Bartonian species from the Anglo- Parisian Basin: Athleta (A.) athleta (Solander in Brander, 1766) and A. (A.) strombiformis (Deshayes, 1835). These species exhibit a biconic shape and have shoulder spines, so generating a superficial resemblance with L. blanfordi, but they differ by their early teleoconch whorls and by their columellar folds. In A. (A.) athleta and A. (A.) strombiformis, the early teleoconch whorls have two main spiral rows of nodules (subsutural and shoulder rows) and one or two abapical rows. This kind of sculpture is typical of Paleogene Athleta (Athleta) species and it is present in other subgenera such as A. (Volutospina), A. (Volutopupa) and A. (Volutocorbis). However, it is absent in Lyrischapa, which has only a single shoulder row during the ontogenetic development of the shell. A. (A.) athleta and A. (A.) strombiformis also bear three strong columellar folds and several weaker folds adapically, whereas in L. blanfordi the number of folds varies from 6 to 8. In L. blanfordi, as in other members of Lyrischapa, these folds are stronger in most specimens than those observed in the two Bartonian Athleta species and other Paleogene Athleta species. Therefore, it is with little uncertainty that we transfer L. blanfordi to the genus Lyrischapa. Stratigraphic range. Lakhra Formation: Jhirak (Vredenburg 1923), Lakhra Dome (this paper). Lyrischapa brevispira sp. nov. (Figs 6G I, 7A D) Etymology. From the Latin brevis (short) and spira (spire). Type locality. Stn 4: Lakhra Dome, Lakhra village section, base of the Lakhra Formation. Type material. Holotype (stn 4: CPAG.RAN.I.18, cast MNHN.F.A50356), paratype 1 (stn 4: CPAG.RAN.I.19, cast MNHN.F.A50357), paratype 2 (stn 4: CPAG.RAN.I.20, cast MNHN.F.A50358). Other material. 18 spm (stn 4: MNHN). Description. Shell conical, very narrow, H 64, D ca. 35 mm (holotype (subadult) H 38.2, D 20.2 mm). Protoconch smooth, very flat, of two whorls, H 10 12, D 10 mm when complete. Transition protoconch/teleoconch not defined. Teleoconch of 3 whorls. Spire very low, occupying 6% of total shell height. Spiral whorls flat, narrow, separated by shallow suture. Sutural ramp extending partially onto preceding whorl. Last whorl narrow, rather conic. Spiral sculpture of obsolete threads, present only on first teleoconch whorl. Second and last whorls lacking spiral sculpture, even on base. Axial sculpture of very short shoulder costae formed by small spinelets. First whorl: 9 10 costae. Second whorl: 8 9 costae. Third whorl: 11 costae. Aperture narrow, occupying 97% of total height, 15% of diameter. Inner lip straight, columella with five strong folds succeeded posteriorly by one weaker fold, one or two secondary folds intercalated between major folds in a few specimens; inner lip callus narrow. Anal sulcus thin, against suture. No shoulder sinus. Outer lip thin, not thickened externally. Siphonal canal short; siphonal notch shallow; siphonal fasciole low, weak. Comparisons. In Lyrischapa haimei and in L. sismondai, spiral sculpture consisting of fine threads persists during the whole ontogeny and covers the last whorl, whereas in L. brevispira, the spiral sculpture is obsolete on the first teleoconch whorl and disappears on later whorls. For this reason, this species can be compared with L. vredenburgi and L. blanfordi, with which it shares obsolete or absent sculpture on the last whorl. L. vredenburgi and L. brevispira have a large protoconch of two whorls, up to 10 mm in width, but the apex is globose in L. vredenburgi, whereas it is flat in L. brevispira. L. brevispira differs from L. vredenburgi by its narrower shape, by its poorly developed spines, by its numerous columellar folds (6 8 instead 5 6) and by its anal sulcus against the suture. In its narrow shape, L. brevispira is similar to L. blanfordi, but it is distinguished by its larger protoconch, a very flat spire and poorly developed spines. Because of its poorly developed shoulder spines on the last whorl, L. brevispira retains a juvenile aspect in its adult stage and could be regarded as a paedomorphic species of Lyrischapa. Stratigraphic range. Lakhra Formation: Lakhra Dome. 16 Zootaxa 0000 (0) 2014 Magnolia Press MERLE ET AL.

17 FIGURE 7. A D. Lyrischapa brevispira sp. nov., A B (paratype 2, CPAG.RAN.I.20, ventral (A) and dorsal (B) views, stn 4), C D (paratype 1, CPAG.RAN.I.19, ventral (C) and dorsal (D) views, stn 4); E J. Athleta (Volutopupa) intercrenatus (Cossmann & Pissarro, 1909), E F (CPAG.RAN.I.22, ventral (E) and dorsal (F) views, stn 4), G (CPAG.RAN.I.21, ventral view, stn 4), H (CPAG.RAN.I.23, ventral view, stn 4), I-J (CPAG.RAN.I.24, ventral view (I) and enlarged view of the spire (J), stn 4). Scale bars: A I = 10 mm; J = 5 mm. EARLY EOCENE VOLUTIDS FROM PAKISTAN Zootaxa 0000 (0) 2014 Magnolia Press 17

18 Subfamily Athletinae Pilsbry & Olsson, 1954 Genus Athleta Conrad, 1853 Type species. Voluta rarispina Lamarck, 1811 (Neogene, Europe) by subsequent designation (Dall 1890). Comments. 1) Many authors consider the gender of Athleta to be feminine, but it is masculine. 2) Darragh (1971) synonymised a number of genera with Athleta, among which were Volutospina Newton, 1906, Volutopupa Dall, 1890 and Volutocorbis Dall, 1890 and recognised only two subgenera, Athleta (s.s.) and A. (Ternivoluta) Martens, However, the difference between the two subgenera used by Darragh (1971) is solely based on the multispiral versus paucispiral protoconch, a developmental difference not now considered to be of taxonomic significance, as in the Harpidae (Merle & Pacaud 2003) or the Muricidae (Merle et al. 2011) for example. The aim of this work is not to revise the genus Athleta and a cladistic analysis would be necessary for this revision. For instance, we prefer to continue to use provisionally Volutopupa (see Pacaud & Pons 2013), Volutocorbis and Volutospina as subgenera, because they are useful to characterize different Paleogene morphotypes. Subgenus Volutospina Newton, 1906 Type species. Conus spinosus Linnaeus, 1758 (Middle Eocene, Paris Basin, France) by original designation. Athleta (Volutospina) noetlingi (Cossmann & Pissarro, 1909) Volutospina noetlingi Cossmann & Pissarro, 1909: 24, pl. 2, figs ? Voluta cithara (non Voluta cithara Lamarck, 1803). d Archiac & Haime, 1853: 325, pl. 32, figs 4 5. Athleta (Neoathleta) noetlingi. Vredenburg 1923: 256; Cotter in Vredenburg 1928: 36 [new combination]. Lyria cossmanni (non Lyria cossmanni Vredenburg, 1923). Iqbal 1972: 65, pl. 19, figs 3, 9. Comments. According to Cossmann & Pissarro (1909), the specimen illustrated on their pl. 2, figs displays a biconic shape with spiny costae delineating a developed shoulder and looks more like a species of Volutospina than a species of Volutopupa Dall, 1890 (type species: Voluta cithara Lamarck, 1803 by original designation). We designate here this specimen as the lectotype of V. noetlingi. Vredenburg (1923) regarded Volutospina intercrenata Cossmann & Pissarro, 1909 as a junior synonym of V. noetlingi, considering that the specimens used to describe V. intercrenata represent juvenile specimens of V. noetlingi. However, the figured material of V. intercrenata does not display spiny costae delineating a developed shoulder, as observed in juvenile specimens of Volutospina species (e.g. the type species). Thus, these specimens do not belong to Volutospina and are not juveniles of V. noetlingi. In the Lakhra Dome, we have not found specimens of V. noetlingi, but a juvenile specimen of A. (V.) noetlingi from Jhirak has been illustrated by Iqbal (1972) under the erroneous name Lyria cossmanni. As in other juvenile specimens of Volutospina species, it has a biconic shape with spiny costae delineating a developed shoulder. Stratigraphic range. Lakhra Formation: Jhirak (Cossmann & Pissarro 1909; Vredenburg 1923). Subgenus Volutopupa Dall, 1890 Type species. Voluta cithara Lamarck, 1803, not Solander & Lightfoot, 1786 [subjective synonym of Buccinum citharoedus Holten, 1802] by original designation (Middle Eocene, Lutetian, Paris Basin, France). Athleta (Volutopupa) intercrenatus (Cossmann & Pissarro, 1909) comb. nov. (Fig. 7E J) Volutospina intercrenata Cossmann & Pissarro, 1909: 24, pl. 2, figs 18 20, pl. 3, figs 1 3. Description. Shell biconic, H 33 37, D mm. Protoconch and first teleoconch whorl unknown; at least 5 teleoconch whorls present. Spire low, occupying 12% of total shell height. Spiral whorls subcarinate, last whorl 18 Zootaxa 0000 (0) 2014 Magnolia Press MERLE ET AL.

19 weakly ventricose. Spiral sculpture on second whorl of one spiral row of small nodules on shoulder angle. From third to fourth whorl, one spiral row of subsutural nodules develops rapidly in addition to shoulder nodules (Fig. 7J); one or two small nodules between shoulder and suture. On last whorl, spiral sculpture displaying: 1) adapically two main spiral rows (subsutural and shoulder rows) with one weaker row abapically; 2) between these rows and mid-whorl, weak, poorly marked cords; 3) from mid-whorl to base, ca. 10 flattened cords, producing an imbricate effect. Axial sculpture of collabral costae extending across whorl, interrupted by spiral sculpture. From second whorl to third whorl: 17 costae; fourth whorl: costae; fifth and last whorl: costae. On last whorl, costae obsolete between adapical rows of nodules, thick and relatively high below shoulder angle, extending across base, becoming almost obsolete towards end of last whorl. Aperture ovate, occupying 61% of total height, 41% of diameter. Inner lip almost straight, columella with 6 10 low folds; inner lip callus very narrow, apparently not spreading far laterally. Outer lip thin, not thickened externally. Siphonal notch not well preserved, apparently weak. Material. 5 spm (stn 4: 4 spm CPAG.RAN.I.21 24, cast MNHN.F.A and 1 spm MNHN). Comments. As seen above with Athleta (Volutopupa) noetlingi, the four specimens of V. intercrenata illustrated by Cossmann & Pissarro (1909) are species of Volutopupa and not species of Volutospina. They are not well preserved, except the specimen illustrated in Cossmann & Pissarro s (1909) pl. 3, fig. 3, which is designated here as the lectotype of Volutospina intercrenata Cossmann & Pissarro, Its early teleoconch whorls displays two strong spiral rows of nodules (subsutural and shoulder rows) and its last whorl has 11 thick, relatively high costae, similar to the spire of the adult specimens of A. (V.) intercrenatus collected in the Lakhra Dome. The oval shape of these adults resembles more that of A. (V.) lyra (Lamarck, 1803) or A. (V.) mutatus (Deshayes, 1835) from the Middle Eocene of the Paris Basin, than it does A. (V.) citharoedus, which displays an inflated and ventricose last whorl. A. (V.) intercrenatus also differs from these species by its strongly developed row of subsutural nodules. Stratigraphic range. Lakhra Formation: Jhirak (Cossmann & Pissarro 1909) and Lakhra Dome (this paper). Athleta (Volutopupa) citharopsis sp. nov. (Fig. 8A D) Etymology. The two first syllables cithar because of the similarity with the type species, combined to the suffix opsis (Greek: appearance). Type locality. Stn 4: Lakhra Dome, Lakhra village section, base of the Lakhra Formation. Type material. Holotype (stn 4: CPAG.RAN.I.25, cast MNHN.F.A50363), paratype 1 (stn 4: CPAG.RAN.I.26, cast MNHN.F.A50364), paratype 2 (stn 4: CPAG.RAN.I.27, cast MNHN.F.A50365). Other material. 3 spm (stn 4: MNHN). Description. Shell of holotype large, H 60, D probably 31 mm (outer lip broken). Protoconch and first teleoconch whorl unknown; at least 6 teleoconch whorls. Spire moderately high, occupying 21% of total shell height. Spiral whorls convex; last whorl ventricose. Spiral sculpture of one row of very small subsutural nodules, appearing on third whorl. From fourth to fifth whorl, a second row of shoulder nodules (Fig. 8D) appears, becoming variably subspinose on penultimate whorl. On last whorl, spiral sculpture displaying: 1) adapically two main spiral rows of nodules (subsutural and shoulder rows); 2) no cords between these rows and mid-whorl; 3) from mid-whorl to base, ca. 15 flattened cords, producing an imbricate effect. Axial sculpture of collabral costae present on second whorl, extending across whorl, apparently without nodules. Third whorl: 15 costae; fourth whorl: 14 costae; fifth whorl: 13 costae; sixth whorl: 11 costae. On last (sixth) whorl, costae obsolete between adapical rows of nodules, thin and sharp below shoulder angle, extending across base, becoming almost obsolete towards end of last whorl; one intercostal ridge appearing at end of last whorl. Aperture ovate, occupying 78% of total height. Inner lip almost straight, columella with 3 to 4 low folds; inner lip callus very wide, parietal callus relatively thick, spreading far laterally, but not extending up to suture. Outer lip broken. Siphonal notch not well preserved, apparently weak. Comparisons. This species differs from Athleta (Volutopupa) intercrenatus by its more ventricose shape, by its less numerous columellar folds and by its smaller subsutural and shoulder nodules. In shape and in its wide parietal callus, it resembles A. (V.) citharoedus, the type species of the subgenus. However, A. (V.) citharopsis can be easily distinguished from A. (V.) citharoedus by its stronger columellar folds and more strongly developed subsutural nodules. Stratigraphic range. Lakhra Formation: Lakhra Dome. EARLY EOCENE VOLUTIDS FROM PAKISTAN Zootaxa 0000 (0) 2014 Magnolia Press 19

20 FIGURE 8. A D. Athleta (Volutopupa) citharopsis sp. nov., A B (holotype, CPAG.RAN.I.25, ventral (A) and dorsal (B) views, stn 4), C (paratype 1, CPAG.RAN.I.26, dorsal view, stn 4), D (paratype 2, CPAG.RAN.I.27, enlarged view of the spire, stn 4); E J. Athleta (Volutocorbis) eugeniae Vredenburg, 1923, E F (CPAG.RAN.I.29, dorsal view (E) and enlarged view of the spire (F), stn 4), G H (CPAG.RAN.I.28, dorsal (G) and ventral (H) views, stn 4), I J (CPAG.RAN.I.30, dorsal (I) and ventral (J) views, stn 4). Scale bars: A E, G J = 10 mm; F = 5 mm. 20 Zootaxa 0000 (0) 2014 Magnolia Press MERLE ET AL.

21 Subgenus Volutocorbis Dall, 1890 Type species. Volutilithes limopsis Conrad, 1860 (Paleocene, Texas, USA) by original designation. Athleta (Volutocorbis) eugeniae Vredenburg, 1923 (Fig. 8E J) Athleta (Volutocorbis) eugeniae Vredenburg, 1923: 259, pl. 15, figs 5, 7. Volutospina sykesi (non Voluta sykesi d Archiac & Haime, 1853). Cossmann & Pissarro, 1909: 23, pl. 2, figs Athleta (Volutocorbis) eugeniae. Cotter in Vredenburg 1928: 36. Volutocorbis eugeniae. Glibert 1960: 49; Ibbal 1972: 63, pl. 18, figs 5 6, pl. 20, figs [new combination]. Volutocorbis engeniae. Iqbal 1969: 60 (misspelling of eugeniae). Description. Shell biconic, slightly fusiform, H 30 38, D mm. Protoconch conical, multispiral, of three whorls. Teleoconch of 5 to 5.5 whorls. Spire moderately high, occupying 18% of total shell height. Spire whorls weakly convex, last whorl oval. Spiral sculpture on spire consisting of first row of subsutural nodules, becoming more developed and slightly spiny on penultimate whorl (Fig. 8F); abapically, three rows of nodules including shoulder row, which is weaker than other two. Shoulder row bears spiny nodules on some specimens. On last whorl, spiral sculpture displaying: 1) subsutural row of nodules; 2) shoulder row of nodules spiny on a few specimens, and 3) abapically, 13 rows of nodulose crenulations, becoming less prominent on base. From second to last whorls, costae interrupted by rows of nodules. On last whorl, costae moderately high, cut into prominent, regular, nodulose crenulations by broad spiral grooves. Third whorl: 16 costae; fourth whorl: costae; fifth and last whorl: costae. Aperture ovate, rather narrow, occupying 55% of total height, 39% of diameter. Inner lip feebly sinuous, columella with one oblique fold succeeded posteriorly by 3 or 4 weak folds; inner lip callus very narrow, not spreading. Outer lip thin, not thickened externally. Siphonal canal slightly elongate; siphonal notch shallow. Material. 33 spm (stn 4: 3 spm CPAG.RAN.I.28 30, cast MNHN.F.A and 30 spm MNHN). Comments. According to Cossmann & Pissarro (1909) and Vredenburg (1923), Athleta (Volutocorbis) eugeniae is closely similar to A. (Volutocorbis) elevatus (Sowerby, 1840) from the Ypresian of northern Europe, particularly in the spiny shape of the subsutural and shoulder nodules. The European species differs in its other adapical spiral elements including flatter cords separated by narrow spiral grooves, producing an imbricate effect, but lacking nodules. A. (V.) rouaulti Cossmann, 1923 from the late Ypresian of southwestern France superficially resembles A. (V.) elevatus and A. (V.) eugeniae, but can easily be distinguished by its subsutural nodules disappearing progressively during growth. Stratigraphic range. Lakhra Formation: Jakhmari, Jhirak (Cossmann & Pissarro 1909; Vredenburg 1923) and Lakhra Dome (this paper). Athleta (Volutocorbis) burtoni Vredenburg, 1923 (Fig. 9A G) Athleta (Volutocorbis) burtoni Vredenburg, 1923: 261, pl. 15, fig. 2. Athleta (Volutocorbis) victoriae Vredenburg, 1923: 260, pl. 15, figs 4. Athleta (Volutocorbis) burtoni. Cotter in Vredenburg 1928: 37. Volutocorbis soriensis Eames, 1952: 109, pl. 4, fig. 95a b. Volutocorbis burtoni. Glibert 1960: 48 [new combination]. Description. Shell biconic, H 22, D 11 mm. Protoconch and two first teleoconch whorls unknown; at least 5 teleoconch whorls. Spire moderately high, occupying 21% of total shell height. Spiral whorls convex, last whorl rather oval. Spiral sculpture on spire of first row a subsutural nodules, becoming more developed apically; abapically, one or two rows of nodules including shoulder row, which is less distinct than other rows (Fig. 9F). On last whorl, spiral sculpture displaying: 1) subsutural row of nodules; 2) shoulder row of nodules; 3) abapically, rows of almost rounded nodules, becoming less prominent on base. From second to last whorls, costae EARLY EOCENE VOLUTIDS FROM PAKISTAN Zootaxa 0000 (0) 2014 Magnolia Press 21

22 interrupted by rows of nodules. On last whorl, costae rather low, cut into prominent small, regular nodules by broad spiral grooves. Third whorl: costae; fourth whorl: costae; fifth and last whorl: costae. Aperture ovate, rather narrow, occupying 75% of total height, 42% of diameter. Inner lip straight, columella with two folds, succeeded posteriorly by one or two weaker folds. Inner lip callus very narrow, not spreading. Outer lip thin, not thickened externally. Siphonal canal short; siphonal notch shallow. Material. 1 spm (stn 1: CPAG.RAN.I.33, cast MNHN.F.A50371); 3 spm (stn 2: 1 spm CPAG.RAN.I.31, cast MNHN.F.A50369; 2 spm MNHN); 1 spm (stn 3: CPAG.RAN.I.32, cast MNHN.F.A50370); 3 spm (Jhirak, Lakhra Formation; Cossmann coll., MNHN.F.J12748, MNHN.F.J ). Comments. Athleta (Volutocorbis) burtoni was originally described by Vredenburg (1923) from the Lakhra Formation. Until now, it was recorded only from the Lakhra formation, but several specimens have been collected in marine sediments from the uppermost Bara Formation (see Geological setting). According to Vredenburg (1923), A. (V.) burtoni can be distinguished from A. (V.) eugeniae. Unfortunately, this author did not give diagnostic characters distinguishing the two species. He compared A. (V.) burtoni with a Bartonian species from the Paris Basin, A. (V.) digitalinus (Lamarck, 1811) [= Buccinum scabriculum sensu Solander in Brander, 1766 partim, non Linnaeus, 1758], because of its sculpture characterized by the presence of rounded nodules. A. (V.) burtoni differs obviously from A. (V.) eugeniae by its smaller size, its more numerous costae (19 20 instead on the last whorl), only two rows of nodules on the spire (subsutural and shoulder nodules) and in having rounded nodules instead crenulations on the last whorl. From the Lakhra Formation at Jhirak, Vredenburg (1923) described a species rather similar to A. (V.) burtoni, A. (V.) victoriae (Vredenburg, 1923). The type specimen, illustrated as Vredenburg s (1923) pl. 15, fig. 4, is designated here as the lectotype, but it is poorly preserved and it seems only wider than specimens of A. (V.) burtoni. Thus, it is difficult to evaluate whether it corresponds to a variation of A. (V.) burtoni or a distinct species. Cox (1930) described a comparable species, A. (V.) daviesi (Cox, 1930) from the Paleocene of the Hangu Formation. The rounded aspect of the nodules of A. (V.) daviesi recalls the European Bartonian species A. (V.) digitalinus, but A. (V.) daviesi differs from A. (V.) burtoni by having two abapical spiral rows of nodules below the shoulder row. A. (V.) soriensis (Eames, 1952) from the Early Eocene of the Zinda Pir section (Western Punjab, Pakistan; Eames 1951b) displays numerous similarities to A. (V.) burtoni, such as rounded nodules, only two rows of nodules (the subsutural and shoulder rows), around costae, small size and a biconic shape. In view of these similarities, we consider that A. (V.) burtoni and A. (V.) soriensis could belong to the same species. Stratigraphic range. Bara Formation (this paper); Lakhra Formation (Cossmann & Pissarro 1909; Vredenburg 1923);? Ghazij Formation (Punjab). Athleta (Volutocorbis) cf. wynnei Cox, 1930 (Fig. 9H L) Athleta (Volutospina) wynnei Cox, 1930: 199, pl. 31, figs 7 8, 11. Description. Shell biconic, ventricose, H ca. 40 mm (spire broken in adult specimen), D 25 mm. Protoconch and first teleoconch whorl unknown; at least 5.5 teleoconch whorls. Spire apparently moderately high. Spire whorls convex, last whorl rather inflated. Spiral sculpture on spire of four or five rows of small nodules, subsutural row better developed than others (Fig. 9J). On last whorl, spiral sculpture displaying: 1) subsutural row of rounded nodules; 2) shoulder row of rounded nodules; 3) from shoulder to base, around 20 almost flat cords separated by thin spiral grooves. From second to penultimate whorl, costae interrupted by rows of nodules. On last whorl, costae obsolete between subsutural and shoulder nodules, thin but relatively low below shoulder angle, extending across base, becoming almost obsolete towards close of last whorl. Second whorl: 15 costae; third whorl: 15 costae; fourth whorl: costae; fifth and last whorl: 19 costae. Aperture ovate, rather wide. Inner lip straight, columella with two folds succeeded posteriorly by two or three smaller, weaker folds; inner lip callus narrow, apparently not spreading anteriorly. Outer lip thin, not thickened externally. Siphonal canal slightly elongate; siphonal notch shallow. Material. 4 spm (stn 4: 3 spm CPAG.RAN.I.34 36, cast MNHN.F.A and 1 spm MNHN). 22 Zootaxa 0000 (0) 2014 Magnolia Press MERLE ET AL.

23 FIGURE 9. A G. Athleta (Volutocorbis) burtoni Vredenburg, 1923, A (CPAG.RAN.I.31, dorsal view, stn 2), B (CPAG.RAN.I.32, ventral view, stn 3), C (CPAG.RAN.I.33, dorsal view, stn 1), D F (MNHN.F.J13433, Cossmann coll., ventral (D) and dorsal (E) views, enlarged view of the spire (F), Jhirak), G (MNHN.F.J13434, Cossmann coll., dorsal view, Jhirak); H L. Athleta (Volutocorbis) cf. wynnei Cox, 1930, H I (CPAG.RAN.I.35, dorsal view (H) and enlarged view of the spire (I), stn 4), J (CPAG.RAN.I.36, ventral view, stn 4), K L (CPAG.RAN.I.34, ventral (K) and dorsal (L) views). Scale bars: A E, G H, J L = 10 mm; F, I = 5 mm. EARLY EOCENE VOLUTIDS FROM PAKISTAN Zootaxa 0000 (0) 2014 Magnolia Press 23

24 Comments. The material collected from the Lakhra Dome includes three juvenile specimens and a broken adult. The juvenile specimens can be compared with a paratype of Athleta (Volutocorbis) wynnei Cox, 1930 (GSI no. 14,700) from the Hangu Formation, which is also a juvenile. With these young specimens of A. (V.) wynnei, the Sindh specimens share close similarities, particularly the elongate shape of the spire, the sculpture on the spire (with four to five rows of small nodules) and the deep crenulations of costae on the last whorl. The adult specimens of A. (V.) wynnei, as for example the holotype (GSI no. 14,698), apparently differ by their more crenulated costae on the last whorl, and by their less ventricose shape. Because of these small differences, the present identification of A. (V.) wynnei from the Lakhra Formation should be confirmed by the examination of more material. Stratigraphic range. Lakhra Formation: Lakhra Dome. Athleta (Volutocorbis) lasharii sp. nov. (Fig. 10A D) Etymology. Dedicated to Doctor Rafique Ahmed Lashari, Centre for Pure and Applied Geology, University of Sindh, Jamshoro, Pakistan. Type locality. Stn 4: Lakhra Dome, Lakhra village section, base of the Lakhra Formation. Type material. Holotype (stn 4: CPAG.RAN.I.37, cast MNHN.F.A50375), one paratype (stn 4: CPAG.RAN.I.38, cast MNHN.F.A50376). Other material. 2 spm (stn 4: MNHN). Description. Shell biconic, slightly fusiform, H 40 48, D ca mm (holotype H 43.5, D 19.6 mm, outer lip not complete. Protoconch and two first teleoconch whorls unknown; at least 5 teleoconch whorls. Spire moderately high, occupying 20% of total shell height. Spire whorls weakly convex, last whorl rather oval. Spiral sculpture on spire of a first row of rounded subsutural nodules, appearing on third whorl and becoming more developed aperturally; abapically, three rows of rounded nodules including shoulder row, which is less distinct than others (Fig. 10C). On last whorl, spiral sculpture displaying: 1) subsutural row of spiny nodules; 2) shoulder row of spiny nodules; 3) between shoulder and mid-whorl, 9 to 10 rows of almost flat cords, separated by narrow spiral grooves; 4) from mid-whorl to base, nodules and cords produce imbricate effect. From second to penultimate whorl, costae interrupted by rows of nodules. On last whorl, costae obsolete between subsutural and shoulder nodules, narrow but relatively high below shoulder angle, extending across base, becoming almost obsolete towards end of last whorl. Second whorl: 12 costae; third whorl: 14 costae; fourth whorl: costae; fifth and last whorl: 9 13 costae. Aperture ovate, rather narrow, occupying 72% of total height, 61% of diameter. Inner lip straight, columella with one fold, succeeded posteriorly by several weaker and smaller folds; inner lip callus very narrow, spreading a little anteriorly. Outer lip thin, not thickened externally. Siphonal canal slightly elongate; siphonal notch shallow. Comparisons. Athleta (Volutocorbis) lasharii sp. nov. differs from the other Sindh species of A. (Volutocorbis) in its larger size, its spiral sculpture on the last whorl characterized by the loss of crenulations or nodules between the shoulder and the mid-whorl and by its axial sculpture of higher, narrow costae. Such sculpture distinguishes it from the A. (V.) digitalinus-type species bearing strongly nodular sculpture on the last whorl (e.g. A. (V.) burtoni, A. (V.) daviesi and A. (V.) soriensis from the Early Paleogene of Pakistan) and the A. (V.) elevatus-type species bearing crenulate sculpture (e.g. A. (V.) eugeniae and A. (V.) wynnei from the Early Paleogene of Pakistan). The type of sculpture displayed in A. (V.) lasharii sp. nov. more resembles that of A. (V.) bicorona (Lamarck, 1803) from the Lutetian of the Paris Basin. A. (V.) bicorona shares a last whorl characterized by the loss of crenulations or nodules between the shoulder and the mid-whorl. Moreover, both species have two rows of spiny nodules developed adapically (the subsutural and shoulder rows). A (V.) bicorona differs, however, from A. (V.) lasharii sp. nov. by its early whorls having two more abapical spiral rows of crenulations. In A. (V.) bicorona, a residual color pattern observed under UV light has been reported (Merle et al. 2008), but the recrystallized shells of A. (V.) lasharii did not reveal any traces of a residual color pattern. Stratigraphic range. Lakhra Formation: Lakhra Dome. 24 Zootaxa 0000 (0) 2014 Magnolia Press MERLE ET AL.

25 FIGURE 10. A D. Athleta (Volutocorbis) lasharii sp. nov., A C (holotype, CPAG.RAN.I.37, dorsal (A) and ventral (B) views, enlarged view of the spire (C), stn 4), D (paratype, CPAG.RAN.I.38, dorsal view, stn 4); E H. Volutilithes welcommei sp. nov., E (paratype 1, CPAG.RAN.I.40, dorsal view, stn 4), F (paratype 2, CPAG.RAN.I.41, juvenile, dorsal view, stn 4), G H (holotype, CPAG.RAN.I.39, ventral (G) and (H) dorsal views. Scale bars: A B, D E, G H = 10 mm; C, F = 5 mm. EARLY EOCENE VOLUTIDS FROM PAKISTAN Zootaxa 0000 (0) 2014 Magnolia Press 25

26 Subfamily Volutilithinae Pilsbry & Olsson, 1954 Genus Volutilithes Swainson, 1831 Type species. Voluta muricina Lamarck, 1803 (Middle Eocene, Paris Basin, France) designated here (Art , ICZN 1999). Comments. Swainson (1831) discussed Voluta muricina Lamarck, 1803, for which he introduced the new genus Volutilithes. His description, the cross-reference to the Tableau encyclopédique (Lamarck 1798, pl. 383, fig. 1) [in which V. muricina is illustrated], his own figure 1 entitled muricina and the comparison with Volutilithes pertusa Swainson, 1831 indicate unambiguously that this author considered V. muricina to be the type species of Volutilithes. Unfortunately, Swainson (1831) designated Voluta musicalis Lamarck, 1803 as the type species (by original designation) of Volutilithes. Voluta musicalis Lamarck, 1803 [also the type species of Pseudaulicina Chavan in Furon & Kouriatchy, 1948] is morphologically very different from the two species (V. muricina and V. pertusa) illustrated by Swainson (1831) to represent his new genus Volutilithes, and Swainson s designation appeared to subsequent authors to be an error (Dall 1906; Wenz 1943; Korobkov 1955). Consequently, the type species was originally misidentified. In addition, Swainson (1840: 318, fig. 81e) increased this confusion by designating Conus spinosus Linnaeus, 1758 (erroneously) as the type species of Volutilithes. Later, Newton (1906) introduced Volutospina for Conus spinosus Linnaeus, 1758 considering correctly that the secondary designation by Swainson (1840) was not correct. On the other hand, Dall (1906: 143), Wenz (1943: 1328), and Korobkov (1955: 310) did not respect the original designation of Swainson (1831) in subsequently designating Voluta muricina as the type species of Volutilithes and this designation cannot be considered to be valid, even if it seems scientifically logical. Following article (ICZN 1999) concerning the case of misidentified type species, we select as type species the species that will, in our judgment, best serve stability and universality. Therefore, we designate here Voluta muricina Lamarck, 1803 as the type species of Volutilithes, because this species, discussed and illustrated by Swainson (1831: pl. 53, fig. 1) was misidentified as Voluta musicalis Lamarck, 1803 in the original designation. Eopsephaea Fischer, 1883 (type species: Voluta muricina Lamarck, 1803 by monotypy) becomes a junior objective synonym of Volutilithes. Volutilithes welcommei sp. nov. (Fig. 10E H) Etymology. Dedicated to Jean-Lou Welcomme. Type locality. Stn 4: Lakhra Dome, Lakhra village section, base of the Lakhra Formation. Type material. Holotype (stn 4: CPAG.RAN.I.39, cast MNHN.F.A50377), paratype 1 (stn 4: CPAG.RAN.I.40, cast MNHN.F.A50378), paratype 2 (stn 4: CPAG.RAN.I.41, cast MNHN.F.A50379). Other material. 4 spm (stn 4: MNHN). Description. Shell biconic, H 38 39, D mm (holotype H 38.5, D 20.1 mm. Protoconch smooth, rounded, bulbous, of 1¾ whorls (Fig. 10F). Transition protoconch/teleoconch not defined. Teleoconch of 5 whorls. Spire moderately high, occupying 24% of total shell height. Two first whorls spire whorls convex, later whorls subcarinate. Last whorl narrow, rather conic. Suture linear, undulating between bases of costae. Axial sculpture of strong but narrow, rounded costae, not corresponding from whorl to whorl. Costae slightly orthocline, extending from suture to suture on spire, angulate at shoulder, extending to base of last whorl. First whorl: 12 costae; second whorl: costae; third whorl: 9 10 costae; fourth whorl: costae; fifth and last whorls: 9 12 costae. No spiral sculpture on spire. Spiral sculpture of fine threads on base of last whorl. Aperture lenticular, occupying 67% of total height, 29% of diameter. No posterior notch. Outer lip slightly thickened externally. Inner lip almost straight anteriorly, sinuous posteriorly. Parietal callus slightly developed, spreading posteriorly. Five oblique columellar folds, the most posterior being weaker. Siphonal canal short, slightly curved dorsally; siphonal fasciole low, weak; siphonal notch very shallow. Comparisons. From the Lakhra Formation, this species can be compared to Volutilithes jhirakensis Vredenburg, 1923 [V. jhirakensis Cossmann & Pissarro, 1909 is a nomen nudum]. V. welcommei is distinguished from V. jhirakensis by its wider shape and lower spire. The costae of V. jhirakensis are elongate, whereas they are 26 Zootaxa 0000 (0) 2014 Magnolia Press MERLE ET AL.

27 angulate in V. welcommei. By its elongate shape, V. jhirakensis recalls the Lutetian species from the Paris Basin, V. torulosus (Deshayes, 1835), whereas the low-spired shape and the angulate costae of V. welcommei are more similar to those of V. torreyensis Givens, 1978 from the early Middle Eocene of California. Stratigraphic range. Lakhra Formation: Lakhra Dome. Volutilithes sindhiensis sp. nov. (Fig. 11A E) Etymology. From Sindh, Pakistan province. Type locality. Stn 4: Lakhra Dome, Lakhra village section, base of the Lakhra Formation. Type material. Holotype (stn 4: CPAG.RAN.I.42, cast MNHN.F.A50380), paratype 1 (stn 4: CPAG.RAN.I.43, cast MNHN.F.A50381), paratype 2 (stn 4: CPAG.RAN.I.44, cast MNHN.F.A50382). Other material. 2 spm (stn 4: MNHN). Description. Shell biconic, H ca. 36, D 19 mm (holotype, H not complete 30.6, D 18.8 mm. Protoconch not preserved. Teleoconch of 5.5 whorls. Spire moderately high, occupying 20% of total shell height. Spire whorls shouldered, carinate. Last whorl conic. Suture linear, undulating between bases of costae. Axial sculpture of strong costae, not corresponding from whorl to whorl. Costae rather orthocline, extending from suture to suture on spire, spiny on the shoulder, extending to base on last whorl. First whorl: sculpture not preserved; second whorl: 12 costae; third whorl: 10 costae; fourth whorl: 11 costae; fifth and last whorls: costae. No spiral sculpture. Aperture lenticular, angulate posteriorly, occupying 62% of total height, 35% of diameter. No posterior notch. Outer lip of aperture slightly thickened externally. Inner lip almost straight anteriorly, sinuous posteriorly. Parietal callus poorly developed, not spreading posteriorly. Three oblique columellar folds observed, but probably more present in complete specimens. Base of siphonal canal not preserved. Comparisons. This third species of Volutilithes from the Lakhra Formation shares similarities in its dimensions with V. welcommei, but it is easily distinguishable by its spiny costae. A young specimen shows that spines on the costae appear very early in ontogeny (second whorl). In comparison, the costae of V. welcommei become angulate only over the two last whorls. The biconic shape and the spiny costae suggest resemblances to species of A. (Volutospina), but most species of A. (Volutospina) exhibit several rows of nodules on their early teleoconch whorls (subsutural and shoulder rows and two abapical rows), which are missing in V. welcommei. Stratigraphic range. Lakhra Formation: Lakhra Dome. Genus Pseudaulicina Chavan in Furon & Kouriatchy, 1948 Type species. Voluta musicalis Lamarck, 1803 (Middle Eocene, Paris Basin, France) by original designation. Pseudaulicina coxi sp. nov. (Fig. 11F I) Etymology. Dedicated to L. R. Cox for his work on the Hangu Formation. Type locality. Stn 4: Lakhra Dome, Lakhra village section, base of the Lakhra Formation. Type material. Holotype (stn 4: CPAG.RAN.I.45, cast MNHN.F.A50383), paratype 1 (stn 4: CPAG.RAN.I.46, cast MNHN.F.A50384), paratype 2 (stn 4: CPAG.RAN.I.47, cast MNHN.F.A50385). Other material. 5 spm (stn 4: MNHN). Description. Shell biconic, H probably 60 mm, to judge from largest fragment (holotype H 32.8 not complete, D 20.7 mm). Largest and most complete specimen: H 40, D 22 mm. Protoconch not preserved. Teleoconch of probably 6 7 whorls. Spire relatively high, occupying 32% of total shell height. Spire whorls shouldered, with low carina formed by shoulder spines. Last whorl moderately wide, excavated at beginning of siphonal canal. Suture linear, with weak undulation between bases of costae. Axial sculpture of strong but rounded costae. Costae slightly orthocline, subvertical on early whorls, with acute shoulder spines increasing in prominence until last whorl. On last whorl, costae obsolete from suture to shoulder spine, thick and relatively high below shoulder angle, extending EARLY EOCENE VOLUTIDS FROM PAKISTAN Zootaxa 0000 (0) 2014 Magnolia Press 27

28 across base, becoming almost obsolete towards end of last whorl. First whorl: not preserved; second and third whorls: 10 costae; fourth and fifth whorls: 9 10 costae; sixth whorl: not preserved. No apparent spiral sculpture. Aperture narrow, acutely angular posteriorly, occupying 55% of total height, 30% of diameter. No posterior notch. Outer lip slightly thickened externally. Inner lip sinuous posteriorly (base not preserved). Parietal callus thin, not spreading posteriorly. Four strong, oblique columellar folds, anterior-most weaker. Siphonal canal not preserved. Comparisons. Because of its proportions, its axial sculpture on the early teleoconch whorls and the aspect of its columella, this species is very close to Pseudaulicina vredenburgi (Cox, 1930) from the Hangu Formation. The specimens described by Cox (1930) do not exceed 40 mm in height, like most of the complete specimens collected in the Lakhra Dome, which are probably juveniles. Cox (1930: 197) stated that the spiral sculpture of P. vredenburgi consists of numerous closely and rather irregularly spaced striae, covering the whole surface, and extending anteriorly well down the stem of the shell. This sculpture is missing in the specimens of P. coxi, which have a smooth surface. According to Cox (1930), P. vredenburgi can be compared to P. mitrata (Deshayes, 1835) from the Lutetian of the Paris Basin and to P. wateleti (Deshayes, 1865) from the Ypresian of the Paris and Aquitaine basins. P. mitrata is probably a variant of the type species of Pseudaulicina, P. musicalis (Lamarck, 1803), bearing a more elongate spire and stronger spiral cords than the more common specimens. P. musicalis differs from P. vredenburgi and P. coxi by having spiral cords and by its angulate shoulder spines. In addition, large specimens of P. musicalis display one or two posterior folds, which are missing in the two Pakistani species. P. wateleti shares with P. vredenburgi and P. coxi acute shoulder spines and four strong columellar folds. The surface of P. wateleti seems smooth as in P. coxi, but a careful examination of the last whorl of several specimens revealed low spiral cords as in P. musicalis. Although these cords are obsolete, they differ from the fine threads of P. vredenburgi. Stratigraphic range. Lakhra formation: Lakhra Dome. Genus Sindhiluta gen. nov. Type species. Sindhiluta lakhraensis nov. sp. Early Eocene, Lakhra Formation (Sindh, Pakistan). Etymology. From Sindh (province of Pakistan) and luta, the two last syllables of Voluta. Gender feminine. Diagnosis. Shape elongate, high-spired. First teleoconch whorls subcarinate; last whorl smooth, poorly terraced at shoulder, not excavated anteriorly. Costae on spire rounded, and orthocline, fading out progressively over last whorl. Outer lip not thickened externally. Four strong, oblique columellar folds. Parietal callus thick. Included species. Sindhiluta lakhraensis sp. nov. (Lakhra Formation, Ypresian, Sindh Province, Pakistan) and Voluta prevosti Rouault, 1850 [= Psephaea (Eopsephaea) atacica Doncieux, 1908; Psephaea (Eopsephaea) rabetensis Doncieux, 1908; Volutilithes ogormani Cossmann, 1923] (Ypresian, Aquitaine Basin, France). Discussion. Pilsbry and Olsson (1954) when introducing the new subfamily Volutilithinae included two genera: Volutilithes and Lapparia Conrad, 1855 (type species: Mitra dumosa Conrad, 1854, Middle Eocene, USA). Subsequent authors such as Pacaud & Le Renard (1996) and Merle et al. (2008) also included Pseudaulicina in this subfamily. Sindhiluta shares characters of the axial sculpture with Volutilithes and columellar characters with Pseudaulicina and Lapparia and therefore is assigned to the Volutilithinae. Rounded and subcarinate costae and a lack of spiral sculpture are characters usually found in species of Volutilithes, such as: V. angustus (Deshayes, 1835) from the Ypresian of the Paris Basin, V. deshayesianus (Rouault, 1850) from the Ypresian of the Aquitaine Basin, and V. torulosus (Deshayes, 1835) and V. costarius (Lamarck, 1803) from the Lutetian of the Paris Basin. However, Sindhiluta differs obviously from Volutilithes in two characters. First, Sindhiluta lacks costae on its last whorls, whereas they are present on Volutilithes. Second, Sindhiluta has four strong columellar folds, whereas members of Volutilithes have only two or three weaker columellar folds. The number and development of the columellar folds are more nearly similar to those of Pseudaulicina than to Volutilithes. Nevertheless, the persistence of the costae on the last whorls, associated with variably developed shoulder spines, an excavated base and the presence of spiral sculpture in most species of Pseudaulicina contrast greatly with the morphology of Sindhiluta. As in Sindhiluta, Lapparia has four strong columellar folds, the most anterior being weaker, but it is distinguished by its axial sculpture, which continues to persist down the shell and finishes with the appearance of acute shoulder spines. 28 Zootaxa 0000 (0) 2014 Magnolia Press MERLE ET AL.

29 FIGURE 11. A E. Volutilithes sindhiensis sp. nov., A B (holotype, CPAG.RAN.I.42, dorsal (A) and ventral (B) views, stn 4), C D (paratype 1, CPAG.RAN.I.43, ventral (C) and dorsal (D) views, stn 4), E (paratype 2, CPAG.RAN.I.44, juvenile, ventral view, stn 4); F I. Pseudaulicina coxi sp. nov., F G (holotype, CPAG.RAN.I.45, ventral (F) and dorsal (G) views, stn 4), H (paratype 2, CPAG.RAN.I.47, juvenile, dorsal view, stn 4), I (paratype 1, CPAG.RAN.I.46, ventral view, stn 4). Scale bars: A D, F G, I = 10 mm; E, H = 5 mm. EARLY EOCENE VOLUTIDS FROM PAKISTAN Zootaxa 0000 (0) 2014 Magnolia Press 29

30 Sindhiluta lakhraensis sp. nov. (Fig. 12A F) Etymology. From the type locality (Lakhra village). Type locality. Stn 4: Lakhra Dome, Lakhra village section, base of the Lakhra Formation. Type material. Holotype (stn 4: CPAG.RAN.I.48, cast MNHN.F.A50386), paratype 1 (stn 4: CPAG.RAN.I.49, cast MNHN.F.A50387), paratype 2 (stn 4: CPAG.RAN.I.50, cast MNHN.F.A50388). Other material. 4 spm (stn 4: MNHN). Description. Shell elongate, H 56 58, D mm (holotype H 53.6 not complete, D 31.9 mm). Protoconch not preserved. Teleoconch of 7 whorls. Spire relatively high, occupying 31% of total shell height. Spiral whorls convex to subcarinate. Last whorl narrow, convex posteriorly, not excavated anteriorly. Suture linear with weak undulation between bases of costae. Axial sculpture of strong but rounded costae on spire. Costae orthocline, subcarinate at shoulder, not spinous. On penultimate whorl, costae decreasing in prominence, becoming obsolete, disappearing on last whorl. First whorl: not preserved; second whorl: 11 costae; third whorl: 10 costae; fourth and fifth whorls: 9 10 costae; sixth whorl: 8 9 costae, becoming obsolete; seventh whorl: no costae. No apparent spiral sculpture. Aperture narrow, lenticular, occupying 63% of total height, 26% of diameter. Outer lip thin, not thickened externally. Parietal callus thick posteriorly. Four strong, oblique columellar folds, anterior-most weaker. Siphonal canal not preserved. Comparisons. From the Ypresian of the Aquitaine Basin, Voluta prevosti Rouault, 1850 [Mitra prevosti Rouault, 1848 is a nomen nudum] is the single Paleogene volutid that shares rounded, subcarinate costae disappearing progressively at the end of growth with Sindhiluta. In addition, as in S. lakhraensis sp. nov., the last whorls of V. prevosti are rounded and not shouldered posteriorly, its base is not excavated, its outer lip is not thickened and a parietal callus is present. Regarding the columellar folds, their numbers vary between three and six, but their construction is rather similar to that of S. lakhraensis sp. nov. The most anterior fold is present in young specimens, but is weaker than the three adapical folds or tends to disappear in adult specimens. In S. lakhraensis sp. nov., this anterior fold is also weaker even if it persists at the end of growth. Adapically, the columella of V. prevosti displays three strong folds as in S. lakhraensis. In its posterior part, the columella displays two weak folds in a few specimens, but they are missing in S. lakhraensis. Considering the large number of shared characters with the Pakistani species, we do not hesitate to assign Voluta prevosti to the genus Sindhiluta. On biogeographic grounds, it seems unlikely that these species, located at two extremities of Tethyan Ocean, belong to a same genus. However, the both extremities of the Tethyan Ocean share numerous common genera and even some common species (Pacaud 1997). In addition, examples of range disjunction in tropical marine area are also known (Lozouet et al. 1994). Stratigraphic range. Lakhra formation: Lakhra Dome. Incertae sedis Subfamily Volutodermatinae Pilsbry and Olsson, 1954 Comments. Sohl (1964) followed by Bouchet et al. (2005: 255) emended the subfamily name Volutoderminae as originally proposed by Pilsbry and Olsson (1954) to Volutodermatinae, see also Art. 29.3, ICZN (1999). Genus Pakiluta gen. nov. Type species. Pakiluta solangii nov. sp. Early Eocene, Lakhra Formation (Sindh, Pakistan). Etymology. From Paki, the two first syllables of Pakistan and luta, the two last syllables of Voluta. Gender feminine. Included species. The type species only. Diagnosis. Shell solid, with coeloconoid spire. Five early teleoconch whorls convex and elongate, followed by shorter whorls. Last whorl moderately wide, convex posteriorly, almost straight in its median part, slightly 30 Zootaxa 0000 (0) 2014 Magnolia Press MERLE ET AL.

31 FIGURE 12. A F. Sindhiluta lakhraensis gen. nov., sp. nov., A B (holotype, CPAG.RAN.I.48, ventral (A) and dorsal (B) views, stn 4), C D (paratype 1, CPAG.RAN.I.49, ventral and dorsal view, stn 4), E F (paratype 2, CPAG.RAN.I.50, dorsal view of the broken specimen showing the columellar folds (E) and enlarged view of the spire (F); G M. Pakiluta solangii gen. nov., sp. nov., G H (holotype, CPAG.RAN.I.51, ventral (G) and dorsal (H) views, stn 4), I J (paratype 3, CPAG.RAN.I.54, spire (I), and enlarged view of the spire (J), stn 5), K L (paratype 2, CPAG.RAN.I.53, spire (K) and enlarged view of the spire (L), stn 4), M (paratype 1, CPAG.RAN.I.52, ventral view, stn 4). Scale bars: A I, K, M = 10 mm; J, L = 5 mm. EARLY EOCENE VOLUTIDS FROM PAKISTAN Zootaxa 0000 (0) 2014 Magnolia Press 31