Première mention d’une tortue bothrémydidée, <i>Sindhochelys ragei</i> n. gen., n. sp., du Paléocène inférieur du Pakistan, implications systématiques et paléobiogéographiques

Nous présentons la découverte de restes de grand chélonien provenant de la base du Paléocène inférieur de la Formation Khadro, affleurant dans la région du Fort Ranikot (Groupe Ranikot, Province du Sindh, Sud du Pakistan). Cette formation a déjà livré le serpent Gigantophis Andrews, 1901 étudié par notre ami Jean-Claude Rage. Deux spécimens, une carapace et un autre fragment de tortue Bothremydidae, famille d’origine gondwanienne, sont décrits. Un nouveau genre Sindhochelys ragei n. gen., n. sp. est nommé en l’honneur de Jean-Claude Rage. C’est la première découverte d’un bothrémydidé dans le sud-ouest du Pakistan. Les relations avec les représentants crétacés et paléocènes de la famille sont examinées. L’association de caractères comme la forme de la carapace, le pattern des écailles plastrales antérieures et la décoration très accusée le caractérisent et, malgré des similitudes, exclut toute affinité avec les Taphrosphyini et le groupe indien Carteremys, les autres bothrémydidés bien définis étant aussi exclus. L’autre fragment, également très décoré, reste indéterminé. La découverte de ces deux spécimens dans un environnement littoral du Paléocène inférieur du Pakistan comble une lacune géographique et stratigraphique dans la connaissance des Bothremydidae, connus depuis le Crétacé inférieur continental d’Afrique et s’étant répartis dans le monde jusqu’au Miocène de la Néotéthys. Une importante diversification est reconnue pendant le Maastrichtien et le Paléocène, avec une dispersion le long des côtes néotéthysienne et peut-être à travers cet océan : Sindhochelys ragei n. gen., n. sp. peut avoir colonisé le sous-continent Indien à cette époque, ou résulter d’une ancienne diversification avant la rupture du Gondwana.We report the discovery of remains of a large chelonian from the base of the early Paleocene Khadro Formation exposed in the Ranikot Fort area (Ranikot Group, Sindh Province, Southern Pakistan). This formation already yielded the snake Gigantophis Andrews, 1901, studied by our friend Jean-Claude Rage. The chelonian specimens consist of a large carapace and a shell fragment of Bothremydidae, a family of Gondwanan origin. A new genus and species, Sindhochelys ragei n. gen., n. sp. is identified from the first specimen and named in honor of Jean-Claude Rage. It is the first report of a Bothremydidae in Southern Pakistan. Its affinities with Cretaceous and Paleocene representatives of the family are discussed. The association of characters such as the shape of the shell, anterior plastral scute pattern and strongly marked decoration characterize the taxon and, despite some similarities, allows excluding close phylogenetic affinities with Taphrosphyini and Carteremys group; other well-documented bothremydids are also excluded. The shell fragment, also strongly decorated, is left undetermined. The discovery of two new littoral bothremydid specimens in the early Paleocene of Pakistan fills a geographic and stratigraphic gap in our knowledge of the family, which is known since the continental early Cretaceous of Africa, diversifying in the world up to the Miocene deposits of the Neotethys. A particular diversification during the Maastrichtian-Paleocene is recognized along the neotethyan coasts, and occasional dispersals across this ocean were possible. Sindhochelys ragei n. gen., n. sp. may have colonized the Indian subcontinent by this time, or may represent an older diversification before the Gondwana breakup.</p

Volutilithes welcommei Merle, Pacaud, Métais, Bartolini, Lashari, Brohi, Solangi, Marivaux & Welcomme, 2014, sp. nov.

Volutilithes welcommei sp. nov. (Fig. 10 E–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. A 50377), paratype 1 (stn 4: CPAG.RAN. I. 40, cast MNHN.F. A 50378), paratype 2 (stn 4: CPAG.RAN. I. 41, cast MNHN.F. A 50379). Other material. 4 spm (stn 4: MNHN). Description. Shell biconic, H 38–39, D 19–20 mm (holotype H 38.5, D 20.1 mm. Protoconch smooth, rounded, bulbous, of 1 ¾ whorls (Fig. 10 F). 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: 10–11 costae; third whorl: 9–10 costae; fourth whorl: 10–11 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. j h i r a ke n s i s by its wider shape and lower spire. The costae of V. jhirakensis are elongate, whereas they are 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.Published as part of Merle, Didier, Pacaud, Jean-Michel, Métais, Grégoire, Bartolini, Annachiara, Lashari, Rafiq A., Brohi, Imdad A., Solangi, Sarfraz H., Marivaux, Laurent & Welcomme, Jean-Loup, 2014, Volutidae (Mollusca: Gastropoda) of the Lakhra Formation (Earliest Eocene, Sindh, Pakistan): systematics, biostratigraphy and paleobiogeography, pp. 101-138 in Zootaxa 3826 (1) on pages 126-127, DOI: 10.11646/zootaxa.3826.1.3, http://zenodo.org/record/22853

Lyriopsis Merle, Pacaud, Métais, Bartolini, Lashari, Brohi, Solangi, Marivaux & Welcomme, 2014, gen. nov.

Genus &lt;i&gt;Lyriopsis&lt;/i&gt; gen. nov. &lt;p&gt; &lt;b&gt;Type species.&lt;/b&gt; &lt;i&gt;Lyria cossmanni&lt;/i&gt; Vredenburg, 1923. Early Eocene from Lakhra Formation (Sindh, Pakistan).&lt;/p&gt; &lt;p&gt; &lt;b&gt;Etymology.&lt;/b&gt; Combination of &lt;i&gt;Lyria&lt;/i&gt; with the suffix &lt;i&gt;opsis&lt;/i&gt; (resembling in appearance). Gender feminine.&lt;/p&gt; &lt;p&gt; &lt;b&gt;Diagnosis.&lt;/b&gt; A ventricose &lt;i&gt;&ldquo; Lyria &rdquo;&lt;/i&gt; 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.&lt;/p&gt; &lt;p&gt; &lt;b&gt;Included species&lt;/b&gt;. &lt;i&gt;Lyria cossmanni&lt;/i&gt; Vredenburg, 1923 (Lakhra Formation, Sindh Province, Pakistan) and &lt;i&gt;Lyria samanaensis&lt;/i&gt; Cox, 1930 (Hangu Formation, Paleocene, Khyber Pakhtunkhwa, Pakistan).&lt;/p&gt; &lt;p&gt; &lt;b&gt;Discussion.&lt;/b&gt; Cossmann &amp; Pissarro (1909), Vredenburg (1923) and Cox (1930) attributed these two species to the genus &lt;i&gt;Lyria&lt;/i&gt; and compared them to three Eocene species from the Paris Basin: &lt;i&gt;Lyria (Lyria) harpula&lt;/i&gt; (Lamarck, 1803), &lt;i&gt;L. (L.) subturgidula&lt;/i&gt; (d&rsquo;Orbigny, 1850) (= &lt;i&gt;Voluta turgidula&lt;/i&gt; Deshayes, 1835, not of Brocchi, 1814) and &lt;i&gt;L. (Pseudolyria) coroni&lt;/i&gt; (Morlet, 1888). According to these authors, strong but narrowly rounded costae and a biconic shape are shared with the Eocene &lt;i&gt;Lyria&lt;/i&gt; species of the Paris Basin and even with many other Cenozoic species of &lt;i&gt;Lyria&lt;/i&gt;. Thus, &lt;i&gt;Lyriopsis&lt;/i&gt; is placed in the tribe Lyriini, but four characters distinguish members of &lt;i&gt;Lyria&lt;/i&gt; from &lt;i&gt;Lyriopsis&lt;/i&gt;: the morphology of the costae, the construction of the columellar folds, the shape of the whorl, and the protoconch. In &lt;i&gt;L. (Lyria)&lt;/i&gt;, the posterior ends of the costae are straight, whereas they are obviously sinuous in &lt;i&gt;Lyriopsis&lt;/i&gt;. Regarding the subgenus &lt;i&gt;Pseudolyria&lt;/i&gt; Martin, 1931 [type species &lt;i&gt;Pseudolyria ventricola&lt;/i&gt; Martin, 1931 by monotypy], the costae are more sinuous than in &lt;i&gt;L. (Lyria)&lt;/i&gt;, but the sinuosity is more anteriorly placed than in &lt;i&gt;Lyriopsis&lt;/i&gt;. Moreover, the outer lip of &lt;i&gt;L. (Pseudolyria)&lt;/i&gt; is crenulated, whereas it is smooth in &lt;i&gt;Lyriopsis&lt;/i&gt;. In &lt;i&gt;L. (Lyria)&lt;/i&gt;, 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 &lt;i&gt;Lyriopsis&lt;/i&gt;. The shape of the whorl is rather depressed, slightly convex and terraced in &lt;i&gt;Lyriopsis&lt;/i&gt; species, whereas the whorls are strongly convex in most species of &lt;i&gt;Lyria (Lyria)&lt;/i&gt;. For these reasons, the placement of &lt;i&gt;Lyriopsis&lt;/i&gt; in the Lyriini is provisional and uncertain.&lt;/p&gt;Published as part of &lt;i&gt;Merle, Didier, Pacaud, Jean-Michel, Métais, Grégoire, Bartolini, Annachiara, Lashari, Rafiq A., Brohi, Imdad A., Solangi, Sarfraz H., Marivaux, Laurent &amp; Welcomme, Jean-Loup, 2014, Volutidae (Mollusca: Gastropoda) of the Lakhra Formation (Earliest Eocene, Sindh, Pakistan): systematics, biostratigraphy and paleobiogeography, pp. 101-138 in Zootaxa 3826 (1)&lt;/i&gt; on page 108, DOI: 10.11646/zootaxa.3826.1.3, &lt;a href="http://zenodo.org/record/228537"&gt;http://zenodo.org/record/228537&lt;/a&gt

Volutilithes Swainson 1831

Genus Volutilithes Swainson, 1831 Type species. Voluta muricina Lamarck, 1803 (Middle Eocene, Paris Basin, France) designated here (Art. 70.3. 2, 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. m u r i c i na 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. 81 e) 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 70.3. 2 (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.Published as part of Merle, Didier, Pacaud, Jean-Michel, Métais, Grégoire, Bartolini, Annachiara, Lashari, Rafiq A., Brohi, Imdad A., Solangi, Sarfraz H., Marivaux, Laurent & Welcomme, Jean-Loup, 2014, Volutidae (Mollusca: Gastropoda) of the Lakhra Formation (Earliest Eocene, Sindh, Pakistan): systematics, biostratigraphy and paleobiogeography, pp. 101-138 in Zootaxa 3826 (1) on page 126, DOI: 10.11646/zootaxa.3826.1.3, http://zenodo.org/record/22853

Athleta (Volutocorbis) lasharii Merle, Pacaud, Métais, Bartolini, Lashari, Brohi, Solangi, Marivaux & Welcomme, 2014, sp. nov.

Athleta (Volutocorbis) lasharii sp. nov. (Fig. 10 A–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. A 50375), one paratype (stn 4: CPAG.RAN. I. 38, cast MNHN.F. A 50376). Other material. 2 spm (stn 4: MNHN). Description. Shell biconic, slightly fusiform, H 40–48, D ca. 20–22 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. 10 C). 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: 12–14 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.Published as part of Merle, Didier, Pacaud, Jean-Michel, Métais, Grégoire, Bartolini, Annachiara, Lashari, Rafiq A., Brohi, Imdad A., Solangi, Sarfraz H., Marivaux, Laurent & Welcomme, Jean-Loup, 2014, Volutidae (Mollusca: Gastropoda) of the Lakhra Formation (Earliest Eocene, Sindh, Pakistan): systematics, biostratigraphy and paleobiogeography, pp. 101-138 in Zootaxa 3826 (1) on page 124, DOI: 10.11646/zootaxa.3826.1.3, http://zenodo.org/record/22853

Volutospina Newton 1906

Subgenus &lt;i&gt;Volutospina&lt;/i&gt; Newton, 1906 &lt;p&gt; &lt;b&gt;Type species.&lt;/b&gt; &lt;i&gt;Conus spinosus&lt;/i&gt; Linnaeus, 1758 (Middle Eocene, Paris Basin, France) by original designation.&lt;/p&gt;Published as part of &lt;i&gt;Merle, Didier, Pacaud, Jean-Michel, Métais, Grégoire, Bartolini, Annachiara, Lashari, Rafiq A., Brohi, Imdad A., Solangi, Sarfraz H., Marivaux, Laurent &amp; Welcomme, Jean-Loup, 2014, Volutidae (Mollusca: Gastropoda) of the Lakhra Formation (Earliest Eocene, Sindh, Pakistan): systematics, biostratigraphy and paleobiogeography, pp. 101-138 in Zootaxa 3826 (1)&lt;/i&gt; on page 118, DOI: 10.11646/zootaxa.3826.1.3, &lt;a href="http://zenodo.org/record/228537"&gt;http://zenodo.org/record/228537&lt;/a&gt

Athleta (Volutopupa) intercrenatus Cossmann & Pissarro 1909, comb. nov.

&lt;i&gt;Athleta (Volutopupa) intercrenatus&lt;/i&gt; (Cossmann &amp; Pissarro, 1909) comb. nov. &lt;p&gt;(Fig. 7 E&ndash;J)&lt;/p&gt; &lt;p&gt; &lt;i&gt;Volutospina intercrenata&lt;/i&gt; Cossmann &amp; Pissarro, 1909: 24, pl. 2, figs 18&ndash;20, pl. 3, figs 1&ndash;3.&lt;/p&gt; &lt;p&gt; &lt;b&gt;Description.&lt;/b&gt; Shell biconic, H 33&ndash;37, D 18&ndash;23 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 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. 7 J); 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: 12&ndash;13 costae; fifth and last whorl: 11&ndash;12 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&ndash;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.&lt;/p&gt; &lt;p&gt; &lt;b&gt;Material.&lt;/b&gt; 5 spm (stn 4: 4 spm CPAG.RAN. I.21&ndash;24, cast MNHN.F. A50359 &ndash;62 and 1 spm MNHN).&lt;/p&gt; &lt;p&gt; &lt;b&gt;Comments.&lt;/b&gt; As seen above with &lt;i&gt;Athleta (Volutopupa) noetlingi&lt;/i&gt;, the four specimens of &lt;i&gt;V. intercrenata&lt;/i&gt; illustrated by Cossmann &amp; Pissarro (1909) are species of &lt;i&gt;Volutopupa&lt;/i&gt; and not species of &lt;i&gt;Volutospina&lt;/i&gt;. They are not well preserved, except the specimen illustrated in Cossmann &amp; Pissarro&rsquo;s (1909) pl. 3, fig. 3, which is designated here as the lectotype of &lt;i&gt;Volutospina intercrenata&lt;/i&gt; Cossmann &amp; Pissarro, 1909. 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 &lt;i&gt;A. (V.) intercrenatus&lt;/i&gt; collected in the Lakhra Dome. The oval shape of these adults resembles more that of &lt;i&gt;A. (V.) lyra&lt;/i&gt; (Lamarck, 1803) or &lt;i&gt;A. (V.) mutatus&lt;/i&gt; (Deshayes, 1835) from the Middle Eocene of the Paris Basin, than it does &lt;i&gt;A. (V.) citharoedus&lt;/i&gt;, which displays an inflated and ventricose last whorl. &lt;i&gt;A. (V.) intercrenatus&lt;/i&gt; also differs from these species by its strongly developed row of subsutural nodules.&lt;/p&gt; &lt;p&gt; &lt;b&gt;Stratigraphic range.&lt;/b&gt; Lakhra Formation: Jhirak (Cossmann &amp; Pissarro 1909) and Lakhra Dome (this paper).&lt;/p&gt;Published as part of &lt;i&gt;Merle, Didier, Pacaud, Jean-Michel, Métais, Grégoire, Bartolini, Annachiara, Lashari, Rafiq A., Brohi, Imdad A., Solangi, Sarfraz H., Marivaux, Laurent &amp; Welcomme, Jean-Loup, 2014, Volutidae (Mollusca: Gastropoda) of the Lakhra Formation (Earliest Eocene, Sindh, Pakistan): systematics, biostratigraphy and paleobiogeography, pp. 101-138 in Zootaxa 3826 (1)&lt;/i&gt; on pages 118-119, DOI: 10.11646/zootaxa.3826.1.3, &lt;a href="http://zenodo.org/record/228537"&gt;http://zenodo.org/record/228537&lt;/a&gt

Pseudaulicina coxi Merle, Pacaud, Métais, Bartolini, Lashari, Brohi, Solangi, Marivaux & Welcomme, 2014, sp. nov.

Pseudaulicina coxi sp. nov. (Fig. 11 F–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. A 50383), paratype 1 (stn 4: CPAG.RAN. I. 46, cast MNHN.F. A 50384), paratype 2 (stn 4: CPAG.RAN. I. 47, cast MNHN.F. A 50385). 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 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.Published as part of Merle, Didier, Pacaud, Jean-Michel, Métais, Grégoire, Bartolini, Annachiara, Lashari, Rafiq A., Brohi, Imdad A., Solangi, Sarfraz H., Marivaux, Laurent & Welcomme, Jean-Loup, 2014, Volutidae (Mollusca: Gastropoda) of the Lakhra Formation (Earliest Eocene, Sindh, Pakistan): systematics, biostratigraphy and paleobiogeography, pp. 101-138 in Zootaxa 3826 (1) on pages 127-128, DOI: 10.11646/zootaxa.3826.1.3, http://zenodo.org/record/22853

Lyrischapa haimei d'Archiac & Haime 1853

&lt;i&gt;Lyrischapa haimei&lt;/i&gt; (d&rsquo;Archiac &amp; Haime, 1853) &lt;p&gt;(Fig. 4 L)&lt;/p&gt; &lt;p&gt; &lt;i&gt;Voluta haimei&lt;/i&gt; d&rsquo;Archiac &amp; Haime, 1853: 325, p1. 31, figs. 26, 27.&lt;/p&gt; &lt;p&gt; &lt;i&gt;Voluta haimei&lt;/i&gt; d&rsquo;Archiac, 1850: 298 (nomen nudum).&lt;/p&gt; &lt;p&gt; &lt;i&gt;Aulicina haimei&lt;/i&gt;. Cossmann &amp; Pissarro 1909: 26, pl. 2, fig. 29 only [new combination]. &lt;i&gt;Volutoconus funiculifer&lt;/i&gt; Cossmann &amp; Pissarro, 1909: 27, pl. 3, figs 10&ndash;12.&lt;/p&gt; &lt;p&gt; &lt;i&gt;Aulica (Aulicina) haimei&lt;/i&gt;. Vredenburg 1923: 269; Cotter &lt;i&gt;in&lt;/i&gt; Vredenburg 1928: 38 [new combination]. &lt;i&gt;Eovasum haimei&lt;/i&gt;. Cox 1930: 186 in part (not pl. 21, figs 1&ndash;4), see (Givens 1991) [new combination]; Davies 1934: 297. &lt;i&gt;Lyrischapa haimei&lt;/i&gt;. Givens 1991: 664, figs 3.6&ndash;3.8 [new combination].&lt;/p&gt; &lt;p&gt; &lt;b&gt;Material.&lt;/b&gt; 1 spm (stn 6: CPAG.RAN. I.8, cast MNHN.F.A50346).&lt;/p&gt; &lt;p&gt; &lt;b&gt;Comments.&lt;/b&gt; In his paper on &lt;i&gt;Lyrischapa&lt;/i&gt;, Givens (1991) gave a revised description of &lt;i&gt;L. haimei&lt;/i&gt;. He recognized two differential characters distinguishing it from &lt;i&gt;L. sismondai&lt;/i&gt; (d&rsquo;Archiac &amp; 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 &lt;i&gt;L. sismondai&lt;/i&gt; is larger than that of &lt;i&gt;L. haimei&lt;/i&gt;. From the middle part of Lakhra Formation at Jhirak, we collected a juvenile specimen of H 22 mm (Fig. 4 L). Although it is a little crushed, it displays these three distinguishing characters of &lt;i&gt;L. haimei&lt;/i&gt;.&lt;/p&gt; &lt;p&gt; &lt;b&gt;Stratigraphic range.&lt;/b&gt; Lakhra Formation: Jhirak (Cossmann &amp; Pissarro 1909; Vredenburg 1923; this paper).&lt;/p&gt;Published as part of &lt;i&gt;Merle, Didier, Pacaud, Jean-Michel, Métais, Grégoire, Bartolini, Annachiara, Lashari, Rafiq A., Brohi, Imdad A., Solangi, Sarfraz H., Marivaux, Laurent &amp; Welcomme, Jean-Loup, 2014, Volutidae (Mollusca: Gastropoda) of the Lakhra Formation (Earliest Eocene, Sindh, Pakistan): systematics, biostratigraphy and paleobiogeography, pp. 101-138 in Zootaxa 3826 (1)&lt;/i&gt; on pages 111-112, DOI: 10.11646/zootaxa.3826.1.3, &lt;a href="http://zenodo.org/record/228537"&gt;http://zenodo.org/record/228537&lt;/a&gt

Athleta (Volutocorbis) burtoni Vredenburg 1923

&lt;i&gt;Athleta (Volutocorbis) burtoni&lt;/i&gt; Vredenburg, 1923 &lt;p&gt;(Fig. 9A&ndash;G)&lt;/p&gt; &lt;p&gt; &lt;i&gt;Athleta (Volutocorbis) burtoni&lt;/i&gt; Vredenburg, 1923: 261, pl. 15, fig. 2. &lt;i&gt;Athleta (Volutocorbis) victoriae&lt;/i&gt; Vredenburg, 1923: 260, pl. 15, figs 4. &lt;i&gt;Athleta (Volutocorbis) burtoni.&lt;/i&gt; Cotter &lt;i&gt;in&lt;/i&gt; Vredenburg 1928: 37. &lt;i&gt;Volutocorbis soriensis&lt;/i&gt; Eames, 1952: 109, pl. 4, fig. 95a&ndash;b.&lt;/p&gt; &lt;p&gt; &lt;i&gt;Volutocorbis burtoni&lt;/i&gt;. Glibert 1960: 48 [new combination].&lt;/p&gt; &lt;p&gt; &lt;b&gt;Description.&lt;/b&gt; 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, 13&ndash;15 rows of almost rounded nodules, becoming less prominent on base. From second to last whorls, costae interrupted by rows of nodules. On last whorl, costae rather low, cut into prominent small, regular nodules by broad spiral grooves. Third whorl: 15&ndash;16 costae; fourth whorl: 14&ndash;19 costae; fifth and last whorl: 19&ndash;20 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.&lt;/p&gt; &lt;p&gt; &lt;b&gt;Material.&lt;/b&gt; 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. J13433 &ndash;34).&lt;/p&gt; &lt;p&gt; &lt;b&gt;Comments.&lt;/b&gt; &lt;i&gt;Athleta (Volutocorbis) burtoni&lt;/i&gt; 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), &lt;i&gt;A. (V.) burtoni&lt;/i&gt; can be distinguished from &lt;i&gt;A. (V.) eugeniae&lt;/i&gt;. Unfortunately, this author did not give diagnostic characters distinguishing the two species. He compared &lt;i&gt;A. (V.) burtoni&lt;/i&gt; with a Bartonian species from the Paris Basin, &lt;i&gt;A. (V.) digitalinus&lt;/i&gt; (Lamarck, 1811) [= &lt;i&gt;Buccinum scabriculum&lt;/i&gt; sensu Solander &lt;i&gt;in&lt;/i&gt; Brander, 1766 &lt;i&gt;partim&lt;/i&gt;, non Linnaeus, 1758], because of its sculpture characterized by the presence of rounded nodules. &lt;i&gt;A. (V.) burtoni&lt;/i&gt; differs obviously from &lt;i&gt;A. (V.) eugeniae&lt;/i&gt; by its smaller size, its more numerous costae (19&ndash;20 instead 14&ndash;15 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 &lt;i&gt;A. (V.) burtoni&lt;/i&gt;, &lt;i&gt;A. (V.) victoriae&lt;/i&gt; (Vredenburg, 1923). The type specimen, illustrated as Vredenburg&rsquo;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 &lt;i&gt;A. (V.) burtoni&lt;/i&gt;. Thus, it is difficult to evaluate whether it corresponds to a variation of &lt;i&gt;A. (V.) burtoni&lt;/i&gt; or a distinct species. Cox (1930) described a comparable species, &lt;i&gt;A. (V.) daviesi&lt;/i&gt; (Cox, 1930) from the Paleocene of the Hangu Formation. The rounded aspect of the nodules of &lt;i&gt;A. (V.) daviesi&lt;/i&gt; recalls the European Bartonian species &lt;i&gt;A. (V.) digitalinus&lt;/i&gt;, but &lt;i&gt;A. (V.) daviesi&lt;/i&gt; differs from &lt;i&gt;A. (V.) burtoni&lt;/i&gt; by having two abapical spiral rows of nodules below the shoulder row. &lt;i&gt;A. (V.) soriensis&lt;/i&gt; (Eames, 1952) from the Early Eocene of the Zinda Pir section (Western Punjab, Pakistan; Eames 1951b) displays numerous similarities to &lt;i&gt;A. (V.) burtoni&lt;/i&gt;, such as rounded nodules, only two rows of nodules (the subsutural and shoulder rows), around 15&ndash;19 costae, small size and a biconic shape. In view of these similarities, we consider that &lt;i&gt;A. (V.) burtoni&lt;/i&gt; and &lt;i&gt;A. (V.) soriensis&lt;/i&gt; could belong to the same species.&lt;/p&gt; &lt;p&gt; &lt;b&gt;Stratigraphic range.&lt;/b&gt; Bara Formation (this paper); Lakhra Formation (Cossmann &amp; Pissarro 1909; Vredenburg 1923);? Ghazij Formation (Punjab).&lt;/p&gt;Published as part of &lt;i&gt;Merle, Didier, Pacaud, Jean-Michel, Métais, Grégoire, Bartolini, Annachiara, Lashari, Rafiq A., Brohi, Imdad A., Solangi, Sarfraz H., Marivaux, Laurent &amp; Welcomme, Jean-Loup, 2014, Volutidae (Mollusca: Gastropoda) of the Lakhra Formation (Earliest Eocene, Sindh, Pakistan): systematics, biostratigraphy and paleobiogeography, pp. 101-138 in Zootaxa 3826 (1)&lt;/i&gt; on pages 121-122, DOI: 10.11646/zootaxa.3826.1.3, &lt;a href="http://zenodo.org/record/228537"&gt;http://zenodo.org/record/228537&lt;/a&gt