Sindhochelys ragei Broin & Métais & Bartolini & Brohi & Lashari & Marivaux & Merle & Warar & Solangi 2021, n. gen., n. sp.

<i>Sindhochelys ragei</i> n. gen., n. sp. <p>(Figs 2-9)</p> <p>urn:lsid:zoobank.org:act: EC548A31-BD5A-478D-83D7-266EE85C9186</p> <p>TYPE MATERIAL. — Holotype, a shell, CPAG-RANKT-V-3. The final repository of the fossil material reported here is the Centre for Pure and Applied Geology (CPAG), University of Sindh, Jamshoro, Pakistan. High-resolution casts are housed at the Muséum national d’Histoire naturelle, Paris.</p> <p> DIAGNOSIS. — For monotypic genus and species. A bothremydid turtle defined by the ornamentation of the shell consisting of raised well protruding polygons but, unlike other ornamented forms such as Taphrosphyini and <i>Carteremys leithii</i>, in having conjoined rather larger irregular polygons without a clear net of granulations, polygons clearly but irregularly protruding and either elongated or regularly polygonal, or open in dichotomic sulci at places. Sharing with <i>Elochelys perfecta</i>, <i>Carteremys</i> group, “ <i>Carteremys</i> ” <i>pisdurensis</i>, and Taphrosphyini the generalized plastral anterior lobe scute pattern of a long intergular, separating narrower and shorter gulars, and widely reaching the pectorals, intergular also separating the thus narrowed humerals; but: humero-intergularo-pectoral sulcus crossing the entoplastron at the 2nd anterior third of its length, its humero-pectoral part posterior to the epi-hyoplastral suture in its median width, but close to it and laterally joining it, contrarily to <i>E. perfecta</i> where it is completely anterior to the suture, contrarily to “ <i>C.</i> ” <i>pisdurensis</i> where it is medially anterior to the suture and lateroposteriorly inclined much posteriorly to the suture, and contrarily to Taphrosphyini and the <i>Carteremys</i> group where it does not cross the suture; intergularo-pectoral sulcus relatively transverse on the entoplastron, as in the <i>Carteremys</i> group, not being medially inclined in a marked V contrarily to <i>E. perfecta</i>, “ <i>C.</i> ” <i>pisdurensis</i> and Taphrosphyini. Bothremydid carapace low with well-inclined bridges but different from other forms, being wider at the suture of peripherals 6-7; displaying a combination of other characters which are distributed in mosaic in the bothremydid groups such as: carapace anteriorly relatively elongated (with regard to primitive forms), at costals 1, neural 1 and anterior peripherals, but moderately, and with an anterior lateral convex contour and without medial nuchal projection; rounded marginal free border of the anterior peripherals; posterior contour slightly protruding towards pygal in a wide convex V; seven relatively narrow neurals; anterior and posterior (overall) plastral lobes not reaching the carapace borders in length; short trapezoid anterior lobe with lateral borders converging only a little towards the very gently convex transverse front, wider at base than the posterior lobe that is long; axillary buttresses laterally beginning at suture peripherals 3 – 4, not particularly thinned, medially overlapping in a curve two thirds of the thoracic rib 2 and medially ending in a rounded extremity on the rib 2 and, contrarily to <i>Taphrosphys sulcatus</i>, rib 2 not close to the costals 1-2 suture towards the median part; rib 1 much anteriorly standing up close to the suture between costals 1 and 2; bridge (between axillary and inguinal notches bottom) longer than the anterior lobe and shorter than the posterior one, with upwards inclined lateral processes, massively shaped around the large (nearly as long as wide) mesoplastra and beside the wide bridge peripherals; posterior lobe with gently convex lateral borders little posteriorly converging. Long marginals 1 overlapping at least the anterior mid-length of the nuchal, and marginals 2 overlap laterally much shortening, in convex curve from peripherals 1 to peripherals 2, this short marginal overlap extending at least to the fourth peripherals (not preserved in situ on the specimen up to the peripherals 8), and with a short ventral marginal overlap; dorsally these short lateroanterior marginals correspond to long pleurals 1, longer in relation to the following pleurals 2 to 4; vertebral 1 a little wider than the nuchal, 2nd to 4th as wide; possibly 4th the longer; and 5th the shorter, but posteriorly the wider. As in various other bothremydids, pectoroabdominal sulcus convex and slightly anterior to the mesoplastra, and curving towards them at its lateral extremity, and not posterior on the hyoplastron all along, contrarily to Taphrosphyini; femorals longer than pectorals, both long in relation to abdominals and anals.</p> <p> ETYMOLOGY. — From <i>chelys</i>, turtle in Latin. Species: in honor to our friend and colleague Jean-Claude Rage. HORIZON AND TYPE LOCALITY. — Lower part of the Khadro Formation, Ranikot Group, early Paleocene (probably early Danian in age), type locality K18-12, in the proximity of Ranikot Fort, Jamshoro District, Sindh Province, Southern Pakistan (Fig. 1).</p> <p>DESCRIPTION (Figs 2-10)</p> <p> <i>Preservation</i></p> <p>The shell has been flattened during the fossilization, and the plastron is fractured at bridges and slightly pushed into the carapace. The carapace is therefore a little wider than when the individual was alive. A great part of the carapace was imbedded in a hard matrix, which could not be removed (Fig. 2), and a part of the plates were dislocated inside the whole assemblage. After the preparation, the elements of broken plates were not in contact, and they are preserved separately; some parts remained in the hard matrix. The photograph (Fig. 3) shows the preserved prepared part with plates in place or in imprints allowing a drawing of the full carapace and plastron (Fig. 4).</p> <i>Dimensions (Table 1)</i> <p> The shell is large (around 70 cm long) and the length/width ratio on (<i>ca.</i> 85 to 90 %) shows it was relatively high for the family, even taking into account the post-mortem flattening. The shell was clearly wider than in the more primitive Pelomedusoides and podocnemidoid forms of Brazil and Morocco. It is neither the smallest nor the largest known bothremydid fossil shell. The oldest and primitive Brazilian bothremydid <i>Cearachelys</i> has a shell length of <i>c.</i> 15 cm (Gaffney <i>et al.</i> 2006). Whereas some Maastrichtian shells from Niger have a length of 120 to 150 cm (Lapparent de Broin <i>et al.</i> 2020). Although much flattened post-mortem, the shell was however rather</p> <p>low and gently rounded dorsally, without dorsal carina, and with well inclined bridges.</p> <i>Carapace (Figs 2-9)</i> <p> The maximum width is at the suture between the peripherals 6 and 7 (Figs 3A; 4A).There are no parallel lateral borders on a short length between the peripherals 4 to 8, unlike most bothremydids: the contour widens from peripherals 2 to 6 and narrows from peripherals 6-7 contact up to the pygal. Anteriorly, the outline of the carapace is semicircular, being just straight at the nuchal area border (Fig. 4A). Posteriorly, the carapace reduces in width with a medial posterior protrusion, forming a wide V.Thus, the carapace was not ovoid in shape, and its anterior part was not anteriorly longer and narrower, in relation to the less long posterior part. The anterior border of the nuchal 1 and most of the border of peripherals 1 is not preserved, but the contour may be reconstructed as transversally straight at the nuchal (Fig.4A) and at medial part of peripherals 1. Indeed, this part does not seem to have been much projected because of the very moderately inclined border of peripherals 2 and 3; this character differs from the condition seen in some taphrosphyine (<i>sensu</i> Lapparent de Broin & Guntupalli 2020) bothremydids such as “ <i>Gafsachelys neurriregularis</i> ” Bergounioux, 1952 from the Ypresian of Gafsa (Tunisia) (Bergounioux 1956, pl. 10). Besides, the median carapace part has no parallel lateral borders, and the posterior part is not rounded as in some bothremydids such as <i>Elochelys perfecta</i> Nopcsa, 1931a (Nopcsa 1931b) from the Campanian</p> <p> of Fuveau-Valdonne (France). The anterior part of the shell is elongated in relation to the primitive condition seen in the oldest bothremydids such as <i>Cearachelys</i> (Gaffney <i>et al.</i> 2006). The maximal length of the costal 1 (at the suture of peripherals 2 and 3) is here as large as the length of the part including the costals 2, 3 and 4 at their mid width (measured following the surface curve). In agreement, the nuchal and peripherals 1 and 2 are also elongated. Although its anterior border is missing, it is clear the nuchal was elongated, but shorter than the costals 1 (Fig. 4A). The free border of peripherals 2 and 3 is not acute; it is rounded as in some of the indetermined Bothremydidae from the Maastrichtian of Upparhatti (India) (Lapparent de Broin & Guntupalli 2020: fig. 7); it is ventrally rounded with a short marginal scute overlap (Figs 3B; 6). The costal 1 is united by concave sutures: 1) anterolaterally to the posterolateral nuchal border and to peripherals 1 to 4, and medially to the neural 1; 2) and by straight sutures to the anterolateral border of the neural 2 and with the costal 2. The first peripheral is the narrowest of all the series, and it is overall narrow posteriorly, at its suture with the costal 1. There are seven neurals. The neural 1 is narrow and elongated in agreement with the costal 1. It is quadrangular, the longer of the series (in agreement with the costal 1 great length). Neurals 2 to 5 (hexagonal with short anterolateral sides) are narrow, and an individual variability in their shape and relative length is observed:1)they have a narrow and nearly equidimensional contact between them but a narrower contact is situated between the neurals 4 and 5; and 2) they have a slightly different length, and a variable maximal width (at the junction of successive costals). Hexagonal neural 6 is shorter than neurals 2 to 5, and pentagonal neural 7 is even shorter, being the shortest in the series: both 6th and 7th have lateral sides which are as long anteriorly and posteriorly. The costals 7 partly posteriorly meet in the mid-line behind neural 7; the costals 8 are fully meeting medially. Costals 2 to 4 are approximately as long laterally as medially, costals 5 are laterally longer, and costals 6 to 8 are laterally longer and less wide (overall the 8th), accompanying the posterior carapace protrusion. The neurals 2 to 5 appear long because they are relatively narrow for their length, particularly the neurals 3 and 4, and evidently their length is in agreement with the costal length, i.e. they are not particularly long in the carapace. The suprapygal is pentagonal as usual in the family when the posterior costals 8 meet in the midline and the pygal is quadrangular, longer than wide in dorsal views (Figs 3A; 9C). In the dorsal views, the posteromedially pointed posterior border of the carapace appears shorter than the anterior one because the carapace is more inclined than the anterior border. On the left, part of the costal 6 and costals 7 and 8 are seen only by their impression: the transection of the iliac bone appears as an oval part in the matrix, between the 7 th and 8 th plates impression (Figs 3A; 9C), as usual below its suture with these plates.</p> <p>It has been possible to separate a fragment with a good part of the neural 1, the left costal 1 and a triangular medial part of the costal 2, which allows to see its ventral aspect (Fig. 5B) and the corresponding impression of the carapace in dorsal view (Fig. 5A). The thoracic rib 2 is extended, below the costal 1 surface, medially from the interval between the neurals 1 and 2 towards laterally the peripheral 4; all along its course, it is rather close to the suture with the costal 2 but not very close, and not linked to it as in some forms. The thoracic rib 1 first extends (from laterally to medially) below the costal 1 along the medial third of the rib 2. Close to the medial border of the costal 1, rib 1 abruptly curves to extend anteromedially towars the level of the mid-length of the neural 1, where it sutured with the apophysis (not preserved) of the thoracic vertebra. The axillary buttress extends medially upon the thoracic rib 2 in a gentle curve, from the suture between the peripherals 3 and 4, and it covers the rib 2 on the two lateral thirds (Fig. 5A, B), making a rounded bulge ventrally (Fig. 5B). It medially ends in a rounded scar which is not dilated. In their median part, the axillary buttress and rib 2 are not as close to the suture between the costals 1 and 2 than in taphrosphyine specimens (Lapparent de Broin & Werner 1998).</p> <i>Plastron (Figs 3B; 4B; 6; 7)</i> <p> It is nearly complete, missing most of the right xiphiplastron and the end of the left xiphiplastron, close to the anal notch, at the insertion of the ischion. The pelvis is concealed by the matrix and the shape of the pubis and ischiatic scars are unknown. However, in posterior view of the shell, the ischiatic contact with the xiphiplastron appears as making a transverse contact: there is no indication of the rounded scar of <i>Taphrosphys sulcatus</i> (Leidy, 1856) and other members of the taphrosphyine tribe (Gaffney 1975: fig. 11; Gaffney <i>et al.</i> 2006) (this tribe corresponding here to the <i>Taphrosphys</i> group</p> <p> of Broin in Antunes & Broin 1988, i.e. Taphrosphyini <i>sensu</i> Lapparent de Broin & Guntupalli 2020). The axillary notches are slightly deformed, better preserved on the right side. The bottom of the right inguinal notch is preserved, and the left one is partly cut by a crack; but its bottom position is indicated by the lateral end of the abdominofemoral sulcus on the hypoplastron, beside the crack, and conform to the right preserved side. The bridge length between the notches bottom was long, reaching approximately 36% of the full length of the plastron. The axillary buttress reaches the boundary between the peripherals 3 and 4 (Fig. 6), and the inguinal buttress reaches 2/3rd of the peripheral 8 (Fig. 3B, left side).</p> <p> The anterior lobe is wide so that it laterally covers rather well the axillary notches space, much occupying the anterior carapace opening in ventral view. It is trapezoidal with a wide transverse anterior border, barely convex, wide at its base and not reaching the anterior nuchal border. The epiplastral symphysis is short and the entoplastron is large in the anterior lobe. It is widely pyriform and its posterior extremity is posterior to the transverse line passing at the bottom of the axillary notches (Fig. 3B, right side). The bridge length between the axillary and inguinal notches is nearly as long as the posterior lobe. The bridge presents the great oblique surfaces of the lateral processes between the main plastral body and the corresponding ventral border of the wide peripherals. The large lateral mesoplastra are hexagonal, a little wider than long, their length representing about one third of the lateral bridge length. The posterior lobe is slightly less wide at its base than the anterior one. Its lateral borders, barely rounded just posterior to the inguinal notch bottom, are gently converging posteriorly towards the mid-line. The posterior lobe does not cover as much the inguinal notches space than in some other bothremydids, but more than in the primitive condition, in which lateral borders are straighter and more posteriorly converging (e.g., <i>Francemys</i> Pérez-García, 2019 [Pérez-García 2019a] and <i>Cearachelys</i> Gaffney, Campos de Almeida & Hirayama, 2001 [Gaffney <i>et al.</i> 2001a]). It is likely that the anal notch was wide and short (doing a short and wide triangle).</p> <p>The plastral plate length formula is: anterior lobe <bridge <posterior lobe length.</p> <i>Shell scutes</i> <p>An important part of the sulci is not preserved. Some are clearly visible, other are estimated (Fig. 4 [dotted lines]). When not visible, their lateral extremity on the border of the element is positioned by the corresponding small incurved border of the plates, and by the sulcus continuation on the inferior face of the plates: this is seen by the border of marginals on the peripheral plates and by the border of plastral scutes on the epi-hyoplastra (Figs 3B; 4B; 6).</p> <p>The carapace (Figs 3; 4) has long marginals 1 on the nuchal. These scutes are estimated as overlapping about the nuchal anterior mid-length (Fig. 4A). Dorsally, the sulci between the two marginals 1 and the vertebral 1, corresponding to the posterior borders of the marginals 1, are slightly concave. On the nuchal, the curve of the sulcus between the vertebral 1,</p> <p> and marginal 1 and the sulcus between the marginal 2 and the vertebral 1 on the peripheral 1 helps identifying the possible position of the missing sulcus between the marginals 1 and 2: this was close to the nuchal-peripheral 1 suture (Fig. 4A [dotted line left side]). Each marginal 1 had to be a rectangular scute and barely wider than long. The marginal 2 was wide, long medially but laterally shorter, from the indicated right curve on the peripheral 1. It was followed by a short marginal 3. The following marginal sulci are mostly missing but from the 7th marginal , on the peripherals 6 and 7, an overlap elongation is estimated, thanks to the preservation of parts of some sulci: as a whole, the marginal dorsal overlap seems to widen at the bridge and slightly narrow down after the bridge (from the peripherals 9 up to the pygal). This medioposterior marginal overlap is longer than that on the anterior peripherals. In conformity, ventrally, the marginals 8-9 sulcus, which is preserved on the left peripheral 8, shows that the ventral marginal overlap is wide at this bridge 8th peripheral, and then the ventral overlap is slightly reduced in length posteriorly. The anterior ventral carapace face indicates a short scute overlap and a rounded thick border (Figs 3B; 6). The ventral scuteskin boundary is marked: anteriorly to the bridge by a thin groove along a low bulge, and posteriorly to the bridge by a protruding rounded half-bulge.The vertebral 1 was wider than the nuchal (thanks to the preserved beginning of the sulcus on the right peripheral 1), the second (its right border being preserved) and third are estimated as equally wide or barely wider than the first, and the fourth seems to be posteriorly narrowed and in contact with the fifth vertebral on the costals 8. If the vertebrals are correctly reconstituted, the longer vertebral is the 4 th, as long as wide, while the 1 st, the 2 nd and 3 rd are wider than long. By the preserved parts of sulci, the vertebrals 1 to 4 seem to be as wide all along, and they are not narrowed at their mutual contact, contrarily to the 4th at its contact with the 5th. From the width of the posterior costals and their contact with the peripherals, it seems the vertebral 5 was posteriorly wider, on the peripheral 10, wider than the other vertebrals. Because of the shorter marginals 2 and 3 in <i>Sindhochelys ragei</i> n. gen., n. sp., the pleural scute 1 was the longest of the series and more than in many derived forms, those which also have an elongated anterior border.</p> <p> The plastral scute sulci are mostly preserved (Figs 3B; 4B; 6). The anterior lobe scute pattern of <i>S. ragei</i> n. gen., n. sp. is estimated as a derived complex of scute arrangement. It is found in several taxa of several different podocnemidoid groups, but the details in the sulci outlines and scute proportions are different in each species, genus or group, and in particular in <i>S. ragei</i> n. gen., n. sp.: this derived “generalized anterior lobe scute pattern” is the intergular, which is wide and long, fully separates the short gulars and humerals, and widely reaches the pectorals. The particularities are here: 1) the position of the humeropectoral sulcus on the hyoplastron; 2) this close but posterior to the epi-hyoplastral suture; and 3) except reaching the lateral extremity of the suture on the left side. It is transversally positioned, following medially (intergular-pectoral sulcus part) to cross the entoplastron in the same direction, slightly posteriorly to the mid-length of the plate; its direction is transverse as a whole and it is not incurved in a V medially on the entoplastron, contrarily to some Taphrosphyini including <i>Taphrosphys sulcatus</i> (in Hay, 1908), the <i>Carteremys</i> group and “ <i>Carteremys</i> ” <i>pisdurensis</i> Jain, 1977 (in Jain 1986) (Lapparent de Broin & Guntupalli 2020), and <i>Elochelys</i> (see Nopcsa 1931b; Gaffney <i>et al.</i> 2006); it is not laterally posteriorly directed towards the hyoplastral border contrarily to “ <i>Carteremys</i&gt

Sindhochelys Broin & Métais & Bartolini & Brohi & Lashari & Marivaux & Merle & Warar & Solangi 2021, n. gen.

Genus Sindhochelys n. gen. urn:lsid:zoobank.org:act: 60DE5DBE-F7CE-4B17-9B04-971BDC06247F TYPE SPECIES AND ONLY SPECIES OF THE GENUS. — Sindhochelysragei n. sp. DIAGNOSIS. — Same as the type species of the genus. ETYMOLOGY. — From Sindh, the province of discovery in Pakistan.Published as part of Métais, Grégoire, Bartolini, Annachiara, Brohi, Imdad Ali, Lashari, Rafiq A., Marivaux, Laurent, Merle, Didier, Warar, Mashooque Ali & Solangi, Sarfraz H., 2021, First report of a bothremydid turtle, Sindhochelys ragei n. gen., n. sp., from the early Paleocene of Pakistan, systematic and palaeobiogeographic implications, pp. 1341-1363 in Geodiversitas 43 (25) on page 1345, DOI: 10.5252/geodiversitas2021v43a25, http://zenodo.org/record/577671

Bothremydidae BAUR 1891

Gen. et sp. indet. (Fig. 10) EXAMINED MATERIAL. — Fragment of a carapace constituted of two right peripherals with adjacent peripherals and costal parts, CPAG-RANKT-V-4. LOCALITY. — Lower part of the Khadro Formation of the Ranikot Group, early Paleocene, locality K18-12, in the proximity of Ranikot Fort, Jamshoro District, Sindh Province, Southern Pakistan (Fig. 1). DESCRIPTION AND COMPARISONS The specimen (17 cm long × 10 cm wide) includes a right bridge portion of carapace with the posterior part of the peripheral 5, the complete peripherals 6 and 7, with a small part of the peripheral 8, sutured to the lateral end of the costals 3 (part), 4 (complete extremity) and 8 (minimal part). In dorsal view (Fig. 10B, C), the peripherals (complete 6 th and 7 th) are wide and display an incurved surface. The free border is thick but neither rounded nor acute. In ventral view (Fig. 10A), the posterior extremity of the bridge is positioned at first third of the 7th peripheral. The sutured border of the plates with the plastron is weakly indented to receive the corresponding weak indentations of the hypoplastron. The fragment was colonized by a perforating animal, such as the pholad bivalve, which nested forming rounded cavities. That occurred once the carapace was dislocated in the marine-littoral water ground, because the cavities are distributed as well on the external face, on the periphero-hypoplastral suture and in the inner bridge cavity, being of various sizes. The fragment is noticeable by its ornamentation, which does not substantially differ from that of S. ragei n. gen., n. sp., being made of wide protruding polygons (Fig. 10C) but which are more marked. The polygons are very elongated and well defined and as elongated as in some plates of S. ragei n. gen., n. sp., (Fig. 8C, E), but more protruding, perhaps because of a better preservation at this location. The only certitude is that it does not correspond to any other decorated bothremydid examined above by comparison with Sindhochelys. Although similar to Sindhochelys by the marked decoration and different from taphrosphyines and other examined bothremydids from France and India, the fragment remains undetermined.Published as part of Métais, Grégoire, Bartolini, Annachiara, Brohi, Imdad Ali, Lashari, Rafiq A., Marivaux, Laurent, Merle, Didier, Warar, Mashooque Ali & Solangi, Sarfraz H., 2021, First report of a bothremydid turtle, Sindhochelys ragei n. gen., n. sp., from the early Paleocene of Pakistan, systematic and palaeobiogeographic implications, pp. 1341-1363 in Geodiversitas 43 (25) on page 1358, DOI: 10.5252/geodiversitas2021v43a25, http://zenodo.org/record/577671

Lyrischapa haimei d'Archiac & Haime 1853

<i>Lyrischapa haimei</i> (d’Archiac & Haime, 1853) <p>(Fig. 4 L)</p> <p> <i>Voluta haimei</i> d’Archiac & Haime, 1853: 325, p1. 31, figs. 26, 27.</p> <p> <i>Voluta haimei</i> d’Archiac, 1850: 298 (nomen nudum).</p> <p> <i>Aulicina haimei</i>. Cossmann & Pissarro 1909: 26, pl. 2, fig. 29 only [new combination]. <i>Volutoconus funiculifer</i> Cossmann & Pissarro, 1909: 27, pl. 3, figs 10–12.</p> <p> <i>Aulica (Aulicina) haimei</i>. Vredenburg 1923: 269; Cotter <i>in</i> Vredenburg 1928: 38 [new combination]. <i>Eovasum haimei</i>. Cox 1930: 186 in part (not pl. 21, figs 1–4), see (Givens 1991) [new combination]; Davies 1934: 297. <i>Lyrischapa haimei</i>. Givens 1991: 664, figs 3.6–3.8 [new combination].</p> <p> <b>Material.</b> 1 spm (stn 6: CPAG.RAN. I.8, cast MNHN.F.A50346).</p> <p> <b>Comments.</b> In his paper on <i>Lyrischapa</i>, Givens (1991) gave a revised description of <i>L. haimei</i>. He recognized two differential characters distinguishing it from <i>L. sismondai</i> (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 <i>L. sismondai</i> is larger than that of <i>L. haimei</i>. 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 <i>L. haimei</i>.</p> <p> <b>Stratigraphic range.</b> Lakhra Formation: Jhirak (Cossmann & Pissarro 1909; Vredenburg 1923; this paper).</p>Published as part of <i>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)</i> on pages 111-112, DOI: 10.11646/zootaxa.3826.1.3, <a href="http://zenodo.org/record/228537">http://zenodo.org/record/228537</a&gt

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 <i>Lyriopsis</i> gen. nov. <p> <b>Type species.</b> <i>Lyria cossmanni</i> Vredenburg, 1923. Early Eocene from Lakhra Formation (Sindh, Pakistan).</p> <p> <b>Etymology.</b> Combination of <i>Lyria</i> with the suffix <i>opsis</i> (resembling in appearance). Gender feminine.</p> <p> <b>Diagnosis.</b> A ventricose <i>“ Lyria ”</i> 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.</p> <p> <b>Included species</b>. <i>Lyria cossmanni</i> Vredenburg, 1923 (Lakhra Formation, Sindh Province, Pakistan) and <i>Lyria samanaensis</i> Cox, 1930 (Hangu Formation, Paleocene, Khyber Pakhtunkhwa, Pakistan).</p> <p> <b>Discussion.</b> Cossmann & Pissarro (1909), Vredenburg (1923) and Cox (1930) attributed these two species to the genus <i>Lyria</i> and compared them to three Eocene species from the Paris Basin: <i>Lyria (Lyria) harpula</i> (Lamarck, 1803), <i>L. (L.) subturgidula</i> (d’Orbigny, 1850) (= <i>Voluta turgidula</i> Deshayes, 1835, not of Brocchi, 1814) and <i>L. (Pseudolyria) coroni</i> (Morlet, 1888). According to these authors, strong but narrowly rounded costae and a biconic shape are shared with the Eocene <i>Lyria</i> species of the Paris Basin and even with many other Cenozoic species of <i>Lyria</i>. Thus, <i>Lyriopsis</i> is placed in the tribe Lyriini, but four characters distinguish members of <i>Lyria</i> from <i>Lyriopsis</i>: the morphology of the costae, the construction of the columellar folds, the shape of the whorl, and the protoconch. In <i>L. (Lyria)</i>, the posterior ends of the costae are straight, whereas they are obviously sinuous in <i>Lyriopsis</i>. Regarding the subgenus <i>Pseudolyria</i> Martin, 1931 [type species <i>Pseudolyria ventricola</i> Martin, 1931 by monotypy], the costae are more sinuous than in <i>L. (Lyria)</i>, but the sinuosity is more anteriorly placed than in <i>Lyriopsis</i>. Moreover, the outer lip of <i>L. (Pseudolyria)</i> is crenulated, whereas it is smooth in <i>Lyriopsis</i>. In <i>L. (Lyria)</i>, 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 <i>Lyriopsis</i>. The shape of the whorl is rather depressed, slightly convex and terraced in <i>Lyriopsis</i> species, whereas the whorls are strongly convex in most species of <i>Lyria (Lyria)</i>. For these reasons, the placement of <i>Lyriopsis</i> in the Lyriini is provisional and uncertain.</p>Published as part of <i>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)</i> on page 108, DOI: 10.11646/zootaxa.3826.1.3, <a href="http://zenodo.org/record/228537">http://zenodo.org/record/228537</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

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

<i>Athleta (Volutopupa) intercrenatus</i> (Cossmann & Pissarro, 1909) comb. nov. <p>(Fig. 7 E–J)</p> <p> <i>Volutospina intercrenata</i> Cossmann & Pissarro, 1909: 24, pl. 2, figs 18–20, pl. 3, figs 1–3.</p> <p> <b>Description.</b> Shell biconic, H 33–37, D 18–23 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: 12–13 costae; fifth and last whorl: 11–12 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 6–10 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.</p> <p> <b>Material.</b> 5 spm (stn 4: 4 spm CPAG.RAN. I.21–24, cast MNHN.F. A50359 –62 and 1 spm MNHN).</p> <p> <b>Comments.</b> As seen above with <i>Athleta (Volutopupa) noetlingi</i>, the four specimens of <i>V. intercrenata</i> illustrated by Cossmann & Pissarro (1909) are species of <i>Volutopupa</i> and not species of <i>Volutospina</i>. 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 <i>Volutospina intercrenata</i> Cossmann & 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 <i>A. (V.) intercrenatus</i> collected in the Lakhra Dome. The oval shape of these adults resembles more that of <i>A. (V.) lyra</i> (Lamarck, 1803) or <i>A. (V.) mutatus</i> (Deshayes, 1835) from the Middle Eocene of the Paris Basin, than it does <i>A. (V.) citharoedus</i>, which displays an inflated and ventricose last whorl. <i>A. (V.) intercrenatus</i> also differs from these species by its strongly developed row of subsutural nodules.</p> <p> <b>Stratigraphic range.</b> Lakhra Formation: Jhirak (Cossmann & Pissarro 1909) and Lakhra Dome (this paper).</p>Published as part of <i>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)</i> on pages 118-119, DOI: 10.11646/zootaxa.3826.1.3, <a href="http://zenodo.org/record/228537">http://zenodo.org/record/228537</a&gt

Athleta (Volutocorbis) burtoni Vredenburg 1923

<i>Athleta (Volutocorbis) burtoni</i> Vredenburg, 1923 <p>(Fig. 9A–G)</p> <p> <i>Athleta (Volutocorbis) burtoni</i> Vredenburg, 1923: 261, pl. 15, fig. 2. <i>Athleta (Volutocorbis) victoriae</i> Vredenburg, 1923: 260, pl. 15, figs 4. <i>Athleta (Volutocorbis) burtoni.</i> Cotter <i>in</i> Vredenburg 1928: 37. <i>Volutocorbis soriensis</i> Eames, 1952: 109, pl. 4, fig. 95a–b.</p> <p> <i>Volutocorbis burtoni</i>. Glibert 1960: 48 [new combination].</p> <p> <b>Description.</b> 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, 13–15 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: 15–16 costae; fourth whorl: 14–19 costae; fifth and last whorl: 19–20 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.</p> <p> <b>Material.</b> 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 –34).</p> <p> <b>Comments.</b> <i>Athleta (Volutocorbis) burtoni</i> 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), <i>A. (V.) burtoni</i> can be distinguished from <i>A. (V.) eugeniae</i>. Unfortunately, this author did not give diagnostic characters distinguishing the two species. He compared <i>A. (V.) burtoni</i> with a Bartonian species from the Paris Basin, <i>A. (V.) digitalinus</i> (Lamarck, 1811) [= <i>Buccinum scabriculum</i> sensu Solander <i>in</i> Brander, 1766 <i>partim</i>, non Linnaeus, 1758], because of its sculpture characterized by the presence of rounded nodules. <i>A. (V.) burtoni</i> differs obviously from <i>A. (V.) eugeniae</i> by its smaller size, its more numerous costae (19–20 instead 14–15 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 <i>A. (V.) burtoni</i>, <i>A. (V.) victoriae</i> (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 <i>A. (V.) burtoni</i>. Thus, it is difficult to evaluate whether it corresponds to a variation of <i>A. (V.) burtoni</i> or a distinct species. Cox (1930) described a comparable species, <i>A. (V.) daviesi</i> (Cox, 1930) from the Paleocene of the Hangu Formation. The rounded aspect of the nodules of <i>A. (V.) daviesi</i> recalls the European Bartonian species <i>A. (V.) digitalinus</i>, but <i>A. (V.) daviesi</i> differs from <i>A. (V.) burtoni</i> by having two abapical spiral rows of nodules below the shoulder row. <i>A. (V.) soriensis</i> (Eames, 1952) from the Early Eocene of the Zinda Pir section (Western Punjab, Pakistan; Eames 1951b) displays numerous similarities to <i>A. (V.) burtoni</i>, such as rounded nodules, only two rows of nodules (the subsutural and shoulder rows), around 15–19 costae, small size and a biconic shape. In view of these similarities, we consider that <i>A. (V.) burtoni</i> and <i>A. (V.) soriensis</i> could belong to the same species.</p> <p> <b>Stratigraphic range.</b> Bara Formation (this paper); Lakhra Formation (Cossmann & Pissarro 1909; Vredenburg 1923);? Ghazij Formation (Punjab).</p>Published as part of <i>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)</i> on pages 121-122, DOI: 10.11646/zootaxa.3826.1.3, <a href="http://zenodo.org/record/228537">http://zenodo.org/record/228537</a&gt