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Shells of Bursa granularis complex. A–D. Bursa granularis (Röding, 1798). A. MHNG 1098/85/1, lectotype of Ranella granifera Röding, 1798 andneotype of Tritonium granulare Röding, 1798, of T. jabick Röding, 1798 and of Biplex rubicola Perry, 1811, from Red Sea, H 64 mm. B. IM-2009–5148,MAINBAZA 2009 (Mozambique Channel), stn DW3168, H: 26 mm. C. UF-423792, UF 2008 (Madagascar), NW side Nosy Komba, H 40 mm. D.IM-2007–43071, Santo Marine Biodiversity Survey 2006 (Vanuatu), stn VM62, H 25 mm. E–I. Bursa affinis (Broderip, 1833). E. NHMUK 1950.11.28.4,lectotype of Ranella affinis Broderip, 1833, supposedly from ‘Annaa,’ Tuamotu Islands, H 62.5 mm. F. IM-2007–43056, Santo Marine Biodiversity Survey2006 (Vanuatu), stn VM02, H 37 mm. G. NHMUK 1967657, lectotype of Ranella livida Reeve, 1844, supposedly from ‘Annaa,’ Tuamotu Islands, H56.8 mm. H. UF-422918, UF 2008 (Federated States of Micronesia), Kosrae Letu, H 25 mm. I. IM-2007–43039, PANGLAO 2004 (Philippines), stn R24,H 41 mm. J. IM-2009–5148, Bursa granularis protoconch detail. K. IM-2007–43056, Bursa affinis protoconch detail. Abbreviations: P1, P3, P5, preponderantprimary cords; numbers indicates the number of denticles.

Shells of Bursa granularis complex. A–D. Bursa granularis (Röding, 1798). A. MHNG 1098/85/1, lectotype of Ranella granifera Röding, 1798 andneotype of Tritonium granulare Röding, 1798, of T. jabick Röding, 1798 and of Biplex rubicola Perry, 1811, from Red Sea, H 64 mm. B. IM-2009–5148,MAINBAZA 2009 (Mozambique Channel), stn DW3168, H: 26 mm. C. UF-423792, UF 2008 (Madagascar), NW side Nosy Komba, H 40 mm. D.IM-2007–43071, Santo Marine Biodiversity Survey 2006 (Vanuatu), stn VM62, H 25 mm. E–I. Bursa affinis (Broderip, 1833). E. NHMUK 1950.11.28.4,lectotype of Ranella affinis Broderip, 1833, supposedly from ‘Annaa,’ Tuamotu Islands, H 62.5 mm. F. IM-2007–43056, Santo Marine Biodiversity Survey2006 (Vanuatu), stn VM02, H 37 mm. G. NHMUK 1967657, lectotype of Ranella livida Reeve, 1844, supposedly from ‘Annaa,’ Tuamotu Islands, H56.8 mm. H. UF-422918, UF 2008 (Federated States of Micronesia), Kosrae Letu, H 25 mm. I. IM-2007–43039, PANGLAO 2004 (Philippines), stn R24,H 41 mm. J. IM-2009–5148, Bursa granularis protoconch detail. K. IM-2007–43056, Bursa affinis protoconch detail. Abbreviations: P1, P3, P5, preponderantprimary cords; numbers indicates the number of denticles.

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Bursa granularis (Röding, 1798) is a tonnoidean gastropod that is regarded as broadly distributed throughout the Indo-Pacific and tropical western Atlantic. Because of its variable shell it has received no less than thirteen names, now all synonymized under the name B. granularis. We sequenced a fragment of the cox1 gene for 82 specimens covering a...

Citations

... The giant triton snail Charonia tritonis (Linnaeus, 1758) is known to prey on crown-of-thorns starfish Acanthaster planci (Linnaeus, 1758), and the over-harvesting of Charonia tritonis in the Indo-Pacific Ocean could lead to outbreaks of crown-of-thorns starfish that could damage the local coral reefs [6]. Tonnoideans are also known for their long planktonic larval stages [7], with the longest larva period of Fusitriton oregonensis (Redfield, 1846) (Cymatiide) recorded as 4.5 years in an aquarium [8]. In Southeast Asia, Tonnoideans are consumed as food, and shells of Bufonaria rana (Linnaeus, 1758) have been used in traditional Chinese medicine to cure ulcers and furuncle carbuncles [9]. ...
... However, it has been shown that this capability was overestimated for some species within the Bursidae family. For example, the Dulcerana granularis (Röding, 1798) was originally considered to be a single species with a worldwide distribution, whereas subsequent systematic analysis revealed that D. granularis was composed of at least four endemic species, forming the genus Dulcerana Oyama, 1964 [7,20]. A similar result was detected in our phylogeny due to the separated positions of two Tut. ...
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The Tonnoidea Suter, 1913 (1825) is a moderately diverse group of large predatory gastropods, the systematics of which remain unclear. In the present study, the complete mitochondrial genomes of nine Tonnoidean species were sequenced. All newly sequenced mitogenomes contain 13 protein-coding genes (PCGs), 22 transfer RNA genes and two ribosomal RNA genes, showing similar patterns in genome size, gene order and nucleotide composition. The ratio of nonsynonymous to synonymous of PCGs indicated that NADH complex genes of Tonnoideans were experiencing a more relaxed purifying selection compared with the COX genes. The reconstructed phylogeny based on the combined amino acid sequences of 13 protein-coding genes and the nucleotide sequences of two rRNA genes supported that Ficidae Meek, 1864 (1840) is a sister to Tonnoidea. The monophylies of all Tonnoidean families were recovered and the internal phylogenetic relationships were consistent with the current classification. The phylogeny also revealed that Tutufa rebuta (Linnaeus, 1758) is composed of at least two different species, indicating that the species diversity within Bursidae Thiele, 1925 might be underestimated. The present study contributes to the understanding of the Tonnoidean systematics, and it could provide important information for the revision of Tonnoidean systematics in the future.
... Th e taxonomy and terminology follow Beu (2010), Kovács & Vicián (2018), Sanders et al. (2019), and Craig et al. (2020) with modifi cations. For sculpture description method of Merle (2001Merle ( , 2005, Sanders et al. (2017), and Landau et al. Description -Medium-sized shells, moderately low spire, slightly concave sutural ramp, globose last whorl. ...
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Middle Eocene Cassidae, Charoniidae, Cymatiidae and Personidae species are described and illustrated from the Hungarian Paleogene Basin. Pseudosassia n. gen. and eight new species are designated: Cassis kalmani n. sp., Pseudosassia gurdoni n. sp., Pseudosassia traceyi n. sp., Monoplex s.l. szakonyii n. sp., Cymatiella dulaii n. sp., Parasassia vargai n. sp., Protoplex ? zsoldosi n. sp., and Personopsis merlei n. sp. Genus Semiranella De Gregorio, 1880 is assigned to the family Charoniidae. Sconsia cyrenaica Stefanini, 1921 is regarded as a junior synonym of Sconsia ambigua (Solander, 1766). Cymatiella tzankovi nom. nov. is proposed as new name for “Eutritonium (Sassia) rutoti” Tzankov, 1940.
... Furthermore, in another neogastropod species complex for which there is a comparable number of protoconch whorls (e.g. Sanders et al., 2017), there is a strict ocean-delimited (e.g. Indian only, Pacific only) distribution. ...
Article
Although raphitomid snails are a dominant component of gastropod communities in deep seas worldwide, their systematics is still largely tentative. We assembled the most complete sampling of Raphitomidae from southeastern Australia to date. Based on morphological and molecular data from this material, we produced a robust phylogenetic framework and used it to delimit genera. For the focus area, our results show a large proportion of undescribed species-and genus-level taxa, 11 of which are formally described herein. We demonstrate that the examination of purely morphological characters rarely suffices for the purpose of accurate genus delimitation. As a result, some traditionally highly diverse raphitomid genera (such as Gymnobela) turn out to be artificial assemblages of several unrelated, mostly undescribed, genus-level lineages. Our data suggest that comparable configurations of shell and radular features, observed at the genus level, commonly do not reflect true phylogenetic relationships. However, our results are inconclusive as to whether homoplasy or conservatism are the drivers of this phenomenon. Accommodating for the inevitable sampling biases, southeastern Australia appears as a possible hotspot for both raphitomid diversity and endemism, when compared with adjacent areas.
... Le Campanile du Miocène de Cap Amparafaka, présentait une surface encroutée et a subi un traitement à l'hydroxyde de potassium (KOH) pour le dégager de la gangue, nous permettant ainsi d'observer, sur une grande partie et très nettement, la sculpture spirale des tours. La méthode analytique pour identifier les homologies structurales des Calliostomatidae (Ikebe, 1942 ;Marshall, 1995), des Architectonicidae et des Mathildidae (Bieler, 1988(Bieler, , 1992(Bieler, , 1995, des Muricidae (Merle, 1999(Merle, , 2001(Merle, , 2005Pacaud, 2018), des Turbinidae (Pacaud et Ledon, 2010), des Rostellariidae (Pacaud et al., 2011) et des Bursidae (Sanders et al., 2017(Sanders et al., , 2019 est ici utilisée. La méthode descriptive du type de sculpture spirale de cette espèce nouvelle est basée sur l'identification des cordons spiraux que nous avons mise en évidence (Pacaud et al., 2014) dans la morphologie des coquilles du genre Campanile. ...
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The species Campanile madagasikara sp. nov. is described from the Upper Miocene of Cap Amparafaka, Madagascar. The genus Campanile is recorded for the first time to Cenozoic in Madagascar. Indeed, the genus has not hitherto been recorded from Madagascar, a part of Cretacous sediments. This is the most recent occurrence of the genus so far recorded from Madagascar. Miocene Campaniles are only found in Southeast Asia, New Zealand, Australia and East Africa. The discovery of theMalagasy species described here suggests the existence of relict areas in the Indian Ocean and the Pacific at the Neogene and even at present in Australia. The identity of the spiral cords is researched by their topological and ontogenetical correspondences, then codified by a terminology. This descriptive method used by Pacaud et al. (2014) allows to clarify the presentation of the characters and to propose structural homologies. Résumé – L’espèce Campanile madagasikara nov. sp. est décrite du Miocène supérieur du Cap Amparafaka à Madagascar. Le genre Campanile est signalé pour la première fois au Cénozoïque à Madagascar. En effet, le genre n’a pas jusqu’ici été rencontré à Madagascar, à part dans les sédiments crétacés. Il s’agit donc de l’occurrence la plus récente du genre jusqu’ici enregistré à Madagascar. Les Campanile au Miocène sont seulement présents en Asie du Sud-Est, en Nouvelle-Zélande, en Australie et en Afrique de l’Est. La découverte de l’espèce malgache décrite ici suggère l’existence d’aires relictes dans l’océan Indien et le Pacifique au Néogène et jusque dans l’actuel en Australie. L’identité des cordons spiraux est recherchée par leur correspondance topologique et ontogénétique, puis codifiée par une terminologie. Cette méthode descriptive utilisée par Pacaud et al. (2014) permet de clarifier la présentation des caractères et de proposer des homologies structurales.
... Bursidae Thiele, 1925 (frog shells) are a moderately diverse tonnoidean family with 59 currently recognized extant species (WoRMS: http://www.marinespecies.org/aphia.php?p=taxdetails&id=22995). Twenty extinct species were previously recognized but this number was downsized recently to eleven (Sanders et al., 2019). Recent bursids have a tropical and subtropical distribution with the southernmost records in eastern South Africa (Sanders et al., 2017), Northern New Zealand (Spencer et al., 2016) and the Walters shoal (https://expeditions.mnhn. fr/campaign/waltersshoal). The northernmost records (Bursa scrobilator (Linnaeus, 1758)) are from Nice, southern France and Savona, northwestern Italy [we consider the specimen from northern Finistère, Britany, France attributed by Delongueville and Scaillet (2000) to Bufonaria rana (Linnaeus, 1758) to be dubious]. ...
... As inferred from their protoconch, similar to all other tonnoideans, they are considered to have a long to extremely long larval stage granting for some of them trans-oceanic dispersal capabilities. However, Sanders et al. (2017) showed that such capabilities were greatly overestimated for one of the most widespread species of Bursidae: Bursa granularis. Once considered to be a single species with a worldwide distribution, it in fact includes at least four species restricted to the Indian, the Pacific and the western Atlantic Oceans, respectively, plus one restricted to southernmost Western Australia, as confirmed by an integrative taxonomy approach combining molecular and morphological characters. ...
... Further morphological characteristics are a spiral ornamentation of the convex part of the whorl composed of six to eight primary cords (sensu Merle, 2005;Sanders et al., 2017) that can be lost during ontogeny. Secondary and tertiary ones can appear later on and sometimes grow to such importance that they can be confused with primary cords (e.g., Dulcerana granularis (Röding, 1798)). ...
Article
With 59 Recent species, Bursidae, known as « frog shells », are a small but widely distributed group of tropical and subtropical gastropods that are most diverse in the Indo-West Pacific. The present study is aimed at reconstructing phylogenetic relationships of bursid gastropods based on extensive and representative taxon sampling. Five genetic markers (cytochrome c oxidase subunit I (cox1), 16s and 12s rRNA mitochondrial genes, 28s rRNA and Histone H3 nuclear gene) were sequenced for over 30 species in every known genus but Crossata. Furthermore, we sequenced the complete mt-genome of 9 species (10 specimens) (Aspa marginata, Marsupina bufo, Korrigania quirihorai, Korrigania fijiensis, Tutufa rubeta, Bursa lamarckii, Lampasopsis rhodostoma (twice), Bufonaria perelegans and Bursa aff. tuberosissima). Our analysis recovered Bursidae as a monophyletic group, whereas the genus Bursa was found to be polyphyletic. The genera Talisman and Dulcerana are resurrected and the genera Alanbeuella gen. nov. and Korrigania gen. nov. are described. Dating analysis using 21 extinct taxa for node and simplified tip calibrations was performed, showing a diversification of the group in two phases. Diversification seems to be linked to tectonic events leading to biodiversity relocation from the western Tethys toward the Indo-Pacific.
... Tonnoideans (Tonnoidea) are a moderately diverse super family of Caenogastropoda and it contains about 361 valid species around the world in 21 families (Strong et al. 2019), which are the most valuable and vulnerable inshore fisheries resources from subtropical to tropical waters. Bursidae is one of the diverse group in Tonnoidea with 54 recent species, among which several are potentially species complexes (Sanders et al. 2017). However, due to limited evaluation of the intraspecific variability and molecular systematic studies, the taxonomy and classification of the Bursidae have been debated and synonyms among species have been reported recently (Ran et al. 2020;Strong et al. 2019). ...
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The common frogsnail Bufonaria rana, is an ecologically and economically important Tonnoideans in China due to valuable nutrition and pharmacological compounds. However, the taxonomy and phylogeny of the Bursidae have been debated and synonyms among Bursidae species have been reported recently. In this study, we report the first complete mitochondrial genome of Bursidae from B. rana. The mitogenome has 15,510 base pairs (69.0% A + T content) and made up of total of 37 genes (13 protein-coding, 22 transfer RNAs and 2 ribosomal RNAs), and a putative control region. This study was the first available complete mitogenomes of Bursidae and will provide useful genetic information for future phylogenetic and taxonomic classification of Tonnoideans.
... Our work on other neogastropod taxa shows that species initially identified by by molecular characters are often characterized by distinctive and recognizable shell characters; examples in this respect include species in the genera Bathytoma (Puillandre et al., 2010a), Crassispira , Lophiotoma and Hemilienardia (Fedosov et al., 2016). Another example is the tonnoidean Bursa granularis, which was previously treated as a single, broadly distributed species, but is now considered to consist of four separate species (Sanders et al., 2017). However, as has been shown by Puillandre et al. (2010b) for Gemmuloborsonia, there are cases where species, which are clearly defined using molecular systematic characters, are not always recognizable on the basis of morphology. ...
Article
According to a recent taxonomic revision by Kantor et al. (2001), the neogastropod genus Exilia Conrad, 1860, comprises ten mostly rare species that live at depths between 200 and 2000 m. Adult Exilia measure between 30 and 90 mm in shell length, and the genus is mostly represented in museum collections by empty shells. The abundance of this genus is low in the wild, but recent expeditions organized by the Muséum national d'Histoire naturelle have yielded several dozen specimens. These new collections include samples preserved for molecular studies. Here, we present the results of the first molecular systematic study of Exilia. Our aim was to investigate the species limits proposed by Kantor et al. (2001) on the basis of shell and anatomical characters. Analysis of DNA sequence data for the cytochrome c oxidase I gene suggests that Exilia hilgendorfi, previously considered to be a single, polymorphic and broadly distributed species, is a complex of at least six species (four of which we sequenced). Two of these species, Exilia cognata n. sp. and E. fedosovi n. sp., are described as new to science. Exilia gracilior, E. claydoni and E. prellei are resurrected from the synonymy of Exilia hilgendorfi; of these three, only the last was sequenced. Exilia vagrans is a well-defined taxon, but our molecular systematic data shows that it consists of two distinct species, which occur sympatrically off Taiwan and are strikingly similar in shell and radular morphology; due to the absence of DNA sequence data from the type locality of E. vagrans (Vanuatu), it is unclear to which of these two species the name would apply. Exilia karukera n. sp., which is conchologically very similar to E. vagrans, was discovered off Guadeloupe, represents the first record of the genus from the Atlantic. For E. elegans, which was previously known only from a single shell, we provide new data including new distributional records (South Africa and the Mozambique Channel), details of the radula and DNA sequence data.
... Furthermore, in another neogastropod species complex for which there is a comparable number of protoconch whorls (e.g. Sanders et al., 2017), there is a strict ocean-delimited (e.g. Indian only, Pacific only) distribution. ...
Article
The practice of species delimitation using molecular data commonly leads to the revealing of species complexes and an increase in the number of delimited species. In a few instances, however, DNA-based taxonomy has led to lumping together of previously described species. Here, we delimit species in the genus Cryptogemma (Gastropoda: Conoidea: Turridae), a group of deep-sea snails with a wide geographical distribution, primarily by using the mitochondrial COI gene. Three approaches of species delimitation (ABGD, mPTP and GMYC) were applied to define species partitions. All approaches resulted in eight species. According to previous taxonomic studies and shell morphology, 23 available names potentially apply to the eight Cryptogemma species that were recognized herein. Shell morphometrics, radular characters and geographical and bathymetric distributions were used to link type specimens to these delimited species. In all, 23 of these available names are here attributed to seven species, resulting in 16 synonymizations, and one species is described as new: Cryptogemma powelli sp. nov. We discuss the possible reasons underlying the apparent overdescription of species within Cryptogemma, which is shown here to constitute a rare case of DNA-based species lumping in the hyper-diversified superfamily Conoidea.
... Strathmann & Strathmann, 2007). Only recently has the molecular systematics of the family Bursidae begun to be investigated (Castelin et al., 2012;Sanders et al., 2017;Strong et al., 2019); the same holds true for Bursa scrobilator (Linnaeus, 1758), one of the most iconic marine gastropods of the Eastern Atlantic and Mediterranean. On the basis of extensive shell-based studies and a molecular phyloge-netic analysis of some Mediterranean and Senegalese specimens, Smriglio et al. (2019) proposed that B. scrobilator consists of two subspecies: B. scrobilator scrobilator (Linnaeus, 1758) and B. s. coriacea (Reeve, 1844). ...
... Sequences were retrieved from the GenBank and Barcode of Life Data System (BOLD) databases by searching for DNA barcodes of specimens belonging to the genus Bursa; the taxonomic identifications in the two databases were cross-checked with the most recent taxonomic literature on the family Bursidae (Castelin et al., 2012;Sanders et al., 2017;Smriglio et al., 2019;Strong et al., 2019 Beu, 1987;B. granularis (Röding, 1798); B. larmacki (Deshayes, 1853); B. quirihorai Beu, 1987;B. ...
... This suggests a more recent split between the two ESUs (2-3 Myr), which would be consistent with ranking the two morphotypes as separate subspecies. We note that the genetic divergence between B. s. scrobilator and B. s. coriacea is roughly one third the minimum genetic distances observed between cryptic species in the B. granularis species complex (6.4%; Sanders et al., 2017), which is a clade of at least four species, for which a maximum calibration age of 9 Myrs has been suggested (Sanders et al., 2019). We suggest that a molecular phylogenetic study based on several loci and incorporating data from the recent review of fossil data by Sanders et al. (2019) should be carried out. ...
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Genetic connectivity plays a crucial role in shaping the geographic structure of species. Our aim in this study was to explore the pattern of genetic connectivity in Bursa scrobilator, an iconic marine caenogastropod with long-lived pelagic larvae. Our study was based on the analysis of DNA sequence data for the 658-bp barcoding fragment of the mitochondrial cytochrome c oxidase subunit I (COI) gene. This is the largest DNA sequence dataset assembled to date for B. scrobilator. These data confirm that the two recently described subspecies B. scrobilator scrobilator (Linnaeus, 1758), from the Mediterranean and Macaronesia, and B. s. coriacea (Reeve, 1844), from West Africa, constitute two evolutionarily significant units (ESUs). We found that for the nominal subspecies, the variation in morphology (shell, radula and gross anatomy) and DNA sequences was not geographically structured, and this agrees with what we would expect in a species with high connectivity at the larval stage. The divergence between the two subspecies cannot be easily explained by isolation by distance, and we would argue that one or more extrinsic factors may have played a role in isolating the two ESUs and maintaining that isolation.
... Furthermore, in other neogastropod species complex for which there is a comparable number of protoconch whorls (e.g. Sanders et al., 2017) the study showed a strict ocean-delimited (e.g. Indian only, Pacific only) distribution. ...
Thesis
Les Turridae constituent un groupe de gastéropodes marins prédateurs venimeux, présents dans presque toute la zone pantropicale et des côtes à plus de 3000 m de profondeur. Etant donné leur faible diversité (environ 200 espèces décrites), relativement à leurs taxons frères au sein des Conoidea, et l’absence apparente d’innovation-clé spécifique à ce groupe, la question de la thèse est la suivante : quelles sont les dynamiques de diversification et les moteurs de cette diversification chez les Turridae ? Toutefois, il n’est possible d’y répondre qu’à condition d’avoir un cadre systématique robuste. Les approches traditionnelles de systématique, basées sur des considérations morpho-anatomiques, sont confrontées à des problèmes de plasticité de la coquille et des organes, que ce soit dans des temps évolutifs courts ou longs. L’outil moléculaire a donc été privilégié pour répondre à cette question.Dans une première partie, un pipeline de taxonomie intégrative a été défini et utilisé pour délimiter des hypothèses d’espèces chez les Turridae. Ce pipeline a résulté en environ 200 hypothèses d’espèces, dont il n’est pas toujours facile de savoir si elles ont déjà été décrites ou non. J’explore cette problématique avec une analyse poussée du genre Cryptogemma. L’acquisition semi-automatisée de données morphométriques a permis d’attribuer avec confiance 23 spécimens porte-noms à 8 espèces délimitées, illustrant la confusion taxonomique qui existe chez les Turridae.Dans une seconde partie, je construis une phylogénie des Turridae en utilisant une approche de « génome-réduit ». Ces approches ont permis de séquencer environ 4000 loci pour 110 hypothèses d’espèce. Les topologies retrouvées mettent en évidence de façon robuste la polyphylie du genre Gemmula, au sein duquel 13 genres potentiellement doivent être décrits.Enfin, j’utilise les résultats combinés de la taxonomie intégrative et la phylogénie, ainsi que des méthodes récentes pour pour étudier la dynamique de diversification de groupe. La famille des Turridae apparaît comme un groupe dont les taux de diversification passés seraient plus importants qu’actuellement. Des liens ont été trouvés entre la diversification et certains traits comme la protoconque, la forme et la taille de la coquille et la bathymétrie. La stratégie de systématique utilisée pendant cette thèse s’est révélée efficace pour permettre à répondre à des questions-clefs liées à la diversification des gastéropodes marins, proposant des résultats inédits.