Malcolm T Sanders
email: malcolm.t.sanders@gmail.com

Didier Merle
email: didier.merle@mnhn.fr

Nicolas Puillandre
email: nicolas.puillandre@mnhn.fr

A review of fossil Bursidae and their use for phylogeny calibration

Keywords: Bursidae, Tonnoidea, node calibration, lectotypifications, new combinations, new genera Bursidae, Tonnoidea, lectotypifications, combinaisons nouvelles, genres nouveaux

 and Aquitanobursa n. gen. (type species: Ranella grateloupi d Orbigny, 1852), containing only fossil species. Lectotypes are designated for Ranella grateloupi d Orbigny, 1852, Ranella morrisi d rchiac & Haime, 1853 and Apollon pelouatensis Cossmann & Peyrot, 1924. Based on this revision of the fossil record, we propose five fossil calibration points that can be used to date molecular phylogenetic trees of Bursidae.

INTRODUCTION

The importance of phylogenetic reconstruction no longer needs to be proven and the advances in molecular phylogeny have provided new possibilities for molluscan systematics, renowned for the difficulty of defining primary homologies based on shell characters [START_REF]Morphology should not be forgotten in the era of genomics a phylogenetic perspective[END_REF]. However, the use of fossils, for which only the shells are available, is unavoidable in modern evolutionary biology because they are the favoured means of calibration for dating molecular phylogenies. A time-tree goes beyond patterns, it gives information regarding tempo of evolution, and allows testing specific hypotheses (e.g., biogeography, diversification rate, etc.). For these reasons time-trees have become used commonly in systematics [START_REF] Donoghue P | Rocks and clocks: calibrating the Tree of Life using fossils and molecules[END_REF]. In the vast majority of cases researchers have used fossils to calibrate molecular trees, providing information to constrain nodes [START_REF] Donoghue P | Rocks and clocks: calibrating the Tree of Life using fossils and molecules[END_REF]. To do so most of them assume that the taxonomy reflects the correct phylogenetic position of the fossils, something that is difficult to justify. Furthermore, it may lead to under-estimation of uncertainty in the resulting node ages if conducted without proper homology hypotheses or phylogenetic assessment [START_REF] Pol D. & Laurin M | Incorporating phylogenetic uncertainty on phylogeny-based paleontological dating and the timing of turtle diversification[END_REF]. Lastly, a wrong node attribution could lead to huge distortions of divergence time [START_REF] Douzery E | Les datations moléculaires à l re de la génomique[END_REF]. Therefore, a review of the fossil record of the considered clade should be done before any analysis. [START_REF] Parham | Crossata (Gastropoda: Bursidae) in the eastern Pacific: a morphologic and paleontologic perspective[END_REF] discussed a set of data required to justify a fossil calibration. They are: 1) a museum numbers for the specimen (s) that demonstrate all the relevant characters and provenance data; 2) an apomorphy-based diagnosis of the specimen (s); 3) an explicit statement on the reconciliation of morphological and molecular data; 4) the locality and stratigraphic level (s) from which the calibrating fossil (s) was/were collected; and 5) a reference to a published radioisotopic age and/or numeric timescale and details of numeric age selection of the specimen (s). From all these criteria, the most difficult to satisfy when selecting fossils to calibrate a molluscan phylogeny is the third one, due to the homology issues cited above.

In this paper we review the fossil record of the Bursidae Thiele, 1925. This tonnoidean family is currently under systematic revision by the authors, who will produce a molecular phylogeny. In order to do it a re-evaluation of calibration points is required.

Bursidae are a moderately diverse group with 59 living species (Worms 2018). Recent bursids have a tropical and sub-tropical distribution with a southernmost record in eastern South Africa (Sanders et al. 2017), northern New Zealand (Spencer et al. 2016) and Walters shoal (Expeditions Muséum 2018). The northernmost record (Bursa scrobilator (Linnaeus, 1758)) is from Nice, southern France and Savona, north-western Italy [START_REF]The Early Pliocene Gastropoda (Mollusca) of Estepona, Southern Spain. Part 5: Tonnoidea, Ficoidea[END_REF]. They are intertidal to subtidal species, most often associated with hard substrates such as rocks, coral or sponges, where they can be locally abundant. They are active predators, feeding on polychaetes, sipunculids and echinoderms [START_REF] Fretter V | Some aspects of the functional anatomy and biology of Cymatium and Bursa[END_REF][START_REF] Taylor | Habitats and diet of predatory gastropods at Addu Atoll, Maldives[END_REF].

Bursidae are readily recognizable by their well-defined posterior exhalant siphon (anal notch) at the top of the outer lip, attached to the suture and by their thick, coarsely sculptured shells with knobs, warts, tubercles and nodules, hence their vernacular name: frog shells. Their size range is great; the smallest species, Bursa lucaensis (Parth, 1991) never exceeds 23 mm in height whereas Tutufa bardeyi (Jousseaume, 1881), with a height up to 450 mm and a width up to 300 mm, has one of the largest and most capacious shell of all gastropods [START_REF] Ho-On | Résultats des Campagnes MUSORSTOM volume 19. Indo-West Pacific Ranellidae, Bursidae and Personidae (Mollusca: Gastropoda), a monograph of the New Caledonian fauna and revisions of related taxa[END_REF]. Most bursids range in size between 40 and 60 mm.

As inferred from their protoconch, they are assumed to have a long to extremely long larval stage granting, for some of them, trans-oceanic dispersal capabilities. However, Sand- et al. 2004, 2009, 2011).

This paper aims to review the fossil record of Bursidae, discussing the exclusion of several species from the family. We will provide a morphological description, i.e., an apomorphy-based diagnosis sensu [START_REF] Parham | Crossata (Gastropoda: Bursidae) in the eastern Pacific: a morphologic and paleontologic perspective[END_REF], for all known extinct genera. In addition, we will list and discuss all the necessary information regarding their use as calibration points in phylogenies. Two new genera are described.

SHELL DESCRIPTION

Shells are described here using the methods of Sanders et al. (2017) adapted from Merle (2001[START_REF]The spiral cords of the Muricidae (Gastropoda, Neogastropoda): importance of ontogenetic and topological correspondences for delineating structural homologies[END_REF] for primary homology definition. Primary cords (P) are spiral ornamentations present from the first teleoconch whorl. Secondary cords (S) are spiral ornamentations appearing later during shell ontogeny. The shoulder cord is designated as P1. Cords of the convex part of the whorl are added from adapical to abapical (P2 to P8). The primary cord of the sutural ramp is designated IP and the secondary cord IS. Secondary cords on the convex part of the whorl are designated S1to S8. Cords of the anterior siphonal canal were not considered.

AbbreviAtions

Specimen repositories

EXCLUSION OF SPECIES FORMERLY INCLUDED IN BURSIDAE

The attribution of the Aptian (Hijochi and Hiraiga formations, Japan) species Hanaibursa aquilana (Parona, 1909) (Fig. 1A, B) to Bursidae by Kase (1984: fig. 16a, b) is most surprising considering that it bears few, if any, bursid characters. When published it was considered to be the oldest known bursid species. The relatively high-spired shell of H. aquilana with weak varix-like ridges every 90° shows greater similarity to batillariid gastropods such as Pyrazopsis Hacobjan, 1972, than with Tonnoidea. Furthermore, the posterior siphonal canal is not homologous to that of bursids, as it appears to be an adaxially extending fold within the outer lip. [START_REF] Riedel | An outline of cassoidean phylogeny (Mollusca, Gastropoda)[END_REF] was the first to notice this resemblance with cerithoideans; we agree with him on this superfamily position. Bursa saundersi Adegoke, 1977 (Fig. 1C,D) was described by Adegoke (1977: 209, pl. 31, figs 27-28) and recognized by [START_REF] Ho-On | Résultats des Campagnes MUSORSTOM volume 19. Indo-West Pacific Ranellidae, Bursidae and Personidae (Mollusca: Gastropoda), a monograph of the New Caledonian fauna and revisions of related taxa[END_REF][START_REF]Neogene tonnoidean gastropods of tropical and South America; a contribution to the Dominican Republic and Panama Paleontology projects and uplift of the Central American Isthmus[END_REF] as a possible Paleocene bursid. With aligned prominent varices and coarse spiral sculpture, the small specimen from the Ewekoro Formation (Selandian of Nigeria) looks a lot like a frog shell. Nevertheless, it lacks the posterior siphonal canal, the key character used to discriminate Bursidae from Ranellidae and Cymatiidae. For this reason, we conclude that this species does not belong to crown-Bursidae; [START_REF] Strong E. E | Frogs and tuns and tritons A molecular phylogeny and revised family classification of the predatory gastropod superfamily Tonnoidea (Caenogastropoda)[END_REF] suggested that this species possibly belongs in a new family along with Halgyrineum louisae (Lewis, 1974) and Ranella neuvillei Cossmann & Peyrot, 1924 (both currently included tentatively in Cymatiidae, following the phylogeny of [START_REF] Strong E. E | Frogs and tuns and tritons A molecular phylogeny and revised family classification of the predatory gastropod superfamily Tonnoidea (Caenogastropoda)[END_REF].

According to [START_REF]Taxonomy of gastropods of the families Ranellidae (= Cymatiidae) and Bursidae. Part 5. Early history of the families, with four new genera and recognition of the Family Personidae[END_REF][START_REF]Neogene tonnoidean gastropods of tropical and South America; a contribution to the Dominican Republic and Panama Paleontology projects and uplift of the Central American Isthmus[END_REF], there is only one longextinct genus within the family Bursidae: Olequahia Stewart, 1926, with five accepted species (Olequahia domenginica (Vokes, 1939), O. lincolnensis (Weaver, 1916), O. lorenzana (Wagner & Schilling, 1923), O. schencki Durham, 1944, and O. washingtoniana (Weaver, 1912)). Specimens presented here (Fig. 2) challenge this classification.

The type species of the genus, Olequahia washingtoniana (Fig. 2C-H), possesses an ornamentation pattern constituted mostly of spiral cords, the primary cords more prominent than the secondary ones. Nodules are consistently present on the shoulder. Two rows of nodules are present adapically; however, they fade into cords after half a whorl, an unusual feature among Bursidae, but found in Galeodea Link, 1807 (Cassidae). The low spire and inflated last whorl of O. washingtoniana are more reminiscent of shells of Cassidae or Tonnidae than of Ranellidae, Cymatiidae or Bursidae. In this species, varices are only present on the first whorl and as a terminal varix. According to [START_REF]Taxonomy of gastropods of the families Ranellidae (= Cymatiidae) and Bursidae. Part 5. Early history of the families, with four new genera and recognition of the Family Personidae[END_REF], pre-terminal varices are entirely lost in Oligocene species.

Varices are an important character because every single species of Bursidae possesses them even though they may be reduced in some cases (e.g. in Aspa marginata (Gmelin, 1791)). According to [START_REF] Strong E. E | Frogs and tuns and tritons A molecular phylogeny and revised family classification of the predatory gastropod superfamily Tonnoidea (Caenogastropoda)[END_REF] tonnoidean varices are most probably a plesiomorphic character (with an apomorphic loss due to varix dissolution during ontogeny in the clade Tonnidae + Cassidae). This lack of a varix on every whorl suggests that Olequahia may not be correctly referred to Bursidae.

However, Olequahia washingtoniana has a well-defined posterior siphonal canal resembling that of Bursidae. This siphonal canal is unusually large, similar to that in species of Crossata Jousseaume, 1881; this led some authors [START_REF]Taxonomy of gastropods of the families Ranellidae (= Cymatiidae) and Bursidae. Part 5. Early history of the families, with four new genera and recognition of the Family Personidae[END_REF][START_REF]Does the morphology of Pleistocene specimens of Crossata californica (Hinds, 1843) elucidate evolutionary patterns?[END_REF] to consider Olequahia as possibly ancestral to Crossata. We consider the posterior siphonal canal of Olequahia to be structurally different from that of Crossata (or any other Bursidae). The siphonal canal of Olequahia is formed by two rather large tubercules, one on the parietal part of the columella and one on the inner part of the outer lip. Columellar tubercules may occur in bursids (e.g. Bursa latitudo Garrard, 1961; see Fig. 3E[black arrow]), although never as large as in O. washingtoniana and, to our knowledge, there is no species of Bursidae with a single large tubercle or denticle inside the outer lip (in bursids there is either no denticle, or denticles are present throughout the outer lip). However, this feature occurs in several cassid species in the genus Galeodea (e.g., Galeodea hoaraui Drivas & Jay, 1989). In the light of our reinterpretation of these characters, we consider that O. washingtoniana is not a species of Bursidae.

Though many clues may indicate that Olequahia washingtoniana belongs in Galeodea, some characters do not point in that direction. The high-spired protoconch of this species (Beu 1988: fig. 1F) is quite different from the short, wide protoconch of one whorl in Galeodea; the columella is curved in Galeodea and straight in Olequahia; and finally, the wavy, frilly outer lip extension of O. washingtoniana is quite unique among tonnoideans. The position of O. washingtoniana in Galeodea, as a genus of Cassidae or even as a new family of Tonnoidea requires further attention but is beyond the scope of this paper.

Olequahia domenginica (Fig. 2A,B) is the oldest known species referred to the genus. The figured syntype has prominent varices every 200° or so over the entire teleoconch and is taller and narrower than its congeners. However, the general ornamentation resembles that of O. washingtoniana. Furthermore, the shell has no anal notch, casting doubts on its kinship with Bursidae. This shell is more reminiscent of a cymatiid such as Monoplex Perry, 1810. It is quite possible that it is more closely related to Monoplex than to O. washingtoniana. Either way, it is not a species of Bursidae. The other syntype of O. domenginica (UCNP 15803, not figured) is slightly larger than the illustrated specimen, but otherwise quite similar. However, the ventral side is not preserved in this specimen.

TACKLING BURSA IN THE FOSSIL RECORD

As currently understood, Bursa Röding, 1798 is very morphologically variable, so variable in fact that Beu (1998: 143) gave the broad generic description that Bursa is a large genus containing species with short to tall spires, weak to coarse sculpture, heavy and thick to light, thin shells, with varices variable in position and with or without a red colour area on the parietal lip One conclusion may be that Bursa is a trash genus, and exists only to classify species that do not fit anywhere else (this status was confirmed by Castelin et al. 2012 [Fig. 4] and [START_REF] Strong E. E | Frogs and tuns and tritons A molecular phylogeny and revised family classification of the predatory gastropod superfamily Tonnoidea (Caenogastropoda)[END_REF]. Being paraphyletic (or polyphyletic), such genera are an impediment for every systematist aiming to describe biodiversity in its historical dimension [START_REF]Strong effects of variation in taxonomic opinion on diversification analyses[END_REF].

To our knowledge there are 14 recognized extinct species of Bursidae, two-thirds of them referred to Bursa (Bursa amphitrites Maury, 1917, B. chipolana Schmelz, 1997, Ranella grateloupi d Orbigny, 1852, Apollon inaequicrenatus Cossmann & Peyrot, 1924, B. sangirana Beu, 2005, R. tuberosa Grateloup, 1833, B. victrix Dall, 1916, R. morrisi d rchiac & Haime, 1853and B. landaui Harzhauser, 2009) and the rest to Marsupina Dall, 1904(Bursa chira Olsson, 1930, Bursa (Marsupina) freya Olsson, 1932, Bursa chira var. yasila Olsson, 1930, Marsupina judensis Beu, 2010, and Gyrineum strongi Jordan, 1936).

Following Beu s (1988Beu s ( , 2010) ) opinion we consider Marsupina freya, M. judensis and M. strongi to be correctly attributed to the extant genus Marsupina.

As we want Bursa to be monophyletic, based on the phylogeny of [START_REF]Speciation patterns in gastropods with long-lived larvae from deep-sea seamounts[END_REF] and considering the position of the type species of the genus (B. bufonia (Gmelin, 1791)), we end up with a narrower definition of Bursa; it contains only B. bufonia and B. lamarckii (Deshayes, 1853), to which we can add B. luteostoma (Pease, 1861), B. rosa (Perry, 1811) and B. tuberosissima (Reeve, 1844), on the basis of their tubular and elongated posterior siphonal canal.

None of the fossil species mentioned above are closely similar to Bursa bufonia and none of them are type species of a previously published name that could be resurrected. So we are left with two options: either erect new genera (as few as possible) or propose phylogenetic hypotheses linking fossils with extant species (preferably type species).

Bursa sangirana Beu, 2005 (Fig. 5E), from the late Pliocene Kalibeng layers (Sangiran, central Java), has a low spire with a warty ornamentation. Its posterior siphonal canal is short; it is very reminiscent of Bursa rhodostoma (G. B. Sowerby II, 1835) (Fig. 3F) and is probably closely related to it. Both species may be included in the separate genus Lampadopsis Jousseaume, 1881 for which B. rhodostoma is the type species. The resurrection of this genus was suggested recently by [START_REF] Strong E. E | Frogs and tuns and tritons A molecular phylogeny and revised family classification of the predatory gastropod superfamily Tonnoidea (Caenogastropoda)[END_REF].

Bursa landaui Harzhauser, 2009 (Fig. 4C,D): this slender shell is only known by two external molds from the Aquitanian of Ras Tipuli, Lindi Bay, Tanzania. It has a very fine granulose ornamentation and a weakly expressed shoulder, slightly reminiscent of Bursa granularis (Röding, 1798) (Fig. 3G) and similar to Bursa condita (Fig. 3A). This may be an indication of the close relationship of the three species, for which there are two or possibly three available generic names: Colubrellina Fischer, 1884and Dulcerana Oyama, 1964. Bufonariella Thiele, 1929 is possibly another available name considering the close resemblance of its type species, Bursa scrobilator (Linnaeus, 1758), to B. granularis. Bursa victrix Dall, 1916: this species is only known from a poorly preserved external mold (USNM 166728) from the late Oligocene of Mascot Point, Flint River, Georgia, United States. The almost evenly inflated whorl profile, with a slightly protruding shoulder angle formed by a row of rounded nodules at approximately the upper third of the whorl height on the spire, the numerous lower rows of nodules extending down over the anterior canal, and the deeply buttressed abapertural face of the varix on the ultimate whorl are all characters in common with Bursa corrugata (Perry, 1811) (Fig. 3D). We follow Beu s (2010) opinion, keeping it as a species separate from B. corrugata pending the discovery of better-preserved specimens.

Among the remaining extinct bursid species, two are strikingly similar: Marsupina chira [START_REF]Contributions to the Tertiary paleontology of northern Peru: Part 3, Eocene Mollusca[END_REF] and M. yasila (2004,2009) and Beu (2010) to the conclusion that the earliest specimens of Aspa belong to another species, for which they considered the valid name to be Aspa subgranulata (d Orbigny, 1852) (Fig. 6). The exact upper range of A. subgranulata and the exact lower range of A. marginata are not well defined due to the use of the name A. depressa (Grateloup, 1833) for either of the two species. This species was originally considered as a transition species by Cossmann & Peyrot (1924), ranging from the Langhian to the Messinian. This species is currently considered a junior synonym of A. marginata (Landau et al. 2004A. marginata (Landau et al. , 2009;;[START_REF]Neogene tonnoidean gastropods of tropical and South America; a contribution to the Dominican Republic and Panama Paleontology projects and uplift of the Central American Isthmus[END_REF]. Further quantitative studies should be conducted to determine the exact ranges of the two (or three) species. Shell biconic, dorsoventrally compressed, shortspired, with 7 primary cords on the convex part of the whorl, all but P1 evenly reduced in variceal and intervariceal intervals of each whorl (P1 hardly more expressed than other cords); posterior siphonal canal short; varices strictly aligned; prominent columellar callus.

SYSTEMATICS

compArisons.

Olssonia n. gen. resembles Marsupina but possesses a lesser number of primary cords on the convex part of the whorl (7 in Olssonia n. gen., 8 in Marsupina). It resembles Aspa but has a much more prominent columellar callus, it is much more granulose, and it has a more sharply defined shoulder and a wider spire angle. Olssonia n. gen. resemble Bufonaria (Fig. 3B), with the same straight anterior siphonal canal, but lacks a spine or blade on the posterior siphonal canal (Fig. 3B; white arrow).

remArks The representatives of this genus are the oldest confirmed Bursidae; as such they can be used to calibrate the node Bursidae. Olssonia chira [START_REF]Contributions to the Tertiary paleontology of northern Peru: Part 3, Eocene Mollusca[END_REF] Olssonia yasila [START_REF]Contributions to the Tertiary paleontology of northern Peru: Part 3, Eocene Mollusca[END_REF] Derived from Aquitaine (Aquitania in Latin), the region where most of the species of the genus occur. species included. Aquitanobursa grateloupi (d Orbigny, 1852) n. comb., Aq. amphitrites (Maury, 1917), n. comb., Aq. inaequicrenata (Cossmann & Peyrot, 1924) n. comb., Aq. morrisi (d rchiac & Haimes, 1853) n. comb., Aq. tuberosa (Grateloup, 1833) n. comb.

diAgnosis.

Shell thick, moderately short-spired, varices slightly displaced, 6 primary cords on the convex part of the whorl, variceal and intervariceal P2 very reduced, axial ridges on the first whorl, parietal callus prominent. Posterior siphonal canal short, well-defined; outer lip with weak projection.

compArison.

With their deeply marked and angular varices and strongly expressed shoulder, Aquitanobursa n. gen. species resemble some deep-sea species of the Bursa s.l. genus such as Bursa quirihorai Beu, 1987 (Fig. 3H); however, Aquitanobursa n. gen. species are much strongly built, with a thicker shell. Atavistic features displayed by deep-sea gastropods are a well-known phenomenon, described recently for Muricidae by Merle (2012). The phylogenetic relationships of Aquitanobursa n. gen. with other Bursidae remain unclear; this genus is possibly a sister group to all Recent Bursidae, but that in unlikely considering that it occurs at the same time as Aspa marginata and Bursa corrugata. A fair assumption would be that it is the sister group to all bursids except Aspa and B. corrugata, but that would need a phylogenetic analysis for confirmation.

Aquitanobursa grateloupi

1852) n. comb. (Fig. 7A) Bursa grateloupi is only known from the lower Miocene of the Aquitaine Basin.

remArks Aquitanobursa grateloupi n. comb. was originally written gratteloupi. This name is obviously based on that of J.P.S. de Grateloup, and in accord with ICZN 32.5.1 the correct spelling is Aquitanobursa grateloupi n. comb. (Maury, 1917) n. comb.

Aquitanobursa amphitrites

(Fig. 5T) Occurs from the Late Miocene to Early Pliocene Gurabo Formation (Tortonian to Piacenzian, according to [START_REF] Denniston R | Caribbean chronostratigraphy refined with U-Pb dating of a Miocene coral[END_REF] in the Dominican Republic and from the early Middle Miocene Buenevara Adentro beds of the Paraguaná Peninsula, Venezuela (Beu 2010). (Schmelz, 1997) n. comb. (Fig. 5F,G) remArks Aquitanobursa chipolana n. comb. seems to have a slightly greater number of primary cords than other species referred to Aquitanobursa n. gen. This species possibly belongs in another genus, for which a new name needs to be created. (Cossmann & Peyrot, 1924) n. comb.

Aquitanobursa chipolana

Bursa (Bufonariella) chipolana

Aquitanobursa inaequicrenata

(Fig. 7B) Five specimens from Brongniart coll., three from le Peloua, Burdigalian, one from Saint-Paul-les-Dax (Cabannes), Burdigalian, one from Saubrigues, Burdigalian; one specimen from Jussieu coll., from Mérignac, Burdigalian; one specimen from Lhomme coll., from Saucats, Aquitanian; all housed in the collection de Paléontologie, MNHN.

geogrAphic And strAtigrAphic occurrence.

Only known from the Atlantic Aquitaine Basin of France (Aquitanian and Burdigalian).

Aquitanobursa morrisi

e, 1853) n. comb. (Fig. 5P-S Aquitanobursa tuberosa [START_REF]Tableau des coquilles fossiles qu rencontre dans les terrains calcaire tertiaires (faluns) des environs de Dax, département des Landes[END_REF] n. comb.

(Fig. 7C)

Ranella tuberosa [START_REF]Tableau des coquilles fossiles qu rencontre dans les terrains calcaire tertiaires (faluns) des environs de Dax, département des Landes[END_REF]: 92, no. 420. Bellardi 1873: 236, pl. 15, fig. 7. Ranella subtuberosa d Orbigny, 1852: 76, no. 1405. Apollon pelouatensis Cossmann & Peyrot, 1924: no. 842, pl. 15, figs 38-39, pl. 17 [START_REF]Révision des gastropoda (Mollusca) du Stratotype de l Aquitanien[END_REF]: 45, pl. 19, figs 3a-3b. Landau et al. 2004 : 68. type locAlity.

Bursa tuberosa

Ranella tuberosa and Apollon pelouatensis both from the Aquitanian of Saint-Paul-Lès-Dax, France. type mAteriAl.

Ranella tuberosa syntypes in Grateloup coll., University of Bordeaux, Talence, France (not seen); Apollon pelouatensis: the specimen MNHN.F.J06123 is here designated as lectotype. Three paralectotypes (MNHN.F.J06124, J06125, J06126). . other mAteriAl exAmined. Two specimens from Hofstetter coll., from le Peloua, Burdigalian, and one from Saubrigues, Burdigalian; 13 specimens from Brongniart coll., from le Peloua, Burdigalian; 15 specimens from Staadt coll., from le Peloua, Burdigalian; one specimen from Lhomme coll., from le Peloua, Burdigalian; two specimens (MNHN.F.A70577, A70578) from Gaas rde Rupelian; all in the collections de paléontologie, MNHN.

geogrAphic And strAtigrAphic occurrence.

Rupelian to middle Miocene of the Aquitaine basin (France) and Po Valley (Italy).

RECOMMENDATIONS FOR CALIBRATION

In regard to the systematic considerations, we can provide some recommendations for calibration of the molecular tree of the Bursidae. Numerical dating for this section is provided by the international geological time scale (Gradstein et al. 2012); a complete stratigraphic distribution chart for the Bursidae is shown in Figure 8. First occurrence and last occurrence data are provided in Appendix 1.

Considering that it is the oldest undisputable taxon included in Bursidae and apparently is not closely related to other clades within the family, the genus Olssonia n. gen. seems to be the best candidate for the calibration of the whole Bursidae. The oldest representative of the genus is O. yasila n. comb. from the early Bartonian (Olsson 1932[START_REF]Taxonomy of gastropods of the families Ranellidae (= Cymatiidae) and Bursidae. Part 5. Early history of the families, with four new genera and recognition of the Family Personidae[END_REF]) for which we propose to use an age of 41.2 Ma. As for all the following node calibrations, we propose to apply a truncated log normal distribution, as it places the highest probability on ages somewhat older than the fossil, diversification being necessarily older than the observed fossil record [START_REF] Ho S | Accounting for calibration uncertainty in phylogenetic estimation of evolutionary divergence times[END_REF].

The genus Aspa does not resemble any other Bursidae closely and, considering its very long stratigraphic history, possibly commencing before the Neogene [START_REF]The Early Pliocene Gastropoda (Mollusca) of Estepona, Southern Spain. Part 5: Tonnoidea, Ficoidea[END_REF], it may very well be a sister group to the rest of the Recent Bursidae. On the other hand, the fossil genus Aquitanobursa n. gen. looks like some Recent species such as B. quirihorai and B. granosa.

If Aspa were to branch at the base of the Bursidae tree, the clade containing every other bursid but excluding Aspa could be calibrated by Aquitanobursa n. gen. The oldest representative of the genus (two specimens MNHN.F.A70577, A70578) is A. tuberosa n. comb. from the Rupelian (Stampian) of the Aquitaine Basin. We propose an age of 27.82 Ma (Rupelian/ Chattian transition) for the node calibration.

Bursa, under the definition suggested in this paper, could be calibrated by the specimen resembling B. rosa from middle-late Miocene of Citalahab, West Java mentioned by Beu (2005: fig. 33). He attributed this specimen to B. rosa. Indeed, this specimen bears the long tubular posterior siphonal canal characteristic of the genus, but it is far less warty and could warrant erecting a new species, becoming thus the oldest species of all Bursa s.s. We propose an age of 9 Ma.

All the species of the former Bursa granularis complex (Sanders et al. 2017) could have their clade calibrated by the three specimens from Citalahab, West Java attributed by Beu (2005) to B. granularis, with an age of 9 Ma as for Bursa s.s. This calibration should not be blindly trusted, because we have not seen the specimens and they were not figured by [START_REF]Neogene fossil tonnoidean gastropods of Indonesia[END_REF]. The relationship of B. landaui to this clade remains too uncertain to be used as a calibration.

The genus Marsupina could be calibrated by the Shoal river formation specimen (USNM 647109), middle Miocene, figured by Beu (2010: pl. 9, fig. 6). This juvenile specimen strikingly resembling M. bufo, but has a more coarsely sculptured shell than later specimens of Marsupina, too much so to be conspecific. We propose an age of 14 Ma for the calibration of crown Marsupina. Strong et al.(2019) proposed to use the large specimen of Tutufa sp. Looking a lot like T. bubo figured by Tomida et al. (2013) as a calibration point for the genus Tutufa, calibrated at 13.5 Ma. We follow their recommendation.

No other node could be calibrated easily based on what we know currently of the systematics of Bursidae.

TIMING OF DIVERSIFICATION OF BURSIDAE BASED ON THE FOSSIL RECORD

Providing a scenario explaining the diversification of a highly dispersive family like the Bursidae is not an easy task, especially without a proper phylogenetic context. What we can say, however, is that the diversification of the Bursidae happened in three phases (Fig. 8).

Strong et al. (2018) estimated (with very low support regarding its relationship with its sister family) the origin of the family at around 113 Ma but, as stated above, we have no trace of morphologically recognizable bursids prior to the middle Eocene (40 Ma) Peruvian fauna. Following this record there is no other trace of Bursidae in the Paleogene. A possible explanation of this lack of fossils is a general diminution of fossil-rich (unlithified) marine outcrops in the late Paleogene [START_REF]Taphonomic Overprints on Phanerozoic Trends in Biodiversity: Lithification and Other Secular Megabiases, in Allison p[END_REF]. This tendency dramatically shifts in the Neogene, as we found an already well diversified fauna throughout the lower Miocene Tethyan realm. In the west (Aquitaine Basin) most species of the genus Aquitanobursa n. gen., Aspa subgranulata and Bursa corrugata occur. In the Mediterranean region (sensu [START_REF]Oligocene and Early Miocene gastropods from Kutch (NW India) document an early biogeographic switch from Western Tethys to Indo-Pacific[END_REF] Bursa corrugata and Bursa ranelloides occur. The presence of B. corrugata in both regions is a clear indication of a connection between the westernmost Tethys and the Atlantic. In the Eastern Proto-Indo-West-Pacific region, Bursidae are represented by Aquitanobursa morrisi n. comb. in the north (Pakistan) and by Bursa landaui in the southwest (Tanzania). The arrival of A. morrisi n. comb. in the Proto-Indo-West-Pacific region was most certainly through the Tethys, as most of the species of the genus are found in the Aquitaine Basin. On the other hand, the arrival B. landaui in the region is more difficult to explain.

The next phase of diversification happened during the middle Miocene with the closure of the Tethys Ocean in the east. Following this closure, we observed a relocation of the main coral biodiversity hotspot from the Mediterranean to the present-day coral triangle [START_REF] Cowman | atlantique européen au Cénozoïque moyen : diversité et évolution des gastéropodes[END_REF]. This hot spot shift could explain the radiation of the coral-dependent bursids (Tutufa, Bufonaria, Bursa s.s.). An eastward colonization of most of those species is probable, although some genera (e.g. Lampadopsis) possibly arrived from the Pacific.

In the west, also during the middle Miocene, Aspa marginata entered the Eastern Atlantic and Lampadopsis rugosa entered the Pacific Ocean from the Caribbean. The genus Aquitanobursa n. gen. entered the Western Atlantic (A. amphitrites n. comb.) and the genus Marsupina appeared. In the Proto-Mediterranean Atlantic region the species Bursa scrobilator appeared.

The high species-level diversity of tonnoideans in general during the middle Miocene was already pointed out by [START_REF]A Revision of the Tonnoidea (Caenogastropoda, Gastropoda) from the Miocene Paratethys and their palaeobiogeographic implications[END_REF].

At the Miocene-Pliocene transition we see the appearance of the last currently admitted genera: Bursina in the Indo-Pacific and Crossata in western America (there is an unconfirmed occurrence of Crossata in the middle Miocene [START_REF]Neogene tonnoidean gastropods of tropical and South America; a contribution to the Dominican Republic and Panama Paleontology projects and uplift of the Central American Isthmus[END_REF]Powell & Berschauer 2017).

Following the Messinian salinity crisis in the Mediterranean Sea, we observe the return of Bursa scrobilator to this domain. In the Recent fauna it is the only bursid species in the Mediterranean. Bursa corrugata returned briefly as well, but disappeared from the Mediterranean during the Pleistocene [START_REF]A Revision of the Tonnoidea (Caenogastropoda, Gastropoda) from the Miocene Paratethys and their palaeobiogeographic implications[END_REF]. It was also during the Pliocene that B. corrugata entered the western Atlantic, from which it reached the Eastern Pacific during the Pleistocene (Beu 2010). Aspa marginata reappeared briefly in the Mediterranean at the beginning of the Pliocene and reached its maximum geographic extension during the late Pliocene/early Pleistocene by entering the Western Atlantic. At present it occurs only in the Eastern Atlantic and at some mid-Atlantic islands.

CONCLUDING REMARKS

Dating biological events such as diversification or speciation in the fossil record remain a daunting task without a proper phylogenetic background, at least as much so as dating phylogenetic trees without proper fossil data. For this reason, reviews such as this one are paramount in order to understand the paleobiodiversity of a taxonomic group reliably. We followed as much as possible the recommendations of [START_REF] Parham | Crossata (Gastropoda: Bursidae) in the eastern Pacific: a morphologic and paleontologic perspective[END_REF] by systematically referring to the literature and by providing characters linking the fossils to extant clades and providing institution specimen numbers when possible. We hope the calibrations provided here will help unveil the Paleogene diversification pattern of the Bursidae, a period from which few fossils are known.

  fig. 1. -Taxa excluded from Bursidae: A, B, Hanaibursa aquilana (Parona, 1909) UMUT MM15655, Hiraiga Formation (Aptian of Japan), reproduced from Kase (1984: pl. 28, fig. 16a); A, dorsal view; B, ventral view; C, D, Bursa saundersi Adegoke, 1977, UIMG 394, holotype, Ewekoro Formation (Selandian of Nigeria), reproduced from Adegoke (1977: pl. 31, figs 27, 28); C, ventral view; D, dorsal view. Scale bar: 2 cm.

  fig. 2. -Taxa excluded from Bursidae. Olequahia Stewart, 1926 species: A, B, Olequahia domenginica (Vokes, 1939): UCMP 15804, syntype of Ranella domenginica, Tejon formation (Bartonian), California, United States; A, dorsal view; B, ventral view; C-H, Olequahia washingtoniana (Weaver, 1912): UCMP 16071 (loc. ID D8044) , in dorsal (C, F), left lateral (D, G) and ventral views (E, H), all from Cowlitz Formation (Priabonian), Cowlitz River, Washington, United States. Scale bar: 2 cm

  fig. 3. -Recent Bursidae: A, IM-2008-5350, Bursa condita (Gmelin,1791), « Nouvelle Calédonie », Jousseaume coll.; B, IM-2007-43072, Bufonaria perelegans Beu, 1987, Santo Marine Biodiversity Survey 2006, Vanuatu; C, IM-2009-11906, Bursa corrugata (Perry, 1811), Dakar'09, Senegal; D, IM-2007-43063, Tutufa rubeta (Linnaeus, 1758), Santo Marine Biodiversity Survey 2006, Vanuatu; E, IM-2007-40338, Bursa latitudo Garrard, 1961, Exploration de la Biodiversité et Isolement en mer du Corail, Nouvelle Calédonie; F, IM-2009-5150, Lampadopsis rhodostoma (G.B. Sowerby II, 1835), MAINBAZA, Mozambique channel; G, IM-2009-5148, Bursa granularis (Röding, 1798), juvenile, MAINBAZA, Mozambique channel; H, IM-2009-11653, Bursa quirihorai Beu, 1987, Exploration de la Biodiversité et Isolement en mer du Corail, New Caledonia. All are ventral views. Scale bars: A-D, 2 cm; E-H, 1 cm.

  fig. 5. -Extinct Bursidae species: A, B, Marsupina judensis Beu, 2010; NMB H 18308, holotype, late Miocene, Punta Judas, Pacific Costa Rica, reproduced from Beu (2010); C, D, "Bursa" landaui Harzhauser, 2009, NHMW 2007z0181/0029 (C) holotype and NHMW 2007z0181/0030 (D), both from the Aquitanian of Ras Tipuli, Lindi Bay, Tanzania, reproduced from Harzhauser (2009); E, Lampadopsis sangirana (Beu, 2005), RGM 456 230, holotype, late Pliocene of Kalibeng layers, Sangiran, central reproduced from Beu (2005); F, G, Aquitanobursa chipolana (Schmelz, 1997) n. comb., USNM 647108, paratype, specimen illustrated by Vokes (1973: text-figs 2a, b), coral reef facies of Chipola Formation (late Early Miocene), loc. TU547, reproduced from Beu (2010); H, I, Marsupina freya (Olsson, 1932), PRI 2312, holotype, "Zorritos Miocene," divide between Quebrada Conchudo Bravo and Quebrada Seca, Mancora, Peru; J, K, Olssonia chira (Olsson, 1930) n. comb., PRI 24257, holotype, Late Eocene/Oligocene of Chira Formation, Quercotilla, Peru; L-O, Olssonia yasila (Olsson, 1930) n. comb., PRI 24254 (L, M) and PRI 24255 (N, O), Middle Eocene Talara Formation, Yasila, Peru; P-S, Aquitanobursa morrisi (d'Archiac & Haime, 1853) n. comb., original representation of one of the syntypes by d'Archiac & Haime (1853) (P, R) and (NHMUK PI TG 27045) lectotype, from "Calcaire grossier jaune de la chaîne d'Hala" (Aquitanian?) Pakistan; T, Aquitanobursa amphitrites (Maury, 1917) n. comb., PRI 28763, holotype, Cercado Fm (Late Miocene), Maury's bluff 3, Cercado de Mao, Dominican Republic, reproduced from Beu (2010). Scale bar: 2 cm.

  fig.6. -Aspa subgranulata(d'Orbigny, 1852). MNHN.F.A70283 (Brongniart coll.), ventral(A), lateral left (B) and dorsal (C) views, Miocene of Le Peloua, Landes, France. Scale bar: 2 cm.

  fig. 7. -Aquitanobursa n. gen. of Aquitaine Basin: A, Aquitanobursa grateloupi (d'Orbigny, 1852) n. comb., MNHN.F.A27211, lectotype from the Aquitanian of Saint-Paul-les-Dax, France; B, Aquitanobursa inaequicrenata (Cossmann & Peyrot, 1924) n. comb., MNHN.F.A70284, Burdigalian of le Peloua, France; Faullummel coll.; C, Aquitanobursa tuberosa (Grateloup, 1833) n. comb., MNHN.F.A70285, Burdigalian of le Peloua, Staadt coll.

)

  Ranella d rchiac & Haime, 1853: 309, pl. XXX, fig. 1, pl XXXI, fig. 3. Ranella elegans Martin 1884: 137 (not G.B. Sowerby 2 nd , 1836 in G.B. Sowerby 1 st & G.B. Sowerby 2 nd , 1832-1841). Apollon morrisi Harzhauser et al. 2009: 360, pl. 1. type locAlity. re grossier jaune de la chaîne d Hala Pakistan. type mAteriAl. Types figured by d rchiac & Haime (1853) previously in UK Geological Survey, now in NHMUK. The complete specimen (NHMUK PI TG 27045) is here designated as the lectotype. One paralectotype (NHMUK PI TG 27046). geogrAphic And strAtigrAphic occurrence. This species is only known by d rchiac & Haime s (1853) specimens for the lower Miocene of Halla Mountain, Sindh, Pakistan.

  , figs 1-2. Bursa (Bufonariella) pelouatensis Beu 1981: 258. Schmelz 1997: 106, pl. 1, fig. 2.

  Bursa chipolanaSchmelz, 1997, and Ranella morrisi d rchiac &[START_REF]Description des animaux fossiles du groupe nummulitique de l Inde[END_REF] are here attributed to a new genus: Aquitanobursa n. gen. These new genera are described below. Bursa species in bold type are taxa that make the genus polyphyletic. Bracket on top indicates the phylogenetic extent of the genus Bursa if we consider the tubular posterior siphonal canal to be one of its synapomorphies, dotted bracket indicates the maximum extent of the genus considering the topology.

						Bursa s.s.
	Tutufa tenuigranosa	Tutufa rubeta	Bursa ranelloides	Bursa latitudo	Bursa rhodostoma	Bursa granularis	Bursa lamarckii	Bursa bufonia	Bursina gnorima	Bursina nobilis	Bufonaria perelegans	Bursa quirihorai	Bursa fijiensis	Bursa fosteri	Bursa awatii

[START_REF]Contributions to the Tertiary paleontology of northern Peru: Part 3, Eocene Mollusca[END_REF]

, for which we propose a new genus, Olssonia n. gen.

In the same fashion, Bursa amphitrites Maury, 1917, B. grateloupi

(d Orbigny, 1842), B. inaequicrenata (Coss- mann & Peyrot, 1923), B. tuberosa (Grateloup, 1833)

, fig. 4. -Phylogeny of the Bursidae. Topology based on the multigene Bayesian analysis of Castelin et al. (2012); black dots (•) represent well supported nodes (posterior probabilities over 0.98) and white dots (○) represent moderately supported nodes (PP over 0.90).

ON THE GENUS ASPA

Aspa is the most abundant fossil bursid. This is possibly due Family bursidAe Thiele, 1925 Genus Olssonia n. gen. to its extremely thick shell. Some authors

[START_REF]Malacofauna pliocenica Umbra[END_REF][START_REF]Middle Miocene (Badenian) gastropods from Korytnica, Poland. Part II[END_REF] Solsona 1998)

suggested that Aspa was monospecific with the extant species Aspa marginata ranging from the earliest Miocene (Aquitanian of Landes or Rupelian, France [Beu 2010]) to the Recent. However, early specimens possess a higher spire, coarser sculpture, a row of nodules at P1 persisting onto the last whorl and an obsolescent lower row of nodules, and larger nodules inside the outer lip than in Recent specimens of A. marginata. This led Landau et al.

  Olssonia n. gen. is a genus restricted to the Eocene to early Miocene of Peru.

	type species. Bursa chira Olsson, 1930.
	derivAtio nominis. Dedicated to the American palaeontologist
	A. A. Olsson.
	species included.	Olssonia chira (Olsson, 1930) n. comb.,
	O. yasila (Olsson, 1930) n. comb.
	distribution.
	Phylum MOLLUSCA Cuvier, 1795
	Class GASTROPODA Cuvier, 1797
	Superfamily tonnoideA Suter, 1913 (1825)
	(Conserved under ICZN Article 40b)

diAgnosis.

  Apollon grateloupi Cossmann & Peyrot 1924: 305, pl. XV, figs 42, 43. type locAlity. Lower Miocene of Aquitaine Basin, Bordeaux and Saint-Paul-lès-Dax, France.

	type mAteriAl.	MNHN.F.A27211 (from Saint-Paul-
	lès-Dax) is here designated as lectotype. One paralectotype
	(MNHN.F.B27595).
	other mAteriAl exAmined. 13 specimens from Brongniart coll.
	from le Peloua, Burdigalian.
	geogrAphic And strAtigrAphic occurrence.

  Bursa (Colubrellina) caelata amphitrites Aguilar in[START_REF]Sedimentología y paleoecología de un estuario del litoral pacifico del istmo centroamericano primordial (Mioceno medio, Costa Rica)[END_REF]: 64, appendix (not Maury, 1917).

	Bursa amphitrites Maury, 1917: 273, pl. 17, fig. 9.	Beu 2010:
	42, pl. 1, figs 1-4, 8.
	Bursa amphitrites E. Vokes 1973: 100 (in part = Marsupina bufo)
	(not Maury, 1917).
	type locAlity. Bluff 3, Cercado de Mao, Dominican Republic;
	Cercado Formation, late Miocene.
	type mAteriAl. Holotype (PRI 28763).
	geogrAphic And strAtigrAphic occurrence.

  Aquitanobursa chipolana n. comb. is recorded only from the Chipola Formation (Burdigalian) on and near the Chipola River, Calhoun Co., Florida, UnitedStates.

		geogrAphic And strAtigrAphic occurrence.
		Schmelz, 1997: 105, pl. 1, figs 1a-c;
	pl. 2, figs 1a-b, 2a-b. Vokes 1997: 212.
	Bursa (Bufonariella) pelouatensis Vokes 1973: 97, text-fig. 2 (not
	Cossmann & Peyrot, 1924).
	Bursa chipolana Beu 2010: 44, pl. 1, figs 5-7, 10.
	type locAlity.	Tulane University locality TU546, Chipola
	Formation (Burdigalian), Tenmile Creek, Florida, United States.
	type mAteriAl.	Bursa (Bufonariella) chipolana, holotype (UF
	73199), figured paratype (USNM 647108), from TU547, west
	bank Chipola River 600 m upstream from Fourmile Creek,
	Calhoun Co. (specimen figured by Vokes 1973: text-figs 2a-b,
	refigured in good quality by Beu 2010: pl. 1, figs 6, 7); figured
	paratype (UF 73200), from TU951, Tenmile Creek, Calhoun
	Co.; nine further paratypes from localities on or near the Chipola
	River listed by Schmelz (1997).

  Burdigal. Lan. Ser. Tortonian Mes. Zan. fig. 8. -Stratigraphic range of the extinct and extant Bursidae. Black arrows indicate total stratigraphic range; grey arrows indicate regional stratigraphic range. Abbreviations: P, Peru; WI, Western Indian Ocean; AB, Aquitaine Basin; CWA, Caribbean, Western Atlantic Ocean; PT, Paratethys, CWT: Central Western Tethys; EA, Eastern Atlantic; CWM, Central Western Mediterranean; WP, Western Pacific Ocean; J, Java, Indo-Pacific; S, Sumatra, Indo-Pacific; EP, Eastern Pacific Ocean. Grey squares regroup accepted genera and putative genera within Bursa.

	Eocene	Oligocene		Miocene		Plio.
	Barto. Priabon. Rupelian	Chattian Aqui.				Pia.
		P Olssonia chira, n. comb.
	P	Olssonia yasila, n. comb.	WI			Aquitanobursa morrisi, n. comb.
						AB			Aquitanobursa grateloupi, n. comb.
					AB				Aquitanobursa inaequicrenata, n. comb.
		?Aquitanobursa chipolana, n. comb.	CWA		CWA	Aquitanobursa amphitrites, n. comb.
				AB	AB	?			Aquitanobursa tuberosa, n. comb. Aspa subgranulata
					?
						PT		CWA
									EA	Aspa marginata
						CWT	CWM
						AB	PT		EP	CWA	Bursa corrugata
					CWT			CWM
							PT		WP	Tritonoranella ranelloides
						CWT		CWA
						J	?		Bufonaria crumena
						J	?		Bufonaria echinata
					S	?			Bufonaria elegans
									J	?	Bufonaria granosa
						J	?		Bufonaria rana
									J	?	Bursina margaricula
									J	?	Bursina thersites
							CWA		Lampadopsis rugosa
									EP
									CWA	Lampadopsis grayana
						CWA			Lampadopsis thomae
			Marsupina freya	P			CWA	EP	Marsupina nana
								?	EP	Marsupina bufo
							?	EP	?	Marsupina judensis
								J	?	Tutufa bubo
						J	?		Tutufa tenuigranosa
						J			Tutufa oyamai
							J	?	Tutufa rubeta
									EP	Crossata californica
									EP	Crossata ventricosa
						PT	CWT	CWM	CWM	Bursa scrobilator
									CWA
									CWA	Bursa cubaniana
				WI	Bursa landaui	J	?	Bursa granularis
							J	?	Bursa rosa
	40	35	30	25	20	15		10	5
		PALEOGENE			NEOGENE		QUAT.

  appenDix 1. -First occurrence and last occurrence data for all species of Bursidae. N.A., not applicable.

				Reference Reference Reference Reference			Reference Reference Reference Reference
				to the to the	to the to the	Last Last		for the for the	for the for the
	Species Species	First occurrence Formation First occurrence Formation	occurrence occurrence stratigraphyoccurrence stratigraphy occurrence	Formation Formation	occurrence occurrence stratigraphy stratigraphy
	Aquitanobursa Bursina	Burdigalian Pliocene	Chipola Tjikeusik	Beu 2010 Beu 2010 Burdigalian Beu 2005 Beu 2005 Extant	Chipola N.A.	Beu 2010 Beu 2010 N.A. N.A.
	chipolana margaricula		formation, [Cikeusik],				formation,
	n. comb. Aquitanobursa Crossata	Tortonian Tortonian	Florida Buenevara Java Tumbez Fm,	Beu 2010 Beu 2010 Zanclean Olsson Olsson 1932Extant	Florida Gurabo N.A.	Beu 2010 Beu 2010 N.A. N.A.
	amphitrites ventricosa		Adentro beds, Quebrada	1932; Beu			Formation,
	n. comb.		Venezuela				Dominican
	Aquitanobursa inaequicrenata n. comb. Marsupina bufo Serravallian Lower Burdigalian Aquitanobursa morrisi n. comb. Burdigalian Aquitanobursa tuberosa Bufonaria crumena Middle Miocene (early n. comb. Rupelian Aquitanian Marsupina freya Chattian/	Le Peloua, Aquitaine Basin, France Tucillal, Zorritos, Peru Shoal River Calcaire grossier de la chaine d'Hala, Pakistan formation, Florida Robba's locality CIJ1, Cijarian, Rica Pacific Costa Aquitaine basin, France Peninsula, Nicoya coast of Malpais, SW	Cossmann & Peyrot 1924 2010 Beu 2010 Gardner Lozouet 1998 D'Archiac & Haime 1853 Vredenburg 1925 Harzhauser et al. 2009 1947 Beu 2005 Beu 2005 Extant Lower Burdigalian Extant Burdigalian Lozouet 1998 Lozouet 1998 Lower Burdigalian Langhian Beu 2010 Beu 2010 Burdigalian/	republic Le Peloua, Aquitaine N.A. Basin, France Calacaire grossier dela chaine d'Hala N.A. Ecuador Aquitaine basin Lozouet Lozouet 1997 N.A. D'Archiac & Haime 1853 N.A. 1998 Basin, SW Progreso Formation, Subibaya Beu 2010 Beu 2010 Lozouet 1998 N.A. Vredenburg 1925 Harzhauser N.A. 1998 Lozouet et al. 2009
	Preangerian) Middle Miocene (early Preangerian) Bufonaria elegans Pleistocene (Early Bufonaria echinata Miocene) judensis Marsupina Late Miocene	western Java Robba's locality Nias Island, Sumatra Costa Rica Pacific coast of western Java of Quepos, CIJ1, Cijarian, 40 km WNW Punta Judas,	Beu 2005 Beu 2005 Extant Beu 2005 Beu 2005 Extant Beu 2010 Beu 2010 Late Miocene Punta Judas, N.A. Costa Rica Pacific coast of of Quepos, N.A. 40 km WNW	N.A. N.A. Beu 2010 Beu 2010 N.A. N.A.
	Bufonaria granosa Bufonaria rana Bursa corrugata Burdigalian Late Pliocene (Piacenzian?) Late Miocene Marsupina nana Zanclean	(Pakistan) Kalibeng layers, Kendeng beds, Jave Aquitaine basin, Costa Rica Pacific coast of Peninsula, Penita, Burica Quebrada La central Java Penita Fm, Sangiran, Group, Charco Azul	Beu 2005 Robba 1996 Cossmann & Landau 2005 et al. Sharaf Beu 2005 Beu 2005 Extant Extant Extant 1942 Beu 2010 Olsson Extant	N.A. N.A. N.A. N.A.	N.A. N.A. N.A. N.A.	N.A. N.A. N.A. N.A.
	Olssonia chira Bursa cubaniana Pleistocene Priabonian n. comb. Olssonia yasila Bartonian n. comb.	France Chira formation, Various Quercotilla, Peru formations talara formation, from Panama, Cuba, Peru	Peyrot 1924 Olsson 1930Stone 1949 Bartonian et al. 2009 Beu 2010 Beu 2010 Extant Olsson 1930Olsson 1930Bartonian	Chira Formation, Olsson 1930Stone 1949 N.A. N.A. Quercotilla, N.A. Peru Talara Formation Olsson 1930Olsson 1930 Peru
	Dominican Late Miocene? Pendopo Oilfield, Beu 2005 Robba 1996 Extant republic and 16 km east of Barbados Bursa granularis Late Miocene Tutufa bubo Citalahab, West Beu 2005 Beu 2005 Extant Talang Akar, Sumatra Java Aquitanobursa Burdigalian Bordeau d'Orbigny d'Orbigny Serravallian Tutufa oyamai Langhian/ Karas, Rembang, Beu 2005 Robba Extant	N.A. N.A. Tauziets N.A.	N.A. N.A. Lozouet N.A.	N.A. N.A. Lozouet N.A.
	grateloupi Bursa grayana Tutufa rubeta	Serravallian Messinian Late Miocene	surroundings, Aquitaine Ngrayong Formation, Basin, France Cercado Java formation, republic Java Dominican Citalahab, West	1852 Beu 2010 Beu 2010 Extant 1852 1996; Sharaf et al. 2005 Beu 2005 Beu 2005 Extant	Aquitaine basin, France N.A. N.A.	1997 N.A. N.A.	1998 N.A. N.A.
	Bursa landaui Tutufa	Aquitanian Miocene	Ras Tipuli Mount Butack,	Harzhauser Beu 2005 Beu 2005 Extant Harzhauser Aquitanian	Ras Tipuli N.A.	Harzhauser N.A.	Harzhauser N.A.
	Bursa rosa tenuigranosa	Late Miocene	Tanzania Citalahab, West Java Rembang, central Java	2009 Beu 2005 Beu 2005 Extant 2009	Tanzania N.A.	2009 N.A.	2009 N.A.
	Bursa rugosa Aspa	Burdigalian Rupelian	Cantaure, Gaas, Landes	Beu 2010 Beu 2010 Extant Beu 2010 Beu 2010 Middle	N.A. ?	N.A. Landau	N.A. Landau
	Bursa scrobilator Langhian/Serra-vallian subgranulata	Venezuela Paratethys Austria and	Landeau et al.	Baluk 1995 Extant Miocene	N.A.	N.A. 2004 et al.	N.A. et al. 2004
	Aspa marginata Burdigalian	Poland Po valley and	2004 Landau	Landau	Extant	N.A.	N.A.	N.A.
	Lampadopsis	Burdigalian	Chipola Paratethys	Beu 2010 Tulane fossil et al. et al.	Extant	N.A.	N.A.	N.A.
	thomae		formation,	2004,	localities 2004,
			Florida	2009	2009

appenDix 1. -Continuation.

Acknowledgements

This project was supported by the Labex BCDiv Diversités biologiques et culturelles funded by the CNRS, MNHN, ANR, Sorbonne Université, IRD and EPHE. We thank Erica Clites (UCMP) for the loan and the photos of the Olequahia material. For providing pictures we are very grateful to Leslie Skibinski (PRI), Philippe Loubry (MNHN) and Alexandre Lardeur. We are grateful to Pierre Lozouet (MNHN) for the donation of the Palaeogene material of the Aquitaine Basin. We are grateful to Alan Beu (GNS Sciences, Lower Hutt, New Zealand) and Bernard Landau (Naturalis Biocenter, Leiden, The Netherlands) for their fruitful comments. For curation of the MNHN.F material we thank Jean-Michel Pacaud (MNHN).