identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03D387DF1931FFF4FF12FAA7FB42FBD1.text	03D387DF1931FFF4FF12FAA7FB42FBD1.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Bursa Roding 1798	<div><p>TACKLING BURSA IN THE FOSSIL RECORD</p><p>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 Strong et al. 2018). Being paraphyletic (or polyphyletic), such genera are an impediment for every systematist aiming to describe biodiversity in its historical dimension (Faurby et al. 2016).</p><p>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 &amp; Peyrot, 1924, B. sangirana Beu, 2005, R. tuberosa Grateloup, 1833, B. victrix Dall, 1916, R. morrisi d’Archiac &amp; Haime, 1853 and 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).</p><p>Following Beu’s (1988, 2010) opinion we consider Marsupina freya, M. judensis and M. strongi to be correctly attributed to the extant genus Marsupina .</p><p>As we want Bursa to be monophyletic, based on the phylogeny of Castelin et al. (2012) 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.</p><p>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).</p><p>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 Strong et al. (2018).</p><p>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, 1884 and 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 .</p><p>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.</p><p>Among the remaining extinct bursid species, two are strikingly similar: Marsupina chira (Olsson, 1930) and M. yasila (Olsson, 1930), for which we propose a new genus, Olssonia n. gen.</p><p>In the same fashion, Bursa amphitrites Maury, 1917, B. grateloupi (d’Orbigny, 1842), B. inaequicrenata (Coss- mann &amp; Peyrot, 1923), B. tuberosa (Grateloup, 1833), Bursa chipolana Schmelz, 1997, and Ranella morrisi d’Archiac &amp; Haime, 1853 are here attributed to a new genus: Aquitanobursa n. gen. These new genera are described below.</p><p>Bursa s.s.</p><p>tenuigranosa rubeta ranelloides latitudo rhodostoma granularis lamarckii bufonia gnorima nobilis perelegans quirihorai fijiensis fosteri awatii</p><p>Tutufa Tutufa Bursa Bursa Bursa Bursa Bursa Bursa Bursina Bursina Bufonaria Bursa Bursa Bursa Bursa</p></div>	https://treatment.plazi.org/id/03D387DF1931FFF4FF12FAA7FB42FBD1	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Sanders, Malcolm T.;Merle, Didier;Puillandre, Nicolas	Sanders, Malcolm T., Merle, Didier, Puillandre, Nicolas (2019): A review of fossil Bursidae and their use for phylogeny calibration. Geodiversitas 41 (5): 247-265, DOI: 10.5252/geodiversitas2019v41a5
03D387DF193FFFF4FC65FBE4FC54FDE0.text	03D387DF193FFFF4FC65FBE4FC54FDE0.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Olssonia Sanders & Merle & Puillandre 2019	<div><p>Genus Olssonia n. gen.</p><p>urn:lsid:zoobank.org:act: 9BB6F0C9-71ED-4F96-BAE8-3BA325B57714</p><p>TYPE SPECIES. — Bursa chira Olsson, 1930 .</p><p>DERIVATIO NOMINIS. — Dedicated to the American palaeontologist A. A. Olsson.</p><p>SPECIES INCLUDED. — Olssonia chira (Olsson, 1930) n. comb., O. yasila (Olsson, 1930) n. comb.</p><p>DISTRIBUTION. — Olssonia n. gen. is a genus restricted to the Eocene to early Miocene of Peru.</p><p>DIAGNOSIS. — 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.</p><p>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).</p><p>REMARKS</p><p>The representatives of this genus are the oldest confirmed Bursidae; as such they can be used to calibrate the node Bursidae .</p></div>	https://treatment.plazi.org/id/03D387DF193FFFF4FC65FBE4FC54FDE0	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Sanders, Malcolm T.;Merle, Didier;Puillandre, Nicolas	Sanders, Malcolm T., Merle, Didier, Puillandre, Nicolas (2019): A review of fossil Bursidae and their use for phylogeny calibration. Geodiversitas 41 (5): 247-265, DOI: 10.5252/geodiversitas2019v41a5
03D387DF193DFFF6FF79FC04FF13FF79.text	03D387DF193DFFF6FF79FC04FF13FF79.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Olssonia chira (Olsson 1930) Sanders & Merle & Puillandre 2019	<div><p>Olssonia chira (Olsson, 1930) n. comb. (Fig. 5L, M)</p><p>Bursa chira Olsson, 1930: 62, pl. 10, figs 5-7,13.</p><p>Marsupina chira – Beu 1988: 71; 2010: 72.</p><p>TYPE MATERIAL. — Holotype (PRI 24257), one paratype (PRI 24263).</p><p>TYPE LOCALITY. — Late Eocene/Early Oligocene of Chira Formation, Quercotilla, Peru.</p><p>GEOGRAPHIC AND STRATIGRAPHIC OCCURRENCE. — Olssonia chira n. comb. seems to occur only on the Late Eocene/Early Oligocene of Peru.</p></div>	https://treatment.plazi.org/id/03D387DF193DFFF6FF79FC04FF13FF79	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Sanders, Malcolm T.;Merle, Didier;Puillandre, Nicolas	Sanders, Malcolm T., Merle, Didier, Puillandre, Nicolas (2019): A review of fossil Bursidae and their use for phylogeny calibration. Geodiversitas 41 (5): 247-265, DOI: 10.5252/geodiversitas2019v41a5
03D387DF193DFFF6FF7AFA46FE54FD4A.text	03D387DF193DFFF6FF7AFA46FE54FD4A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Olssonia yasila (Olsson 1930) Sanders & Merle & Puillandre 2019	<div><p>Olssonia yasila (Olsson, 1930) n. comb. (Fig. 5N, O)</p><p>Bursa chira var. yasila Olsson, 1930: 63, pl. 10, figs 3, 4.</p><p>Marsupina yasila – Beu 1988: 71.</p><p>TYPE LOCALITY. — Middle Eocene (Bartonian) Talara Formation, Yasila, Peru</p><p>TYPE MATERIAL. — Holotype (PRI 24254), with two paratypes (PRI 24255 and PRI 24262).</p><p>GEOGRAPHIC AND STRATIGRAPHIC OCCURRENCE. — Known only from the Bartonian of Peru.</p></div>	https://treatment.plazi.org/id/03D387DF193DFFF6FF7AFA46FE54FD4A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Sanders, Malcolm T.;Merle, Didier;Puillandre, Nicolas	Sanders, Malcolm T., Merle, Didier, Puillandre, Nicolas (2019): A review of fossil Bursidae and their use for phylogeny calibration. Geodiversitas 41 (5): 247-265, DOI: 10.5252/geodiversitas2019v41a5
03D387DF193DFFF0FEBCF861FDA2F8FC.text	03D387DF193DFFF0FEBCF861FDA2F8FC.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Aquitanobursa Sanders & Merle & Puillandre 2019	<div><p>Genus Aquitanobursa n. gen.</p><p>TYPE SPECIES. — Ranella grateloupi d’Orbigny, 1852 .</p><p>DERIVATIO NOMINIS. — Derived from Aquitaine (Aquitania in Latin), the region where most of the species of the genus occur.</p><p>SPECIES INCLUDED. — Aquitanobursa grateloupi (d’Orbigny, 1852) n. comb., Aq. amphitrites (Maury, 1917), n. comb., Aq. inaequicrenata (Cossmann &amp; Peyrot, 1924) n. comb., Aq. morrisi (d’Archiac &amp; Haimes, 1853) n. comb., Aq. tuberosa (Grateloup, 1833) n. comb.</p><p>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.</p><p>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.</p><p>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.</p><p>Aquitanobursa grateloupi (d’Orbigny, 1852) n. comb. (Fig. 7A)</p><p>Ranella gratteloupi [sic] d’Orbigny, 1852: 76.</p><p>Ranella leucostoma – Grateloup 1833: 91, pl VI (not Lamarck, 1822).</p><p>Ranella semigranosa – Grateloup 1845: pl. XXIX, fig. 6 (not Lamarck, 1822).</p><p>urn:lsid:zoobank.org:act: 69CEE556-A552-4FB2-AE6A-2FCB4374C21D</p><p>Apollon grateloupi – Cossmann &amp; Peyrot 1924: 305, pl. XV, figs 42, 43.</p><p>TYPE LOCALITY. — Lower Miocene of Aquitaine Basin, Bordeaux and Saint-Paul-lès-Dax, France.</p><p>TYPE MATERIAL. — MNHN.F.A27211 (from Saint-Paullès-Dax) is here designated as lectotype. One paralectotype (MNHN.F.B27595).</p><p>OTHER MATERIAL EXAMINED. — 13 specimens from Brongniart coll. from le Peloua, Burdigalian.</p><p>GEOGRAPHIC AND STRATIGRAPHIC OCCURRENCE. — Bursa grateloupi is only known from the lower Miocene of the Aquitaine Basin.</p><p>REMARKS</p><p>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.</p></div>	https://treatment.plazi.org/id/03D387DF193DFFF0FEBCF861FDA2F8FC	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Sanders, Malcolm T.;Merle, Didier;Puillandre, Nicolas	Sanders, Malcolm T., Merle, Didier, Puillandre, Nicolas (2019): A review of fossil Bursidae and their use for phylogeny calibration. Geodiversitas 41 (5): 247-265, DOI: 10.5252/geodiversitas2019v41a5
03D387DF193BFFF0FF3AFCC4FD67FF67.text	03D387DF193BFFF0FF3AFCC4FD67FF67.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Aquitanobursa amphitrites (Maury 1917) Sanders & Merle & Puillandre 2019	<div><p>Aquitanobursa amphitrites (Maury, 1917) n. comb. (Fig. 5T)</p><p>Bursa amphitrites Maury, 1917: 273, pl. 17, fig. 9. — Beu 2010: 42, pl. 1, figs 1-4, 8.</p><p>Bursa amphitrites – E. Vokes 1973: 100 (in part = Marsupina bufo) (not Maury, 1917).</p><p>Bursa (Colubrellina) caelata amphitrites – Aguilar in Seyfried et al. 1985: 64, appendix (not Maury, 1917).</p><p>TYPE LOCALITY. — Bluff 3, Cercado de Mao, Dominican Republic ; Cercado Formation, late Miocene.</p><p>TYPE MATERIAL. — Holotype (PRI 28763).</p><p>GEOGRAPHIC AND STRATIGRAPHIC OCCURRENCE. — Occurs from the Late Miocene to Early Pliocene Gurabo Formation (Tortonian to Piacenzian, according to Denniston et al. 2008) in the Dominican Republic and from the early Middle Miocene Buenevara Adentro beds of the Paraguaná Peninsula, Venezuela (Beu 2010).</p></div>	https://treatment.plazi.org/id/03D387DF193BFFF0FF3AFCC4FD67FF67	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Sanders, Malcolm T.;Merle, Didier;Puillandre, Nicolas	Sanders, Malcolm T., Merle, Didier, Puillandre, Nicolas (2019): A review of fossil Bursidae and their use for phylogeny calibration. Geodiversitas 41 (5): 247-265, DOI: 10.5252/geodiversitas2019v41a5
03D387DF193BFFF0FF3BFA46FC0BF83F.text	03D387DF193BFFF0FF3BFA46FC0BF83F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Aquitanobursa chipolana (Schmelz 1997) Sanders & Merle & Puillandre 2019	<div><p>Aquitanobursa chipolana (Schmelz, 1997) n. comb. (Fig. 5F, G)</p><p>Bursa (Bufonariella) chipolana Schmelz, 1997: 105, pl. 1, figs 1a-c; pl. 2, figs 1a-b, 2a-b. — Vokes 1997: 212.</p><p>Bursa (Bufonariella) pelouatensis – Vokes 1973: 97, text-fig. 2 (not Cossmann &amp; Peyrot, 1924).</p><p>Bursa chipolana – Beu 2010: 44, pl. 1, figs 5-7, 10.</p><p>TYPE LOCALITY. — Tulane University locality TU546, Chipola Formation (Burdigalian), Tenmile Creek, Florida, United States.</p><p>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) .</p><p>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, United States.</p><p>REMARKS</p><p>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.</p></div>	https://treatment.plazi.org/id/03D387DF193BFFF0FF3BFA46FC0BF83F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Sanders, Malcolm T.;Merle, Didier;Puillandre, Nicolas	Sanders, Malcolm T., Merle, Didier, Puillandre, Nicolas (2019): A review of fossil Bursidae and their use for phylogeny calibration. Geodiversitas 41 (5): 247-265, DOI: 10.5252/geodiversitas2019v41a5
03D387DF193BFFF0FC5EFD85FC73FF44.text	03D387DF193BFFF0FC5EFD85FC73FF44.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Aquitanobursa inaequicrenata (Cossmann & Peyrot 1924) Sanders & Merle & Puillandre 2019	<div><p>Aquitanobursa inaequicrenata</p><p>(Cossmann &amp; Peyrot, 1924) n. comb. (Fig. 7B)</p><p>Apollon inaequicrenatus Cossmann &amp; Peyrot, 1924: 311 pl. 15, figs 44-45.</p><p>Bursa (Bufonariella) inaequicrenata – Beu 1981: 258.</p><p>Bursa inaequicrenata – Landau et al. 2004: 68; 2009: 76.</p><p>TYPE LOCALITY. — Burdigalian of Le Peloua, near Saucats (Gironde), Aquitaine Basin, France.</p><p>TYPE MATERIAL. — Apollon inaequicrenatus, holotype (MNHN.F. J06127 Cossmann coll.) from the Burdigalian of Le Peloua and one paratype (MNHN.F. J06128 Cossmann coll.).</p><p>OTHER MATERIAL EXAMINED. — 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.</p><p>GEOGRAPHIC AND STRATIGRAPHIC OCCURRENCE. — Only known from the Atlantic Aquitaine Basin of France (Aquitanian and Burdigalian).</p></div>	https://treatment.plazi.org/id/03D387DF193BFFF0FC5EFD85FC73FF44	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Sanders, Malcolm T.;Merle, Didier;Puillandre, Nicolas	Sanders, Malcolm T., Merle, Didier, Puillandre, Nicolas (2019): A review of fossil Bursidae and their use for phylogeny calibration. Geodiversitas 41 (5): 247-265, DOI: 10.5252/geodiversitas2019v41a5
03D387DF1938FFEDFF3FFF2AFD28FCE3.text	03D387DF1938FFEDFF3FFF2AFD28FCE3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Aquitanobursa tuberosa (Grateloup 1833) Sanders & Merle & Puillandre 2019	<div><p>Aquitanobursa tuberosa (Grateloup, 1833) n. comb . (Fig. 7C) Ranella tuberosa Grateloup, 1833: 92, no. 420. — Bellardi 1873: 236, pl. 15, fig. 7. Ranella subtuberosa d’Orbigny, 1852: 76, no. 1405. Apollon pelouatensis Cossmann &amp; Peyrot, 1924: no. 842, pl. 15, figs 38-39, pl. 17, figs 1-2.</p><p>Bursa (Bufonariella) pelouatensis – Beu 1981: 258. — Schmelz 1997: 106, pl. 1, fig. 2. Bursa tuberosa – Lozouet et al. 2001: 45, pl. 19, figs 3a-3b. — Lan- dau et al. 2004: 68. TYPE LOCALITY. — 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 peloua- tensis: 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 “Lagouarde”, 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</p><p>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, Beu 1988) 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 (Ho &amp; Phillips 2009). The genus Aspa does not resemble any other Bursidae closely and, considering its very long stratigraphic history, possibly commencing before the Neogene (Landau et al. 2004), 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 representa- tive 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.</p><p>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 spe- cies of all Bursa s.s. We propose an age of 9 Ma.</p><p>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 Beu (2005). 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, fig- ured by Beu (2010: pl. 9, fig. 6). This juvenile specimen strik- ingly 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 .</p><p>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 hap- pened 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 (Hendy 2011). This tendency dramatically shifts in the Neogene, as we found an already well diversified fauna throughout the lower Miocene Tethyan realm.</p><p>In the west (Aquitaine Basin) most species of the genus Aquitanobursa n. gen., Aspa subgranulata and Bursa corrugata occur. In the “Mediterranean region” (sensu Harzhauser et al. 2002) 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.</p><p>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 (Leprieur et al. 2016). 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.</p><p>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.</p><p>The high species-level diversity of tonnoideans in general during the middle Miocene was already pointed out by Landau et al. (2009).</p><p>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 (Beu 2010; Powell &amp; Berschauer 2017).</p><p>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 (Landau et al. 2009). 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.</p></div>	https://treatment.plazi.org/id/03D387DF1938FFEDFF3FFF2AFD28FCE3	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Sanders, Malcolm T.;Merle, Didier;Puillandre, Nicolas	Sanders, Malcolm T., Merle, Didier, Puillandre, Nicolas (2019): A review of fossil Bursidae and their use for phylogeny calibration. Geodiversitas 41 (5): 247-265, DOI: 10.5252/geodiversitas2019v41a5
