taxonID	type	description	language	source
03D387DF1931FFF4FF12FAA7FB42FBD1.taxon	description	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’Archiac & 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). Following Beu’s (1988, 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 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. 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).	en	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
03D387DF1931FFF4FF12FAA7FB42FBD1.taxon	description	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. 3 D). 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 (Olsson, 1930) and M. yasila (Olsson, 1930), 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), Bursa chipolana Schmelz, 1997, and Ranella morrisi d’Archiac & Haime, 1853 are here attributed to a new genus: Aquitanobursa n. gen. These new genera are described below. Bursa s. s. tenuigranosa rubeta ranelloides latitudo rhodostoma granularis lamarckii bufonia gnorima nobilis perelegans quirihorai fijiensis fosteri awatii Tutufa Tutufa Bursa Bursa Bursa Bursa Bursa Bursa Bursina Bursina Bufonaria Bursa Bursa Bursa Bursa	en	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.taxon	materials_examined	TYPE SPECIES. — Bursa chira Olsson, 1930. DERIVATIO NOMINIS. — Dedicated to the American palaeontologist A. A. Olsson.	en	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.taxon	description	SPECIES INCLUDED. — Olssonia chira (Olsson, 1930) n. comb., O. yasila (Olsson, 1930) n. comb.	en	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.taxon	distribution	DISTRIBUTION. — Olssonia n. gen. is a genus restricted to the Eocene to early Miocene of Peru.	en	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.taxon	diagnosis	DIAGNOSIS. — Shell biconic, dorsoventrally compressed, shortspired, with 7 primary cords on the convex part of the whorl, all but P 1 evenly reduced in variceal and intervariceal intervals of each whorl (P 1 hardly more expressed than other cords); posterior siphonal canal short; varices strictly aligned; prominent columellar callus. 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. 3 B), with the same straight anterior siphonal canal, but lacks a spine or blade on the posterior siphonal canal (Fig. 3 B; white arrow).	en	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.taxon	discussion	REMARKS The representatives of this genus are the oldest confirmed Bursidae; as such they can be used to calibrate the node Bursidae.	en	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.taxon	materials_examined	TYPE MATERIAL. — Holotype (PRI 24257), one paratype (PRI 24263). TYPE LOCALITY. — Late Eocene / Early Oligocene of Chira Formation, Quercotilla, Peru. GEOGRAPHIC AND STRATIGRAPHIC OCCURRENCE. — Olssonia chira n. comb. seems to occur only on the Late Eocene / Early Oligocene of Peru.	en	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.taxon	materials_examined	TYPE LOCALITY. — Middle Eocene (Bartonian) Talara Formation, Yasila, Peru TYPE MATERIAL. — Holotype (PRI 24254), with two paratypes (PRI 24255 and PRI 24262). GEOGRAPHIC AND STRATIGRAPHIC OCCURRENCE. — Known only from the Bartonian of Peru.	en	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.taxon	materials_examined	TYPE SPECIES. — Ranella grateloupi d’Orbigny, 1852. DERIVATIO NOMINIS. — Derived from Aquitaine (Aquitania in Latin), the region where most of the species of the genus occur.	en	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.taxon	description	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’Archiac & Haimes, 1853) n. comb., Aq. tuberosa (Grateloup, 1833) n. comb.	en	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.taxon	diagnosis	DIAGNOSIS. — Shell thick, moderately short-spired, varices slightly displaced, 6 primary cords on the convex part of the whorl, variceal and intervariceal P 2 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. 3 H); 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.	en	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.taxon	description	(Fig. 7 A)	en	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.taxon	description	Apollon grateloupi – Cossmann & Peyrot 1924: 305, pl. XV, figs 42, 43.	en	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.taxon	materials_examined	TYPE LOCALITY. — Lower Miocene of Aquitaine Basin, Bordeaux and Saint-Paul-lès-Dax, France. TYPE MATERIAL. — MNHN. F. A 27211 (from Saint-Paullès-Dax) is here designated as lectotype. One paralectotype (MNHN. F. B 27595).	en	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.taxon	description	GEOGRAPHIC AND STRATIGRAPHIC OCCURRENCE. — Bursa grateloupi is only known from the lower Miocene of the Aquitaine Basin.	en	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.taxon	discussion	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.	en	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.taxon	materials_examined	TYPE LOCALITY. — Bluff 3, Cercado de Mao, Dominican Republic; Cercado Formation, late Miocene. TYPE MATERIAL. — Holotype (PRI 28763). 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).	en	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.taxon	materials_examined	TYPE LOCALITY. — Tulane University locality TU 546, Chipola Formation (Burdigalian), Tenmile Creek, Florida, United States. TYPE MATERIAL. — Bursa (Bufonariella) chipolana, holotype (UF 73199), figured paratype (USNM 647108), from TU 547, west bank Chipola River 600 m upstream from Fourmile Creek, Calhoun Co. (specimen figured by Vokes 1973: text-figs 2 a-b, refigured in good quality by Beu 2010: pl. 1, figs 6, 7); figured paratype (UF 73200), from TU 951, Tenmile Creek, Calhoun Co.; nine further paratypes from localities on or near the Chipola River listed by Schmelz (1997). 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.	en	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.taxon	discussion	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.	en	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.taxon	description	Apollon inaequicrenatus Cossmann & Peyrot, 1924: 311 pl. 15, figs 44 - 45.	en	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.taxon	materials_examined	TYPE LOCALITY. — Burdigalian of Le Peloua, near Saucats (Gironde), Aquitaine Basin, France. TYPE MATERIAL. — Apollon inaequicrenatus, holotype (MNHN. F. J 06127 Cossmann coll.) from the Burdigalian of Le Peloua and one paratype (MNHN. F. J 06128 Cossmann coll.). 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.	en	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.taxon	description	GEOGRAPHIC AND STRATIGRAPHIC OCCURRENCE. — Only known from the Atlantic Aquitaine Basin of France (Aquitanian and Burdigalian).	en	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.taxon	description	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. 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. 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. 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 Landau et al. (2009). 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 & 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 (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.	en	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
