Cuneolina, d'Orbigny, 1839

Cuneolina View in CoL ex gr. pavonia d’Orbigny, 1846

Figure 17 View Fig

1839 Cuneolina pavonia n. sp. – d’Orbigny, p. 151; Nomen nudem – no description.

T 1846 Cuneolina pavonia – d’Orbigny, p.253, figs. 50- 52; Turonian (reinterpreted as Cenomanian), western France.

1850 Cuneolina conica n. sp. – d’Orbigny, p. 186; Turonian (reinterpreted as Cenomanian), western France.

1900 Cuneolina conica – Schlumberger, p. 461-462, pl 8, figs. 8-10; Turonian (reinterpreted as Cenomanian), western France.

1947 Cuneolina walteri n. sp. – Cushman & Applin, p. 30, pl. 10, figs 4-5; middle Cenomanian, Florida

1948b Cuneolina pavonia var. parva var. nov. – Henson, pp. 624-627, pl. XIV, figs. 1-6; pl. XVII, figs. 7-12; pl. XVIII, figs. 12-14; Santonian (variously reinterpreted as Albian or Turonian), Egypt.

1961 “Cuneolinas” – Cuvillier, pl. XLII, fig. 1; “ Lower Senonian ”, Aquitaine, France .

1962 Cuneolina pavonia parva – Sartoni & Crescenti, p. 278-279, pl. 31, fig. 2; pl. 32; pl. 47, figs. 4-6; Albian – intra-Late Cretaceous, southern Italy.

1964 Cuneolina pavonia – Loeblich & Tappan, figs. 193/1-2; Cenomanian, western France.

1964 Cuneolina sp. – Bozorgnia & Banafti, pl. LXXVII, fig. 2; late Albian – Cenomanian, central Iran.

1964 Cuneolina sp. – Bozorgnia & Banafti, pl. LXXX, fig. 2; Cenomanian, Iranian Zagros.

1964 Cuneolina cf. hensoni – Bozorgnia & Banafti, pl. LXXIX, fig. 1; Cenomanian, Iranian Zagros.

? 1965 Cuneolina pavonia parva – Gollesstaneh, p. 165- 166, pl. 24, figs. 1-4; Early Cretaceous (precise age uncertain), Iranian Zagros.

1965 Cuneolina walteri – Applin & Applin, pl. 2, figs. 1- 2; middle Cenomanian, Florida .

1967 Cuneolina conica – Bismuth et al., pl. 12, fig. 8; Cenomanian, Tunisia.

1967 Cuneolina pavonia – Neumann, pl. 53, fig. 1; Cenomanian, western France.

1968 Cuneolina conica – Gendrot, p. 676-677, pl. 4, fig. 16; Santonian, southern France.

1968 Cuneolina pavonia – Gendrot, p. 676-677, pl. 4, figs. 17-19; Santonian, southern France.

1969 Cuneolina sp. – Sampò, pl. XXXVII, fig. 16; pl. XLII, figs. 13, 17; Aptian, Cenomanian, Iranian Zagros.

1973 Cuneolina pavonia-parva – Berthou, p. 2, fig. 3; late Cenomanian, Portugal.

1974 Cuneolina pavonia – Saint-Marc, p. 220, pl. 2, fig. 1; Albian – Turonian, Lebanon.

1974 Cuneolina gr. pavonia – Bignot & Poisson, pl. 1, fig. 1; pl. 2, fig. 1; Cenomanian, Turkish Taurides.

? 1976 Iraqia simplex Henson – Kalantari , pl. 15, fig. 10; Albian, Iranian Zagros.

1976 Cuneolina cf. pavonia – Kalantari, pl. 18; pl. 22, fig. 5; Cenomanian, Iranian Zagros.

1976 Cuneolina gr. pavonia – Decrouez, p. 79-82, pl. 14, figs. 3-4, pl. 17, figs. 1, 5; Late Cretaceous, Greece [ fide Cvetko Tešović et al., 2001 ]

1977 Cuneolina pavonia – Rey et al., p. 381, pl. 3, figs. 8-9; late Albian, Portugal.

1977 Cuneolina pavonia parva – Velić, pl. 29, figs. 1-3, 5; early Albian, Croatia.

1977 Cuneolina pavonia – Velić, pl. 29, figs. 4, 6-7; early Albian, Croatia.

1977 Cuneolina ex. gr. pavonia parva – Velić, pl. 31, figs. 2-4; late Albian, Croatia.

1978 Cuneolina gr. pavonia – Decrouez, pl. 1, figs. 2-3; early – late Cenomanian, Greece.

1978 Cuneolina sp. – Luperto Sinni & Richetti, pl. 46, figs. 1-11; Santonian – Maastrichtian, southern Italy.

1979 Cuneolina pavonia parva Sartoni & Crescenti [sic] – Bachmann & Risch, pl. 8, figs. 7-9; early Cenomanian, Greece.

1981 Cuneolina pavonia – Bismuth et al., pl. 1, figs. 7-8; late Cenomanian, Tunisia.

1981 Cuneolina pavonia – Saint-Marc, pl. 1, fig. 4; Albian – Turonian, Lebanon.

1982 Cuneolina gr. pavonia – Altıner & Decrouez, pl. 3, figs. 13-14; Aptian – Cenomanian, Turkish Taurides.

1982 Cuneolina pavonia – Mouty & Saint-Marc, pl. 1, fig. 9; latest Aptian – Cenomanian, Syria.

1984 Cuneolina gr. pavonia – Chiocchini et al., pl. 5, figs. 1-3; mid-Cretaceous, Italy.

1985 Cuneolina conica – Bilotte, pl. 5, fig. 3; Cenomanian, French Pyrenees.

1985 Cuneolina gr. pavonia-parva – Bilotte, pl. 5, fig. 4; late Cenomanian, French Pyrenees.

1988 Cuneolina gr. pavonia – Kuss & Schlagintweit, p. 83, pl. 18, fig. 5; pl. 20, figs. 9-10; latest Aptian – early Cenomanian, Sinai, Egypt.

1988 Cuneolina pavonia – Sartorio & Venturini, p. 109, 112, 113; Albian-Santonian; Italy.

1988 Cuneolina pavonia – Sartorio & Crescenti, p.113; “Lower Senonian”, southern Italy.

1990 Cuneolina pavonia – Weidich & Al-Harithi, p. 604, pl. 3, fig. 6; pl. 4, fig. 24; late Albian – Cenomanian, Jordan.

1990 Cuneolina pavonia – Šribar & Pleničar, pl. 4, fig. 5; late Turonian, Slovenia.

1991 Cuneolina pavonia parva – Schlagintweit, p. 36, pl. 11, figs. 13-16; late Aptian – early Albian, Austria.

1991 Cuneolina walteri – Scott & Gonzalez-Leon, p. 58, fig. 5O; Albian, Mexico.

1992 Cuneolina ex gr. pavonia – Schlagintweit, p. 336- 337, text-fig. 6; pl. 2, figs. 1-6; Cenomanian – Coniacian, Austria. Demonstrates the initial chambers are planispiral rather than trochospiral.

1993 Cuneolina pavonia – Grötsch et al., fig. 5B-E; late Albian, Slovenia.

1995 Cuneolina sp. cf. C. pavonia – Arnaud-Vanneau & Sliter, p. 554, pl.4, figs. 1-5; late Albian, Mid-Pacific seamounts.

1995 Cuneolina parva – Arnaud-Vanneau & Sliter, p.554, pl, 5, figs. 6-9; late Albian, Mid-Pacific seamounts.

1998 Cuneolina pavonia parva – Whittaker et al., p. 28- 29; pl. 4, figs. 1-2; pl. 45, figs. 7-8; pl. 46, figs. 1-6; Type material of Henson (1948b) of uncertain age (see text), Egypt .

1999 Cuneolina walteri – Scott & Finch, fig. 4A-C; late Albian, Honduras.

2000 Cuneolina pavonia – Aguilera-Franco, p. 160; middle – late Cenomanian, Mexico.

2000 Cuneolina conica – Aguilera-Franco, p. 161; middle – late Cenomanian, Mexico.

2000 Cuneolina pavonia – Benedetti et al., fig. 54; Late Cretaceous, Montenegro.

2001 Cuneolina pavonia – Cvetko Tešović et al., p. 601- 602, fig. 6B-C; Campanian, Brac Island, Croatia.

2004 Cuneolina gr. pavonia – Menegatti, p. 2/24-2/25, pl. 7, fig. 5; pl. 8, figs. 2-3, 6; pl. 9, fig. 5; Cenomanian, Dubai.

? 2005 Cuneolina pavoni [sic] – Vaziri et al., pl. 2, figs 5- 7; Campanian – Maastrichtian, central Iran. [Fragmentary specimens].

2006 Cuneolina pavonia – Husinec & Sokač, fig. 9 H- L; Albian, Croatia.

2006 Cuneolina pavonia – Taslı et al., fig. 6L; middle – late Cenomanian, Turkish Taurides.

2009 Cuneolina pavonia – Sari et al., pl. 3, fig. 7; middle Cenomanian – Coniacian, Turkish Taurides.

? 2009 Cuneolina pavonia – Shirazi et al., pl. 1, figs. 10- 11; Albian – Cenomanian, Iranian Zagros [Fragmentary specimens].

? 2009 Nakkadyia awadi n. gen, n. sp. – Ismail et al., 402-403; text-fig. 4; pl. 3, figs. 1-5; pl. 4, figs. 2-4; Cenomanian, Egypt.

2010 Cuneolina pavonia – Spalluto & Caffau, fig. 13AB; Albian – Cenomanian, southern Italy.

Non 2011 Cuneolina pavonia – Shirazi et al., pl. 2, fig. 16; Albian, Iranian Zagros. [=Indeterminate textulariid].

2011 Cuneolina parva – Filkorn & Scott, p. 186, figs. 4.9-4.12; late Albian, Mexico.

? 2011 Cuneolina pavonia – Roozbahani, pl. 1, fig. 11; Albian, central Iran. Indeterminate fragment.

? 2011 Cuneolina walteri – Roozbahani, pl. 2, fig. 3; Albian, central Iran. Indeterminate fragment.

2012 Cuneolina pavonia – Ghanem et al., fig. 6c/3, 8, 11; fig. 6d/1-2; Albian – Cenomanian, Syria.

2012 Cuneolina pavonia – Chiocchini et al., pl. 96; middle Cenomanian, southern Italy.

2012 Cuneolina sp. 1 – Chiocchini et al., pl. 82; early Cenomanian, southern Italy.

2012 Cuneolina pavonia – Orabi et al., fig. 5A, C; late Cenomanian, Egypt.

2012 Cuneolina parva – Orabi et al., fig. 5B; late Cenomanian, Egypt.

2012 Cuneolina cylindrica Henson – Orabi et al. , fig.5 E; late Cenomanian, Egypt. [May be an oblique and/or partial section].

2012 Cuneolina pavonia – Spalluto, fig. 4a; Albian – Cenomanian, southern Italy.

2013 Cuneolina parva – Ghanem & Kuss, fig. 10/13; fig. 12/13; Albian – Cenomanian, Syria.

2013 Cuneolina pavonia – Ghanem & Kuss, fig. 11/2-4; Albian – Cenomanian, Syria.

2013 Dicyclina sp. – Ghanem & Kuss, fig. 12/30; Albian – Cenomanian, Syria.

2013 Cuneolina pavonia – Omaña et al., pl. 1, fig. 2; pl. 5, fig. 5; late Cenomanian, Mexico.

? 2013 Cuneolina aff. C. parva – Hfaiedh et al., fig. 12/L- N; Aptian, Tunisia.

2013 Cuneolina hensoni – Shanin & Elbaz, pl. 1, fig. 44; late Cenomanian, Sinai .

2013 Cuneolina pavonia – Shanin & Elbaz, pl. 1, figs. 45-46; late Cenomanian, Sinai .

2014b Cuneolina pavonia – Omidvar et al., fig. 4/3; late Cenomanian – intra-middle Turonian (as interpreted following Simmons et al., 2024a), Iranian Zagros.

2014 Dicyclina schlumbergeri – Afghah et al., Fig. 11F View Fig ; early Cenomanian(?), Iranian Zagros.

2015 Cuneolina conica – Albrich et al., fig. 13 H, O; fig. 14 C-D; Campanian, Spain.

2015 Cuneolina pavonia – Solak et al., fig. 9A; Cenomanian – Coniacian, Turkish Taurides.

2016 Cuneolina pavonia – Rikhtegarzadeh et al., pl. 3, fig. 1; Cenomanian, Iranian Zagros.

Non 2016 Cuneolina pavonia – Ghaseminia et al., fig. 4L; Cenomanian, Iranian Zagros. [Indeterminate textulariid].

2017 Cuneolina pavonia – Ahmadi et al., pl. 1, fig. 11; Albian – Cenomanian (probably late Albian – early Cenomanian), Iranian Zagros.

2017 Cuneolina pavonia – BouDagher-Fadel et al., fig. 11/8-10; Albian – early Cenomanian, Tibet.

2017 Cuneolina pavonia – Hamedanian et al., pl. 1/F; Aptian, Iranian Zagros.

? 2017 Cuneolina gr. pavonia – Koç, fig. 6/G1-2; Aptian, Turkish Taurides.

2018 Cuneolina pavonia – Luger, p. 66-67, pl. 6, figs. 1, 2, 5; late Albian – Early Cenomanian, Somalia.

2019 Cuneolina pavonia – Omaña et al., p. 709, fig. 10ab, middle – late Cenomanian, Mexico.

2019 Cuneolina parva – Omaña et al., p. 709-711, fig. 10c-f; middle – late Cenomanian, Mexico.

? 2019 Cuneolina pavonia – Kiarostami et al., pl. 1K; Cenomanian, Iranian Zagros.

2019 Cuneolina pavonia – Saeedi Razavi et al., pl. 2, fig. 10; Cenomanian, Iranian Zagros.

2019 Cuneolina pavonia – Solak et al., fig., 8R-T; fig. 10T; middle – late Cenomanian, late Campanian, Turkish Taurides.

2019 Cuneolina pavonia – Taslı & Solak, fig. 10/5; late Albian, Turkish Taurides.

2019 Cuneolina ex gr. pavonia – Özkan & Altıner, fig. 9/14; early – middle Cenomanian (as interpreted by Simmons et al., 2020), south-eastern Türkiye.

2020 Cuneolina sp. – Haftlang et al., fig. 2/24; Santonian, Iranian Zagros.

? 2020 Cuneolina pavonia – Randazzo et al., fig. 9R; Albian – Cenomanian, Sicily.

2020 Cuneolina pavonia – Solak et al., fig. 6T?; fig. 11N- O; fig. 14S; Albian?, late Cenomanian – Turonian, Turkish Taurides.

2021 Cuneolina parva – Solak et al., p. 678, figs. 5.1-5.3; Albian, Turkish Taurides.

2021 Cuneolina pavonia – Sinanoglu, p. 276-277, pl. 1, figs. 15-16; Maastrichtian, southern Türkiye.

2021 Cuneolina pavonia – Saedi Razavi et al., pl. 1, fig. 8; late Cenomanian (as interpreted herein), Iranian Zagros.

2021 Cuneolina sp. – Özkan, fig. 11/13; Campanian, southeast Türkiye.

2021 Cuneolina pavonia – Dousti-Mohajer et al., pl. 1j; Cenomanian, Iranian Zagros.

? 2021 Cuneolina pavonia – Nagm et al., fig. 4E; late Cenomanian, Egypt.

2021 Cuneolina parva – Solak, pl. 2A-C; middle – late Cenomanian, Turkish Taurides.

2021 Cuneolina pavonia – Solak, pl. 3E-F; late Cenomanian, Turkish Taurides.

2021 Cuneolina pavonia – Gholamalian & Fanati Rashidi, pl. 3, fig. 1; Cenomanian, Iranian Zagros.

2021 Dicyclina schlumbergeri – Radmacher et al., pl. 1, figs. 8-9; Albian-Santonian Guatemala.

Non 2022 Cuneolina pavonia – Esfandyari et al., fig. 25c; Cenomanian, Iranian Zagros. [Probably Praetaberina bingstani ].

? 2022 Cuneolina pavonia – Dousti-Mohajer et al., fig. 4l; Cenomanian, Iranian Zagros. [Test is possibly annular, thus suggesting Dicyclina ].

2023 Cuneolina gr. C. pavonia – Solak & Taslı, fig. 10C, K; late Aptian – Albian, Turkish Taurides.

2025 Cuneolina pavonia – Salmouna et al., fig. 15D; Turonian, Tunisia.

2025 Cuneolina pavonia ( ? parva ) – Messaoud et al., fig. 8 (f-g); Turonian, Jordan.

Reference Images: Schlagintweit (1992); Whittaker et al. (1998).

Taxonomy/Identity: As noted by Scott & Gonzalez-Leon (1991), discrimination among the species of Cuneolina (currently composed of 16 accepted species; WoRMS.org – Hayward et al., 2025) is difficult because many were defined by exterior features and the corresponding features or internal features are not known in the same detail for each species. Furthermore, thin-sections usually reveal only partial sections through the specimens.

As can be understood from the long synonymy list (which is only partial and does not include unillustrated records of which there are many), relatively large fan-shaped cuneolinids are common components of Cretaceous Neotethyan carbonate sediments. They are known from Central America (e.g., Filkorn & Scott, 2011), the peri-Mediterranean (e.g. Solak & Taslı, 2023), the Middle East (e.g., Saint-Marc, 1974), Tibet (e.g., BouDagher-Fadel et al., 2017), and guyots in the Pacific (e.g., Arnaud-Vanneau & Sliter, 1995). These have been described under a variety of names such as Cuneolina pavonia d’Orbigny (the type species of Cuneolina ), Cuneolina parva Henson and Cuneolina conica d’Orbigny. Some authors maintain that separate species can be distinguished (e.g. Arnaud-Vanneau & Sliter, 1995; Ghanem & Kuss, 2013), others place them into synonymy (with C. pavonia the senior synonym) (e.g. Saint-Marc, 1974; Luger, 2018), and there are advocates to use a “group” concept (e.g. Altıner & Decrouez, 1982; Simmons & Hart, 1987; Kuss & Schlagintweit, 1988; Schlagintweit, 1992; Solak & Taslı, 2023). Pending a full taxonomic review and evaluation of intra-specific variation, we feel that use of “ Cuneolina ex gr. pavonia ” is the best solution for the moment and includes a group of similar taxa that are hard to separate from one another in many thin-sections. This includes C. pavonia , C. parva , C. conica , and Cuneolina walteri Cushman & Applin. This grouping does not include Cuneolina compressa Schlagintweit, 1988 that is smaller, with a thinner wall, and with finer septa and radial partitions.

The following description by Kuss & Schlagintweit (1988) is a useful summary of the concept of “ Cuneolina ex gr. pavonia ”: “ …represents a highly developed flabelliform Cuneolina with a convex base [apertural face]. The primary chambers of the test are divided by radial partitions into chamberlets, forming narrow rectangular shapes with a height approximately twice width. Secondary subepidermal partitions (both horizontal and vertical) are developed. Unilocular proloculus large, measuring about 0.11 mm in diameter. Size (in mm): height: 0.75 – 2.0 mm; thickness 0.27 – 0.3 mm. The angle of inclination [apical angle] of the test varies too greatly to use the width of individual tests as a characteristic measurement.”

A description of C. conica is similar (Albrich et al., 2015): “ Fan-shaped shell, biserially arranged, with low and broad chambers occupying an opening angle [apical angle] of about 70º. The number of chambers is about 20. The shell varies from 0.7mm to 1.2 mm in length. The exoskeleton consists of well-developed beams and rafters. The number of beams per chamber can reach about 20 in the last chamber ”.

C. pavonia View in CoL was mentioned by d’Orbigny in 1839 and a description subsequently provided by him in 1846 based on material from Ile Madam, France. Although originally considered as Turonian by d’Orbigny, Loeblich & Tappan (1964) regarded the types as Cenomanian. Schlumberger (1900) subsequently described the species from the Santonian of Spain.

C. parva View in CoL was originally described as a subspecies (because of a smaller size) of C. pavonia View in CoL by Henson (1948b) but has often been regarded as a species in its own right (e.g. Arnaud-Vanneau & Sliter, 1995). It was first described from Egypt from a section considered to be Santonian, but subsequently regarded as Albian (Arnaud-Vanneau & Sliter, 1995) or Turonian ( Whittaker et al., 1998).

C. walteri View in CoL was described from the middle Cenomanian of Florida (Cushman & Applin, 1947; Applin & Applin, 1965), with a supposed relatively small size: Length up to 1.00 mm; breadth up to 1.40 mm; thickness 0.30 mm. The indistinct sutures are said to be a distinguishing feature, and it has a markedly flaring test. Nonetheless, Filkorn & Scott (2011) regarded it as synonymous with C. parva View in CoL (see also Omaña et al., 2019), in turn noting that this is hardly distinguishable from C. pavonia View in CoL .

The Early Cretaceous species Cuneolina hensoni Dalbiez, 1958 View in CoL is similar to C. ex gr. pavonia View in CoL but has a much coarser internal structure with broad rectangular, almost square chamberlets. There are a number of other Cretaceous species of cuneolinids. These include Cuneolina axinoides Arnaud-Vanneau, 1980 View in CoL ; Vercorsella arenata Arnaud-Vanneau, 1980 View in CoL ; Scythiolina camposaurii ( Sartoni & Crescenti, 1962) View in CoL ; Vercorsella laurentii ( Sartoni & Crescenti, 1962) ; Vercorsella scarsellai ( De Castro, 1963) View in CoL ; and Cuneolina sliteri Arnaud-Vanneau & Premoli-Silva, 1995 View in CoL and other species assigned to Scythiolina Neagu 1997 View in CoL and Histerolina Neagu 1997 View in CoL . All are smaller and less flabelliform than C. ex gr. pavonia View in CoL , and with fewer radial partitions and typically no or few secondary subepidermal partitions. Those assigned to Vercorsella View in CoL have a slit-like aperture including the earliest species, V. halleinensis Schlagintweit & Gawlick, 2005 View in CoL from the late Tithonian – early Berriasian of Austria, also recorded by Hosseini & Conrad (2008) from the Berriasian of Iran. The poorlyknown Late Cretaceous Cuneolina cylindrica Henson, 1948b View in CoL is characterised by an acute apical angle leading to a narrow, cylindrical test shape.

Random sections of C. ex gr. pavonia can be confused with the discoidal genus Dicyclina (see separate species entries) that has annular chambers. As mentioned by Brönnimann et al. (1983), Dicyclina represents a more specialized form and is regarded as the final stage in the evolution of a flabelliform Cuneolina . Nakkadyia awadi introduced by Ismail et al. (2009) from the Cenomanian of Egypt might be an intermediate form between Cuneolina and Dicyclina (and is considered as a taxon of uncertain status by WoRMS.org – Hayward et al., 2025). Given that it is not truly annular (i.e. it is said to be fan-shaped) it is tentatively regarded as a synonym of Cuneolina ex gr. pavonia pending further research. As noted by Schlagintweit (1992) and Cvetko Tešović et al. (2001) the embryonic apparatus of advanced Cuneolina ex gr. pavonia comprises a globular eccentric proloculus surrounded by low and wide chambers (i.e. with a subembryonic zone similar to that of Dicyclina schlumbergeri ). However, that of D. schlumbergeri is larger and more complex. Axial sections also have a similarity in the nature of septa and subepidermal partitions.

Confident Stratigraphic Range: Aptian – Maastrichtian. Common records throughout this range.

Uncertain Stratigraphic Range: not applicable.

The long range of C. ex gr. pavonia is well established. Sartoni & Crescenti (1962) introduced a biozone of Cuneolina pavonia parva for the Albian – Cenomanian and of Cuneolina pavonia parva and Dicyclina schlumbergeri for the Turonian – Senonian (intra-Late Campanian) period of the southern Apennines, Italy. However, according to Chiocchini et al. (2012) the range of Cuneolina pavonia in the Apennines is restricted to within the late Cenomanian. The oldest specimens of C. ex gr. pavonia appear to be Aptian (Altıner & Decrouez, 1982; Mouty & Saint-Marc, 1982; Kuss & Schlagintweit, 1988, Schlagintweit et al., 2016; Solak & Taslı, 2023 and – possibly – Gollesstaneh, 1965 and Koç, 2017) and the taxon has long been regarded as long-ranging throughout the mid-Cretaceous and into the upper part of the Late Cretaceous ( Saint-Marc, 1974; Velić, 2007). Barremian records (e.g. Soklić, 2019) are not confirmed by illustration. It appears to range throughout the Late Cretaceous to the Maastrichtian (Caus & Cornella, 1983; Sinanoğlu et al., 2020), giving a total long range of Aptian – Maastrichtian.

Locally, the inception or extinction of C. ex gr. pavonia may have stratigraphic significance (e.g. Aguilero-Franco, 2000, 2003), but given the long range of the taxon these will be facies-controlled events.

Geographic Distribution: Very widespread from the Caribbean/Central America region (including mid-Pacific seamounts), and through Neotethys as far east as Somalia and Tibet.