Dicyclina schlumbergeri Munier-Chalmas, 1877

Simmons, Michael, Bidgood, Michael, Consorti, Lorenzo & Schlagintweit, Felix, 2025, A Review Of The Identity And Biostratigraphy Of Cenomanian “ Larger ” Benthic Foraminifera: Part 2 - The Order Loftusiida (Excluding The Suborder Orbitolinina), Acta Palaeontologica Romaniae 21 (1), pp. 103-192 : 154-159

Dicyclina schlumbergeri Munier-Chalmas, 1877

Figure 22

T 1877 Dicyclina schlumbergeri n. gen., n. sp. Munier-Chalmas , p. 30-31; Senonian, southern France (type material lost) [probably Coniacian – Santonian see Gendrot, 1968] .

1904 Dicyclina schlumbergeri – Schlumberger & Choffat, p. 148-149, text-figs. 1-2; Senonian, southern France [probably Coniacian – Santonian see Gendrot, 1968].

? 1948b Dicyclina schlumbergeri – Henson, pl. XIV, fig. 9; Maastrichtian, Qatar & Iraq [fragment only].

? 1956 Dicyclina schlumbergeri – Bonet, p. 406-7, pls. 5- 7; Cenomanian, Mexico [fragments only].

? 1961 Dicyclina cf. qatarensis – Hamaoui, pl. 1, figs. 2- 6; Cenomanian, Israel.

Non L 1964 Dicyclina schlumbergeri – Loeblich & Tappan, p. C303, fig. 209 (1a-1b); middle Cenomanian, France [lectotype invalid fide Cherchi & Schroeder, 1990a = D. simplex ].

? 1966 Dicyclina schlumbergeri – Luperto Sinni, pl. 13, fig. 3; Santonian?, southern Italy [= Dicyclina sp. ].

? 1967 Dicyclina schlumbergeri – Luperto Sinni, p. 176- 177, pl. 10, fig. 1; pl. 13, fig. 1; late Cenomanian – Santonian, southern Italy [= Dicyclina sp. ].

1967 Dicyclina cf. schlumbergeri – Neumann, pl. 55, fig. 3; pl. 56, figs. 1-2; early Senonian, southern France.

Non 1967 Dicyclina schlumbergeri – Neumann, pl. 55, fig. 2; Cenomanian, western France [from the type locality/strata of D. simplex ].

? 1967 Dicyclina schlumbergeri – Neumann, pl. 55, fig. 1; Santonian, Spain [= Dicyclina sp. ].

1968 Dicyclina schlumbergeri – Gendrot, p. 677-678, pl. V, figs. 14-16; Coniacian – Santonian , southern France. [fig. 15 designated as a Neotype by Cherchi & Schroeder, 1990a] .

? 1970 Dicyclina sp. – Hamaoui & Saint-Marc, pl. 40, fig. 14; late Cenomanian, Lebanon.

1976 Dicyclina schlumbergeri – Luperto-Sinni, p. 308- 309, pl. 33, figs. 1-6; pl. 34, figs. 1; Senonian, southern Italy.

1978 Dicyclina schlumbergeri – Luperto-Sinni & Richetti, pl. 47, figs. 1-7; pl. 48, figs. 1-6; pl. 49, figs. 1- 6; Santonian – Maastrichtian, southern Italy.

? 1984 Dicyclina schlumbergeri – Bilotte, pl. 5, fig. 2; middle Cenomanian, Pyrenees [= Dicyclina sp. ].

1984 Dicyclina schlumbergeri – Bilotte, pl. 15, fig. 2; early Campanian, Pyrenees.

? 1984 Dicyclina schlumbergeri – Bilotte, pl. 25, fig. 5; Maastrichtian, Pyrenees [= Dicyclina sp. ].

? 1987 Dicyclina schlumbergeri – Simmons & Hart, pl. 10.4, fig. 5; middle-late Cenomanian, Oman [= D. simplex or D. schlumbergeri , possibly Cuneolina ex. gr. pavonia ].

? 1987 Dicyclina schlumbergeri – Al-Rifaiy & Cherif, pl. II, fig. 11; Turonian, Jordan [= Dicyclina sp. ].

1987 Dicyclina schlumbergeri – Loeblich & Tappan, pl. 157, fig. 7 (non 8-10); Senonian, France.

Non 1987 Dicyclina schlumbergeri – Loeblich & Tappan, pl. 157, figs. 8-10; Cenomanian, France [ fide Cherchi & Schroeder, 1990a = D. simplex ].

? 1988 Dicyclina schlumbergeri – Sartorio & Venturini, p. 114 lower image, p. 115 upper image, p. 121; Senonian, southern Italy.

? 1988 Dicyclina schlumbergeri – Sartorio & Venturini, p. 115 lower image; upper Senonian, Tunisia.

Non 1989 Dicyclina schlumbergeri – Rosales-Dominguez, fig. 11B; Albian-early Cenomanian; southwest USA [= Cuneolina sp. ].

? 1990 Dicyclina schlumbergeri – Šribar & Pleničar, pl. 4, fig. 4; late Turonian, southwest Slovenia [indeterminate fragment].

N 1990a Dicyclina schlumbergeri – Cherchi & Schroeder, figs. 5-11; Coniacian, France & Sardinia.

? 1992 Dicyclina schlumbergeri – Kalantari, pl. 77; Cenomanian, Iranian Zagros [= Dicyclina sp. ].

1994 Dicyclina schlumbergeri – Chiocchini et al., pl. 23, fig. 16; Senonian, central Italy.

Non 1998 Dicyclina schlumbergeri – Charrière et al., fig. 4 (3a, 6); late Cenomanian, Morocco [= D. simplex ].

2000 Dicyclina schlumbergeri – Aguilera-Franco, p. 161; middle-late Cenomanian, southern Mexico.

2001 Dicyclina schlumbergeri – Cvetko Tešović et al., p. 602, fig. 6D-H; Campanian, Brač Island, Croatia.

? 2003 Dicyclina schlumbergeri – Polavder, fig. 5 (17- 18); Santonian-Campanian, former Yugoslavia [= Dicyclina sp. ].

Non 2004 Dicyclina schlumbergeri – Khosrow Tehrani & Afghah, pl. 2, fig. 4 [= Broeckinella arabica Henson fide Schlagintweit & Rashidi, 2016 ].

? 2005 Dicyclina schlumbergeri – Vaziri et al., pl. 2, fig. 4; Santonian-Maastrichtian, central Iran [indeterminate fragment].

? 2006 Dicyclina schlumbergeri – Taslı et al., fig. 7U; Coniacian-Santonian, southern Türkiye [= Dicyclina sp. ]. 2007 Dicyclina schlumbergeri – Velić, pl. 23, figs. 1-4; Santonian-Campanian, Adriatic platform.

? 2008 Dicyclina schlumbergeri – Schlüter et al., fig. 3E; late Campanian, southern Italy [= Dicyclina sp. ].

Non 2008 Dicyclina schlumbergeri – Khosrow Tehrani et al., pl. 1, fig. 6 [= Broeckinella arabica Henson fide Schlagintweit & Rashidi, 2016 ].

? 2008 Dicyclina schlumbergeri – Ahmadi et al., pl. 3, fig. 1; Cenomanian, Iranian Zagros [best considered as Dicyclina sp. ].

Non 2009 Dicyclina schlumbergeri – Sari et al., pl. 3, figs. 8-9; age uncertain, reworked, Turkish Taurides [= D. simplex fide Schlagintweit & Rashidi, 2018 ].

? 2010 Dicyclina sp. – Cavin et al., fig. 9F; late Cenomanian-middle Turonian, Morocco. [image captioned as Dicyclina sp. but referred to as D. schlumbergeri in text (p. 405)].

Non 2011 Dicyclina schlumbergeri – Vaziri, fig. 2D, F; Santonian-Maastrichtian, central Iran [most likely Cuneolina sp. ].

2011 Dicyclina schlumbergeri – Khosrotehrani et al., pl. 1, fig. C; Santonian, Iranian Zagros.

? 2011 Dicyclina schlumbergeri – Amer, pl. 19, fig. 1; Cenomanian, western Iraq [best considered as Dicyclina sp. ].

2012 Dicyclina schlumbergeri – Chiocchini et al., pl. 134, figs. 1-8; Santonian, central Italy.

Non 2012 Dicyclina schlumbergeri – Rahimpour-Bonab et al., fig. 8G; Cenomanian – Turonian, Iranian Zagros [= D. sampoi ].

? 2012 Dicyclina shlumbergeri [sic] – Kiarostami et al., pl. 2, fig. 10; Cenomanian, Iranian Zagros [= Dicyclina sp. ].

? 2012 Dicyclina schlumbergeri – Omaña et al., fig. 5(1); late Cenomanian, Mexico [= Dicyclina / Cuneolina sp. ].

Non 2013 Dicyclina schlumbergeri – Rahimpour-Bonab et al., fig. 8U; Turonian [?], Iranian Zagros [= D. sampoi ].

? 2013 Dicyclina schlumbergeri – Al-Dulaimi et al., fig. 10(1); late Cenomanian; southern Iraq [= Dicyclina sp. ].

2013 Dicyclina schlumbergeri – Omaña et al., pl. 1, figs. 2, 4; pl. 4, figs. 2, 4; late Cenomanian, Mexico.

? 2013 Dicyclina schlumbergeri – Shahin & Elbaz, p. 276, pl. 2, figs. 3-4; late Cenomanian, Egypt [= Dicyclina / Cuneolina sp. ].

? 2014a Dicyclina schlumbergeri – Omidvar et al., pl. 2, fig. P; Cenomanian, Iranian Zagros [= Dicyclina sp. ].

Non 2014b Dicyclina schlumbergeri – Omidvar et al., fig. 3(10); Cenomanian-Turonian, Iranian Zagros. [= D. sampoi Cherchi & Schroeder fide Schlagintweit & Rashidi, 2018 ].

Non 2014 Dicyclina schlumbergeri – Afghah et al., fig. 11(f); early Cenomanian, Iranian Zagros [= Cuneolina ex. gr. pavonia ].

Non 2014 Dicyclina schlumbergeri – Afghah & Yaghmour, pl. 2, fig.3; pl. 3, fig. 2; Maastrichtian, Iranian Zagros [perhaps Dicyclina sp. , but embryonic apparatus too simple to be D. schlumbergeri ].

? 2014 Dicyclina schlumbergeri – Shahin & Elbaz, fig. 6/6; late Cenomanian, Egypt [= Dicyclina sp. ].

? 2015 Dicyclina schlumbergeri – Frijia et al., fig. 9C; Coniacian-Campanian, central Italy [= Dicyclina sp. ].

Non 2015 Dicyclina schlumbergeri – Babazadeh & Dehej, fig. 7 (i, j, k); early Albian, central Iran. [= Balkanica balkhanica Mamontova fide Schlagintweit, 2024 ].

Non 2016 Dicyclina schlumbergeri – Ghaseminia et al., fig. 4E; Coniacian – Santonian, Iranian Zagros [indeterminate fragment that cannot be assigned to a genus].

Non 2016 Dicyclina schlumbergeri – Kazemzadeh & Loftpoor, pl. 2, fig. 6; Cenomanian, Iranian Zagros [indeterminate fragment, but unlikely to be Dicyclina sp. ].

? 2016 Dicyclina schlumbergeri – Rikhtegarzadeh et al., pl. 1, figs. 3-5; Cenomanian, Iranian Zagros. [= Dicyclina / Cuneolina sp. ].

? 2016 Dicyclina schlumbergeri – Dehghani et al., pl. 2, fig. 7; Campanian-Maastrichtian, Iranian Zagros [= Dicyclina sp. ].

? 2016 Dicyclina schlumbergeri – Assadi et al., fig. 6(b2); Cenomanian-Turonian, Iranian Zagros [indeterminate fragment].

? 2017 Dicyclina schlumbergeri – Consorti et al., fig. 3A; late Santonian? – middle Campanian, Italy [= Dicyclina sp. ].

? 2017 Dicyclina schlumbergeri – Rikhtegarzadeh et al., pl. 1, fig. 3; Cenomanian, Iranian Zagros [axial section that may be D. simplex or D. schlumbergeri ].

? 2017 Dicyclina schlumbergeri – Koç, fig. 10D; Santonian-early Campanian, central Taurides, Türkiye [= Dicyclina sp. ].

? 2017 Dicyclina schlumbergeri [sic] – Jamalpour et al., pl. 2d; Cenomanian, Iranian Zagros [= Dicyclina sp. ].

2018 Dicyclina schlumbergeri – Schlagintweit & Rashidi, fig. 7a-f; late Maastrichtian, Iranian Zagros.

Non 2019 Dicyclina schlumbergeri – Saedi Razavi et al., pl. 1, fig. 3; Cenomanian, Iranian Zagros [= D. sampoi ].

Non 2018 Dicyclina schlumbergeri – Omidi et al., pl. 2, fig. 9; Cenomanian, Iranian Zagros [= D. simplex ].

2019 Dicyclina schlumbergeri – Omaña et al., p. 711, fig. 10 (g-j); middle-late Cenomanian, Mexico.

? 2019 Dicyclina schlumbergeri – Özkan & Altıner, fig. 9 (1-2); Cenomanian, southeast Türkiye [= Dicyclina sp. ].

Non 2019 Dicyclina schlumbergeri – Alloul, fig. III.9C; Cenomanian, southwest Algeria. [= D. qatarensis ].

2019 Dicyclina schlumbergeri – Villalonga et al., p. 29, pl. 12(a-f); middle Campanian, Spanish Pyrenees.

? 2020 Dicyclina schlumbergeri – Sinanoğlu et al., pl. 1, fig. 19; Maastrichtian, Turkish Arabian Plate [axial section only].

? 2020 Dicyclina schlumbergeri – Consorti, fig. 3B; Late Cretaceous, central Italy [= Dicyclina sp. ].

? 2020 Cuneolina pavonia d’Orbigny – Solak et al. , fig. 11N; late Cenomanian, western Taurides, Türkiye [probably best regarded as Dicyclina / Cuneolina sp. ].

? 2020 Dicyclina schlumbergeri – Fabbi et al., fig. 5 (l, n); Santonian, Italy [= Dicyclina sp. ].

2021 Dicyclina schlumbergeri – Bagherpour et al., fig. 12h; Coniacian-Santonian, Iranian Zagros.

Non 2021 Dicyclina schlumbergeri – Gholamalian & Fanati Rashidi, pl. 3, figs 4-5; Cenomanian, Iranian Zagros [= indeterminate foraminifera, not Dicyclina ].

? 2021 Dicyclina schlumbergeri – Saedi Razavi et al., pl. 1, fig. 3; Cenomanian, Iranian Zagros [axial section of D. simplex or D. schlumbergeri ].

2021 Dicyclina schlumbergeri – Özkan, fig. 11(12); Campanian, Turkish Arabian Plate.

? 2021 Dicyclina schlumbergeri – Sinanoğlu, p. 277, pl. 1, fig. 19; Maastrichtian, Turkish Arabian Plate.

? 2021 Dicyclina schlumbergeri – Dousti-Mohajer et al., pl. 2, fig. u; Cenomanian, Iranian Zagros [= Dicyclina sp. ].

Non 2021 Dicyclina schlumbergeri – Radmacher et al., pl. 1, figs. 8 (cf.), 9; Albian-Santonian, Guatemala [= Cuneolina pavonia ].

? 2021 Dicyclina schlumbergeri – Shapourikia et al., fig. 8f; Turonian [?], Iranian Zagros [= Dicyclina / Cuneolina sp. ].

? 2022 Dicyclina schlumbergeri – Dousti-Mohajer et al., fig. 4k; Cenomanian, Iranian Zagros [= Dicyclina sp. ].

? 2022 Dicyclina shlumbergri (sic) – Al-Dulaimi et al., pl. 1D; middle-late Cenomanian, southern Iraq [effectively indeterminate].

? 2022 Dicyclina cf. schlumbergeri – Ghanbarloo et al., fig. 4k;?fig. 6d; Maastrichtian, Iranian Zagros [= Dicyclina sp. ].

? 2022 Dicyclina schlumbergeri – Esfandyari et al., fig. 25a; Cenomanian-Turonian, Iranian Zagros [= Dicyclina sp. ].

2023a Dicyclina schlumbergeri – Mehrabi et al., fig. 11 (C-D); Santonian, Persian Gulf.

? 2023b Dicyclina schlumbergeri – Mehrabi et al., fig. 6(R); Turonian, Iranian Zagros [= Dicyclina sp. ].

? 2023b Dicyclina schlumbergeri – Schlagintweit et al., fig. 3E, H, L; Campanian, Croatia [= Dicyclina sp. ].

Non 2023 Dicyclina schlumbergeri – Al-Salihi & Ibrahim, pl. 1, fig. E, pl. 3, fig. f; middle – late Cenomanian, southern Iraq [? = D. sampoi and D. qatarensis ].

? 2024 Dicyclina schlumbergeri – Božović et al., fig. 3(c); Turonian – Santonian, Montenegro [= Dicyclina sp. ].

Reference Images: Luperto-Sinni & Richetti (1978), pl. 47, figs. 1-7; pl. 48, figs. 1-6; pl. 49, figs. 1-6; Cvetko Tešović et al., (2001) figs. 6D- H; Schlagintweit & Rashidi (2018) fig. 7a-f; Villalonga et al. (2019) pl. 12, figs. a-f.

Taxonomy/Identity: First described from the Senonian of the Étang de Berre, Bouches-du-Rhône province of France by Munier-Chalmas (1877), the original types of D. schlumbergeri are said to be lost (Cherchi & Schroeder, 1990a). The species was later recorded from the Île Madame, Charente-Maritime department by Schlumberger & Choffat (1904) from which population Loeblich & Tappan (1964) designated a lectotype. This, however, is not valid according to Cherchi & Schroeder (1990a) because the specimen does not come from the type locality, nor, in fact, does it belong to D. schlumbergeri (it is D. simplex ).

Cherchi & Schroeder (1990a) chose a neotype based on an axial section previously illustrated by Gendrot (1968) from the Coniacian of the Étang de Berre locality .

Cvetko Tešović et al. (2001) noted that specimens of D. schlumbergeri from the Campanian of Croatia indicated a value of 5-7 annular chambers per mm of radius (see also Table 5). They also described the structure of the embryonic apparatus, situated in the centre of the test, noting that is composed of an irregular ellipsoidal or globular embryonic chamber followed by a subembryonic zone. The upper part of the embryonic chamber shows a well-developed layer of subepidermal chambers (chamberlets), which are sometimes subdivided by very short septula. The dimensions of the embryonic apparatus range from 0.82 – 1.2 mm, which is larger than type material from the Coniacian of France: 0.65 – 0.95 mm (Cherchi & Schroeder, 1990a). Thus, they speculated this was a consequence of the phylogenetic evolution of D. schlumbergeri during the Late Cretaceous.

Following Cherchi & Schroeder (1990a), Cvetko Tešović et al. (2001) noted that the wall of D. schlumbergeri can display a pseudokeriothecal texture. The fine size of these features led them to believe that these served to facilitate exchange of gases rather than house algal symbionts. The features are most likely (or similar to) the “egg holder” structures of Hottinger (2006; fig. 40) which facilitate gas-exchange of the housed symbiont and thus are not mutually exclusive.

Specimens from the late Maastrichtian of the Iranian Zagros by Schlagintweit & Rashidi (2018; fig. 7a, b & e) show that the approximate number of radial elements per quadrant of D. schlumbergeri is approximately 20-?25. These are useful biometric data not mentioned in the type descriptions, and confirm the qualitative assessment of differences between D. schlumbergeri and D. qatarensis by Henson (1948b), the former species having lower annular and radial element densities than the latter (see below).

D. simplex is smaller than D. schlumbergeri and the post-embryonic disc is also, relatively, the thickest of all the Dicyclina species. D . sampoi has characteristically hook-shaped septa in transverse section (even “S”-shaped in some views) compared with the more curved, “C”- shaped septa of other species, and with subepidermal partitions confined to attachment to the inside of the outer wall and not both the wall and septa like D. schlumbergeri . This results in a smoother outline to the wall, without the depressed sutures seen in D. simplex and D. schlumbergeri . However, there are specimens with the embryonic apparatus of D. sampoi that show rafters attached to the septal wall (see Omidvar et al., 2014b; fig. fig. 3.10 and Solak et al. 2020; fig. 10U), thus the clear secondary partitions in the subembryonic zone of the embryonic apparatus of D. sampoi may be the best evidence for clear identity since in D. schlumbergeri they are absent or, at best, rare. It is probable that (along with the rather convoluted taxonomic history), specimens not clearly showing these features which otherwise distinguish species of D. qatarensis , D. simplex and D. sampoi are often “by default” labelled as D. schlumbergeri .

Confident Stratigraphic Range: middle – late Cenomanian (local range in Mexico only); common records from Coniacian – Maastrichtian elsewhere.

Uncertain Stratigraphic Range: Not applicable although Turonian records could be anticipated and exist in unconfirmed records, although we know of none.

Despite the many records of D. schlumbergeri in the published literature, only a relatively small proportion are illustrated, and of those only a few can be confidently identified. Therefore, confident assessment of stratigraphic range is based a small amount of material.

The published stratigraphic range of D. schlumbergeri is confused by imprecise use of the species name. Prior to the introduction of D. simplex and D. sampoi by Cherchi & Schroeder (1990a, b) almost all records of Dicyclina were assigned to D. schlumbergeri and even today the name continues to be applied with a lack of precision. Cherchi & Schroeder (1990a) suggested that as the phylogenetic descendant of D. simplex , D. schlumbergeri might be a Coniacian and younger species. This notion has been followed by a number of authors including Cvetko Tešović et al. (2001), Velić (2007), Chiocchini et al. (2012), Schlagintweit & Rashidi (2018) and Schlagintweit & Yazdi-Moghadam (2021). This phenomenon was first hinted at by Henson (1948b) with his separation of D. qatarensis (supposedly Cenomanian) and D. schlumbergeri (supposedly younger) (see also archive records in Le Blanc, 2015).

This appears to be partially correct (i.e., that verified records from the Mediterranean region and Arabian Plate are Coniacian – Maastrichtian), but there are several verified records from middle – late Cenomanian in Mexico only (Aguilera-Franco, 2000; Omaña et al., 2013, 2019). This suggests the species arose in the Caribbean and then migrated eastwards to the Mediterranean and Arabian Plate where it replaced other Dicyclina species by the Coniacian, or that there is a more complex pattern of evolution with homeomorphy and repetition of phylogeny from a Cuneolina ancestor. As noted above, Cvetko Tešović et al. (2001) suggested that the embryonic apparatus of D. schlumbergeri increases in size through its Coniacian – Maastrichtian evolution.

Geographic Distribution: Validated records of this species have a wide stratigraphic distribution from Mexico, across much of the Mediterranean, and from the Arabian Plate. It is also reported from Florida (Applin & Applin, 1965). Interestingly, the occurrences from Mexico are distinctly older (Cenomanian) than those elsewhere which are Coniacian – Maastrichtian.

Figures

Fig. 22. Representative illustrations of Dicyclina schlumbergeri: a. Axial section, Gendrot (1968, pl. V, fig. 15, France); b. Axial section through embryo, Cvetko Tešović et al. (2001, fig. 6H, Campanian, Croatia); c. Axial section through post-embryonic chambers, Cherchi & Schroeder (1990b, fig. 24, Sardinia); d. Schematic view of b, Cherchi & Schroeder (1990b, fig. 10b); e. Oblique equatorial section through embryo and initial annular chambers, Schlagintweit & Rashidi (2018, fig. 7a, Maastrichtian, Iranian Zagros).