Nummofallotia? apula Luperto-Sinni, 1968

SIMMONS, MICHAEL & BIDGOOD, MICHAEL, 2023, “ Larger ” Benthic Foraminifera Of The Cenomanian. A Review Of The Identity And The Stratigraphic And Palaeogeographic Distribution Of Non-Fusiform Planispiral (Or Near-Planispiral) Forms, Acta Palaeontologica Romaniae 19 (2), pp. 39-169 : 108-110

Nummofallotia? apula Luperto-Sinni, 1968

Reference Illustration & Description

Luperto-Sinni in Schroeder & Neumann (1985), pl. 48, p. 100-101 (but see below for issues regarding stated magnifications). The species is also well illustrated by Saint-Marc (1970, pl. 2, figs. 10-15; 1974a, pl. XIII, figs. 13- 19).

Nummofallotia was introduced by Barrier & Neumann (1959) with the Late Cretaceous (Coniacian - Maastrichtian) taxon Nonionina cretacea Schlumberger as type species. The test is lenticular, planispirally enrolled, periphery rounded to subangular, with a globular proloculus followed by whorls of regularly enlarging chambers. Adults are involute with septa that are slightly oblique and slope backwards at the periphery. An umbo (also sometimes called umbilical “plug” or “button”) of radial fibrous calcite is a distinctive feature in most specimens ( Hottinger & Caus, 2009).

Luperto-Sinni (1968) introduced a new species, Nummofallotia apula , with type material from the “Senonian” of southern Italy, which is smaller and with fewer whorls than N. cretacea . Originally described as possessing a single-layered wall, Bilotte & Decrouez (1979) subsequently stated that the test wall of N. apula is, in its entirety, made up of two layers: an inner layer dark microgranular, and a clear outer layer hyaline-radiated, thicker in the axial zone (see for example illustrations by Saint-Marc, 1970, 1974a), leading them to introduce a new genus, Murgeina , with N. apula as the type species. Although accepted by, for example, Loeblich & Tappan (1988), the necessity to introduce this genus was subsequently questioned by Luperto-Sinni in Schroeder & Neumann (1985) who noted that the two layered wall structure is a variable, inconsistent feature, and that otherwise N. apula conforms perfectly to the nature of Nummofallotia . That said, in 1998, she introduced a further new species of Nummofallotia , Nummofallotia cenomana , which alongside its supposedly very small size (see below), a distinguishing feature was said to be a consistent two layered wall structure. Therefore, there is some debate if Murgeina is a valid genus. A detailed taxonomic revision of all species of Nummofallotia / Murgeina is required using pristine material. This is outside the scope of this primarily stratigraphic/biogeographic review and therefore we tentatively retain N. apula within Nummofallotia as “ N.? apula ”.

N.? apula is a small biumbonate form, with maximum dimensions (diameter) of around 0.3 – 0.5 mm (holotype 0.32 mm), a thickness of around 0.15 – 0.25 mm (holotype 0.18mm) a large, 0.06-0.08 mm (holotype 0.06 mm), globular proloculus in macrospheric forms, followed by 3 whorls with 16-20 quadrangular chambers in the last whorl. It is distinctively smaller than N. cretacea , that has typical diameters of around 0.6 – 0.8 mm (although possibly as small as 0.36 mm ( Bilotte & Decrouez, 1979) and note that if the illustrations of Barrier & Neumann (1959) are correctly scaled, then specimens can be 2.0 – 3.0 mm in diameter, although this must be judged unlikely), thickness of 0.4 – 0.5 mm, 5 - 8 whorls, with more than 20 chambers in the last whorl (e.g., Luperto-Sinni, 1968). Although the same specimens of N.? apula are illustrated by Luperto-Sinni (1968) and Luperto-Sinni in Schroeder & Neumann (1985), including the types, there are small differences in size based on the magnifications provided.

N. cenomana was described as a distinctively small (stated diameter dimensions: 0.09 – 0.1 mm) species ( Luperto-Sinni, 1998), known from the Cenomanian of southern Italy. If the figures provided by Luperto-Sinni (1998) are taken on face value, using the magnifications as given, the diameters are larger (c. 0.23 mm), but still distinctly small compared to N.? apula . Luperto-Sinni (1998) implied that N. cenomana was the Cenomanian form of Nummofallotia , whilst N.? apula was the “Senonian” form. However, she did not discuss further the various Cenomanian records of N.? apula (see below), which conform to the type description of this species. Thus, the notion that N.? apula does not occur in the Cenomanian is rejected. Other than its type description, the only other mentions of N. cenomana in the literature are from the Iranian Zagros (e.g., Schlagintweit & Yazdi-Moghadam, 2020, as “ N.? cenomana ”), where the specimens would be better assigned to N.? apula , based on size and morphology. The types of N. cenomana need to be examined and re-illustrated as part of a taxonomic re-evaluation of Nummofallotia / Murgeina , and this taxon is not considered further herein. (As this paper was going to press, Schlagintweit et al. (2023) published a review of this species (as Murgeina apula ) with illustration from the Cenomanian Sarvak Formation of the Iranian Zagros.) Nummofallotia kastomonica Özgen Erdem is an upper Maastrichtian species described from northern Turkey ( Özgen Erdem, 2001). It differs from N. apula by virtue of its large size (stated diameter 0.43 – 1.02 mm; a possible expression of Cope’s Rule?), tighter coiling, greater number of whorls and chambers, straight septa, and smaller umbo.

Stratigraphic Distribution

Middle Cenomanian –?Maastrichtian.

N.? apula was originally described from the “upper Senonian” (possibly Maastrichtian but probably no younger than middle Campanian) of southern Italy ( Luperto-Sinni, 1968). It appears to have an unusually long stratigraphic range almost throughout the Late Cretaceous (e.g., Luperto-Sinni & Ricchetti, 1978; Perugini, 2006; Velić, 2007; Sari et al., 2009), but its FAD lies within the Cenomanian.

Unillustrated records of N.? apula from the Barremian of the Iranian Zagros ( Abyat et al., 2016) must be discounted, as probably should the comment by Keshavarzi et al. (2021) that the oldest occurrence of N.? apula marks the base of the Sarvak Formation in the Iranian Zagros which would place it close to the Albian/Cenomanian boundary ( Bromhead et al., 2022). No illustration is provided. A Lower Cenomanian record by Radoičić et al. (2010) can be reassessed as middle Cenomanian based on associated fauna. Finally, a record of N.? apula from the Yamama Formation of southern Iraq ( Al-Hassani & Al-Dulaimi, 2021) which is Berriasian – Valanginian in age (although the authors describe the material as “early Aptian”) illustrates a fragmentary specimen of Lenticulina sp. or Epistomina sp. Although many records are from the late Cenomanian, Bachmann et al. (2003 - unillustrated) using graphic correlation between sections in Egypt positioned the FAD at or close to the early-middle Cenomanian boundary (following Saint-Marc, 1978; Luperto-Sinni in Schroeder & Neumann, 1985). Chiocchini (2008a - unillustrated) showed a short range for the species in the lowest part of the “late” Cenomanian (Chiocchini used only early and late subdivisions of the Cenomanian which would suggest this range FAD is approximately within the chronostratigraphic middle Cenomanian). Ghanem & Kuss (2013) showed the range of this species (although they illustrated it as cf.) extending into the (upper) middle Cenomanian of Syria, calibrated by planktonic foraminifera. A similar FAD was placed by Schlagintweit (1992) from Austria, and Simone et al. (2012) from Italy, although without independent calibration. Bravi et al. (2004) plausibly illustrate this species from the middle Cenomanian of Central Italy.

Cenomanian Paleogeographic Distribution

Caribbean/North Africa? - Neotethys.

References in Luperto-Sinni in Schroeder & Neumann (1985) indicate Cenomanian records from Lebanon ( Hamaoui & Saint-Marc 1970; Saint-Marc 1970, 1974 a, 1981), Greece ( Decrouez, 1975, 1977; Charvet et al., 1976; see also Fleury, 1971), and Serbia ( Radoičić 1974a). In addition, records confirmed by definite or plausible illustration include Italy ( Benedetti et al., 2000), south-east Turkey (Özcan & Altiner 2019); and the Iranian Zagros ( Sartorio & Venturini, 1988; Jamalpour et al. 2018, Kiarostami et al. 2019, Mohajer et al. 2021a, 2022a, b; an illustration by Mohseni & Javanmard, 2020 is uncertain). Illustrations from Tunisia ( Bismuth et al. 1981) and Syria ( Ghanem & Kuss 2013 – as cf.), may be more compatible with N. cenomana based on their small size (diameter 0.2 – 0.3 mm). Another record from Tunisia ( Abdallah et al., 1995) is more likely Charentia cuvillieri Neumann.

Cenomanian records from other locations but unconfirmed by illustration include Cuba (Diaz Otero, 1985); Mexico ( Michaud et al., 1984; Hernández-Romano et al., 1997; Aguilera-Franco, 2000; Aguilera-Franco & Allison, 2004); Morocco (El-Kadiri et al., 2003, Piuz & Meister, 2013); Egypt ( Bachmann et al., 2003; an illustration by Orabi, 1992 is uncertain); Iraq ( Hamaoui & Brun, 1974; Bernaus & Masse, 2007; Mahdi et al., 2013), Jordan (Schulze 2003, Schulze et al., 2004) and the Oman Mountains ( Rabu 1993; Al-Balushi & Macquaker, 2011; Piuz & Meister, 2013).

Localities from strata younger than Cenomanian are not included herein.