◄ Carnets Geol. 24 (12) ►
Outline:
[1. Introduction]
[2. Systematic palaeontology]
[3. Discussion]
[4. Conclusions] and ...
[Bibliographic references]
Lerchenauerstr. 167, 80935 Munich (Germany)
Published online in final form (pdf) on November 30, 2024
DOI
10.2110/carnets.2024.2412
[Editor: Bruno
R.C. Granier; language editor:
Stephen Carey]
The Coskinolinidae are a family of Middle Jurassic to Paleogene, typically orbitoliniform, Larger Benthic Foraminifera (LBF) characterized by a pseudokeriothecal wall structure. The type genus, Coskinolina Stache, 1875, lacks an exoskeleton (i.e., beams, rafters), resulting in an undivided marginal chamber zone. However, other taxa within the family, such as the Upper Cretaceous Lepinoconus Cruz-Abad et al., 2017, or the Paleogene Coleiconus Hottinger & Drobne, 1980, are genera with vertical partitions (beams) that subdivide the marginal chamber zone. In the currently accepted classification of agglutinated benthic foraminifera, the nature of the exoskeleton is a criterion for distinction at subfamily level, e.g., Cyclamminidae Marie, 1941. Thus, here, the presence or absence of an exoskeleton in the marginal chamber lumen is treated as a criterion at the subfamily level within the classification of the Coskinolinidae, leading to a division into two subfamilies: the Coskinolininae Moullade, 1965 (lacking an exoskeleton), and the newly proposed Coleiconinae subfam. nov. (with an exoskeleton of vertical partitions), as exemplified by Coleiconus Hottinger & Drobne, 1980.
• Larger Benthic Foraminifera (LBF);
• taxonomy;
• classification;
• exoskeleton
Schlagintweit F. (2024).- A new subfamily classification of the Coskinolinidae Moullade, 1965, Middle Jurassic-Paleogene Larger Benthic Foraminifera.- Carnets Geol., Madrid, vol. 24, no. 12, p. 179-186. DOI: 10.2110/carnets.2024.2412
Une nouvelle classification des Coskinolinidae Moullade, 1965, Foraminifères Benthiques de Grande Taille de l'intervalle Jurassique moyen-Paléogène.- La famille des Coskinolinidae regroupe des Foraminifères Benthiques de Grande Taille (FBGT), typiquement orbitoliniformes, datant du Jurassique moyen au Paléogène et caractérisés par une structure de paroi pseudokériothéque. Le générotype, Coskinolina Stache, 1875, ne possède pas d'exosquelette (poutres/rayons et/ou poutrelles/mezzanines) ; par conséquent, la zone marginale des chambres est indivisée. Cependant, d'autres genres de la famille, tels que le Lepinoconus Cruz-Abad et al., 2017, du Crétacé supérieur ou Coleiconus Hottinger & Drobne, 1980, du Paléogène, présentent des séparations verticales (poutres/rayons) subdivisant la zone marginale des chambres. Dans la classification actuelle des foraminifères benthiques agglutinés, la présence ou l'absence d'un exosquelette est un critère permettant la distinction au niveau de la sous-famille, e.g., les Cyclamminidae Marie, 1941. Ainsi, la présence ou l'absence d'un exosquelette dans le lumen des chambres marginales est utilisée ici comme critère taxinomique au niveau de la sous-famille dans la classification de la famille des Coskinolinidae, conduisant à sa subdivision en deux sous-familles : les Coskinolininae Moullade, 1965 (sans exosquelette), et les Coleiconinae subfam. nov. (avec un exosquelette formé de séparations verticales), illustrée par Coleiconus Hottinger & Drobne, 1980.
• Foraminifères Benthiques de Grande Taille (FBGT) ;
• taxonomie ;
• classification ;
• exosquelette
With first representatives as old as Middle(?)-Late Jurassic (Cuvillier et al., 1968; Bordea S. & Bordea J., 1987; Kamoun & Peybernès, 1993), orbitoliniform agglutinated conical Larger Benthic Foraminifera are widespread constituents in Cretaceous-Paleogene shallow-water carbonates (e.g., Schroeder, 1962; Hottinger & Drobne, 1980; Moullade et al., 1985). With respect to these taxa, those displaying a pseudokeriothecal wall (e.g., Hottinger, 2006) contrast with the non-canaliculate wall texture of the Orbitolinidae, as previously noted by Douglass (1960) and Schroeder in Schroeder et al. (1975). In the original description of the Coskinolinidae, Moullade (1965) included forms having pillars in the central zone with undivided (Lituonella Schlumberger in Schlumberger & Douvillé, 1905) and subdivided marginal zones (Coskinolina Stache, 1875) as well as a group having a reticulate central zone represented by Kilianina Pfender, 1933, Abrardia Neumann & Damotte, 1960, Coskinolinoides Keijzer, 1942, and Orbiqia Mamgain & Jagannatha Rao, 1962. With Lituonella representing a junior synonym of Coskinolina (Schubert, 1912; Douglass, 1960; Hofker, 1966; Schroeder, 1974) and the other four genera originally included in the Coskinolinidae now assigned to other families (Loeblich & Tappan, 1987; Kaminski, 2014), only Coskinolina remains from Moullade's original family concept. Loeblich's and Tappan's 1987 classification included five genera: Coleiconus Hottinger & Drobne, 1980, Coskinolina, Coskinon Hottinger & Drobne, 1980, Lituonelloides Henson, 1948, and Pseudolituonella Marie, 1955 (see also Kaminski, 2014). Later, Cantabriconus Schlagintweit et al., 2017, and Lepinoconus Cruz-Abad et al., 2017, were added to the family. Serra-Kiel et al. (2016a) included Fallotella Mangin, 1954, in the Coskinolinidae without any remarks or discussion, a view that is rejected here since a pseudokeriothecal wall structure is seemingly absent (Mangin, 1954; Hottinger & Drobne, 1980; Loeblich & Tappan, 1987). The same can be stated for the inclusion of Barattolites Vecchio & Hottinger, 2007, in the Coskinolinidae by BouDagher-Fadel (2018), which is inconsistent with the original description of Barattolites by Vecchio and Hottinger (2007) as a member of the Orbitolinidae Martin, 1890 (see Serra-Kiel et al., 2016b).
Herein a taxonomic update of the Coskinolinidae is provided including a new subfamily concept.
The classification of the agglutinated benthic foraminifera follows Kaminski (2014). The proposed new suprageneric classification does not affect the generic diagnoses (Loeblich & Tappan, 1987, and taxa established later), and therefore they are not repeated herein. Table 1 shows a comparison of the two subfamilies Coskinolininae and Coleiconinae including common features and differences.
Table 1: New taxonomic concept of the Coskinolinidae Moullade, 1965, as presented herein: diagnoses, included genera, and differences. Further comments about genera in the text. Abbreviations: fo = foramen, pi = pillar. Left: Coskinolina sistanensis Schlagintweit & Hadi, 2018, from the Eocene of Iran (unpublished image). Right: Coleiconus zansi Robinson, 1993, from the Eocene of Jamaica (from Robinson & Wright, 1993, Fig. 7.4).
| Family | Coskinolinidae Moullade, 1965, emended | |
| Common features | Orbitoliniform morphology; no horizontal partitions (rafters); thick wall with pseudokeriotheca | |
| Diagnosis | Test conical, early stage trochospiral, then becomes uniserial and rectilinear with broad, low chambers, interior subdivided by pillars and/or radial partitions; wall agglutinated, single-layered, with pseudokeriotheca; aperture basal, cribrate; marginal foramina/apertures may be present (Loeblich & Tappan, 1987) emend. herein | |
| Subfamilies | Coskinolininae Moullade, 1965, emended | Coleiconinae subfam. nov. |
| Diagnosis | Coskinolinidae with undivided marginal zone (no rafters, no beams) | Coskinolinidae with marginal zone subdivided by beams forming alcoves; intercalary beams may be present; beams either aligned or alternating between subsequent chambers |
| Genera included | Coskinolina Stache,
1875, Lituonelloides Henson, 1948, Pseudolituonella Marie, 1954, Parurgonina Cuvillier et al., 1968, Coskinon Hottinger & Drobne, 1980, Cantabriconus Schlagintweit et al., 2017 |
Coleiconus Hottinger
& Drobne, 1980,
|
| Transverse sections | 
Coskinolina Stache |
Coleiconus Hottinger & Drobne |
Phylum Foraminifera (Orbigny, 1826) Pawlowski et al., 2013
Order Textulariida (Delage & Hérouard, 1896) Kaminski, 2004
Order Loftusiida Kaminski & Mikhalevich in Kaminski, 2004
Suborder Orbitolinina Martin, 1890
Superfamily Coskinolinoidea Moullade, 1965
Family Coskinolinidae Moullade, 1965, emended herein
Type genus: Coskinolina Stache, 1875.
Subfamily Coskinolininae Moullade, 1965
Remarks: A subfamily Coskinolininae was proposed by Cimerman (1969) (without diagnosis, included genera, etc.) for biserial textulariids including the genus Pavonitina Schubert, 1914, morphologically completely different from Coskinolina. The Coskinolininae sensu Cimerman (1969) is invalid due to the priority of the Coskinolininae Moullade, 1965 (article 23 ICZN).
Included genera: Coskinolina Stache, 1875; Lituonelloides Henson, 1948; Pseudolituonella Marie, 1954; ? Parurgonina Cuvillier et al., 1968; ? Coskinon Hottinger & Drobne, 1980; ? Cantabriconus Schlagintweit et al., 2017.
Coskinolina Stache, 1875
Type species: Coskinolina liburnica Stache, 1875.
Remarks: The genus includes only Paleogene species (Hottinger & Drobne, 1980; Vicedo et al., 2014; Schlagintweit & Hadi, 2018). The stratigraphic range of Coskinolina is Thanetian to Priabonian (Vicedo et al., 2014; Serra-Kiel et al., 2016b).
Lituonelloides Henson, 1948
Type species: Lituonelloides compressus Henson, 1948.
Remarks: Lituonelloides is a monospecific genus with its only species being L. compressus Henson from the Maastrichtian Simsima Formation of Qatar (see Whittaker et al., 1998). It has been included in the Coskinolinidae by Loeblich and Tappan (1987). Particularly significant, the illustration (drawing) of the holotype provided in Fig. 2a by Henson (1948) shows an undivided marginal zone, one of the characteristic features of the Coskinolininae (see also Pl. 64, figs. 1-2 in Whittaker et al., 1998). The presence of a pseudokeriothecal wall is not evidenced from the poor material available of just one thin-section image in the original description. The drawing in Fig. 2b of Henson (1948) might possibly exhibit endoskeletal pillars aligned between subsequent chambers, a feature not mentioned previously. Lituonelloides still represents a 'poorly known genus' (Whittaker et al., 1998).
Pseudolituonella Marie, 1954
Type species: Pseudolituonella reicheli Marie, 1954.
Remarks: In his 'Note sur Pseudolituonella', Reiss (1959) drew attention to the microgranular nature of the wall, being formed of a single layer, imperforate and non-lamellar (see also Prestat, 1985; Loeblich & Tappan 1987). Serra-Kiel et al. (2016b) established the new species, Pseudolituonella robineti, from the Eocene of Yemen displaying a pseudokeriothecal wall. Serra-Kiel et al. (2016b) were well aware that the wall structure of their Eocene species differed from the one reported for the type species P. reicheli (from the Cenomanian of France) and that an emendation might require study of type specimens of the Cenomanian species. In fact, the presence of a pseudokeriothecal wall structure in P. reicheli has been demonstrated in specimens from the Cenomanian Sarvak Formation of SW Iran (Schlagintweit & Yazdi-Moghadam, 2020).
Parurgonina Cuvillier et al., 1968
Type species: Urgonina (Parurgonina) caelinensis Cuvillier et al., 1968.
Remarks: The type species was originally described as the orbitolinid Urgonina (Parurgonina) caelinensis by Cuvillier et al. (1968) from the Upper Jurassic of Italy. The presence of a pseudokeriothecal wall structure noted by Schroeder et al. (1975, p. 319) 'indicates that this form does not belong to the Orbitolinidae'. On the other hand, Schroeder et al. (1975) did not provide any information on the family status of Parurgonina. The adult stage was indicated as consisting of uniserial chambers (Cuvillier et al., 1968; Schroeder et al., 1975; Loeblich & Tappan, 1987). Loeblich and Tappan (1987) assigned Parurgonina, in my opinion incorrectly, to the Chrysalidinidae Neagu, 1968. Loeblich & Tappan (1987, p. 185) defined the Chrysalidinidae as having a 'test triserial, later biserial' while Parurgonina is defined as possessing a main 'uniserial stage' (p. 186), a contradiction in itself. Septfontaine (1988, p. 248) created the Parurgonininae as a subfamily within the family Valvulinidae Berthelin, 1880, claiming a trochospiral coiled test throughout (eight or more chambers in the adult stage) (= Jurassic 'praevalvulinids' in Septfontaine, 2020, tab. 8.4). In the current classification of Kaminski (2014), the Parurgonininae were given family status (Parurgoninidae). It appears that Parurgonina shares characters of both Chrysalidinidae (trochospiral 'helicospiral' early stage with valvular toothplate) and Coskinolinidae (uniserial adult part). The undivided marginal chamber, marginal foramina and the pseudokeriothecal wall are present in both families (e.g., Hottinger & Drobne, 1980; De Castro, 1981; Banner et al., 1991; Septfontaine, 2020). Parurgonina is herein placed tentatively within the subfamily Coskinolininae Moullade, 1965. Accordingly, the family Parurgoninidae Septfontaine, 1988, with its only genus Parurgonina, would then become obsolete (see also Kaminski, 2014). It is worth mentioning in this context that specimens of Parurgonina caelinensis from the Upper Jurassic of Greece were named 'Coskinolines' by Bassoullet and Guernet (1970) (Schroeder in Schroeder et al., 1975). The stratigraphic range of Parurgonina is late middle Bajocian to Valanginian (Kamoun & Peybernès, 1993; Bassoullet, 1997; Schlagintweit et al., 2023).
Genus Coskinon Hottinger & Drobne, 1980
Type species: Coskinolina (Coskinon) rajkae Hottinger & Drobne, 1980.
Remarks: The Paleogene genus Coskinon was originally described by Hottinger and Drobne (1980) as a subgenus of Coskinolina later established as an independent genus within the Coskinolinidae by Loeblich and Tappan (1987) and then treated as a member of the Pfenderinidae Smout & Sugden, 1962, by Di Carlo et al. (2010). The presence of pseudokeriotheca in Coskinon has not been demonstrated (Hottinger & Drobne, 1980; Di Carlo et al., 2010). As a consequence, Coskinon is herein kept with some reservations in the Coskinolinidae.
Genus Cantabriconus Schlagintweit et al., 2017
Type species: Cantabriconus reocinianus Schlagintweit et al., 2017.
Remarks: The genus includes the two species C.
reocinianus
Schlagintweit et al., 2017
(upper Aptian-lower Albian of
northern Spain; Fig. 1 
) and Cantabriconus? meridionalis
Schlagintweit & Bucur, 2020
(lower Aptian of Romania). Note
that Cantabriconus is considered a
trochospiral pfenderinid or valvulinid morphotype by Septfontaine
(2020).
In fact, some genera of typical orbitoliniform morphology that posses a
pseudokeriothecal wall structure and a central zone with variously shaped
pillars have been assigned to the Pfenderinidae Smout & Sugden,
1962, namely Conicopfenderina Septfontaine
in Kaminski, 2000 (type species Lituonella
mesojurassica Maync, 1972), and Moulladella
Bucur & Schlagintweit, 2018
(type species Meyendorffina
(Paracoskinolina) jourdanensis
Foury & Moullade, 1966). While Conicopfenderina displays an undivided marginal zone as one
characteristic feature of the Coskinolininae, it has been placed into the
Palaeopfenderininae Septfontaine, 1988. According to Septfontaine
(1988), the Eocene genus Lituonella
(= Coskinolina) 'should
not be used for Middle Jurassic orbitoliniform foraminifera, as it has a
different phylogenetic history' (Kaminski, 2000, p. 215). This
highlights the problem that stratigraphically markedly separated taxa of identical
morphology and inner structure are recorded from the
Coskinolininae. An example is the Cenomanian Pseudolituonella
reicheli Marie, 1954, and the distinctly larger Eocene homeomorph Pseudolituonella robineti Gallardo-Garcia & Serra-Kiel,
2016, which are separated not only by a long interval of time, but also two mass extinctions
(at the Cenomanian/Turonian and Cretaceous/Paleogene boundaries). Phylogenetically they
might well be distinct, but morphologically they are near-identical. Further discussion of this principal problem in
foraminiferal taxonomy is beyond the scope of this article.
|
|
Figure 1: Cantabriconus reocinianus Schlagintweit et al., 2017, from the upper Aptian-lower Albian of northern Spain. A) Tangential section (Ramírez del Pozo collection, unpublished). B) Subaxial section (from Schlagintweit et al., 2017, Fig. 6E). C-D) Transverse sections (from Schlagintweit et al., 2017, Figs. 4G, 6G). |
Subfamily Coleiconinae subfam. nov.
Diagnosis: See Table 1.
Type genus: Coleiconus Hottinger & Drobne, 1980.
Included genera: Coleiconus Hottinger & Drobne, 1980; Lepinoconus Cruz-Abad et al., 2017; Ebrahimiella Yazdi-Moghadam & Schlagintweit, 2021.
Coleiconus Hottinger & Drobne, 1980
Type species: Coleiconus zansi Robinson, 1993.
Remarks: In axial sections, the two Paleogene taxa, Coskinolina and Coleiconus, are not distinguishable from each other. For Coleiconus Hottinger & Drobne, 1980, and Coskinolina Stache, 1875, Loeblich and Tappan (1987) noted a 'wall with keriothecal structure', nowadays termed 'pseudokeriotheca' (Hottinger, 2006, p. 29). The principal difference between the two genera, clearly visible in transverse sections, is an undivided marginal zone in Coskinolina versus a subdivided with one order of beams in Coleiconus (Table 1). Coleiconus includes two species, C. zansi Robinson, 1993, and C. christianaensis Robinson, 1993. The recently established species C. minimus Babazadeh, 2022, is considered a nomen dubium by Hadi and Schlagintweit (2024).
Lepinoconus Cruz-Abad et al., 2017
Type species: Lepinoconus chiochinii Cruz-Abad et al., 2017.
Remarks: The taxon is monospecific with the type species Lepinoconus chiocchinii Cruz-Abad et al., 2017, described from the Campanian of southern Italy. This steeply conical taxon is characterized by an exoskeleton of beams and intercalary beams (one between two beams) aligned between subsequent chambers. The absence of marginal foramina (apertures) in Lepinoconus is noteworthy, as other genera within the Coskinolinidae exhibit this feature.
Ebrahimiella Yazdi-Moghadam & Schlagintweit, 2021
Type species: Valdanchella dercourti Decrouez & Moullade, 1974.
Remarks: Ebrahimiella is monospecific with the type species being 'Valdanchella' dercourti Decrouez & Moullade, 1974, from the upper Albian-Cenomanian of Greece. Schroeder (1985, p. 62) noted the presence of a pseudokeriothecal wall-structure in "V." dercourti concluding that 'it is neither a species of Valdanchella Canérot and Moullade nor a representative of the Orbitolinidae but a form that shows some affinities with Coskinolina Stache'. Schroeder (1985), however, did not assign the Greek species to either an existing genus or a new genus. Yazdi-Moghadam and Schlagintweit (2021) introduced the new genus Ebrahimiella for the Greek taxon having beams and intercalary beams (alternating between subsequent chambers) and a pseudokeriothecal wall structure (Schroeder, 1985) rarely visible due to an often relatively thin wall.
The family Coskinolinidae as summarized herein includes nine genera, among which six or seven clearly display a pseudokeriothecal wall structure (Douglass, 1960; Hottinger & Drobne, 1980; Schroeder, 1985; Serra-Kiel et al., 2016b; Schlagintweit & Yazdi-Moghadam, 2020; Mitchell et al., 2020). The definition of the family provided by Moullade (1965, p. 4033, translated from the French) includes a 'monolamellar wall structure, grossly granular, fibrous-like (? perforated)'. In the monograph of Loeblich and Tappan (1987, p. 155) the wall of the Coskinolinidae is defined as 'agglutinated, of granular calcite, single layered'. It appears however that a pseudokeriothecal wall is common to all coskinolinids and is herein integrated into the definition of the wall structure as provided by Loeblich and Tappan (1987). Note that this type of wall structure is not restricted to the Coskinolinidae (e.g., Schlagintweit & Yazdi-Moghadam, 2022). The Coleiconinae exhibit both a pseudokeriotheca and homogeneous (imperforate) radial partitions thus refuting the statement of Septfontaine (1981, 2020, p. 138) that the two features are incompatible with each other. The presence/absence of an exoskeleton subdividing the marginal zone of the chambers is taken as a criterion of subfamily status allowing the distinction of forms lacking an exoskeleton, the Coskinolininae (referring to the type genus), and a second group, Coleiconinae subfam. nov., with vertical partitions (i.e., beams, and in the case of Ebrahimiella Yazdi-Moghadam & Schlagintweit, 2020, also with intercalary beams). Note that the different type of exoskeletons are used as a subfamilial criterion in the Cyclamminidae Marie, 1954 (Septfontaine, 1981, 1988; Loeblich & Tappan 1987).
The new subfamily Coleiconinae incorporates all coskinolinids exhibiting a marginal zone subdivided by vertical partitions (beams, optional with intercalary beams). The new taxonomic concept presented here is in accordance with the hierarchical order of structural features in agglutinated Larger Benthic Foraminifera. Generic features within this subfamily include the arrangement of the main partitions (alternating or aligned), the order of beams and intercalary beams, and the presence or absence of marginal apertures or foramina. The subfamily Coskinolininae is retained for those genera lacking a subdivided marginal zone.
The author expresses his sincere gratitude to the reviewers, François Le Coze (Saint-Étienne) and Mike Simmons (London), for their insightful remarks, and to Bruno Granier (Brest) for his meticulous editing.
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| Nomenclatural note: Life Sciences Identifier (LSID) https://zoobank.org/References/86D9B493-1246-4427-915E-2AF0EC0286BC Family
Group |
