◄ Carnets Geol. 21 (2) ►
Outline:
[1. Introduction]
[2. Discussion on some taxa of the family Chitinoidellidae]
[3. Systematics]
[4. Evolution of Chitinoidellidae and discussion]
[5. Conclusion] and ... [Bibliographic references]
Université Moulay Ismaïl, Faculté des Sciences, Département
de Géologie, BP 11201, Zitoune, Meknès (Morocco)
Published online in final form (pdf) on February 15, 2021
DOI 10.2110/carnets.2021.2102
[Editor: Bruno Granier; language editor:
Stephen Eagar]
Several new genera and species of the family Chitinoidellidae Trejo, 1975, were erected by Pop (1997, 1998a, 1998b). Some of these taxa are justified, but others are inadequately defined, and require revision. I discuss herein the non-validity of some taxa and propose a new systematic classification and an evolutionary framework for the family Chitinoidellidae, with two subfamilies: 1) Dobeninae, which include small-sized chitinoidellids, with the genera Borziella Pop, 1997, Carpathella Pop, 1998a, Daciella Pop, 1998a (amended), Dobenilla n. gen., and Popiella Reháková, 2002, and 2) Bonetinae, which includes larger-sized chitinoidellids, with the genera Bermudeziella n. gen., Bonetilla n. gen., and Furrazolaia n. gen.. These two families are separated in time. Small species of the subfamily Dobeninae characterise the Dobeni Subzone (Ponti ammonite Zone) and disappear immediately before the occurrence of the larger specimens of the subfamily Bonetinae, which characterise the Boneti Subzone (Microcanthum p.p. ammonite Zone).
• Chitinoidellidae;
• Dobeninae;
• Bonetinae;
• systematic revision;
• Jurassic;
• lower/upper Tithonian boundary
Benzaggagh M. (2021).- Systematic revision and evolution of the Tithonian family Chitinoidellidae Trejo, 1975.- Carnets Geol., Madrid, vol. 21, no. 2, p. 27-53.
Révision systématique et évolution de la famille tithonienne des Chitinoidellidae Trejo, 1975.- Plusieurs nouveaux genres et espèces de la famille des Chitinoidellidae Trejo, 1975, ont été créés par Pop (1997, 1998a, 1998b). Certains de ces taxons sont justifiés, mais d'autres sont mal définis et nécessitent une révision. Je discute ici la non-validité de certains taxons et je propose une nouvelle classification systématique et un cadre évolutif pour la famille des Chitinoidellidae, avec deux sous-familles : 1) Dobeninae, qui regroupe des chitinoïdelles de petite taille, avec les genres Borziella Pop, 1997, Carpathella Pop, 1998a, Daciella Pop, 1998a (amendé), Dobenilla n. gen. et Popiella Reháková, 2002, et 2) Bonetinae, qui regroupe des chitinoïdelles de plus grande taille, avec les genres Bermudeziella n. gen., Bonetilla n. gen. et Furrazolaia n. gen.. Ces deux sous-familles sont séparées dans le temps. Les espèces de petite taille de la sous-famille des Dobeninae caractérisent la Sous-Zone à Dobeni (Zone à Ponti des ammonites) et disparaissent immédiatement avant l'apparition des spécimens de plus grande taille de la sous-famille des Bonetinae qui, quant à eux, caractérisent la Sous-Zone à Boneti (Zone à Microcanthum p.p. des ammonites).
• Chitinoidellidae ;
• Dobeninae ;
• Bonetinae ;
• révision systématique ;
• Jurassique ;
• limite Tithonien
inférieur/ supérieur
Chitinoidellids (Pop, 1997) or calpionellids with microgranular lorica constitute a small, but interesting, microplanktonic fossil group of the lower/upper Tithonian boundary interval. They have often been reported in pelagic deposits from both, southern and northern, Tethysian margins, at least from Iran to Mexico, and more recently in the southwestern Pacific margin, Neuquén Basin of western Argentina (Kietzmann, 2017; Kietzmann et al., 2018).
Chitinoidellids have long been considered as a single genus, Chitinoidella Doben, 1963, within the family Calpionellidae Bonet, 1956. Trejo (1975) erected the family Chitinoidellidae, with a single genus: Chitinoidella. Borza (1969, 1984), Grandesso (1977), Borza & Michalík (1986), and Benzaggagh & Atrops (1995) distinguished within the genus Chitinoidella two groups: Chitinoidella gr. dobeni Borza and Chitinoidella gr. boneti Doben, which can be clearly distinguished based on their lorica size and their stratigraphic range. The first group is restricted to the late early Tithonian time period, and includes chitinoidellids with small sized-lorica, with four classical species: Chitinoidella dobeni Borza, 1966; Ch. colomi Borza, 1966; Ch. slovenica Borza, 1969; and Ch. tithonica Borza, 1969. The second group is restricted to the early late Tithonian time period, and includes chitinoidellids with larger sized-lorica, with four classical species: Ch. boneti Doben, 1963; Ch. cristobalensis (Furrazola-Bermúdez, 1965); Ch. cubensis (Furrazola-Bermúdez, 1965); and Ch. bermudezi (Furrazola-Bermúdez, 1965). Each of the two groups defines a chitinoidellid subzone: Dobeni and Boneti respectively, and they correspond to excellent markers of the lower/upper Tithonian boundary interval.
Pop (1997, 1998a, 1998b), in his revision of the chitinoidellids from Romania, erected several new genera and species. Some of these taxa are legitimate; but some others are inadequately defined leading to confusion in the generic and specific attributions of several illustrated specimens in later published works. Because of these confusions and in order to state in a clear and objective way the definition of certain genera and species of chitinoidellids, it becomes necessary to revise the systematics of this microfossil group.
Based on the study of my own material and the revision of the growing number of illustrated chitinoidellid specimens in numerous published work, I discuss herein the non-validity of certain taxa and I propose a new systematic classification and evolutionary patterns of the family Chitinoidellidae around the lower/upper Tithonian boundary, with two subfamilies: Dobeninae, which includes small-sized chitinoidellids of the Dobeni Subzone, with the genera: Borziella Pop, 1997, Carpathella Pop, 1998a, Daciella Pop, 1998a (amended), Dobenilla n. gen., Popiella Reháková, 2002, and Bonetinae, which includes larger-sized chitinoidellids of the Boneti Subzone, with the genera: Bermudeziella n. gen., Bonetilla n. gen., and Furrazolaia n. gen.
It should be noted that the proposed subdivision of the family Chitinoidellidae into two subfamilies is based on the lorica size and the stratigraphic range of the two chitinoidellid groups. According to Borza (1969, Abb. 4), Borza (1984), Grandesso (1977, Fig. 1), Benzaggagh (1988), Benzaggagh & Atrops (1995, Fig. 3), Benzaggagh (2000), Benzaggagh et al. (2010), Reháková (2002, Figs. 2-4), Petrova et al. (2012, Fig. 4), and Lakova & Petrova (2013, Pls. 1, 5), small chitinoidellid species of the dobeni group disappear at the top of the Dobeni Subzone immediately before the occurrence of the larger chitinoidellid species of the boneti group, at the base of the Boneti Subzone. As a result, species of the two groups cannot belong to the same genera.
Dobeniella Pop, 1997
Pop (1997) erected the genus Dobeniella for chitinoidellids showing, according to this author, a composite collar consisting of two pieces: the outer one corresponds to the extension of the lorica wall; the inner is of variable shape: rounded, comma-like, lens-like or irregular.
However, while Pop's drawings (Pop, 1997, Fig. 1.1-3) unambiguously illustrate the composite collar of the so-called genus Dobeniella, none of the figured specimens by the author [e.g., op. cit., Dobeniella cubensis (Furrazola-Bermúdez), Fig. 2, photos 5-6, D. bermudezi (Furrazola-Bermúdez), Fig. 2, photos 7-8, and D. tithonica (Borza), Fig. 2, photo 9] clearly show such a composite collar with two separated pieces. Most of the illustrated specimens show a partially or totally undivided, thick collar and, on several specimens, the thickening can affect the whole lorica wall (op. cit., Fig. 2, photos 5, 9).
It should be noted that the notion of composite collar for the chitinoidellids was first reported by Furrazola-Bermúdez (1965), who gave drawings with composite collar for his two new species: Tintinnopsella cubensis and T. bermudezi [op. cit.,
Lam. 1, figs. 1.a-c and 2.a-c, respectively]. However, the illustrated specimens of the two aforementioned species [op. cit.,
Lam. 2, figs. 1-8; Lam. 3, fig. 1; Lam. 5, figs. 1-2] all show either a collar of the same thickness as the lorica wall
[op. cit., Lam. 2, Fig. 6. (Fig. 3.A 
herein), Lam. 2, figs. 7-8 (Fig. 3.B herein), Lam. 5, fig. 2], or slightly thicker than the lorica wall [op. cit.,
Lam. 3, fig. 1] or an undivided globular collar [op. cit., Lam. 2, figs. 1 (Fig. 3.M herein), 3-5, and Lam. 5, fig. 1].
Analysis of the most figured specimens of Tintinnopsella bermudezi by Furrazola-Bermúdez (1965), shows that the collar piece in comma shape, corresponds to spots of micrite or ion oxides of diagenetic origin jointed to the collar and giving the impression of a composite collar [e.g., op. cit., specimen Lam. 2, fig. 8, with two small diagenetic subtriangular pieces on both sides of the collar; specimen Lam. 5, fig. 2, with two strange thick elongated black diagenetic pieces on both sides of the collar; also the specimens of Tintinnopsella cubensis (Lam. 2, fig. 1; Lam. 5, fig. 1) showing a small black diagenetic piece jointed to the collar, especially on the right side, giving the impression of a composite collar].
Note what Ferrazola-Bermúdez (1965, p. 17) wrote about the collar of Tintinnopsella bermudezi: "unas commas (...), lo que constituye el caracter distinctivo de esta especie. Tal estructura presenta una textura y una composición differentes a las del resto del material que compone la loriga, incluso presenta una coloración pardo-rojiza, distincta a la de las paredes de la loriga".
Moreover, none of the subsequently illustrated specimens later assigned by authors to the genus Dobeniella, clearly show a double collar, consisting of two separate pieces, like, for example, the collar of the genus Remaniella of the family Calpionellidae. There are often thickenings restricted to the collar, or a part of it, or sometimes affecting the whole lorica wall.
Such thickenings, often irregular, perhaps reflect palaeo-environmental factors or, alternatively diagenetic processes, such as secondary precipitations of micrite or iron oxides around the collar or the whole lorica wall. These thickenings can be clearly seen in several illustrated specimens, among others: Pop (1997, Fig. 2, photos 5, 9, 12), Pop (1998b, Pl. 1, figs. 1-2, 4-8, 20, 23, 25-30), Sallouhi et al. (2011, Pl. 1.2, 1.5-7, 1.9, 1.11, 1.13), and Lakova & Petrova (2013, Pl. 1.7-8, 1.15, 1.25-26; Pl. 5.27, 5.29, 5.31-32).
From the above remarks, it becomes obvious to consider the genus Dobeniella Pop (1997) as invalid, because: 1) the notion of composite collar is not evident on any figured specimens attributed to the genus Dobeniella and 2) the chitinoidellids assigned to this genus included specimens of different size and belonging to two distinct chitinoidellid groups: dobeni and boneti [e.g., Dobeniella tithonica from the Dobeni Subzone, and D. cubensis and D. bermudezi from the Boneti Subzone].
Longicollaria Pop, 1997
Longicollaria Pop is defined as a genus including specimens with long collar, equal to one-half of the total lorica length, and showing distal thickening and acute aboral pole ending with a caudal appendage. This genus so defined includes specimens with heterogeneous collar shape, arched or straight. It also includes specimens of small to larger lorica from both the dobeni and boneti groups, such as: small specimens with long collar of Chitinoidella dobeni or Ch. colomi from the Dobeni Subzone; and larger specimens of Ch. cristobalensis, Ch. bermudezi or Ch. insueta Řehánek, 1986, from the Boneti Subzone.
Cubanella Pop, 1997
Cubanella Pop (1997) is defined as a genus including specimens with a long cylindrical to sub-cylindrical collar, often equal to one-half of the total lorica length, terminated by a long caudal appendage.
This genus, like the two previous genera, includes specimens with long collar and caudal appendage belonging to the two different chitinoidellid groups: for instance, some specimens of Ch. colomi or Ch. dobeni from the Dobeni Subzone, and some others of Ch. cristobalensis of the Boneti Subzone.
This generic name was already occupied by an insect taxon (Fennah, 1948), resulting in its later replacement by the following genus, Almajella Pop (1998b).
Almajella Pop, 1998b
Pop (1998b, p. 11) erected the genus Almajella to replace the genus Cubanella Pop 1997, but the author does not give any definition of this new genus (Almajella) and the reason for this change (pre-occupied generic name). Note also that both genera were erected on the same holotype (that is, type specimen of Chitinoidella cristobalensis). Consequently, it is logical to consider the genus Almajella Pop, 1998b, as invalid.
Cylindrella Pop, 1997
According to the author, this genus is characterised by a more or less an oval lorica bowl, with rounded aboral pole and a long cylindrical collar, longer than the total length of the lorica bowl. The single species of this genus, Chitinoidella insueta Řehánek, 1986, was subsequently assigned to the genus Aninella Pop 1998b (genus without any definition) by Pop (1998b), then to the genus Longicollaria by Reháková (2002) and Kowal-Kasprzyk (2014). Actually, considering its straight and long collar, its wide and rounded aboral pole, without caudal appendage, Ch. insueta cannot indeed belong to the genus Longicollaria or even the genus Aninella Pop 1998b, since this author did not provide any definition for his new genus.
Daciella Pop, 1998a
Daciella Pop (1998a) is defined as a genus showing characteristic swellings located below the collar, on each side of the aperture, likewise the swellings of the genus Crassicollaria. But, while Pop's drawings (Pop, 1998a, Fig. 1.2-5) perfectly illustrate the so-called swellings, the figured specimens by Pop (1998a, Fig. 2, photos 6-20; 1998b, PI. I, figs. 5-15) rarely show the presence of such swellings, which can be considered as a constant morphological feature. There are rather irregular thickenings, often of diagenetic origin, of a part or the whole lorica wall. In addition, most of Pop's figured specimens (Pop, 1998a, 1998b) are of poor quality and do not illustrate in a clear way the presence of such swellings. Actually, most of the specimens attributed to the genus Daciella are indeed characterised by an elongated oval-shaped lorica often with an acute aboral pole and a wide aperture with or without preoral swellings.
Note that the specimens qualified as Daciella banatica (Pop, 1998a, Fig. 2, photos 7, 10) are rather specimens of Chitinoidella slovenica and Ch. tithonica, respectively, and the drawings (Pop, 1998a, Fig. 1.2-5) of the four new chitinoidellid species: Daciella banatica, D. almajica, D. danubica, and D. svinitensis, were perfectly modeled on the basis of the four species of the genus Crassicollaria: Cr. brevis, Cr. intermedia, Cr. parvula, and Cr. massutinia, also respectively.
From that, Pop (1998a, p. 822) assumed that the four species of the genus Daciella gave rise to the four species of the genus Crassicollaria. According to Remane (1971), Benzaggagh & Atrops (1995), and Benzaggagh (2020), the four species of the genus Crassicollaria appeared gradually in time and derived from a single ancestor of primitive Crassicollaria.
Chitinoidella lubimovae Furrazola-Bermúdez & Kreisel, 1973, and
Ch. pinarensis Furrazola-Bermúdez & Kreisel, 1973
herein): this specimen is a typical Ch. boneti with a collar extended on both sides by a strange, thick elongated concretion, most likely of iron oxides from the matrix, and two other, thin and long black, lines extending the extremity of the collar on both sides. herein): this figure shows a specimen with lorica and collar shapes similar to those of the holotype of Tintinnopsella cristobalensis Furrazola-Bermúdez, 1965 (Fig. 4.AL herein).Consequently, these two species, Chitinoidella lubimovae Furrazola-Bermúdez & Kreisel, 1973, and Ch. pinarensis Furrazola-Bermúdez & Kreisel, 1973, are invalid.
As discussed above, the current systematics of the family Chitinoidellidae suffers from several drawbacks and makes the correct specific and generic attributions difficult and the use of chitinoidellids as an excellent microfossil marker group for the lower/upper Tithonian boundary interval. Consequently, the systematics of this fossil group requires revision. Based on the lorica size and the stratigraphic distribution, two groups of chitinoidellids can be distinguished, each one having a subfamily rank, designated herein as: Dobeninae and Bonetinae.
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Figure 1: Longitudinal schematic sections of the revised chitinoidellid species. A-D. Dobenilla dobeni; A-B. Dobenilla dobeni var. dobeni, B. with thick collar; C-D. Dobenilla dobeni var. longidobeni, D. with thick collar; E-H. Dobenilla colomi; E-F. Dobenilla colomi var. colomi, F. with thick collar; G-H. Dobenilla colomi var. longicolomi; I-J. Borziella slovenica, I. with annular constriction; K-L. Borziella tithonica, K. with annular constriction; M. Carpathella longirumanica; N. Carpathella rumanica; O-P. Daciella banatica, P. with thick collar; Q-T. Daciella danubica, with variable collar shape; U. Popiella oblongata; V-W. Bermudeziella bermudezi, W. with thick collar; X-Y. Bermudeziella cubensis; Z. Bonetilla boneti; AA. Bonetilla carthagensis; AB. Bonetilla curva; AC. Bonetilla elongata; AD. Bonetilla germanica; AE. Bonetilla lehegarati; AF. Bonetilla miniboneti; AG. Bonetilla popi; AH. Bonetilla praeboneti; AI. Bonetilla sphaerica; AJ. Bonetilla svinitensis; AK. Furrazolaia cristobalensis; AL. Furrazolaia insueta. |
A - Dobeninae n. subfam.
Type species: Chitinoidella dobeni sp. nov. - Borza (1966), Pl. IX, figs. 1-2.
Derivatio nominis: The name of this subfamily derives from Chitinoidella dobeni, which is the most common and typical chitinoidellid species of the Dobeni Subzone.
Diagnosis: This subfamily includes chitinoidellids with small-sized lorica, ca 40 μm-long, and variable shape of the collar.
Occurrence: Species of the subfamily Dobeninae are exclusively restricted to the Dobeni Subzone (late early Tithonian). They are known on both margins of the Tethys Realm: southern Europe, North Africa, and on the southwestern Pacific margin, western Argentina.
Dobenilla n. gen.
Type species: Chitinoidella dobeni sp. n. - Borza (1966), Taf. IX, figs. 1-2.
Derivatio nominis: The name of this genus derives from Chitinoidella dobeni, which is the most common and typical chitinoidellid species of the Dobeni Subzone.
Remark: Specimens of this genus were often attributed to the genera: Dobeniella Pop, Longicollaria Pop, Almajella Pop, and Cubanella Pop.
Diagnosis: Conical to slightly oval-shaped lorica, with arched to slightly straight collar, equal to or greater than one-third of the total lorica length, with or without a distal thickening; rounded to acute aboral pole, with or without a caudal appendage.
Dobenilla dobeni (Borza, 1966) n. comb.
(Figs. 1.A-D , 2.A-R )
Holotype: Chitinoidella dobeni sp. n. - Borza (1966), Taf. IX, figs. 1-2.
Amended Diagnosis: Conical to slightly oval-shaped lorica, with arched collar, equal to or greater than one-third of the total lorica length, with or without a distal thickening; rounded to acute aboral pole, with or without a caudal appendage.
1966 Chitinoidella dobeni sp. n. - Borza, Taf. IX, figs. 1-12; Taf. X, figs. 1-3.
1969 Chitinoidella dobeni Borza - Borza, Taf. LXV, figs. 5, 6 (Fig. 2.I herein), 7, 8 (Fig. 2.A herein), 12-13, 16.
1969 Chitinoidella colomi Borza - Borza, Taf. LXVI, figs. 4 (Fig. 2.K herein), 5, 7.
1977 Chitinoidella dobeni Borza - Grandesso, Tav. II, fig. 12 (Fig. 2.B herein).
1986 Chitinoidella dobeni Borza - Borza & Michalík, Pl. II, fig. 13.
1991 Chitinoidella dobeni - Cecca & Řehánek, Fig. 2.7.
1991 Chitinoidella sp. - Cecca & Řehánek, Fig. 2.8.
1993 Chitinoidella dobeni Borza - Řehánek & Cecca, Pl. 1, fig. 7 (Fig. 2.C herein).
1993 Chitinoidella sp. - Řehánek & Cecca, Pl. 1, fig. 8.
1995 Chit. dobeni - Benzaggagh & Atrops, Fig. 4.1 (Fig. 2.D herein).
1995 Forme de passage Chit. dobeni-Chit. colomi - Benzaggagh & Atrops, Fig. 4.2 (Fig. 2.L herein).
1997 Dobeniella cubensis (Furrazola-Bermúdez) - Pop, Fig. 2, photos 5-6.
1997 Longicollaria dobeni (Furrazola-Bermúdez) - Pop, Fig. 2, photos 12 (Fig. 2.M herein), 13.
1997 Chitinoidella dobeni Borza - Ivanova, Pl. 1, fig. 20.
1998b Longicollaria dobeni (Borza) - Pop, Pl. I, figs. 1-4.
1998b Dobeniella bermudezi (Furrazola-Bermúdez) - Pop, Pl. I, figs. 23-24.
1998b Dobeniella cubensis (Furrazola-Bermúdez) - Pop, Pl. I, fig. 29.
2000 Chitinoidella dobeni Borza - Benzaggagh, Pl. 5, fig. 1.
2000 Chitinoidella colomi Borza - Benzaggagh, Pl. 5, fig. 2 (Fig. 2.J herein).
2002 Dobeniella tithonica (Borza) - Reháková, Fig. 3.11 (Fig. 2.E herein).
2002 Dobeniella colomi (Borza) - Reháková, Fig. 4.1-3.
2002 Dobeniella dobeni (Borza) - Reháková, Fig. 4.5 (Fig. 2.H herein).
2009 Dobeniella tithonica Borza - Boughdiri et al., Fig. 8.17
2009 Dobeniella cubensis Borza - Boughdiri et al., Fig. 8.18.
2009 Longicollaria dobeni Borza - Boughdiri et al., Fig. 8.21.
2010 Chitinoidella colomi Borza - Benzaggagh et al., Fig. 8.1 (Fig. 2.R herein).
2010 Chitinoidella dobeni Borza - Benzaggagh et al., Fig. 8.2.
2011 Longicollaria dobeni (Borza) - Sallouhi et al., Pl. 1.1 (Fig. 2.N herein), 1.2.
2011 Dobeniella tithonica (Borza) - Sallouhi et al., Pl. 1.4.
2011 Dobeniella cubensis (Furrazola-Bermúdez) - Sallouhi et al., Pl. 1.7 (Fig. 2.F herein), 1.8.
2011 Dobeniella bermudezi (Furrazola-Bermúdez) - Sallouhi et al., Pl. 1.11.
2012 Dobeniella colomi (Borza) - Jach et al., Fig. 12.H.
2012 Dobeniella dobeni (Borza) - Jach et al., Fig. 12.I.
2012 Longicollaria dobeni (Borza) - Petrova et al., Fig. 4.2 (Fig. 2.O herein).
2013 Longicollaria dobeni (Borza) - Lakova & Petrova, Pl. 1, figs. 11-12.
2013 Dobeniella colomi (Borza) - Lakova & Petrova, Pl. 1, figs. 13 (Fig. 2.P herein), 14, 15 (Fig. 2.G herein), 16 (Fig. 2.Q herein); Pl. 5, fig. 14.
2013 Longicollaria dobeni (Borza) - Lakova & Petrova, Pl. 5, figs. 18-20.
2014 Longicollaria dobeni (Borza) - Kowal-Kasprzyk, Fig. 3.R.
2015 Chitinoidella dobeni Borza - Benzaggagh et al., Fig. 5.A-B.
2015 Chitinoidella colomi Borza - Benzaggagh et al., Fig. 5.C.
2016 Longicollaria dobeni (Borza) - Michalík et al., Fig. 8.A.
2017 Longicollaria dobeni - Petrova et al., Fig. 3.1.
2017 Daciella almajica - Petrova et al., Fig. 3.7.
2017 Daciella banatica - Petrova et al., Fig. 3.10.
2017 Daciella danubica - Petrova et al., Fig. 3.11.
2018 Dobeniella colomi (Borza) - Kowal-Kasprzyk, Fig. 9.N.
2018 Longicollaria dobeni (Borza) - Kowal-Kasprzyk, Fig. 9.O.
2018 Dobeniella cf. tithonica (Borza) - Kowal-Kasprzyk, Fig. 9.P.
2019 Longicollaria dobeni - Petrova et al., Fig. 4.1-4.
2020 Chitinoidella dobeni Borza - Benzaggagh, Fig. 6.A-B.
2020 Chitinoidella colomi Borza - Benzaggagh, Fig. 6.C.
Description: Polymorph species, with conical to slightly oval-shaped lorica, as high as wide, to slightly higher than wide; convex lorica walls, with maximum width immediately below the collar; arched collar equal to, or greater than, one-third of the total lorica length, showing in axial section two parallel to slightly convergent branches, with or without distal thickening; rounded to acute aboral pole, with or without a caudal appendage. The junction between the collar and the lorica bowl is marked by a constriction or, rarely, a small shoulder.
Variability: This species shows variability in shape of the lorica bowl, from conical to slightly oval elongated; shape of the aboral pole, rounded to slightly acute, with or without caudal appendage; junction between the collar and the lorica bowl, in form of a constriction or a small shoulder; and especially in length of the collar, equal to, or greater than, one-third of the total lorica length, with or without a distal thickening. The length of the collar allows to distinguish two varieties:
, 2.A-H );, 2.I-R ; Holotype, Fig. 2.I ).Occurrence: This species is known in the Dobeni Subzone of the central West Carpathians, western Balkans, Poland, Slovakia, Bulgaria, Romania, Serbia, Italy, northern Tunisia, and northern Morocco.
Dobenilla colomi (Borza, 1966) n. comb.
(Figs. 1.E-H , 2.S-AC )
Holotype: Chitinoidella colomi sp. nov. - Borza, 1966, Taf. X, figs. 4-5.
Amended diagnosis: Conical to slightly oval-shaped lorica, with rectilinear to slightly arched collar, equal to or greater than one-third of the total lorica length, with or without a distal thickening; acute aboral pole, with or without a caudal appendage.
1966 Chitinoidella colomi sp. n. - Borza, Taf. X, figs. 4-7.
1969 Chitinoidella dobeni Borza - Borza, Taf. LXV, figs. 10-15.
1969 Chitinoidella colomi Borza - Borza, Taf. LXVI, figs. 1 (Fig. 2.S herein), 2, 3 (Fig. 2.T herein), 8.
1977 Forma di transition tra Chitinoidella dobeni Borza e Chitinoidella colomi Borza - Grandesso, Tav. II, fig. 11 (Fig. 2.U herein).
1977 Chitinoidella colomi Borza - Grandesso, Tav. II, Fig. 10 (Fig. 2.Y herein).
1995 Chit. colomi - Benzaggagh & Atrops, Fig. 4.3.
1997 Cubanella cristobalensis (Furrazola-Bermúdez) - Pop, Fig. 2, photos 10 (Fig. 2.V herein), 11 (Fig. 2.Z herein).
1998b Dobeniella cubensis (Furrazola-Bermúdez) - Pop, Pl. I, figs. 27, 29.
1998b Almajella cristobalensis (Furrazola-Bermúdez) - Pop, Pl. I, figs. 30-31.
1999 Chitinoidella dobeni Borza - Lakova et al., Pl. 1, fig. 1.
2002 Dobeniella dobeni (Borza) - Reháková, Fig. 4.6.
2009 Dobeniella colomi Borza - Boughdiri et al., Fig. 8.19 (Fig. 2.W herein).
2011 Dobeniella tithonica (Borza) - Sallouhi et al., Pl. 1.3.
2011 Dobeniella bermudezi (Furrazola-Bermúdez) - Sallouhi et al., Pl. 1.12.
2011 Dobeniella colomi (Borza) - Sallouhi et al., Pl. 1.14, 15 (Fig. 2.AB herein), 16.
2012 Dobeniella colomi (Borza) - Petrova et al., Fig. 4.6-8.
2013 Dobeniella colomi (Borza) - Lakova & Petrova, Pl. 5, figs. 14, 15 (Fig. 2.AC herein), 20.
2018 Daciella danubica Pop - Kowal-Kasprzyk, Fig. 9.L-M.
2019 Dobeniella colomi, Petrova et al., Fig. 4.25 (Fig. 2.X herein), 4.26-28.
2020 Chitinoidella colomi Doben - Benzaggagh, Fig. 6.D (Fig. 2.AA herein).
Description: Polymorph species, with conical to slightly oval-shaped lorica, as high as wide, to slightly higher than wide; convex lorica walls, with maximum width located immediately below the collar; rectilinear to slightly arched collar, equal to or greater than one-third of the total lorica length, showing in axial section, two parallel or divergent branches, with or without a distal thickening. The junction between the collar and the lorica bowl is often marked by a more or less developed shoulder; acute aboral pole, often with a more or less developed caudal appendage.
Variability: This species shows variability in shape of the lorica bowl, from conical to slightly oval elongated; shape of the aboral pole, acute, with or without a caudal appendage; more or less developed shoulder; and, especially, in length of the collar equal to or great than one-third of the total lorica length, with or without a distal thickening, and its shape, rectilinear to more or less divergent. The length of the collar allows distinguishing two varieties:
, 2.S-W );, 2.X-AC ; Holotype, Fig. 2.Y ).Comparison: Dobenilla colomi (Borza) is distinguished from Dobenilla dobeni (Borza) by its rectilinear collar, often forming a well-developed shoulder with the lorica bowl; and its acute aboral pole, often with a more or less long caudal appendage.
Occurrence: This species is known in the Dobeni Subzone of the central West Carpathians, western Balkans, Slovakia, Romania, Bulgaria, Serbia, Poland, Italy, northern Tunisia, and northern Morocco.
Borziella Pop, 1997
Type species: Chitinoidella slovenica sp. nov. - Borza (1969), Taf. LXVI, fig. 9 (Fig. 2.AD herein).
Borziella slovenica (Borza, 1969)
(Figs. 1.I-J , 2.AD-AI )
Holotype: Chitinoidella slovenica sp. nov. - Borza (1969), Taf. LXVI, fig. 9 (Fig. 2.AD herein).
1969 Chitinoidella slovenica sp. nov. - Borza, Taf. LXVI, figs. 10-13, 14 (Fig. 2.AE herein), 15-16.
1977 Chitinoidella cf. tithonica Borza - Grandesso, Tav. II, fig. 7 (Fig. 2.AF herein).
1977 Chitinoidella slovenica Borza - Grandesso, Tav. II, figs. 8-9.
1986 Chitinoidella slovenica Borza - Borza & Michalík, Pl. II, fig. 14.
1995 Chit. slovenica - Benzaggagh & Atrops, Fig. 4.4 (Fig. 2.AG herein).
1998a Daciella banatica sp. nov. - Pop, Fig. 2, photo 7.
1998b Daciella banatica sp. nov. - Pop, Pl. I, figs. 11-12.
1998b Borziella slovenica (Borza) - Pop, Pl. I, figs. 16-17.
2000 Chitinoidella slovenica Borza - Benzaggagh, Pl. 5, fig. 3.
2002 Borziella slovenica (Borza) - Reháková, Fig. 2.9-10, 2.11 (Fig. 2.AH herein), 2.12.
2009 Borziella slovenica Borza - Boughdiri et al., Fig. 8.20.
2010 Chitinoidella slovenica Borza - Benzaggagh et al., Fig. 8.4.
2011 Borziella slovenica (Borza) - Sallouhi et al., Pl. 1.18, 20-21.
2013 Borziella slovenica (Borza) - Lakova & Petrova, Pl. 1, figs. 9-10; Pl. 5, figs. 6-7.
2017 Borziella slovenica (Borza) - Kietzmann, Figs. 4.16 (Fig. 2.AI herein), 4.17-18, 5.7-8.
2018 Borziella slovenica (Borza) - Kietzmann et al., Fig. 3.a.
2018 Borziella slovenica (Borza) - Kowal-Kasprzyk, Fig. 9.Q.
2019 Longicollaria dobeni - Petrova et al., Fig. 4.5-8.
2020 Chitinoidella slovenica Borza - Benzaggagh, Fig. 6.E.
Description: Conical to slightly oval-shaped lorica, as high as wide to slightly higher than wide; convex lorica walls; short collar outwardly deflected, forming or not a well-marked annular constriction with the lorica bowl; rounded to slightly acute aboral pole, without caudal appendage.
Variability: This species shows variability mainly in presence (Figs. 1.I , 2.AD, 2.AF-AH ) or not (Figs. 1.J , 2.AE, 2.AI ) of a more or less developed annular constriction below the collar.
Occurrence: This species is known in the Dobeni Subzone of the central West Carpathians, western Balkans, Bulgaria, Poland, Slovakia, Romania, Italy, northern Tunisia, northern Morocco, and western Argentina.
Borziella tithonica (Borza, 1969) n. comb.
(Figs. 1.K-L , 2.AJ-AN )
Holotype: Chitinoidella tithonica Borza - Borza (1969), Taf. LXVII, fig. 1 (Fig. 2.AJ herein).
1969 Chitinoidella tithonica sp. nov. - Borza, Taf. LXVII, fig. 2.
1986 Chitinoidella cf. tithonica Borza - Řehánek, Pl. 1, Fig. 6.
1995 Chit. tithonica - Benzaggagh & Atrops, Fig. 4.5 (Fig. 2.AK herein).
1997 Dobeniella tithonica (Borza) - Pop, Fig. 2, photo 9.
1998a Daciella banatica sp. nov. - Pop, Fig. 2, photos 7, 10.
1998b Daciella danubica sp. nov. - Pop, PI. I, Fig. 6.
1998b Dobeniella tithonica (Borza) - Pop, PI. I, fig. 25.
2000 Chitinoidella tithonica Borza - Benzaggagh, Pl. 5, fig. 4.
2002 Dobeniella tithonica (Borza) - Reháková, Fig. 3.10, 3.12 (Fig. 2.AL herein).
2009 Dobeniella tithonica (Borza) - Michalík et al., Fig. 3.9 (Fig. 2.AM herein).
2010 Chitinoidella tithonica Borza - Benzaggagh et al., Fig. 8.3.
2011 Dobeniella tithonica (Borza) - Sallouhi et al., Pl. 1.5 (Fig. 2.AN herein).
2014 Daciella danubica (Pop) - Kowal-Kasprzyk, Fig. 3.Q.
2019 Dobeniella tithonica - Petrova et al., Fig. 4.29.
2020 Chitinoidella tithonica Borza - Benzaggagh, Fig. 6.F.
Description: Conical to oval-shaped lorica, higher than wide; convex lorica walls; wide aperture surrounded by a thick and short half-crescentic collar, deflected outwards and forming a right or slightly obtuse angle with the lorica wall. Its junction with the lorica bowl is marked or not by a more or less developed annular constriction; acute aboral pole, with or without a short caudal appendage.
Variability: This species shows variability mainly in presence (Figs. 1.K , 2.AJ, 2.AL-AN ) or not (Figs. 1.L , 2.AK ) of a well-developed annular constriction below the collar.
Comparison: Borziella tithonica (Borza) differs from Borziella slovenica (Borza) by its thick and short half-crescentic collar forming almost a right angle with the lorica bowl.
Occurrence: This species is known in the Dobeni Subzone of the central West Carpathians, Slovakia, Romania, Poland, northern Tunisia, and northern Morocco.
Carpathella Pop, 1998a
Type species: Carpathella rumanica sp. nov. - Pop (1998a), Fig. 2, photo 2 (Fig. 2.AR herein).
Carpathella longirumanica n. sp.
Holotype: Chitinoidella sp. - Borza (1969), Taf. LXIX, fig. 5 (Fig. 2.AO herein).
Derivatio nominis: The name of the species derived from the elongated shape of the lorica.
Diagnosis: Cylindrical elongated-shaped lorica, higher than wide, with a straight collar, forming a well-marked shoulder with the lorica bowl; wide and rounded aboral pole.
1997 Cylindrella insueta (Řehánek) - Pop, Fig. 2, photo 16.
2017 Carpathella rumanica Pop - Kietzmann, Fig. 5.9, 5.11 (Fig. 2.AP herein), 5.13.
2018 Carpathella rumanica Pop - Kietzmann et al., Fig. 3.f.
Description: Cylindrical elongated-shaped lorica, higher than wide, rectilinear to slightly convex lorica walls; straight collar, forming a well-marked shoulder with the lorica bowl; wide and rounded aboral pole, without caudal appendage.
Occurrence: This species is known in the Dobeni Subzone of Slovakia, Romania, and western Argentina.
Carpathella rumanica Pop, 1998a
Holotype: Carpathella rumanica sp. nov. - Pop (1998a), Fig. 2, photo 2 (Fig. 2.AQ herein).
1997 Borziella slovenica (Borza) - Pop, Fig. 2, photos 14 (Fig. 2.AR herein), 15.
1998a Carpathella rumanica sp. nov. - Pop, Fig. 2, photos 1, 3-5.
1998a Daciella danubica sp. nov. - Pop, Fig. 2, photo 16.
1998b Carpathella rumanica Pop - Pop, Pl. I, figs. 18-20.
2002 Carpathella rumanica Pop - Reháková, Fig. 2.13, 2.14 (Fig. 2.AS herein), 2.15-16.
2002 Longicollaria dobeni (Borza) - Reháková, Fig. 4.4 (Fig. 2.AT herein).
2013 Longicollaria dobeni (Borza) - Lakova & Petrova, Pl. 1, figs. 9-10.
2017 Carpathella rumanica Pop - Kietzmann, Fig. 5.12.
2017 Daciella danubica - Petrova et al., Fig. 3.12.
Description: Rounded to slightly oval-shaped lorica, as high as wide to slightly higher than wide; straight, more or less elongated collar, forming a well-marked shoulder with the lorica bowl; wide and rounded aboral pole, without caudal appendage.
Variability: This species shows variability mainly in length of the collar.
Comparison: Carpathella rumanica Pop differs from Carpathella longirumanica n. sp. by its rounded and less elongated lorica.
Occurrence: This species is known in the Dobeni Subzone of the central West Carpathians, western Balkans, Romania, Bulgaria, northern Tunisia, western Cuba, and western Argentina.
Daciella Pop, 1998a
Type species: Daciella banatica sp. nov. - Pop (1998a), Fig. 2, photo 6 (Fig. 2.AV herein).
Amended diagnosis: Oval elongated-shaped lorica, higher than wide; wide aperture with or without preoral swellings, followed or not by a short collar; acute aboral pole, with or without a short caudal appendage.
Daciella banatica Pop, 1998a
(Figs. 1.O-P , 2.AU-AY )
Holotype: Daciella banatica sp. nov. - Pop (1998a), Fig. 2, photo 6 (Fig. 2.AU herein).
Amended diagnosis: Oval elongated-shaped lorica, with L/W ratio around 1.5; wide aperture with or without preoral swellings; rounded to acute aboral pole.
1969 Chitinoidella sp. - Borza (1969), Taf. LXIX, figs. 3 (Fig. 2.AV herein), 6.
1998a Daciella danubica sp. nov. - Pop, Fig. 2, photos 15, 18.
1998a Daciella banatica sp. nov. - Pop, Fig. 2, photo 9.
1998b Daciella danubica sp. nov. - Pop, Pl. I, fig. 7.
1998b Daciella banatica sp. nov. - Pop, Pl. I, fig. 10.
1998b Almajella cristobalensis (Furrazola-Bermúdez) - Pop, Pl. I, fig. 33.
2002 Daciella danubica Pop - Reháková, Fig. 2.17-18, 2.19 (Fig. 2.AW herein), 2.20.
2011 Daciella danubica Pop - Sallouhi et al., Pl. 1.17 (Fig. 2.AX herein).
2012 Dobeniella colomi (Borza) - Petrova et al., Fig. 4.6, 4.7 (Fig. 2.AY herein), 4.8.
2012 Daciella banatica Pop - Petrova et al., Fig. 4.9.
2012 Daciella danubica Pop - Petrova et al., Fig. 4.10-11.
2013 Daciella danubica Pop - Lakova & Petrova, Pl. 1, figs. 1, 3-5; Pl. 5, figs. 9-13.
2013 Daciella sp. - Lakova & Petrova, Pl. 5, fig. 16.
2019 Daciella banatica Pop - Petrova et al., Fig. 4.13-16.
2019 Daciella danubica Pop - Petrova et al., Fig. 4.17-20.
Description: Polymorph species, with oval elongated-shaped lorica, higher than wide, with L/W ratio around 1.5; wide aperture, with or without preoral swellings, followed or not by a short straight or outwardly deflected collar, with or without a distal thickening; convex lorica walls; rounded to acute aboral pole, with or without a short caudal appendage.
Occurrence: This species is known in the Dobeni Subzone of the central West Carpathians, western Balkans, Serbia, Bulgaria, Romania, Slovakia, and northern Tunisia.
Daciella danubica Pop, 1998a
(Figs. 1.Q-T , 2.AZ-BH )
Holotype: Daciella danubica sp. nov. - Pop (1998a), Fig. 2, photo 14 (Fig. 2.AZ herein).
Amended diagnosis: Oval elongated-shaped lorica, with L/W ratio around 2; wide aperture with or without preoral swellings; acute aboral pole.
1969 Chitinoidella sp. - Borza, Taf. LXVI, Fig. 6. (Fig. 2.BD herein).
1969 Chitinoidella sp. - Borza, Taf. LXVIX, Fig. 4.
1997 Chitinoidella elongata sp. nov. - Pop, Fig. 2, photo 4 (Fig. 2.BA herein).
1997 Dobeniella bermudezi (Furrazola-Bermúdez) - Pop, Fig. 2.7-8.
1998a Daciella danubica sp. nov. - Pop, Pl. I, Fig. 1, photo 16.
1998a Daciella almajica sp. nov. - Pop, Fig. 2, photos 11 (Fig. 2.BG herein), 13.
1998a Daciella svinitensis sp. nov. - Pop, Fig. 2, photo 20.
1998b Daciella danubica sp. nov. - Pop, Pl. I, fig. 5.
1998b Daciella svinitensis sp. nov. - Pop, Pl. I, fig. 9.
1998b Daciella almajica sp. nov. - Pop, Pl. I, fig. 14.
1998b Aninella insueta (Řehánek) - Pop, Pl. I, fig. 21.
1998b Dobeniella bermudezi (Řehánek) - Pop, Pl. I, figs. 22-23.
2011 Daciella svinitensis Pop - Sallouhi et al., Pl. 1.19 (Fig. 2.BF herein).
2002 Daciella svinitensis Pop - Reháková, Fig. 3.1-3.
2012 Daciella svinitensis Pop - Petrova et al., Fig. 4.3, 4.5.
2013 Daciella danubica Pop - Lakova & Petrova, Pl. 1, figs. 1 (Fig. 2.BC ), 2 (Fig. 2.BE ); Pl. 5, figs. 9 (Fig. 2.BB herein), 10-13.
2013 Daciella svinitensis Pop - Lakova & Petrova, Pl. 1, figs. 6-8; Pl. 5, figs. 1-2, 3 (Fig. 2.BH herein), 4-5.
2019 Daciella almajica - Petrova et al., Fig. 4.9-12.
2019 Daciella svinitensis - Petrova et al., Fig. 4.21-24.
Description: Polymorph species, with oval elongated-shaped lorica, higher than wide, with L/W ratio around 2; wide aperture, with or without preoral swellings, extended or not by a short collar of variable shape: straight, arched or outwardly deflected, with or without distal thickening; acute aboral pole, with or without a short caudal appendage.
Comparison: Daciella danubica Pop differs from Daciella banatica Pop mainly by its more elongated lorica.
Occurrence: This species is known in the Dobeni Subzone of the central West Carpathians, western Balkans, Romania, Slovakia, Serbia, Bulgaria, and northern Tunisia.
Popiella Reháková, 2002
Popiella oblongata Reháková, 2002
Holotype: Popiella oblongata sp. nov. - Reháková (2002), Fig. 4.10.
1986 Chitinoidella sp. - Řehánek, Pl. 1, fig. 5 (Fig. 2.BJ herein).
2002 Popiella oblongata sp. nov. - Reháková, Fig. 4.10, 4.11 (Fig. 2.BI herein), 4.12.
2011 Popiella oblongata Reháková - Sallouhi et al., Pl. 1.22 (Fig. 2.BK herein).
2020 Popiella oblongata Reháková - Benzaggagh, Fig. 6.G (Fig. 2.BL herein).
Description: Cylindrical elongated-shaped lorica, higher than wide, with straight and parallel walls; wide aperture without collar; rounded to slightly acute aboral pole.
Occurrence: This species is known in the Dobeni Subzone of the central West Carpathians, Slovakia, northern Tunisia, and northern Morocco.
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Figure 2: Main genera and species of the subfamily Dobeninae (Chitinoidellidae) from the Dobeni Subzone (late early Tithonian). A-R. Dobenilla dobeni (Borza): A-H. Dobenilla dobeni (Borza) var. dobeni n. var.: A. Chitinoidella dobeni Borza, excerpt of Borza, 1969, Taf. LXV, fig. 8; B. Chitinoidella dobeni Borza, excerpt of Grandesso, 1977, Tav. II, fig. 12; C. Chitinoidella dobeni Borza, excerpt of Řehánek & Cecca, 1993, Pl. 1, fig. 7; D. Chitinoidella dobeni, excerpt of Benzaggagh & Atrops, 1995, Fig. 4.1; E. Dobeniella tithonica (Borza), excerpt of Reháková, 2002, Fig. 3.11; F. Dobeniella cubensis (Furrazola-Bermúdez), excerpt of Sallouhi et al., 2011, Pl. 1.7; G. Dobeniella colomi (Borza), excerpt of Lakova & Petrova, 2013, Pl. 1, fig. 15; H. Longicollaria dobeni (Borza), excerpt of Reháková, 2002, Fig. 4.5; I-R. Dobenilla dobeni (Borza) var. longidobeni n. var.: I. Chitinoidella dobeni Borza, excerpt of Borza, 1969, Taf. LXV, Fig. 6 (Holotype); J. Chitinoidella colomi Borza, excerpt of Benzaggagh, 2000, Pl. 5, fig. 2; K. Chitinoidella colomi Borza, excerpt of Borza, 1969, Taf. LXVI, fig. 4; L. Forme de passage Chitinoidella dobeni- Chitinoidella colomi, excerpt of Benzaggagh & Atrops, 1995, Fig. 4.2; M. Longicollaria dobeni (Borza), excerpt of Pop, 1997, Fig. 2.12; N. Longicollaria dobeni (Borza), excerpt of Sallouhi et al., 2011, Pl. 1.1; O. Longicollaria dobeni (Borza), excerpt of Petrova et al.,
2012, Fig. 4.2; P. Dobeniella colomi (Borza), excerpt of Lakova & Petrova, 2013, Pl. I, fig. 13; Q. Dobeniella colomi (Borza), excerpt of Lakova & Petrova, 2013, Pl. I, fig. 16; R. Chitinoidella colomi Borza, excerpt of Benzaggagh et al., 2010, Fig.8.1. S-AC. Dobenilla colomi (Borza): S-W. Dobenilla colomi (Borza) var. colomi n. var.: S. Chitinoidella colomi Borza, excerpt of Borza, 1969, Taf. LXVI, fig. 1; T. Chitinoidella colomi Borza, excerpt of Borza, 1969, Taf. LXVI, fig. 3; U. Forma di transition tra Chitinoidella dobeni Borza Chitinoidella colomi Borza, excerpt of Grandesso, 1977, Tav. II, fig. 11. V. Cubanella cristobalensis (Furrazola-Bermúdez), excerpt of Pop, 1997, Fig. 2.10; W. Dobeniella colomi Borza, excerpt of Boughdiri et al., 2009, Fig. 8.19; X-AC. Dobenilla colomi (Borza) var. longicolomi n. var.: X. Dobeniella colomi, excerpt of Petrova et al.,
2019, Fig. 4.25; Y. Chitinoidella colomi Borza, excerpt of Grandesso, 1977, Tav. II, fig. 10 (Holotype); Z. Dobeniella cristobalensis (Furrazola-Bermúdez), excerpt of Pop, 1997, Fig. 2, photo 11; AA. Chitinoidella colomi Borza, excerpt of Benzaggagh, 2020, Fig. 5.20; AB. Dobeniella colomi (Borza), excerpt of Sallouhi et al., 2011, Pl. 1.15; AC. Dobeniella colomi (Borza), excerpt of Lakova & Petrova, 2013, Pl. 5, fig. 15. AD-AI. Borziella slovenica (Borza): AD. Chitinoidella slovenica Borza, excerpt of Borza, 1969, Taf. LXVI, fig. 9 (Holotype); AE. Chitinoidella slovenica Borza, excerpt of Borza, 1969, Taf. LXVI, fig. 14 (Paratype); AF. Chitinoidella slovenica Borza, excerpt of Grandesso, 1977, Tav. II, fig. 8; AG. Chitinoidella slovenica Borza, excerpt of Benzaggagh & Atrops, 1995, Fig. 4.4; AH. Borziella slovenica (Borza), excerpt of Reháková, 2002, Fig. 2.11; AI. Borziella slovenica (Borza), excerpt of Kietzmann, 2017, Fig. 4.16. AJ-AN. Borziella tithonica (Borza); AJ. Chitinoidella tithonica Borza, excerpt of Borza, 1969, Taf. LXVII, fig. 1 (Holotype); AK. Chitinoidella tithonica, excerpt of Benzaggagh & Atrops, 1995, Fig. 4.5; AL. Dobeniella tithonica (Borza), excerpt of Reháková, 2002, Fig. 3.12; AM. Dobeniella tithonica (Borza), excerpt of Michalík et al., 2009, Fig. 3.9; AN. Dobeniella tithonica (Borza), excerpt of Sallouhi et al., 2011, Pl. 1.5. AO-AP. Carpathella longirumanica n. sp.: AO. Chitinoidella sp., excerpt of Borza, 1969, Taf. LXIX, fig. 5 (Holotype); AP. Carpathella rumanica Pop, excerpt of Kietzmann, 2017, Fig. 5.11 (Paratype). AQ-AT. Carpathella rumanica Pop: AQ. Carpathella rumanica Pop, excerpt of Pop, 1998a, Fig. 2.2 (Holotype); AR. Borziella slovenica (Borza), excerpt of Pop, 1997, Fig. 2.14; AS. Carpathella rumanica Pop, excerpt of Reháková, 2002, Fig. 2.14; AT. Longicollaria dobeni (Borza), excerpt of Reháková, 2002, Fig. 4.4. AU-AY. Daciella banatica Pop: AU. Daciella banatica Pop, excerpt of Pop, 1998a, Fig. 2, photo 6 (Holotype); AV. Chitinoidella sp., excerpt of Borza, 1969, Taf. LXIX, fig. 3; AW. Daciella danubica Pop, excerpt of Reháková, 2002, Fig. 2.19; AX. Daciella danubica Pop, excerpt of Sallouhi et al., 2011, Pl. 1.17; AY. Dobeniella colomi (Borza), excerpt of Petrova et al.,
2012, Fig. 4.7; AZ-BH. Daciella danubica Pop: AZ. Daciella danubica Pop, excerpt of Pop, 1998a, Fig. 2.14 (Holotype); BA. Chitinoidella elongata Pop, excerpt of Pop, 1997, Fig. 2.4; BB. Daciella danubica Pop, excerpt of Lakova & Petrova, 2013, Pl. 5, fig. 9; BC. Daciella danubica Pop, excerpt of Lakova & Petrova, 2013, Pl. 1, fig. 1; BD. Chitinoidella sp., excerpt of Borza, 1969, Taf. LXVI, Fig. 6; BE. Daciella danubica Pop, excerpt of Lakova & Petrova, 2013, Pl. 1, fig. 2; BF. Daciella svinitensis Pop, excerpt of Sallouhi et al., 2011, Pl. 1.19; BG. Daciella almajica Pop, excerpt of Pop, 1998a, fig. 2, photo 11; BH. Daciella svinitensis Pop, excerpt of Lakova & Petrova, 2013, Pl. 5, fig. 3. BI-BL. Popiella oblongata Reháková: BI. Popiella oblongata Reháková, excerpt of Reháková, 2002, Fig. 4.11; BJ. Chitinoidella sp., excerpt of Řehánek, 1986, Pl. I, fig. 5; BK. Popiella oblongata Reháková, excerpt of Sallouhi et al., 2011, Pl. 1.22. BL. Popiella oblongata Reháková, excerpt of Benzaggagh, 2020, Fig. 6.G). |
B - Bonetinae n. subfam.
Type species: Chitinoidella boneti sp. nov. - Doben (1963), Taf. 6, Abb. 1-5.
Derivatio nominis: The name of this subfamily derives from Chitinoidella boneti, which is the most common and typical chitinoidellid species of the Boneti Subzone.
Diagnosis: This subfamily includes chitinoidellids of larger size, ca 80 μm-long, with variable size and shape of the collar.
Occurrence: Species of the subfamily Bonetinae range from the base of the Boneti Subzone to the base of the Crassicollaria Zone (AO or Chitinoidellids/Primitive Calpionellids Subzone, Benzaggagh, 2020). They are known on both margins of the Tethysian Realm, from Iran to Mexico and in the southwestern Pacific margin (western Argentina).
Bermudeziella n. gen.
Type species: Tintinnopsella bermudezi sp. nov. - Furrazola-Bermúdez (1965), Lam. 2, Fig. 6. (Fig. 3.A herein).
Remark: Specimens of this genus were often attributed to the genera: Dobeniella Pop, Longicollaria Pop, Almajella Pop, and Cubanella Pop.
Derivatio nominis: This genus is dedicated to Dr. Furrazola-Bermúdez who discovered and described three chitinoidellid species of the boneti group: Tintinnopsella cubensis, T. bermudezi and Calpionella cristobalensis.
Diagnosis: Oval elongated-shaped lorica, with arched collar, with or without a partial or total distal thickening.
Bermudeziella bermudezi (Furrazola-Bermúdez, 1965) n. comb.
(Figs. 1.V-W , 3.A-L )
Holotype: Tintinnopsella bermudezi sp. nov. - Furrazola-Bermúdez (1965), Lam. 2, Fig. 6. (Fig. 3.A herein).
Amended diagnosis: Conical to oval elongated-shaped lorica, with arched collar, with or without a partial or total distal thickening.
1965 Tintinnopsella bermudezi sp. nov. - Furrazola-Bermúdez, Lam. 2, figs. 7-8 (Fig. 3.B herein); Lam. 3, fig. 1; Lam. 5, fig. 2.
1969 Chitinoidella cubensis (Furrazola-Bermúdez) - Borza, Taf. LXVIII, figs. 14 (Fig. 3.C herein), 15.
1969 Chitinoidella bermudezi (Furrazola-Bermúdez) - Borza, Taf. LXVIX, fig. 1.
1973 Chitinoidella bermudezi (Furrazola) - Furrazola-Bermúdez & Kreisel, Lam. 1, figs. 3-4, 5 (Fig. 3.F herein).
1977 Chitinoidella cubensis (Furrazola-Bermúdez) - Grandesso, Tav. II, fig. 4 (Fig. 3.D herein).
1977 Chitinoidella bermudezi (Furrazola-Bermúdez) - Grandesso, Tav. II, fig. 5 (Fig. 3.E herein).
1997 Chitinoidella bermudezi Furrazola-Bermúdez - Grün & Blau, Pl. I, Fig. 6.
2002 Dobeniella bermudezi (Furrazola-Bermúdez) - Reháková, Fig. 3.7-8, 3.9 (Fig. 3.G herein).
2010 Chitinoidella cristobalensis (Furrazola-Bermúdez) - Benzaggagh et al., Fig. 8.9.
2010 Dobeniella cubensis (Furrazola-Bermúdez) - Pszczółkowski & Myczyński, Fig. 12.11.
2011 Dobeniella bermudezi (Furrazola-Bermúdez) - Sallouhi et al., Pl. 1.13 (Fig. 3.H herein).
2012 Chitinoidella cubensis (Furrazola-Bermúdez) - Benzaggagh et al., Fig. 6.D (Fig. 3.I herein).
2013 Dobeniella bermudezi (Furrazola-Bermúdez) - Lakova & Petrova, Pl. 1, fig. 26; Pl. 5, fig. 27 (Fig. 3.J herein).
2013 Dobeniella cubensis (Furrazola-Bermúdez) - Lakova & Petrova, Pl. 5, figs. 30-31.
2017 Dobeniella cf. pinaraensis (Furrazola-Bermúdez & Kreisel) - Kietzmann, Fig. 5.14 (Fig. 3.K herein).
2017 Dobeniella colomi - Petrova et al., Fig. 3.16-17.
2017 Dobeniella bermudezi - Petrova et al., Fig. 4.7 (Fig. 3.L herein).
2017 Almajella cristobalensis - Petrova et al., Fig. 4.12.
2019 Dobeniella bermudezi - Petrova et al., Fig. 5.10-11.
2019 Dobeniella cubensis - Petrova et al., Fig. 5.15.
2019 Almajella cristobalensis - Petrova et al., Fig. 5.17, 5.19.
2020 Chitinoidella bermudezi (Furrazola-Bermúdez) - Benzaggagh, Fig. 6.AA.
Description: Conical to oval elongated-shaped lorica, higher than wide; convex lorica walls, with maximum width towards the mid-lorica bowl; wide aperture, surrounded by a more or less developed arched collar, showing in longitudinal section two parallel or divergent branches, with or without distal thickening, forming with the lorica bowl a small constriction; rounded to slightly acute aboral pole, with or without a caudal appendage.
Comparison: Bermudeziella bermudezi differs from Dobenilla dobeni by its larger lorica, with maximum width towards the mid-lorica bowl, and its collar often shorter than one-third of the total lorica length; and its stratigraphic range.
Occurrence: This species is known in the Boneti Subzone of the central West Carpathians, western Balkans, Bulgaria, Slovakia, Italy, Iran, northern Tunisia, northern Morocco, Cuba, and western Argentina.
Bermudeziella cubensis (Furrazola-Bermúdez, 1965) n. comb.
(Figs. 1.X-Y , 3.M-S )
Holotype: Tintinnopsella cubensis sp. nov. - Furrazola-Bermúdez (1965), Lam. 2, fig. 1; Lam. 5, fig. 1 (same specimen (Fig. 3.M herein).
Amended diagnosis: Conical to oval elongated-shaped lorica, with short arched collar, forming a globular piece.
1965 Tintinnopsella cubensis sp. nov. - Furrazola-Bermúdez, Lam. 2, figs. 2-5.
1969 Chitinoidella cubensis (Furrazola-Bermúdez) - Borza, Taf. LXVIII, fig. 16 (Fig. 3.N herein).
1973 Chitinoidella cubensis (Furrazola) - Furrazola-Bermúdez & Kreisel, Lam. 1, figs. 6 (Fig. 3.Q herein), 7.
1989 Chitinoidella cf. cubensis (Furrazola-Bermúdez) - Cecca et al., Pl. 6, fig. 3 (Fig. 3.O herein).
1989 Chitinoidella boneti Doben - Cecca et al., Pl. 6, Fig. 6.
1995 Chit. cubensis - Benzaggagh & Atrops, Fig. 4.8 (Fig. 3.P herein).
1997 Chitinoidella boneti Doben - Ivanova, Pl. I, fig. 18.
2000 Chitinoidella cubensis (Furrazola-Bermúdez) - Benzaggagh, Pl. 5, fig. 8.
2002 Dobeniella cubensis (Furrazola-Bermúdez) - Reháková, Fig. 3.4, 3.6.
2013 Dobeniella cf. cubensis (Furrazola-Bermúdez) - Lakova & Petrova, Pl. 1, figs. 24, 25 (Fig. 3.R herein).
2013 Dobeniella cubensis (Furrazola-Bermúdez) - Lakova & Petrova, Pl. 5, fig. 29 (Fig. 3.S herein).
2017 Dobeniella cubensis - Petrova et al., Fig. 4.11.
2019 Dobeniella cubensis - Petrova et al., Fig. 5.13-14.
2020 Chitinoidella cubensis (Furrazola-Bermúdez) - Benzaggagh, Fig. 6.N-O.
Description: Conical to oval elongated-shaped lorica, higher than wide; convex lorica walls, with maximum width towards the mid-lorica bowl; wide aperture surrounded by a small arched collar forming a globular piece; rounded to acute aboral pole, with or without a caudal appendage.
Variability: This species shows variability in the collar shape, forming a more or less developed globular piece; as well as the aboral pole, rounded acute, with or without a caudal appendage.
Comparison: Bermudeziella cubensis differs from the other species of the subfamily Bonetinae by its short collar forming a globular piece. It differs from Dobenilla dobeni by its larger lorica, with maximum width towards the mid-lorica bowl; its shorter collar and its stratigraphic range.
Occurrence: This species is known in the Boneti Subzone of the central West Carpathians, western Balkans, Bulgaria, Slovakia, southeastern France, northern Morocco, and Cuba.
Bonetilla n. gen.
Type species: Chitinoidella boneti sp. nov. - Doben (1963), Taf. 6, Abb. 3-4 (Fig. 4.H herein).
Derivatio nominis: The name of this genus derives from Chitinoidella boneti, which is the most common and typical chitinoidellid species of the Boneti Subzone.
Diagnosis: Oval elongated-shaped lorica; wide aperture, surrounded by an outwards deflected collar, forming a variable angle with the lorica bowl.
Bonetilla boneti (Doben, 1963) n. comb.
Holotype: Chitinoidella boneti sp. nov. - Doben (1963), Taf. 6, Abb. 3-4 (Fig. 4.H herein).
Remarks: Doben (1963) gave in his Taf. 6, four specimens of Chitinoidella boneti n. sp., with two different lorica shape. The specimen, figured in Taf. 6, Abb. 3 (op. cit.,
"holotype", Fig. 4.H herein), shows an elongated lorica with a L/W ratio equal to 2, a maximum width at the lower third of the lorica bowl, a well-marked preoral narrowing, and a more developed outwardly deflected collar. This specimen is close to the holotype of Chitinoidella elongata Pop (1997, Fig. 3). The specimens of Taf. 6, Abb. 1 (the two specimens) and Abb. 2 (op. cit.,
"Paratypoide", Fig. 3.T herein) display a wide lorica, with L/W ratio around 1.5, maximum width located at the mid-lorica bowl, and a wide aperture, surrounded by shorter outwardly deflected collar. These two specimens of Chitinoidella boneti should be regarded as two separate species. Although the majority of Chitinoidella boneti figured by authors are consistent with the paratypes
(op. cit., Taf. 6, Abb. 1-2, 5) but because they are not consistent with holotype
(op. cit., Taf. 6, Abb. 1-2, 5), one should consider that they belong to a discrete species (i.e., Bonetilla germanica n. sp.).
Diagnosis: Oval to elongated oval-shaped lorica, with more or less convex walls; wide aperture, surrounded by a curved outwardly deflected collar, forming an obtuse angle with the lorica bowl.
1969 Chitinoidella boneti Doben - Borza, Taf. LXVII, fig. 14; Taf. LXVIII, figs. 8 (Fig. 4.K herein), 9, 10 (Fig. 4.I herein), 11-13.
1985 Sturiella cf. oblonga Borza - Řehánek, Pl. II, fig. 10.
1991 Chitinoidella boneti Doben - Altiner & Özkan, Pl. 3, figs. 1, 2 (Fig. 4.O herein).
2006 Chitinoidella boneti Doben - Grabowski & Pszczółkowski, Fig. 7.B.
2010 Longicollaria insueta (Řehánek) - Pszczółkowski & Myczyński, Fig. 12.9 (Fig. 4.L herein).
2011 Chitinoidella boneti Doben - Sallouhi et al., Pl. 1.24 (Fig. 4.J herein).
2012 Chitinoidella bermudezi (Furrazola-Bermúdez) - Cecca et al., Fig. 3.A (Fig. 4.M herein).
2012 Chitinoidella boneti Doben - Cecca et al., Fig. 3.B.
2012 Chitinoidella boneti Doben - Benzaggagh et al., Fig. 6.A.
2012 Chitinoidella cristobalensis (Furrazola-Bermúdez) - Benzaggagh et al., Fig. 6.C.
2012 Chitinoidella bermudezi (Furrazola-Bermúdez) - Benzaggagh et al., Fig. 6.E.
2012 Dobeniella bermudezi (Furrazola-Bermúdez) - Petrova et al., Fig. 4.17
2013 Chitinoidella boneti Doben - Lakova & Petrova, Pl. 1, fig. 18 (Fig. 4.N herein); Pl. 5, fig. 21.
2017 Longicollaria dobeni - Petrova et al., Fig. 3.2.
Description: Oval to elongated-shaped lorica, higher than wide; with a L/W ratio greater than 1.5; sub-parallel to slightly convex lorica walls, with maximum width located between the median and the lower thirds of the lorica bowl; wide aperture, surrounded by a curved outwardly deflected collar, extending the lorica wall, and forming with this latter an obtuse angle (> 90°); rounded to slightly acute aboral pole, without caudal appendage.
Comparison: Bonetilla boneti (Doben, 1963) differs from Bonetilla curva n. sp. by its more elongated lorica with sub-parallel to slightly convex walls.
Occurrence: This species is known in the Boneti Subzone of the western Balkans, Bulgaria, Slovakia, Poland, Italy, Turkey, Iran, and western Cuba.
Bonetilla carthagensis (Sallouhi et al., 2011) n. comb.
(Figs. 1.AA , 3.AE-AM )
Holotype: Chitinoidella carthagensis sp. nov. - Sallouhi et al. (2011), Pl. 1.28 (Fig. 3.AE herein).
Amended diagnosis: Oval elongated-shaped lorica, higher than wide; convex to sub-parallel lorica walls, with maximum width located between the middle and the upper third of the lorica bowl; wide aperture surrounded by a half-crescentic collar, outwardly deflected and forming an annular constriction with the lorica bowl; rounded to acute aboral pole, with or without a short caudal appendage.
1969 Chitinoidella boneti Doben - Borza, Taf. LXVII, figs. 3, 4 (Fig. 3.AG herein), 5-6, 8, 12 (Fig. 3.AM herein).
1975 Chitinoidella cf. Ch. boneti - Lugo, Lam. I, figs. 4-6 (same specimen); Lam. III, figs. 5 (Fig. 3.AH herein), 8.
1997 Chitinoidella sp. 1 - Grün & Blau, Pl. I, fig. 7 (Fig. 3.AI herein).
2011 Chitinoidella carthagensis sp. nov. - Sallouhi et al., Pl. 1.26-27.
2011 Chitinoidella carthagensis sp. nov. - Sallouhi et al., Pl. 1.29 (Fig. 3.AF herein).
2011 Chitinoidella hegarati sp. nov. - Sallouhi et al., Pl. 1.30-31.
2012 Chitinoidella sp. 2 Sallouhi, Boughdiri & Cordey - Petrova et al., Pl. 4.13.
2014 Chitinoidella carthagensis Sallouhi, Boughdiri & Cordey - Kowal-Kasprzyk, Fig. 3.D (Fig. 3.AJ herein).
2014 Dobeniella tithonica (Borza) - Kowal-Kasprzyk, Fig. 3.M.
2015 Chitinoidella boneti Doben - Benzaggagh et al., Fig. 5.E (Fig. 3.Ak herein).
2017 Chitinoidella hegarati Sallouhi, Boughdiri & Cordey - Kietzmann, Figs. 4.11, 4.12 (Fig. 3.Al herein), 5.1, 5.4.
2017 Chitinoidella boneti Doben - Kietzmann, Figs. 4.10, 5.1
2018 Chitinoidella hegarati Sallouhi, Boughdiri & Cordey - Kietzmann et al., Fig. 3.d.
2018 Chitinoidella carthagensis Sallouhi, Boughdiri & Cordey - Kowal-Kasprzyk, Fig. 9.C.
2019 Chitinoidella carthagensis - Petrova et al., Fig. 5.25-30.
2019 Chitinoidella hegarati - Petrova et al., Fig. 5.31-33, 5.35-36.
2020 Chitinoidella carthagensis Sallouhi et al. - Benzaggagh, Fig. 6.Q-S.
Description: Elongated oval-shaped lorica, higher than wide, with a L/W ratio equal or greater than 1.5; convex to sub-parallel lorica walls, with maximum width located between the median and the upper thirds of the lorica bow, wide aperture surrounded by a short half-crescentic collar, outwardly deflected and forming a right to slightly obtuse angle with the lorica wall. Its junction with the lorica bowl is marked by an annular constriction; rounded to acute aboral pole, with or without a short caudal appendage.
Variability: This species shows variability in more or less elongated lorica; shape of the aboral pole, rounded, acute, with or without a caudal appendage; and more or less developed collar.
Comparison: Bonetilla carthagensis is distinguished from Bonetilla germanica n. sp. by the presence of a constriction below the collar and its maximum width often located towards the upper third of the lorica bowl.
Occurrence: This species is known in the Boneti Subzone of the central West Carpathians, Slovakia, Serbia, Poland, Italy, northern Tunisia, northern Morocco, southeastern Mexico, and western Argentina.
Bonetilla curva n. sp.
(Figs. 1.AB , 3.AN-AT )
Holotype: Chitinoidella boneti Doben - Borza (1969), Taf. LXVII, fig. 12 (Fig. 3.AN herein).
Derivatio nominis: The name of the species derives from the curved shape of the collar.
Diagnosis: Rounded to slightly oval-shaped lorica; wide aperture surrounded by a curved and outwardly deflected collar, forming an obtuse angle with the lorica bowl.
1969 Chitinoidella boneti Doben - Borza, Taf. LXVII, figs. 10, 13, 15-16.
1977 Chitinoidella boneti Doben - Grandesso, Tav. II, fig. 3.
1989 Chitinoidella boneti Doben - Cecca et al., Pl. 6, figs. 1, 2 (Fig. 3.AO herein), 4, 7.
2002 Chitinoidella elongata Pop - Reháková, Fig. 2.1 (Fig. 3.AP herein).
2006. Chitinoidella boneti Doben - Grün & Blau, Pl. I, fig. 5 (Fig. 3.AQ herein).
2007 Chitinoidella boneti Doben - Andreini et al., Pl. 1.5a (Fig. 3.AR herein).
2012 Chitinoidella boneti Doben - Petrova et al., Fig. 4.14-15.
2013 Chitinoidella boneti Doben - Lakova & Petrova, Pl. 5, fig. 23 (Fig. 3.AS herein).
2017 Dobeniella bermudezi - Petrova et al., Fig. 4.8 (Fig. 3.AT herein).
2019 Dobeniella bermudezi - Petrova et al., Fig. 5.8.
2020 Chitinoidella bermudezi Furrazola-Bermúdez - Benzaggagh, Fig. 6.L.
Description: Rounded to oval-shaped lorica, as high as wide, or slightly higher than wide; regularly convex lorica walls, with maximum width towards the mid-lorica bowl; wide aperture surrounded by a curved and outwardly deflected collar, extending the lorica wall and forming with this later an obtuse angle (> 90°); rounded to slightly acute aboral pole, without caudal appendage.
Comparison: Bonetilla curva n. sp. differs from the other species of the genus Bonetilla by its long and curved collar, in continuity with the lorica wall and forming with this later an obtuse angle. It differs from Borziella slovenica by its larger size, the lack of an annular constriction below the collar, and its stratigraphic range.
Occurrence: This species is known in the Boneti Subzone of the central West Carpathians, western Balkans, Slovakia, Serbia, Poland, Bulgaria, Italy, southeastern France, northern Tunisia, and northern Morocco.
Bonetilla elongata (Pop, 1997) n. comb.
(Figs. 1.AC , 3.AU-AV , 4.A )
Holotype: Chitinoidella elongata sp. nov. - Pop (1997), Fig. 2, photo 3 (Fig. 3.AU herein).
1969 Chitinoidella sp. - Borza, Taf. LXVIX, fig. 7.
1998b Chitinoidella elongata Pop - Pop, Pl. I, fig. 38.
2002 Chitinoidella elongata Pop - Reháková, Fig. 2.5 (Fig. 3.AV herein), 2.6-8.
2013 Chitinoidella elongata Pop - Lakova & Petrova, Pl. 1, fig. 20.
2014 Chitinoidella elongata Pop - Kowal-Kasprzyk, Fig. 3.F (Fig. 4.A herein), 3.G.
2016 Chitinoidella elongata Pop - Michalík et al., Fig. 8.C.
2017 Chitinoidella elongata - Petrova et al., Fig. 3.25-26.
2018 Chitinoidella elongata Pop - Kowal-Kasprzyk, Fig. 9.D-E.
2019 Chitinoidella elongata - Petrova et al., Fig. 5.6.
2020 Chitinoidella elongata Pop - Benzaggagh, Fig. 6.M.
Description: Oval elongated-shaped lorica, higher than wide, with a L/W ratio around 2, with maximum width located at the lower third of the lorica bowl; rectilinear to slightly convex lorica walls, converging towards the aperture; wide aperture, with a curved outwardly deflected collar of small to medium size, forming an obtuse angle (> 90°) with the lorica bowl; rounded to acute aboral pole, terminated or not by a more or less developed caudal appendage.
Comparison: Bonetilla elongata differs from Bonetilla germanica n. sp. by its elongated lorica, with a greater L/W ratio; its maximum width located at the lower third of the lorica bowl, and its rectilinear to slightly convex lorica walls converging towards the aperture.
Occurrence: This species is known in the Boneti Subzone of the central West Carpathians, western Balkans, Romania, Slovakia, Serbia, Poland, Bulgaria, and southeastern France.
Bonetilla germanica n. sp.
(Figs. 1.AD , 3.T-AD )
Holotype: Chitinoidella boneti sp. nov. - Doben (1963), Taf. 6, Abb. 2 (Fig. 3.T herein).
Derivatio nominis: The name of the species derives from the Bavarian Alps (Germany) where this species was first discovered and described by Doben (1963).
Diagnosis: Oval elongated-shaped lorica, with a L/W ratio around 1.5; wide aperture surrounded by a half-crescentic collar, outwardly deflected; acute aboral pole, with or without a caudal appendage.
1963 Chitinoidella boneti sp. nov. - Doben, Taf. 6, Abb. 1 (the two specimens), 2, 5.
1965 Tintinnopsella sp. - Furrazola-Bermúdez, Lam. 3, fig. 3.
1965 Tintinnopsella carpathica (Murgeanu & Filipescu) - Furrazola-Bermúdez, Lam. 4, fig. 2 (Fig. 3.Z herein).
1969 Chitinoidella boneti Doben - Borza, Taf. LXVII, figs. 7, 9.
1969 Chitinoidella boneti Doben - Borza, Taf. LXVIII, fig. 7.
1973 Chitinoidella boneti Doben - Furrazola-Bermúdez & Kreisel, Lam. I, figs. 1, 2 (the two specimens).
1973 Chitinoidella lubimovae sp. nov. - Furrazola-Bermúdez & Kreisel, Lam. II, figs. 2, 3 (Fig. 3.AC herein), 4.
1975 Chitinoidella cf. Chitinoidella boneti - Lugo, Lam. I, figs. 7-9; Lam. II, figs. 1-9; Lam. III, figs. 1-2.
1977 Forma di transizione tra Chitinoidella boneti Doben e Praetintinnopsella andrusovi Borza - Grandesso, Tav. I, fig. 10.
1977 Chitinoidella boneti Doben - Grandesso, Tav. I, figs. 11 (Fig. 3.V herein), 12.
1977 Chitinoidella boneti Doben - Grandesso, Tav. II, fig. 1.
1986 Chitinoidella boneti Doben - Borza & Michalík, Pl. II, figs. 15-16.
1995 Chitinoidella boneti Doben - Olóriz et al., Lam. I, fig. 3.
1995 Chit. boneti - Benzaggagh & Atrops, Fig. 4.6 (Fig. 3.W herein).
1995 Chit. aff. cristobalensis (Furrazola-Bermúdez) - Benzaggagh & Atrops, Fig. 4.9 (Fig. 3.AB herein).
1997 Chitinoidella boneti Doben - Pop, Fig. 2, photos 1 (Fig. 3.X herein), 2.
1997 Chitinoidella boneti Doben - Grün & Blau, Pl. I, figs. 1-4.
1998b Chitinoidella boneti Doben - Pop, Pl. I, figs. 34, 36, 39.
1998b Praetintinnopsella andrusovi Borza - Pop, Pl. I, fig. 42.
2000 Chitinoidella boneti Doben - Benzaggagh, Pl. 5, Fig. 6.
2000 Chitinoidella aff. cristobalensis (Furrazola-Bermúdez) - Benzaggagh, Pl. 5, fig. 11.
2002 Chitinoidella boneti Doben - Reháková, Fig. 2.2 (Fig. 3.Y herein), 2.4.
2006 Chitinoidella boneti Doben - Boughdiri et al., Fig. 2.2.
2007 Longicollaria dobeni (Borza) - Andreini et al., Pl. I.2a.
2007 Dobeniella bermudezi (Furrazola-Bermúdez) - Andreini et al., Pl. I.6a, 7a.
2007 Dobeniella cubensis (Furrazola-Bermúdez) - Andreini et al., Pl. I.8a.
2009 Chitinoidella sp. 1 - Boughdiri et al., Fig. 8.22.
2010 Chitinoidella boneti Doben - Benzaggagh et al., Fig. 8.6-7.
2012 Chitinoidella boneti Doben - Benzaggagh et al., Fig. 6.B.
2013 Chitinoidella boneti Doben - Lakova & Petrova, Pl. 5, figs. 22-23.
2013 Dobeniella cubensis (Furrazola-Bermúdez) - Lakova & Petrova, Pl. 5, fig. 32
2014 Chitinoidella boneti Doben - Kowal-Kasprzyk, Fig. 3.A, 3.C.
2015 Chitinoidella boneti Doben - Benzaggagh et al., Fig. 5.D, 5.F.
2017 Chitinoidella boneti Doben - Kietzmann, Fig. 3.22-24.
2017 Chitinoidella boneti Doben - Kietzmann, Fig. 3.c.
2018 Chitinoidella sp. indet. - Kietzmann et al., Fig. 9.K.
2018 Chitinoidella boneti Doben - Kowal-Kasprzyk, Fig. 9.A-B.
2018 Chitinoidella popi Sallouhi et al. - Kowal-Kasprzyk, Fig. 9.I, 9.J (Fig. 3.AD herein).
2018 Chitinoidella sp. indet. - Kowal-Kasprzyk, Fig. 9.K.
2019 Chitinoidella boneti - Petrova et al., Fig. 5.1, 5.3-4.
2019 Chitinoidella elongata - Petrova et al., Fig. 5.7.
2020 Chitinoidella boneti Doben - Benzaggagh, Fig. 6.I, 6.J (Fig. 3.AA herein), 6.K, 6.AB.
Description: Oval elongated-shaped lorica, higher than wide, with a L/W ratio around 1.5; regularly convex lorica walls, with maximum width at the mid-lorica bowl; wide aperture, with a half-crescentic collar, outwardly deflected and forming a right to more or less obtuse angle with the lorica bowl; acute aboral pole, with or without a caudal appendage.
Variability: This species shows variability in size of the lorica, and its shape, more or less elongated; the more or less developed collar and the shape of the aboral pole, acute, with or without a more or less developed caudal appendage.
Occurrence: This species is the most common of the subfamily Bonetinae. It is known in the Boneti Subzone of the central West Carpathians, western Balkans, Poland, Romania, Slovakia, Italy, southeastern France, southeastern Spain, Iran, northern Tunisia, northern Morocco, Cuba, southeastern Mexico, and western Argentina.
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Figure 3: Main genera and species of the subfamily Bonetinae (Chitinoidellidae) from the Boneti Subzone (early late Tithonian). A-L. Bermudeziella bermudezi (Furrazola-Bermúdez): A. Tintinnopsella bermudezi Furrazola-Bermúdez, excerpt of Furrazola-Bermúdez, 1965, Lam. 2, Fig. 6. (Holotype); B. Tintinnopsella bermudezi Furrazola-Bermúdez, excerpt of Furrazola-Bermúdez, 1965, Lam. 2, fig. 8; C. Chitinoidella cubensis (Furrazola-Bermúdez), excerpt of Borza, 1969, Taf. LXVIII, fig. 14; D. Chitinoidella cubensis (Furrazola-Bermúdez), excerpt of Grandesso, 1977, Tav. II, fig. 4; E. Chitinoidella bermudezi (Furrazola-Bermúdez), excerpt of Grandesso, 1977, Tav. II, fig. 5; F. Chitinoidella bermudezi (Furrazola-Bermúdez), excerpt of Furrazola-Bermúdez & Kreisel, 1973, Lam. I, fig. 5; G. Dobeniella bermudezi (Furrazola-Bermúdez), excerpt of Reháková, 2002, Fig. 3.9; H. Dobeniella bermudezi (Furrazola-Bermúdez), excerpt of Sallouhi et al., 2011, Pl. 1.13; I. Chitinoidella cubensis (Furrazola-Bermúdez), excerpt of Benzaggagh et al.,
2012, Fig. 6.D; J. Dobeniella bermudezi (Furrazola-Bermúdez), excerpt of Lakova & Petrova, 2013, Pl. 5, fig. 27; K. Dobeniella cf. pinaraensis (Furrazola-Bermúdez & Kreisel), excerpt of Kietzmann, 2017, Fig. 5.14; L. Dobeniella bermudezi, excerpt of Petrova et al.,
2017, Fig. 4.7. M-S. Bermudeziella cubensis (Furrazola-Bermúdez): M. Tintinnopsella cubensis Furrazola-Bermúdez, excerpt of Furrazola-Bermúdez, 1965, Lam. 2, fig. 1 (Holotype); N. Chitinoidella cubensis (Furrazola-Bermúdez), excerpt of Borza, 1969, Taf. LXVIII, fig. 16; O. Chitinoidella cf. cubensis? (Furrazola-Bermúdez), excerpt of Cecca et al.,
1989, Pl. 6, fig. 3; P. Chitinoidella cubensis, excerpt of Benzaggagh & Atrops, 1995, Fig. 4.8; Q. Chitinoidella cubensis (Furrazola), excerpt of Furrazola-Bermúdez & Kreisel, 1973, Lam. 1, Fig. 6; R. Dobeniella bermudezi (Furrazola-Bermúdez), excerpt of Lakova & Petrova, 2013, Pl. 1, fig. 25, recrystallised wall; S. Dobeniella cubensis (Furrazola-Bermúdez), excerpt of Lakova & Petrova, 2013, Pl. 5, fig. 29. T-AD. Bonetilla germanica n. sp.: T. Chitinoidella boneti Doben, excerpt of Doben, 1963, Taf. 6, Abb. 2 (Holotype); U. Chitinoidella boneti Doben, excerpt of Furrazola-Bermúdez & Kreisel, 1973, Lam. I, fig. 1; V. Chitinoidella boneti Doben, excerpt of Grandesso, 1977, Tav. I, fig. 11; W. Chitinoidella boneti, excerpt of Benzaggagh & Atrops, 1995, Fig. 4.6; X. Chitinoidella boneti Doben, excerpt of Pop, 1997, Fig. 2, photo 2; Y. Chitinoidella boneti Doben, excerpt of Reháková, 2002, Fig. 2.2; Z. Tintinnopsella carpathica (Murgeanu & Filipescu), excerpt of Furrazola-Bermúdez, 1965, Lam. 4, fig. 2; AA. Chitinoidella boneti Doben, excerpt of Benzaggagh, 2020, Fig. 8.J; AB. Chitinoidella aff. cristobalensis (Furrazola-Bermúdez), excerpt of Benzaggagh & Atrops, 1995, Fig. 4.9; AC. Chitinoidella lubimovae Furrazola-Bermúdez & Kreisel, excerpt of Furrazola-Bermúdez & Kreisel, 1973, Lam. I, fig. 3, see comment in the text; AD. Chitinoidella popi Sallouhi et al., excerpt of Kowal-Kasprzyk, 2018, Fig. 9.J. AE-AM. Bonetilla carthagensis (Sallouhi et al.): AE. Chitinoidella carthagensis Sallouhi et al., excerpt of Sallouhi et al., 2011, Pl. 1.28 (Holotype); AF. Chitinoidella carthagensis Sallouhi et al., excerpt of Sallouhi et al., 2011, Pl. 1.29 (Paratype); AG. Chitinoidella boneti Doben, excerpt of Borza, 1969, Taf. LXVII, Fig. 6; AH. Chitinoidella cf. Ch. boneti, excerpt of Lugo, 1975, Lam. III, fig. 5; AI. Chitinoidella sp. 1, excerpt of Grün & Blau, 1997, Pl. I, fig. 7; AJ. Chitinoidella carthagensis Sallouhi et al., excerpt of Kowal-Kasprzyk, 2014, Fig. 3.D; AK. Chitinoidella boneti Doben, excerpt of Benzaggagh et al.,
2015, Fig. 5.E; AL. Chitinoidella hegarati Sallouhi et al., excerpt of Kietzmann, 2017, Fig. 4.12. AM. Chitinoidella boneti Doben, excerpt of Borza, 1969, Taf. LXVII, fig. 4; AN-AT. Bonetilla curva n. sp.: AN. Chitinoidella boneti Doben, excerpt of Borza, 1969, Taf. LXVII, fig. 12; AO. Chitinoidella boneti Doben, excerpt of Cecca et al.,
1989, Pl. 6, fig. 2; AP. Chitinoidella boneti Doben, excerpt of Grün & Blau, 2006, Pl. I, fig. 5; AQ. Chitinoidella boneti Doben, excerpt of Reháková, 2002, Fig. 2.1; AR. Chitinoidella boneti Doben, excerpt of Petrova et al.,
2012, Fig. 4.15; AS. Chitinoidella boneti Doben, excerpt of Lakova & Petrova, 2013, Pl. 5, fig. 23; AT. Chitinoidella boneti Doben, excerpt of Petrova et al.,
2017, Fig. 4.8; AU-AV. Bonetilla elongata (Pop): AU. Chitinoidella elongata Pop, excerpt of Pop, 1997, Fig. 2, photo 3 (Holotype); AV. Chitinoidella elongata Pop, excerpt of Reháková, 2002, Fig. 2.5. |
Bonetilla lehegarati n. sp.
(Figs. 1.AE , 4.B-G )
Remarks on Chitinoidella hegarati Sallouhi et al., 2011: The holotype (Pl. 1.30) and the paratype (Pl. 1.31) of Chitinoidella hegarati Sallouhi et al., 2011, both show an elongated lorica, with an acute aboral pole; they are very close to the holotype (Pl. 1.28) of Chitinoidella carthagensis of the same authors.
I designate their specimen, illustrated in Pl. 1.25, which has a rounded lorica and a rounded aboral pole, as the type of Chitinoidella lehegarati n. sp.
Holotype: Chitinoidella hegarati sp. nov. - Sallouhi et al. (2011), Pl. 1.25 (Fig. 4.D herein).
Derivatio nominis: This species is dedicated to the late Dr Gérard Le Hégarat, Lyon, who introduced me to the study of the calpionellids, for his key contribution to the study of Berriasian calpionellids and ammonites from the Vocontian Through in southeastern France.
Diagnosis: Rounded to slightly oval-shaped lorica, almost as high as wide, or slightly higher than wide, showing an annular constriction below the collar, and a short outwardly deflected collar; wide rounded to slightly acute aboral pole, without caudal appendage.
1998b Chitinoidella boneti Doben - Pop, Pl. 1.35 (Fig. 4.D herein).
1995 Chit. bermudezi - Benzaggagh & Atrops, Fig. 4.7 (Fig. 4.C herein).
2000 Chitinoidella bermudezi (Furrazola-Bermúdez) - Benzaggagh, Pl. 5, fig. 7.
2007 Borziella slovenica (Borza) - Andreini et al., Pl. 1.1a.
2007 Chitinoidella boneti Doben - Andreini et al., Pl. 1.3a (Fig. 4.E herein), 4a.
2010 Chitinoidella cubensis (Furrazola-Bermúdez) - Benzaggagh et al., Fig. 8.10 (Fig. 4.F herein).
2012 Borziella slovenica (Borza) - Petrova et al., Fig. 4.12.
2014 Chitinoidella hegarati Sallouhi, Boughdiri & Cordey - Kowal-Kasprzyk, Fig. 9.I-J.
2017 Borziella slovenica - Petrova et al., Fig. 3.4-5.
2017 Chitinoidella hegarati Sallouhi, Boughdiri & Cordey - Petrova et al., Fig. 4.5-6.
2018 Chitinoidella hegarati Sallouhi, Boughdiri & Cordey - Kowal-Kasprzyk, Fig. 9.G, 9.H (Fig. 4.G herein).
2019 Chitinoidella hegarati - Petrova et al., Fig. 5.34.
Description: Rounded to slightly oval-shaped lorica, as high as wide or slightly higher than wide; regularly convex lorica walls, with maximum width towards the mid-lorica bowl; wide aperture surrounded by a short half-crescentic collar, outwardly deflected and forming a right to slightly obtuse angle with the lorica bowl. Its junction with the lorica bowl is marked by an annular constriction; rounded to slightly acute aboral pole, without caudal appendage.
Comparison: Bonetilla lehegarati n. sp. is distinguished from Bonetilla germanica n. sp. by its rounded lorica, almost as high as wide; its rounded aboral pole and the presence of a constriction below the collar. It differs from Bonetilla carthagensis by having a rounded lorica, almost as high as wide, with rounded aboral pole, without caudal appendage.
Occurrence: This species is known in the Boneti Subzone of Slovakia, Romania, Serbia, Poland, Bulgaria, Italy, northern Tunisia, and northern Morocco.
Bonetilla miniboneti n. sp.
(Figs. 1.AF , 4.P-R )
Holotype: Chitinoidella boneti Doben - Borza (1969), Taf. LXVIII, Fig. 6. (Fig. 4.P herein).
Derivatio nominis: The specific name of this species derives from the similarity of its lorica with that of the paratype of of Chitinoidella boneti Doben, 1963, but differing in smaller size, rounded lorica shape, and low L/W ratio.
Diagnosis: Rounded-shaped lorica of medium size, almost as high as wide; wide aperture surrounded by a short half-crescentic collar; rounded to slightly acute aboral pole.
1969 Chitinoidella boneti Doben - Borza, Taf. LXVIII, figs. 1-5.
1977 Chitinoidella boneti Doben - Grandesso, Tav. II, fig. 2 (Fig. 4.Q herein).
2017 Chitinoidella boneti Doben - Lakova et al., Fig. 1.A (Fig. 4.R herein).
2019 Chitinoidella boneti - Petrova et al., Fig. 4.39-42.
Description: Rounded to slightly oval-shaped lorica of medium size, as high as wide, or slightly higher than wide, with a L/W ratio around 1.2; regularly convex lorica walls, with maximum width located at the mid-lorica bowl; wide aperture, surrounded by a short half-crescentic collar, forming a right to slightly obtuse angle with the lorica bowl; rounded to slightly acute aboral pole, without caudal appendage.
Comparison: Bonetilla miniboneti n. sp. is distinguished from Bonetilla germanica n. sp. by its smaller size, more rounded lorica, with a low L/W ratio and its rounded aboral pole without caudal appendage.
Occurrence: This species is known in the Boneti Subzone of Slovakia, Bulgaria and Italy.
Bonetilla popi (Sallouhi et al., 2011) n. comb.
(Figs. 1.AG , 4.S-W )
Holotype: Chitinoidella popi sp. nov.- Sallouhi et al. (2011), Pl. 1.32 (Fig. 4.S herein).
1975 Chitinoidella cf. Ch. boneti - Lugo, Lam. I, fig. 12, 13 (Fig. 4.T herein).
2009 Chitinoidella sp. 2 - Boughdiri et al., Fig. 8.23.
2010 Chitinoidella boneti Doben - Ben Abdesselam-Mahdaoui et al., Pl. II, fig. 1 (Fig. 4.U herein).
2011 Chitinoidella popi sp. nov., Sallouhi et al., Pl. 1.33.
2011 Chitinoidella cf. elongata Pop - Sallouhi et al., Pl. 1, fig. 34.
2011 Chitinoidella sp. 1 - Sallouhi et al., Pl. 1, fig. 34 (distorted specimen).
2014 Chitinoidella popi Sallouhi, Boughdiri & Cordey - Kowal-Kasprzyk, Fig. 3.K (Fig. 4.V herein), 3.L.
2015 Chitinoidella boneti Doben - Benzaggagh et al., Fig. 5.G.
2017 Chitinoidella elongata Pop - Kietzmann, Figs. 4.15, 5.5 (Fig. 4.W herein), 5.6.
2017 Chitinoidella sp. - Petrova et al., Fig. 4.1-2.
2018 Chitinoidella elongata Pop - Kietzmann et al., Fig. 3.e.
2019 Chitinoidella popi - Petrova et al., Fig. 5.21-24.
Description: Oval elongated-shaped lorica, higher than wide, with a L/W ratio > 2; regularly convex lorica walls, with maximum width towards the mid-lorica bowl; wide aperture surrounded by a short half-crescentic collar, outwardly deflected, forming a right to slightly obtuse angle with the lorica bowl; acute aboral pole, often with a more or less developed caudal appendage.
Comparison: Bonetilla popi is distinguished from Bonetilla elongata by its regular convex lorica walls, with maximum width at the mid-lorica bowl, and its shorter collar, forming a lesser obtuse angle with the lorica wall.
Occurrence: This species is known in the Boneti Subzone of Bulgaria, Poland, northern Tunisia, southeastern Mexico, and western Argentina.
Bonetilla praeboneti n. sp.
(Figs. 1.AH , 4.X-Y )
Holotype: Chitinoidella aff. boneti - Benzaggagh & Atrops (1995), Fig. 4.12 (Fig. 4.X herein).
Derivatio nominis: The species is named with reference to the primitive form of the Chitinoidella boneti group from the base of the Boneti Subzone.
Diagnosis: Rounded to slightly oval-shaped lorica, with reduced or undeveloped collar; rounded to slightly acute aboral pole.
2000. Chitinoidella aff. boneti Doben - Benzaggagh, Pl. 5, fig. 5.
2010 Chitinoidella aff. boneti Doben - Benzaggagh et al., Fig. 8.5 (Fig. 4.Y herein).
2020 Chitinoidella aff. boneti Doben - Benzaggagh, Fig. 6.H.
Diagnosis: Rounded to oval-shaped lorica, slightly higher than wide; regularly convex lorica walls, with maximum width towards the mid-lorica bowl; wide aperture with reduced or lacking collar; rounded to slightly acute aboral pole, without caudal appendage.
Comparison: Bonetilla praeboneti n. sp. differs from Bonetilla germanica n. sp. by its reduced or undeveloped collar.
Occurrence: This species is known in the base of the Boneti Subzone in several sections of northern Morocco.
Bonetilla sphaerica n. sp.
(Figs. 1.AI , 4.Z-AF )
Holotype: Chitinoidella sp. - Cecca et al.
(1989), Pl. 6, fig. 5 (Fig. 4.Z herein).
Derivatio nominis: The name of this species derives from the spherical shape of the lorica.
Diagnosis: Spherical-shaped lorica, as high as wide; wide aperture, with a short half-crescentic collar, outwardly deflected; wide and rounded aboral pole.
1998b Chitinoidella boneti Doben - Pop, Pl. I, fig. 37 (Fig. 4.AA herein).
1999 Chitinoidella boneti Doben - Lakova et al., Pl. 1, fig. 2 (Fig. 4.AB herein).
2011 Chitinoidella sp. 2 - Sallouhi et al., Pl. 1.36 (Fig. 4.AC herein).
2013 Chitinoidella hegarati Sallouhi, Boughdiri & Cordey - Lakova & Petrova, Pl. I, fig. 19.
2013 Dobeniella cf. cubensis (Furrazola-Bermúdez) - Lakova & Petrova, Pl. 1, figs. 23 (Fig. 4.AD herein), 24.
2014 Chitinoidella boneti Doben - Kowal-Kasprzyk, Fig. 3.B.
2014 Dobeniella tithonica (Borza) - Kowal-Kasprzyk, Fig. 3.N (Fig. 4.AE herein).
2014 Dobeniella cubensis (Furrazola-Bermúdez) - Kowal-Kasprzyk, Fig. 3.O.
2017 Chitinoidella sp. - Strzeboński et al., Fig. 6.T (Fig. 4.AF herein).
2019 Chitinoidella bermudezi - Petrova et al., Fig. 5.2.
Description: Spherical-shaped lorica, as high as wide, to slightly higher than wide, with a L/W ratio around 1.2; regularly convex lorica walls; wide aperture, with a short crescentic collar, outwardly deflected, and forming a right to slightly obtuse angle with the lorica bowl; wide and rounded aboral pole, without caudal appendage.
Comparison: Bonetilla sphaerica n. sp. is distinguished from all other species of the genus Bonetilla by its spherical lorica, short collar, and its wide and rounded aboral pole.
Occurrence: The species is known in the Boneti Subzone of the western Balkans, Romania, Bulgaria, Poland, southeastern France, northern Tunisia, and western Argentina.
Bonetilla svinitensis (Pop, 1998b) n. comb.
(Figs. 1.AJ , 4.AG-AK )
Holotype: Daciella svinitensis sp. nov. - Pop (1998a), Fig. 2, photo 19 (Fig. 4.AG herein).
Diagnosis: Elongated cylindrical-shaped lorica, higher than wide; wide aperture with a short half-crescent collar outwardly deflected; rounded to slightly acute aboral pole.
1998b Daciella svinitensis sp. nov. - Pop, Pl. 1, fig. 8.
2012 Chitinoidella elongata Pop. - Petrova et al., Fig. 4.16.
2014 Chitinoidella elongata Pop - Kowal-Kasprzyk, Fig. 3.H.
2014 Popiella oblongata Reháková - Kowal-Kasprzyk, Fig. 3.T.
2017 Chitinoidella elongata Pop - Kietzmann, Fig. 4.14 (Fig. 4.AI herein).
2017 Popiella oblongata - Petrova et al., Fig. 3.27-28.
2017 Daciella sp. - Petrova et al., Fig. 3.29, 3.30 (Fig. 4.AJ herein).
2017 Longicollaria insueta Pop - Petrova et al., Fig. 4.17 (Fig. 4.AYH herein).
2018 Chitinoidella sp. indet. - Kowal-Kasprzyk, Fig. 9F.
2018 Popiella oblongata Reháková - Kowal-Kasprzyk, Fig. 9.S (Fig. 4.AK herein).
2019 Daciella sp. - Petrova et al., Fig. 4.31-32.
2019 Longicollaria insueta - Petrova et al., Fig. 4.34.
2019 Popiella oblongata - Petrova et al., Fig. 4.37-38.
Description: Elongated cylindrical-shaped lorica, higher than wide, with parallel to sub-parallel walls; wide aperture, with a short half-crescentic collar, outwards deflected, forming a right to slightly obtuse angle with the lorica bowl; rounded to slightly acute aboral pole, without caudal appendage.
Comparison: Bonetilla svinitensis differs from Bonetilla elongata and Bonetilla popi by its cylindrical lorica, with parallel to sub-parallel walls, and a shorter collar; it differs from Bonetilla boneti by its rectilinear lorica walls and its shorter collar; and from Popiella oblongata by its larger size and the presence of a short collar.
Occurrence: This species is known in the Boneti Subzone of Romania, Slovakia, Poland, Bulgaria, and western Argentina.
Furrazolaia n. gen.
Holotype: Calpionella cristobalensis sp. nov. - Furrazola-Bermúdez (1965), Lam. 3, fig. 5 (Fig. 4.AL herein).
Remark: Specimens of this genus were often attributed to the genera: Dobeniella Pop, Longicollaria Pop, Almajella Pop, and Cubanella Pop.
Derivatio nominis: This genus is dedicated to Dr. Furrazola-Bermúdez.
Diagnosis: Conical to slightly elongated oval-shaped lorica; wide aperture, with a long rectilinear to slightly arched collar, often forming a shoulder with the lorica bowl; acute aboral pole, with or without a caudal appendage.
Furrazolaia cristobalensis (Furrazola-Bermúdez, 1965) n. comb.
(Figs. 1.AK , 4.AL-AR )
Holotype: Calpionella cristobalensis sp. nov. - Furrazola-Bermúdez (1965), Lam. 3, fig. 5 (Fig. 4.AL herein).
1965 Calpionella cristobalensis sp. nov. - Furrazola-Bermúdez, Lam. 3, figs. 5-8; Lam. 5, fig. 3.
1973 Chitinoidella cristobalensis (Furrazola) - Furrazola-Bermúdez & Kreisel, Lam. I, fig. 8.
1973 Chitinoidella pinaraensis sp. nov. - Furrazola-Bermúdez & Kreisel, Lam. II, fig. 5 (Fig. 4.AO herein).
1977 Chitinoidella cristobalensis (Furrazola-Bermúdez) - Grandesso, Tav. II, Fig. 6. (Fig. 4.AM herein).
1995 Chit. cristobalensis - Benzaggagh & Atrops, Fig. 4.10 (Fig. 4.AN# herein), 4.11.
1997 Chitinoidella boneti Doben - Ivanova, Pl. 1, fig. 17.
2000 Chitinoidella cristobalensis (Furrazola-Bermúdez) - Benzaggagh, Pl. 5, figs. 9-10.
2002 Dobeniella cubensis (Furrazola-Bermúdez) - Reháková, Fig. 3.5.
2010 Chitinoidella cristobalensis (Furrazola-Bermúdez) - Benzaggagh et al., Fig. 8.8.
2010 Longicollaria dobeni (Borza) - Pszczółkowski & Myczyński, Fig. 12.10 (Fig. 4.AP herein).
2010 ?Dobeniella pinarensis (Furrazola-Bermúdez & Kreisel) - Pszczółkowski & Myczyński, Fig. 12.12.
2011 Dobeniella cubensis (Furrazola-Bermúdez) - Sallouhi et al., Pl. 1.10.
2011 Cubanella cristobalensis (Furrazola-Bermúdez) - Sallouhi et al., Pl. 1.23 (Fig. 4.AQ herein).
2013 Almajella cristobalensis (Furrazola-Bermúdez) - Lakova & Petrova, Pl. 1, fig. 22.
2017 Dobeniella cf. pinaraensis (Furrazola-Bermúdez & Kreisel) - Kietzmann, Fig. 5.15 (Fig. 4.AR herein).
2017 Dobeniella tithonica - Petrova et al., Fig. 3.18.
2019 Dobeniella lubimovae - Petrova et al., Fig. 5.12.
2020 Chitinoidella cristobalensis (Furrazola-Bermúdez) - Benzaggagh, Fig. 6.P.
2020 Chitinoidella bermudezi (Furrazola-Bermúdez) - Benzaggagh, Fig. 6.Z.
Description: Conical to elongated oval-shaped lorica, as high as wide, or slightly higher than wide, with maximum width located immediately below the collar; convex lorica walls, strongly converging towards the aboral pole; wide aperture surrounded by a rectilinear to slightly arched collar, equal to or slightly lesser than one-third of the total lorica length, showing in axial section two parallel or divergent branches of the same thickness as that of the lorica wall, forming a more or less developed shoulder; acute aboral pole, often with a caudal appendage.
Variability: This species shows variability in more or less developed shoulder at the junction between the collar and the lorica bowl; length and shape of the collar; and presence or not of a caudal appendage.
Comparison: Furrazolaia cristobalensis differs from Bermudeziella cubensis and Bermudeziella bermudezi in having a conical lorica, with maximum width located immediately below the collar; a longer collar, with the same thickness as that of the lorica wall; the presence of a shoulder at the junction between the collar and the lorica bowl, and a more acute aboral pole, often with a caudal appendage. It differs from Dobenilla longicolomi n. sp. by its larger size; its collar of the same thickness as that of the lorica wall; and its stratigraphic range.
Occurrence: This species is known in the Boneti Subzone of the central West Carpathians, western Balkans, Bulgaria, Italy, Northeast Spain, Iran, northern Tunisia, northern Morocco, Cuba, and western Argentina.
Furrazolaia insueta (Řehánek, 1986) n. comb.
(Figs. 1.AL , 4.AS-AU )
Holotype: Chitinoidella insueta sp. nov. - Řehánek (1986), Pl. 1, fig. 2 (Fig. 4.AS herein).
Amended diagnosis: Cylindrical elongated-shaped lorica, with a long collar, equal to or greater than one-half of the total lorica length; wide and rounded aboral pole.
1969 Chitinoidella sp. - Borza, Taf. LXVIX, fig. 7.
1986 Chitinoidella insueta sp. nov. - Řehánek, Pl. 1, figs. 3-4.
2002 Longicollaria insueta (Řehánek) - Reháková, Fig. 4.7-9.
2014 Longicollaria insueta (Řehánek) - Kowal-Kasprzyk, Fig. 3.S.
2016 Cylindrella insueta (Řehánek, 1986) - Granier et al., Fig. 2.A
2017 Dobeniella pinarensis - Petrova et al., Fig. 4.14.
2017 Longicollaria insueta - Petrova et al., Fig. 4.16 (Fig. 4.AU herein).
2017 Dobeniella cf. pinaraensis (Furrazola-Bermúdez & Kreisel) - Kietzmann, Fig. 4.20 (Fig. 4.AV herein).
2018 Longicollaria cf. insueta (Řehánek) - Kowal-Kasprzyk, Fig. 9.R.
2018 Dobeniella cf. pinaraensis (Furrazola-Bermúdez & Kreisel) - Kietzmann et al., Fig. 3.b.
2019 Dobeniella sp. - Petrova et al., Fig. 4.33.
2019 Longicollaria insueta- Petrova et al., Fig. 4.36.
2020 Chitinoidella insueta Řehánek - Benzaggagh, Fig. 6.T (Fig. 4.AT herein).
Description: Cylindrical elongated-shaped lorica, with a L/W ratio > 2, often wider at its posterior side; rectilinear to divergent collar, between one-half to two-thirds of the total lorica length, forming a constriction or a slight shoulder with the lorica bowl; wide and rounded aboral pole, without caudal appendage.
Comparison: This species differs from all other species of chitinoidellids by its longer and rectilinear collar forming a short shoulder with the lorica bowl.
Occurrence: This species is known in the Boneti Subzone of the central West Carpathians, Slovakia, Poland, Bulgaria, southeastern France, northern Morocco, and western Argentina.
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Figure 4: Main genera and species of the subfamily Bonetinae (Chitinoidellidae) from the Boneti Subzone (early late Tithonian). A. Bonetilla elongata (Pop), excerpt of Kowal-Kasprzyk, 2014, Fig. 3.F; B-G. Bonetilla lehegarati n. sp.: B. Chitinoidella hegarati Sallouhi et al., excerpt of Sallouhi et al., 2011, Pl. 1.25 (Holotype); C. Chitinoidella bermudezi, excerpt of Benzaggagh & Atrops, 1995, Fig. 4.7; D. Chitinoidella boneti Doben, excerpt of Pop, 1998b, Pl. 1.35; E. Borziella slovenica (Borza), excerpt of Andreini et al., 2007, Pl. 1.1a; F. Chitinoidella cubensis (Furrazola-Bermúdez), excerpt of Benzaggagh et al., 2010, Fig. 8.10; G. Chitinoidella hegarati Sallouhi et al., excerpt of Kowal-Kasprzyk, 2018, Fig. 9.H. H-O. Bonetilla boneti (Doben, 1963): H. Chitinoidella boneti Doben, excerpt of Doben, 1963, Taf. 6, Abb. 3 (Holotype); I. Chitinoidella boneti Doben, excerpt of Borza, 1969, Taf. LXVIII, fig. 10; J. Chitinoidella boneti Doben, excerpt of Sallouhi et al., 2011, Pl. 1.24; K. Chitinoidella boneti Doben, excerpt of Borza, 1969, Taf. LXVII, fig. 8; L. Longicollaria insueta (Řehánek), excerpt of Pszczółkowski & Myczyński, 2010, Fig. 12.9; M. Chitinoidella bermudezi (Furrazola-Bermúdez), excerpt of Cecca et al.,
2012, Fig. 3A; N. Chitinoidella boneti Doben, excerpt of Lakova & Petrova, 2013, Pl. 1, fig. 18; O. Chitinoidella boneti Doben, excerpt of Altiner & Özkan, 1991, Pl. 3, fig. 2. P-R. Bonetilla miniboneti n. sp.: P. Chitinoidella boneti Doben, excerpt of Borza, 1969, Taf. LXVIII, Fig. 6. (Holotype); Q. Chitinoidella boneti Doben, excerpt of Grandesso, 1977, Tav. II, fig. 2; R. Chitinoidella boneti Doben, excerpt of Lakova et al.,
2017, Fig. 1.A. S-W. Bonetilla popi (Sallouhi et al.): S. Chitinoidella popi Sallouhi et al., excerpt of Sallouhi et al., 2011, Pl. 1.32 (Holotype); T. Chitinoidella cf. Ch. boneti, excerpt of Lugo, 1975, Lam. I, fig. 2; U. Chitinoidella boneti Doben, excerpt of Ben Abdesselam-Mahdaoui et al., 2010, Pl. II, fig. 1; V. Chitinoidella popi Sallouhi et al., excerpt of Kowal-Kasprzyk, 2014, Fig. 3.K; W. Chitinoidella elongata Pop, excerpt of Kietzmann, 2017, Fig. 5.5. X-Y. Bonetilla praeboneti n. sp.: X. Chitinoidella aff. boneti, excerpt of Benzaggagh & Atrops, 1995, Fig. 4.12 (Holotype); Y. Chitinoidella aff. boneti Doben, excerpt of Benzaggagh et al., 2010, Fig. 8.5. Z-AF. Bonetilla sphaerica n. sp.: Z. Chitinoidella sp., excerpt of Cecca et al.,
1989, Pl. 6, fig. 5 (Holotype); AA. Chitinoidella boneti Doben, excerpt of Pop, 1998b, Pl. I, fig. 37; AB. Chitinoidella boneti Doben, excerpt of Lakova et al.,
1999, Pl. 1, fig. 2; AC. Chitinoidella sp. , excerpt of Sallouhi et al., 2011, Pl. 1.36; AD. Dobeniella cf. cubensis (Furrazola-Bermúdez), excerpt of Lakova & Petrova, 2013, Pl. 1, fig. 23; AE. Dobeniella cf. cubensis, excerpt of Kowal-Kasprzyk, 2014, Fig. 3.N; AF. Chitinoidella sp., excerpt of Strzeboński et al.,
2017, Fig. 6.T. AG-AK. Bonetilla svinitensis (Pop): AG. Daciella svinitensis Pop, excerpt of Pop, 1998a, Fig. 2, photo 19 (Holotype); AH. Longicollaria insueta Pop, excerpt of Petrova et al.,
2017, Fig. 4.17; AI. Chitinoidella elongata Pop, excerpt of Kietzmann, 2017, Fig. 4.14; AJ. Daciella sp., excerpt of Petrova et al.,
2017, Fig. 3.30; AK. Popiella oblongata Reháková, excerpt of Kowal-Kasprzyk, 2018, Fig. 9.S. AL-AR. Furrazolaia cristobalensis (Furrazola-Bermúdez): AL. Calpionella cristobalensis Furrazola-Bermúdez, excerpt of Furrazola-Bermúdez, 1965, Lam. 3, fig. 5 (Holotype); AM. Chitinoidella cristobalensis (Furrazola-Bermúdez), excerpt of Grandesso, 1977, Tav. II, Fig. 6; AN. Chitinoidella cristobalensis, excerpt of Benzaggagh & Atrops, 1995, Fig. 4.10; AO. Chitinoidella pinaraensis Furrazola-Bermúdez & Kreisel, excerpt of Furrazola-Bermúdez & Kreisel, 1973, Lam. II, fig. 5, see comment in the text; AP. Longicollaria dobeni (Borza), excerpt of Pszczółkowski & Myczyński, 2010, Fig. 12.10; AQ. Cubanella cristobalensis (Furrazola-Bermúdez), excerpt of Sallouhi et al., 2011, Pl. 1.23; AR. Dobeniella cf. pinaraensis (Furrazola-Bermúdez & Kreisel), excerpt of Kietzmann, 2017, Fig. 5.15. AS-AU. Furrazolaia insueta (Řehánek): AS. Chitinoidella insueta Řehánek, excerpt of Řehánek, 1985, Pl. 1, fig. 2 (Holotype); AT. Chitinoidella insueta Řehánek, excerpt of Benzaggagh, 2020, Fig. 6.T ; AU. Longicollaria insueta, excerpt of Petrova et al.,
2017, Fig. 4.16; AV. Dobeniella cf. pinaraensis (Furrazola-Bermúdez & Kreisel), excerpt of Kietzmann, 2017, Fig. 4.20. |
According to Énay & Geyssant (1975), chitinoidellids appear at the top of the Fallauxi ammonite Zone and disappear in the upper part of the Microcanthum Zone. Borza (1969, Abb. 4), Grandesso (1977, Fig. 1) and Borza (1984, Tab. 1) reported the exclusive occurrence of small-sized chitinoidellids in the lower part of the Chitinoidella Zone and the exclusive occurrence of larger-sized chitinoidellids in the upper part of this same zone. On the basis of such a distribution, Borza (1984, p. 544) and Borza & Michalík (1986, p. 137) divided the Chitinoidella Zone into two subzones, Dobeni and Boneti. These two subzones were correlated with the ammonite zones (Benzaggagh & Atrops, 1995; Benzaggagh et al., 2010). The first one corresponds to the Ponti Zone; the second to the Microcanthum p.p. Zone.
The stratigraphic distribution of the chitinoidellids in two groups and the two defined subzones were later demonstrated in several published works and from several remote localities of the Tethysian Realm. Amongst others: Reháková (2002, Figs. 2-4), whose illustrated specimens of small-sized chitinoidellids all are from the Dobeni Subzone, except Popiella oblongata? and all larger specimens belonging to the Boneti Subzone; also Petrova et al. (2012), whose small-sized chitinoidellids (op. cit., Figs. 4.1-11) are from the Dobeni Subzone and those of larger size (op. cit., Figs. 4.12-17) are from the Remanei Subzone; and Lakova & Petrova (2013), whose small specimens (op. cit., Pl. 1.1-16, Pl. 5.1-20) are from the Dobeni Subzone and larger specimens (op. cit., Pl. 1.17-26, Pl. 5.21-32) are from the Boneti Subzone to Praetintinnopsella Zone. See also Kowal-Kasprzyk (2018) and Petrova et al. (2019).
Note that, in several sections of the External Rif, Morocco (see Benzaggagh, 2000), small chitinoidellids disappear at the top of the Dobeni Subzone, with the exception probably of Borziella slovenica, which gave rise at the base of the Boneti Subzone to a single species with a larger lorica, similar to that of Chitinoidella boneti, but without collar, or with a very atypical collar, qualified as Chitinoidella aff. boneti (Benzaggagh & Atrops, 1995). This species gave rise, gradually, from the lower part of the Boneti Subzone to a different larger species of the boneti group.
Based on the resemblance of the lorica shape between some species of the Dobeni and Boneti subzones, several authors (Pop, 1997, 1998a, 1998b; Reháková, 2002; Sallouhi et al., 2011) considered that the evolution of the chitinoidellids around the Dobeni-Boneti subzonal boundary interval had gradually transitioned from small species of the dobeni group to larger species of the boneti group.
This phylogenetic interpretation is difficult to reconcile because:
1) chitinoidellid species of the Boneti Subzone appear gradually and after the disappearance of the small species of the Dobeni Subzone (Fig. 5 );
2) the base of the Boneti Subzone is marked by the presence of a single species of chitinoidellids (Bonetilla praeboneti n. sp.); and
3) intermediate forms between the two chitinoidellid groups are notably absent, both at the top of the Dobeni and at the base of the Boneti subzones.
It seems very likely that these similarities between certain species of the two chitinoidellid groups, both in the collar and lorica shapes, represent rather morphological convergence than a gradual anagenetic evolution.
Note that the morphological convergences between species of calpionellids (sensu lato) widely separated in time and without any phylogenetic relationship are very common and recurrent at numerous stratigraphic levels.
Among several examples, we can report the strong likeness between the lorica of:
Actually, these morphological resemblances between loricae of some calpionellid species widely separated in time and without any phylogenetic relationship show that, because of their very simple bell-like lorica, calpionellids had often replicated during their evolution at several stratigraphic levels, lorica and collar shapes of some already extinct species.
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Figure 5: Stratigraphic distribution and evolution of the family Chitinoidellidae (subfamilies Dobeninae and Bonetinae) around the early/late Tithonian boundary interval. For the distribution and the evolution of chitinoidellids and calpionellids at the base of the Crassicollaria Zone, see Benzaggagh & Atrops (1995), Benzaggagh et al. (2010) and Benzaggagh (2020). |
As demonstrated above, specimens of the family Chitinoidellidae split into two groups: specimens of small-sized lorica of the dobeni group and specimens of larger-sized lorica of the boneti group. These two groups, which are separated in time, correspond to two subfamilies: Dobeninae and Bonetinae, respectively. Based on the lorica and the collar shapes, five genera (including one new genus) are characterised within the subfamily Dobeninae, which are: Borziella Pop, Carpathella Pop, Daciella Pop, Dobenilla n. gen., and Popiella Reháková respectively, and three new genera within the subfamily Bonetinae: Bermudeziella n. gen., Bonetilla n. gen., and Furrazolaia n. gen. respectively. A total of twenty-four species (seven of them are new) are described and illustrated by several specimens from previous work (see Figs. 2 - 3 - 4 ), namely: Dobenilla dobeni (Borza), with two new varieties (Dobenilla dobeni var. dobeni n. var. and Dobenilla dobeni var. longidobeni n. var.), Dobenilla colomi (Borza), with two new varieties (Dobenilla colomi var. colomi n. var. and Dobenilla colomi var. longicolomi n. var.), Borziella slovenica (Borza), Borziella tithonica (Borza), Carpathella longirumanica n. sp., Carpathella rumanica Pop, Daciella banatica Pop, Daciella danubica Pop, and Popiella oblongata Reháková, of the subfamily Dobeninae respectively; and Bermudeziella bermudezi (Furrazola-Bermúdez), Bermudeziella cubensis (Furrazola-Bermúdez), Bonetilla boneti (Doben), Bonetilla carthagensis (Sallouhi et al.), Bonetilla curva n. sp., Bonetilla elongata (Pop), Bonetilla germanica n. sp., Bonetilla lehegarati n. sp., Bonetilla miniboneti n. sp., Bonetilla popi (Sallouhi et al.), Bonetilla praeboneti n. sp., Bonetilla sphaerica n. sp., Bonetilla svinitensis (Pop), Furrazolaia cristobalensis (Furrazola-Bermúdez), and Furrazolaia insueta (Řehánek) of the subfamily Bonetinae respectively.
The invalidity of certain chitinoidellid taxa is discussed and a new evolutionary pattern of the family Chitinoidellidae around the lower/upper Tithonian boundary interval is proposed.
This evolutionary pattern is characterised by the disappearance of the small chitinoidellids of the dobeni group at the top of the Dobeni Subzone, followed by a renewal of the chitinoidellid microfauna at the lower part of the Boneti Subzone from a single species (Bonetilla praeboneti), which gave rise to the different larger species of the boneti group, from the middle and the upper parts of the Boneti Subzone.
The resemblance in the lorica and collar shapes of some chitinoidellid species of the Dobeni Subzone with those of the Boneti Subzone, most likely represents a morphological convergence, rather than a gradual transition from small species of the dobeni group to larger species of the boneti group.
The present study was conducted within the framework of a scientific project "Appui à la Recherche, 2013" of Moulay Ismail University of Meknes, Morocco. The author wishes to express his gratitude to Dr Bruno Granier, Editor-in-Chief of "Carnets Geol.", for his valuable scientific remarks on the first version of the manuscript. He thanks the reviewers, Dr Iskra Lakova, Dr Andrzej Pszczółkowski and Dr Daniela Reháková, who accepted to report on this work and whose careful analyses of the original manuscript, numerous constructive remarks and suggestions greatly contributed to make its scientific quality better. The author wishes to express his most sincere gratitude to Dr Bruno Ferré for his valuable advice, highlighting suggestions and linguistic skills. Finally, the author is grateful to Dr Christopher Toland who carefully revised the English text of the manuscript.
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