Class Conodonta Pander, 1856
Order Ozarkodinida Dzik, 1976
Superfamily Gondolelloidea Lindström, 1970
Family Gladigondolellidae Ishida and Hirsch, 2011
Genus Gladigondolella Müller, 1962
Type species Polygnathus tethydis Huckriede, 1958
Figs. 4.19, 6.1–6.11
3.1–3.7, 3.9–3.28. Paragondolella bulgarica Budurov and Stefanov. KTM3 and KTM4 sections: 3.1–3.3. KTM3-9861; 3.5–3.6. KTM3-9874; 3.10. KTM3-9883; 3.4, 3.9, 3.11, 3.13, 3.17. KTM3-9891; 3.7. KTM3-0144; 3.12, 3.14. KTM4-0150; 3.16, 3.22. KTM4-0154; 3.15, 3.18, 3.19, 3.23. KTM4-0158; 3.20, 3.21. KTM4-0160; 3.23. KTM4-0171; 3.24. KTM4-0172; 3.25. KTM4-0175; 3.26. KTM4-0177; 3.27. KTM4-0180; 3.28. KTM4-0191; 3.29. KTM4-0198; 3.8. Nicoraella kockeli (Tatge), KTM4/3-0221.
Paragondolella hirschii n. sp. Kılıç and Budurov. KTM3 and KTM4 sections: 4.1, 4.2. KTM4-0144 (holotype); 4.3, 4.6. kept in Bulgarian Academy of Sciences; 4.4, 4.5. KTM3-9891; 4.7, 4.8. KTM4-0150; 4.9, 4.10. KTM4-1054; 4.11–4.18. Paragondolella praecornuta n. sp. Kılıç, Budurov, Petrunova, Mirăuţă; 4.11, 4.15 are from Gedik (1975; pl. 1 figs. 10, 11); 4.12, 4.13, 4.17 kept in Bulgarian Academy of Sciences; 4.14, 4.16. KTM4-0154 (holotype); 4.18. KTM4-0180; 4.19. Gladigondolella sp. A, KTM4-0144; 4.20–4.21, 4.24–4.25. Cornudina oezdemirae Gedik; 4.20. KTM3-9801; 4.21. KTM3-9811; 4.24. KTM3-9824; 4.25. KTM3-9829; 4.22. Neostrachanognathus tahoensis Koike, KTM3-9841; 4.23. Cornudina cf. latidentata Kozur and Mostler, KTM3-9832. Figs. 5.20–5.25 were also used in Kılıç et al. (2018b).
5.1–5.2, 5.5–5.6. Chiosella timorensis (Nogami). KTM3 and KTM5 sections: 5.1, 5.2. KTM3-9832; 5.5. KTM3-9841; 5.6. KTM3-9857; 5.7. Neogondolella sp., KTM5-98931; 5.3–5.4, 5.8–5.9. Paragondolella hanbulogi (Budurov and Stefanov), KTM5-98893; 5.10–5.15. Paragondolella aegaea (Bender); 5.10, 5.13. KTM3-9832; 5.12. KTM3-9841; 5.11. KTM3-9861; 5.14. KTM3-9879; 5.15. KTM3-9849; 5.16, 5.19–5.21. P. ebruae n. sp.; 5.16, 5.19. KTM3-9861 (holotype); 5.20. KTM3-9878; 5.21. KTM3-9883; 5.17, 5.18. Paragondolella sp. A, KTM3-9891.
Description: Gondolelloid platform conodont element with a basal blade and a keel, extending over the entire element. The amygdaloid basal cavity is not terminal, being a little farther away from the posterior end, with a posterior keel developed. The platform varies in width from a mid-lateral rib to a wide, flat platform, and a free blade in some species.
Gladigondolella tethydis radiated in the Tethys area from Anisian to Julian. In the course of the Middle Triassic G. malayensis evolved from G. tethydis through expansion of the platform.
Stratigraphic range: Gladigondolella ranges from the end of the Early Triassic to the late early Carnian; the genus consists of the Spathian G. carinata, Anisian–Julian G. tethydis and late Ladinian–early Carnian G. malayensis. The present unique specimen can only be determined as G. sp. A.
Gladigondolella tethydis (Huckriede, 1958)
Gladigondolella multielement apparatus (lacking P1 element). 6.1, 6.12. Ozarkodinid P2 element; 6.2. Prioniodinid S2 element; 6.3, 6.4. Roundyid S0 element; 6.5. Anastrophognathid S0 element; 6.7, 6.8. Lonchodinid M element; 6.6, 6.9–6.11. Prioniodinid S elements; 6.1, 6.5. KTM4-0150; 6.2, 6.6, 6.9. KTM4-0158; 6.3, 6.4. KTM4-0172; 6.7. KTM4-0191; 6.8. KTM4/2-0206; 6.10. KTM4/2-0209; 6.11. KTM5-98901; 6.12. KTM5-98929.
1958 Polygnathus tethydis n. sp.; Huckriede, pl. 12, figs. 38a, 38b; pl. 13, figs. 2–5
1960 Polygnathus tethydis; Spasov and Ganev, p. 85, pl. 1, figs. 26, 27; pl. 2, fig. 16.
1960 Polygnathus tethydis; Budurov, p. 117, pl. 2, figs. 28–30, 33.
1965 Polygnathus tethydis; Mosher and Clark, p. 563, pl. 66, fig. 13.
1966 Gladigondolella tethydis; Ishii and Nogami, pl. 1, figs. 1, 2.
1968 Gladigondolella tethydis; Nogami, p. 123, pl. 9, figs. 1–10; pl. 11, figs. 5, 6.
1968 Gladigondolella tethydis; Mosher, p. 937, pl. 116, figs. 1, 2, 5, 8.
1969 Gladigondolella tethydis; Hirsch, Pl. 1, figs. 1–3.
1970 Gladigondolella tethydis; Bender, p. 505, pl. 2, figs. 2–6.
1973 Gladigondolella tethydis; Budurov, p. 802, pl. 1, figs. 4–7.
1975 Gladigondolella tethydis; Gedik, p. 120, pl. 3, figs. 15–17.
1984 Gladigondolella tethydis; Farabegoli et al., fig. 4, part e.
1992 Gladigondolella tethydis; Kozur and Mostler, pl. 2, figs. 19, 20.
2001 Gladigondolella tethydis; Orchard et al., pl. 1, figs. 20, 21.
2007a Gladigondolella tethydis; Orchard et al., fig. 6, part 35.
2007b Gladigondolella tethydis; Orchard et al., fig. 5, parts 29, 30.
2015 Gladigondolella tethydis; Lehrmann et al., fig. 6, parts 26, 27.
2016 Gladigondolella tethydis; Karádi and Dulai, fig. 2.
2019a Gladigondolella tethydis; Chen et al., fig. 5, parts 5a–5c.
Remarks: In the multielement apparatus of Gladigondolella tethydis, platform and non-platform elements occur usually in a 1 : 4 ratio. But in the Kocaeli Triassic, this ratio does not apply; only ramiform elements were recovered. Among the ramiform elements found, Kılıç (2004) regarded Anastrophognathus sagittalis as S0 element in addition to Roundya (Hibbardella) magnidentata. Other elements recovered include Cratognathodus kochi, Cypridodella (Lonchodina) spp. and Prioniodina spp..
Differing not merely in appearance at the level of P1 (platform), each element of the apparatus is morphologically distinct from its correspondent element in Gondolellidae. The multi- element widely corresponds with one of the "conodont sets" that Huckriede (1958) first observed in the Alps. In the reconstruction of an octomembrate Gladigondolellid apparatus, Hirsch (1994) regarded Cratognathodus kochi as an additional ozarkodiniform element, next to Ozarkodina saginata. In Ishida and Hirsch's (2011) view, the elements of the Gladigondolellid apparatus consist of the discrete forms Gladigondolella tethydis (P1), Ozarkodina saginata (P2a), "Cratognathodus" kochi (P2b), Lonchodina venusta (M), Roundya lautissima (S0), an enantiognathid (pars petrae-viridis sensu Koike, 1999) (S1), Lonchodina spengleri (S2), Enantiognathus petrae-viridis (S3) and Hindeodella multihamata (=H. pectiniformis) (S4).
The position of the cratognathodiform element in the Gladigondolellid apparatus remains debatable. Koike (1999, p. 236) recalls the statement of Kozur and Mostler (1972, p. 19) that the so-called Cratognathodus pectiniform element (P1) is an immature Gladigondolella.
Koike (1999) regards the cratognathodiform element as the P1 element of the octomembrate species Cratognathodus multihamatus, whose other elements are supposedly Gladigondolellid. According to Orchard (2005), Gladigondolella and Cratognathodus are multi-element genera of Gladigondolellinae Hirsch 1994. Kolar-Jurkovšek et al. (2005) held the P1 element of the genus Cratognathodus for a cavital neospathodiform. This calls to mind a phylogenetic relationship between Cratognathodus and Gladigondolella of the kind that exists within the lineage of alternating neospathodiform and paragondolelliform taxa, sensu Hirsch (1994). In Orchard (2005), the Cratognathodus multi-element from the Spathian of Oman shows transitional features with gondolellid elements, suggesting the possible early Gladigondolellid phylogeny from a neospathodiform stock. However, Gladigondolella saginata, from the Spathian of Chios, has a cratognathodiform element.
Koike (1999, p. 237–238, fig. 2) based his generic segregation of Cratognathodus from Gladigondolella on the apparently low correlation coefficient of their occurrence within the Taho Formation of SW Japan. Here, Cratognathodus multihamatus occurs in the early Anisian Timorensis Zone and G. tethydis covers the late Anisian (Bulgarica Zone), while they occur together in the early Carnian (Nodosus Zone). On the other hand, in the late Anisian to late Ladinian Schreyeralm Formation of the Feuerkogel Section (northern Calcareous Alps, Austria), Mosher (1968, p. 913, table 2) recorded high ratios of both Cratognathodus kochi and G. tethydis at several levels. Similar ratios occur within the samples of Nogami (1968) and Koike (1982) from Malaysia. Therefore, in our opinion, the steady distribution of ozarkodiniform P2a and cratognathodiform P2b elements in samples containing the Gladigondolella apparatus strongly suggests the coeval nature of these forms alternating in the same apparatus-position of distinct multi- element sets. In a similar case, Lambert et al. (2007) interpreted distinctive dimorphic characters in detail of the P2 element of Jinogondolella, a Middle Permian gondolellid in which an asymmetric pair or two complete pairs may represent different individuals, as indicating sexual dimorphism.
Koike (1999) proposed an octomembrate Cratognathodus apparatus of 15 elements; the single pairs of segminate P1, angulate P2, breviform digyrate S2, extensiform digyrate S2, bipennate S3, and S3 elements, and a single unpaired alate S1 element. P1 elements characterized by relatively broad cusp with expanded basal cavity and large discrete denticles. Mosher (1968) established the genus Cratognathodus, including four form species: Prioniodina kochi Huckriede, Cratognathodus posterognathus and two unidentified species, which are both characterized by the presence of a strong broad cusp with a widely expanded basal cavity, and relatively small number of discrete denticles. Cratognathodus kochi and C. posterognathus are respectively identical with the P1 and P2 elements of the C. multihamatus apparatus. Among three specimens illustrated as C. kochi by Mosher (1968), one specimen (pl. 113, fig. 4) is not a typical P1 element of the C. multihamatus apparatus. Mosher (1968), however, regarded the form species Prioniodina kochi Huckriede as the type species of his genus Cratognathodus. Kozur and Mostler (1972) claimed that the genus Cratognathodus created by Mosher (1968) is not a valid taxon because the holotype and all other specimens previously described as the form species C. kochi are immature forms of the "gladigondolelliform" P1 elements of Gladigondolella tethydis (Huckriede). Based on Koike's (1999) observation on P1 elements of G. tethydis from the Taho Formation and other limestone formations in Japan, the immature forms of the element are characterized by a narrow platform-like anterior process and gradually increasing denticles in length toward the anterior as observed in the mature forms. The immature forms of the P1 elements of G. tethydis can be, therefore, easily distinguished from the form species Cratognathodus kochi. The P1 element of C. multihamatus represents various features in the shape and size of the cusp and denticulation on the anterior process. The holotype of the form species C. kochi (Huckriede, pl. 2, fig. 11) possessing a short broad cusp and subequal denticles is safely assigned within the range of morphologic variation of the P1 element of C. multihamatus and agrees well with the specimens illustrated in Koike (1999, figs. 3, 16, 26).
Stratigraphic range: Aegean (Anisian)–Julian (Carnian) of Turkey, Austria, Romania, Bulgaria, Greece, China, USA and Canada.
Gladigondolella sp. A
Type locality: Kocaeli Peninsula (Turkey), between Orta Dere and Kurt Dere.
Type stratum: Upper part of Kücükburun Member of Tepeköy Formation, sample KTM3-0144.
Age: Early Pelsonian.
Diagnosis: The massive platform of the P1 element is thick, flat-shaped, bilaterally convex and symmetrical. Keel is elevated and elongated with narrow amygdaloid basal pit in the posterior third of the unit. Linear carina with at least a dozen triangular, rather low denticles. Straight basal margin. Apparatus similar to that of Gladigondolella tethydis.
Comparisons: The carina of Gladigondolella sp. A is higher and more fused than that of most other species of Gladigondolella. The platform is also wider than that of G. carinata or G. arcuata.
Material: One single P1 element. Gladigondolellid elements are very abundant in sample KTM3-0144 and are likely to belong to this species.
Occurrence: Limited to the Kücükburun Member of the Tepeköy Formation (Kocaeli Peninsula, NW Turkey).
Family Gondolellidae Lindström, 1970
Subfamily Neogondolellinae Hirsch, 1994
Genus Neogondolella, Bender and Stoppel, 1965
Type species Gondolella mombergensis Tatge, 1956
Original diagnosis (Bender and Stoppel, 1965, p. 243): The holotype of the type species is unornamented segminiplanate P1 element with a strong, partly fused carina of variable height ending in a commonly pronounced (sub)terminal cusp. Neogondolella was introduced for smooth forms that had formerly been included in Gondolella Stauffer and Plummer (Bender and Stoppel, 1965, p. 243; after Orchard, 2005).
Diagnosis: Narrow and long, slightly asymmetric platform; broadest near the middle and gradually narrowing towards both ends. A posterior platform constriction may be present. Blade with 8–16 anteriorly fused denticles, which decrease rapidly in height. The carina is lowest in the middle of the element, and rises again towards the posterior. At the posterior end, the cusp may be terminal or a posterior denticle may be present. Strongly elevated basal field occupies the lower side. Pit, surrounded by a small, rectangular loop, located below the cusp and near the posterior end. Narrow and high keel.
Remarks: The multielement Neogondolella apparatus has been reconstructed by Orchard and Rieber (1999), and further examples are given by Orchard (2005) with modifications by Goudemand et al. (2011). In the paper by Orchard (2007), the apparatus described as "Neogondolella inclinata" is now referred to as N. liardensis. The multielement apparatus of Neogondolella is characterized by the presence of an S4 element with a bifurcated anterior process, and an S0 element whose anterior processes bifurcate either at the cusp, or several denticles to the anterior.
Stratigraphic range and occurrence: Austria, Bulgaria, Turkey, Germany, Hungary, Japan, Nevada (USA), Slovenia, Southern Alps, Spain and Canada. Middle upper parts of the Illyrian to lower parts of the Fassanian; in terms of ammonoid zonation corresponds to the upper part of the trinodosus Zone to curionii Zone. The range extends back as far as Spathian in North America.
Remarks: Narrow, lanceolate platforms with weakly fused denticles that increase in size to the posterior, culminating in a large, terminal cusp are often assigned to N. cf. haslachensis (Tatge), the original stratigraphic range and occurrence of which is the Fassanian (early Ladinian) Upper Muschelkalk (Zone 4) in Germany (Franz et al., 2015). The present specimen is possibly a juvenile P1 element.
Genus Nicoraella Kozur, 1980
Type species Ozarkodina kockeli Tatge, 1956
Remarks: A large blade, no platform, and a large gondolelloid basal groove with a large "cavital" pit characterises the Late Permian–Early Triassic Neospathid lineage, from which the Anisian Gondolelloid conodonts derive.
The two Nicoraella species, N. germanica and N. kockeli (Kozur and Mostler, 1972; Tatge, 1956) are index fossils of the upper Bithynian and the Pelsonian.
It is difficult to ascertain common homeomorphy in conodonts, including in the genus Neospathodus (Clark et al., 1981). It is useful to recognize the appearance of the successively distinct looking proteromorphs in its phylogeny, using distinct genera, as e.g. Neospathodus (Clark et al., 1981), Merrillina Kozur, Nicoraella Kozur, Mosherella Kozur, Neocavitella Budurov and Sudar, Misikella Kozur and Mock, all belonging in the same Darwinian anagenesis, each of them representing the successive atavistic reversals that took place over time. The rather punctuated reappearances of Neospathid morphs at certain intervals during the Middle and Late Triassic are atavistic reversals, interpretable as environmental stress-related retrogradations (Kılıç et al., 2018a).
Partial reconstructions of Nicoraella have been attempted by Kozur and Mock (1991), Kozur (1993), Kolar-Jurkovšek et al. (2005), Sun et al. (2009), Kolar-Jurkovšek and Jurkovšek (2010) and Chen et al. (2019a). These studies differed in their identification of S4 elements, and were conflicting in their interpretation of the morphology and presence of a P2 element. The discovery of fused clusters of Nicoraella from South China has enabled the resolution of many of these issues (Huang et al., 2019a, b). The Nicoraella apparatus has been demonstrated to possess an alate S0 element, breviform digyrate S1, S2 and M elements, bipennate S3 and S4 elements, and a carminate P2 element in addition to the distinctive P1 element.
Occurrence: Higher lower Anisian (upper Bithynian, Anagymnotoceras ismidicus Zone) to the top of the Pelsonian (top of the Balatonites balatonicus Zone).
Nicoraella kockeli (Tatge, 1956)
1956 Ozarkodina kockeli n. sp.; Tatge, p. 137, pl. V, figs. 13, 14.
1960 Ozarkodina kockeli; Budurov, p. 16, pl. V, figs. 15–18.
1975 Neospathodus kockeli; Gedik, p. 137, pl. IV, figs. 18, 19.
1975 Neospathodus kockeli; Trammer, pl. XXII, figs. 1–3.
1975 Neospathodus cf. N. kockeli; Hirsch, p. 94–96, pl. 1, figs. 1–4.
1980 Nicoraella kockeli (Tatge); Kozur, p. 127.
1992 Nicoraella kockeli; Kozur and Mostler, pl. 2, fig. 5.
1999 Nicoraella cf. kockeli; Narkiewicz, pl. 1, fig. 3.
2009 Nicoraella kockeli; Zhang et al., fig. 3, part 12.
2010 Nicoraella kockeli; Götz and Gast, fig. 3, part 2.
Description: The P1 elements of Nicoraella kockeli have a large blade with up to six denticles, which increase in size posteriorly to the large, sub-terminal cusp. A short posterior process bears only one or two denticles, much shorter than the cusp. In lateral view, these denticles are posteriorly inclined. Basal cavity is large and elliptical.
Occurrence in Kocaeli: Pelsonian; KTM4/2 and KTM4/3 measured stratigraphic sections.
Stratigraphic Range and Occurrence: Pelsonian of Germany, Poland, Turkey, Greece, Israel, Nevada, China.
Genus Paragondolella Mosher, 1968
Type species Paragondolella excelsa Mosher, 1968
Figs. 3.1–3.28; 4.1–4.18; 5.3, 5.4, 5.8, 5.9, 5.16–5.21
Diagnosis: The species of Paragondolella have a platform that is rounded or blunt at the posterior end, while the anterior end has a short free blade measuring 1/5 to 1/3 of the total length of the unit in adult specimens. The denticles of the blade are fused to two-thirds of their height, commonly forming a curve in their lateral outline, ending in one, sometimes two, free, low, and conical to node-like denticles. Platform margins are quite thick, may be upturned or flat, and are often sculptured (Budurov and Sudar, 1990) but never bear any nodes. The lower side of the species belonging to Paragondolella is characterized by a shallow basal groove or keel that is wide open, broad or narrow, the end of which may be pointed, squared or rounded, but never bifurcated (Orchard, 2013). The basal field ends in a gently rounded loop that may also vary from rectangular-angular to even sub-triangular. The subterminal to backward shifted pit, according to the original diagnosis of Mosher (1968), is wide but shallow, with no protruding edges above the level of the basal field, or loop.
Remarks: Mosher (1968), Orchard (2013), Golding et al. (2014) and Chen et al. (2016) emphasized the high-crested blade-carina that characterizes all growth stages of this genus as typified by its Middle Triassic type species Paragondolella excelsa. The relatively flat, oval platform is a distinctive feature of P. excelsa, that is not exhibited by some other Middle Triassic species such as P. hanbulogi. Likewise, the distinctive polygonal basal pit seen in adult specimens of Paragondolella is seen in many conodont genera throughout time and is therefore not diagnostic. Many Late Carnian species have been referred to as Paragondolella (e.g., Mazza et al., 2012; Rigo et al., 2007; Kozur, 2003), as the subterminal basal pit precludes their assignment to Metapolygnathus (contra Orchard, 1991a, b; Hayashi, 1968). However, Orchard(2014, 2013) instead referred these late Carnian species to a variety of new genera including Parapetella and Quadralella. The absence of the elements belonging to the Paragondolella multielement apparatus (Orchard, 2005) in the late Carnian is evidence that this genus was restricted to the Middle Triassic and lowest Carnian (Chen et al., 2016; Orchard, 2007).
Comparisons: Objecting to Mosher's (1968) statement that the earliest growth stages in Paragondolella lack a platform, Kozur and Mostler (1971) claimed that this is also the case in the earliest growth stages of Neogondolella mombergensis, the type species of Neogondolella. Sweet pointed out that Paragondolella has morphologic characters indistinguishable from those of Neogondolella (Klapper et al., 1973; Sweet, 1970). Nicora (1977) considered the genus Paragondolella as a junior synonym of Neogondolella. However, Paragondolella differs from the genus Neogondolella by the tendency of developing a free blade. Additionally, it has a higher, arched carina, a deeper basal pit, and a broad basal field and a narrow loop that is commonly sub-rectangular.
Stratigraphic range and occurrence: Anisian to lower Carnian of Austria, Hungary, Poland, Italy, Turkey, Slovenia, Croatia, Bulgaria, Greece, India, China, Japan, British Columbia (Canada), and Nevada (USA).
Paragondolella aegaea (Bender 1970)
1970 Neogondolella aegaea Bender, p. 516, pl. 3, figs. 21–26; pl. 4, fig. l.
1975 Neogondolella aegaea Gedik, p. 31, pl. 2, figs. 1–11.
Description: A species with a high carina and a narrow, thin platform. The first two thirds of the unit are straight, only the rear part is slightly bent down. The carina consists of high, laterally compressed and largely fused denticles, which are almost vertical at the proximal end and increase in inclination towards the back. The last or penultimate denticle is generally slightly larger than the others and should be the main denticle. It is almost straight, only towards the end can it be slightly curved to the side.
The platform tapers to the front of the carina; it reaches its maximum width towards the end of the second third of its length. It is thin and almost straight, its edges are hardly rolled up, so that the surface of the carina and the surface of the platform are almost perpendicular to each other. On the outside it extends a little further back than on the inside, where it usually stops at the main denticle, only rarely does it go beyond the end of the carina.
The keel, especially in juvenile specimens, is pedestal-like and is traversed by a furrow which at the end, under the main denticle, widens into a triangular and relatively large basal pit.
Remarks: According to Bender (1970), this form is derived from Spathognathodus gondolelloides Bender, 1970, which appears to be a more recent synonym of Neospathodus timorensis (Nogami, 1968). On the other hand, it leads to Neogondolella unilobata above, with which it is connected by transitional forms. It can be easily distinguished from all other Neogondolella species by its high, fused carina.
Kozur and Mostler (1972, p. 787), put the lower limit (First Appearance Datum, FAD) of Gondolella? aegaea in the early Anisian (Lenotropites caurus- and Anagymnotoceras varium Zone). Important conodont species in the aegaea zone are "Gondolella" timorensis that still occurs at the immediate base of the zone and Gladigondolella tethydis that appears in the upper Spathian. Since 1968 there were reprints of Benderʼs work dated 1967. The volume appeared in 1970 (Kozur and Mostler, 1972, p. 780).
Since Kozur (1980), the taxon aegaea has been replaced by Neogondolella regale, a species that was established in North America by Mosher (1970) in the subrobustus, caurus and varium zone. Mosher (1970) also referred more evolved forms to N. regale that are thinner, have a free blade and an upturned platform that surrounds the terminal denticle. Doing so, Mosher (1970) gave a very loose definition of the species, allowing considerable morphologic latitude in topotypic specimens. Nicora (1977) noted however that some older specimens (from the subrobustus Zone) differ substantially from Anisian specimens by having a large, flat platform and a lower carina, and should be referred to as another genus (Goudemand et al., 2012). To this species, also belongs Neogondolella mombergensis, in Mosher (1968; p. 937, pl. 116, fig. 15).
Recently, the genus Magnigondolella (Golding and Orchard, 2018) was introduced to encompass Anisian conodonts previously referred to as Neogondolella regalis (Mosher, 1970). The new genus shows a wide variety of platform morphologies, including Spathian forms within its range. The Spathian collections in North America suggest that the genus Magnigondolella does originate in the Early Triassic, with a possible ancestry in Borinella. On the other hand, the early Anisian Paragondolella aegaea, in Kocaeli, does only show resemblance to species identified in North America (Dr. Martyn Golding, personal communication, September 2020). In Nevada (USA), British Columbia (Canada), and possibly Svalbard, M. regalis extends from Spathian to Bithynian.
Material: 66 specimens.
Paragondolella praecornuta n. sp. (Kılıç, Budurov, Petrunova and Mirăuţă)
1972 Neogondolella cornuta n. sp.; Budurov and Stefanov, p. 839–840, pl. 3, figs. 9–15, 20–22
1975 Neogondolella cf. huckriedei Budurov and Sudar; Gedik, p. 20, pl. 1, fig. 4
1975 Neogondolella unilobata n. sp.; Gedik, p. 20, pl. 1, figs. 10–11
Derivation of name: Because of stratigraphic precedence and morphological resemblance with Neogondolella
cornuta Budurov and Stefanov, 1972.
Holotype: The specimen shown on Figs. 4.14 and 4.16 denoted as KTM3-0154 (kept in Department of Geological Engineering, Faculty of Engineering, Balikesir University; Figs. 4.12, 4.13, 4.17 kept in Bulgarian Academy of Sciences; Figs. 4.11 and 4.15 are from Gedik, 1975, pl. 1, figs. 10, 11).
Type locality: Southern part of the Kocaeli Peninsula (Turkey). KTM3 measured stratigraphic section.
Type stratum: Dolomitic limestone bed, Kocaeli Peninsula; KTM3 measured stratigraphic section, sample KTM3-0154.
Diagnosis: Specimens referred to as Paragondolella praecornuta n. sp. are conspicuously bowed laterally and bilaterally symmetrical. The major feature of this species is its long free anterior blade. The platform is typically confined to the posterior half of the unit, and is up-curved marginally. The carina carries a series of fused denticles that increase in length and width towards the anterior end of the free blade. The cusp, which is the posterior denticle at all growth stages, is broad, separate, inclined, and circular in transverse section.
Description: Paragondolella praecornuta n. sp. is characterized by a straight or slightly curved element and an isolated posterior cusp. The short posterior platform that completely surrounds the terminal cusp has slightly raised edges and a weak pre-terminal constriction. The carina has more than 12 denticles, the anterior 4–5 being free and massive, while the cusp is free, tall, slightly backwards inclined or upright. The basal groove widens posteriorly and the terminally located pit is small.
Material: More than 100 specimens from the KTM3 measured stratigraphic section, Kocaeli Peninsula (Turkey, collection AMK), and from the Golobardo Section (Bulgaria, collection KB).
Stratigraphic range and occurrence: Paragondolella praecornuta n. sp. is uncommon in the lower part of the Pelsonian (early late Anisian) to Illyrian of Turkey and Bulgaria.
Paragondolella hanbulogi Sudar and Budurov, 1979
Figs. 5.3, 5.4, 5.8, 5.9
1979 Paragondolella hanbulogi n. sp.; Sudar and Budurov, p. 50, 51, pl. 1, figs. 9, 10; pl. 2, figs. 1–9; pl. 3, figs. 1–12.
1984 Gondolella hanbulogi; Farabegoli et al., fig. 4, part e; fig. 5, part c.
1987 Neogondolella hanbulogi; Dagys, p. 10, pl. 2, figs. 6, 7.
1999 Paragondolella hanbulogi; Meco, pl. 1, figs. 3, 4.
1999 Paragondolella cf. hanbulogi; Narkiewicz, pl. 1, figs. 5, 6.
2000 Paragondolella hanbulogi; Márquez-Aliaga et al., figs. 6, 3–4.
2003 Gondolella hanbulogi; Kovács, pl. C-4, parts 1–5; pl. C-5, part 7; pl. C-8, parts 1–4.
2016 Paragondolella hanbulogi; Karádi and Dulai, fig. 3.
Description: The platform is flat, broadest in the middle and with rounded posterior end. The thin to slightly thickened platform edges gently bend upwards, giving the unit its characteristic flat outline. The anterior blade rises up sharply. The carina has about 16 fused denticles, except for their free tips. The narrow basal field ends in a small, rectangular loop that meets the keel at an obtuse angle (Márquez-Aliaga et al., 2000).
Remarks: Transitional features have been observed in some specimens of Paragondolella hanbulogi from Turkey. Some specimens suggest a transition from P. bulgarica, by the slight inclination of the cusp and a more angular outline of the posterior platform end; and others suggest a transition to P. excelsa, by the larger rounded loop and basal groove.
Discussion: The narrow platform and high carina of Paragondolella suggest its possible evolutionary derivation from Chiosella, although there is a stratigraphic gap between the genera (Chen et al., 2016). The small and rectangular- shaped loop and narrow platform also resemble that of Neogondolella, whereas the high carina is similar to that of Magnigondolella; derivation of Paragondolella from either of these genera is also possible.
Stratigraphic range and occurrence: Pelsonian to middle Illyrian (upper half of the balatonicus Zone to trinodosus Zone). Bulgaria, Turkey, Hungary, Slovenia, Albania, Austria, southern Alps and Spain (CCR).
Paragondolella bulgarica Budurov and Stefanov, 1975
Figs. 3.1–3.7, 3.9–3.28
1975 Paragondolella bulgarica Budurov and Stefanov; in Budurov and Stefanov, June 1975, pl. 1, figs. 1–23.
unilobata Gedik; in Gedik, August 1975; pl. 1, figs. 9–25.
1980 Paragondolella bulgarica; Budurov, p. 773, pl. 57, fig. 12.
1980 Gondolella bulgarica; Kozur, table 2, figs. 1, 2.
1980 Neogondolella bulgarica; Pisa et al., p. 817, pl. 61, figs. 1, 2, 3, 5, 7.
1984 Gondolella bulgarica; Farabegoli et al., fig. 4, parts a, b.
1992 Paragondolella bulgarica; Kozur and Mostler, pl. 1, figs. 13–15; pl. 2, figs. 4, 15.
2000 Paragondolella bulgarica; Márquez-Aliaga et al., fig. 6, parts 1–2.
2003 Gondolella bulgarica; Kovács, pl. C1, parts 1–6; pl. C2, parts 1–4; pl. C3, parts 1–3.
2006 Paragondolella bulgarica; Sun et al., p. 626, pl. 1, figs. 16, 17.
2010 Paragondolella bulgarica; Götz and Gast, fig. 3, part 3.
2015 Paragondolella bulgarica; Lehrmann et al., fig. 6, parts 6–8.
Original diagnosis (Budurov and Stefanov, 1975): "Platform strongly curved; high curved blade with denticles; narrow grooved keel with a small, prominent basal pit."
Description: The medium-wide platform has slightly upturned margins and honey-comb ornamentation. The high carina bears 15–20 almost discrete denticles, with at least a third of the length of the denticles being free. The last denticle is higher, sharp and inclined toward the posterior end. The posterior and anterior ends of the element are downturned to produce the high arched shape. On the lower side, a narrow grooved keel ends at the basal pit, which is small, rounded and located posteriorly. The platform tapers posteriorly where often it is upturned and appears as a denticle in lateral view. The species always shows a free blade that is characterized by 2 to 4 elliptical, laterally compressed denticles (Marquez-Aliaga et al., 2000; Nicora, 1977; Budurov, 1975).
Comparison: With its free blade and well-developed posterior platform, the more curved Paragondolella bulgarica differentiates itself from the straight profiled carina and more massive platform in species of Magnigondolella.
Material: Over 300 specimens from KTM3 and KTM4 sections.
Stratigraphic range and occurrence: P. bulgarica marks the base of the Bithynian and ranges up into the Pelsonian, disappearing before the base of the Illyrian. P. bulgarica is very abundant in the Kocaeli Peninsula. It occurs also in Bulgaria, Hungary, Slovenia, Spain, China and N. America.
Paragondolella hirschii n. sp. Kılıç and Budurov
Derivation of name: Dedicated to Prof. Dr. Francis Hirsch (Naruto, Japan) for his outstanding work on Triassic conodonts and stratigraphy.
This species was established after intensive discussion with late Prof. Kiril Budurov.
Holotype: The specimen shown in Figs. 4.1 and 4.2 (kept in Department of Geological Engineering, Faculty of Engineering, Balikesir University).
Type locality: South of Gebze, between the Orta Dere and the Kurt Dere.
Type stratum: KTM3 measured stratigraphic section. From the sample KTM3-9891 (kept in Department of Geological Engineering, Faculty of Engineering, Balikesir University).
Diagnosis: Platform conodont; the carina bears small and thin fused denticles, the 4 to 5 anterior denticles of the carina being prominent. The main cusp is a strong and isolated denticle at the posterior-end of the platform.
Remarks: The carina of this platform conodont bears small and thin fused denticles, the 4 to 5 anterior denticles of the carina being prominent. The main cusp is a strong and isolated denticle at the posterior-end of the platform.
Material: 186 specimens, from Gebze (Turkey) and from the Golobardo Mountains (Bulgaria).
Occurrence: Turkey and Bulgaria.
Stratigraphic range and occurrence: Early Pelsonian (Anisian) of Turkey and Bulgaria.
Paragondolella ebruae n. sp. Kılıç
Figs. 5.16, 5.19–5.21
Derivation of name: Dedicated to the author's wife, Geological Engineer Ebru Kılıç.
Holotype: The specimen shown in Figs. 5.16, 5.19 (kept in Department of Geological Engineering, Faculty of Engineering, Balikesir University).
Type locality: South of Gebze, between Orta Dere and Kurt Dere.
Type stratum: KTM3 measured stratigraphical section, from sample KTM3-0160.
Diagnosis: The platform element of this species has a carina that bears big and fused denticles, characteristically with 3 to 4 large, broad denticles located anterior of the two most posterior denticles. In lateral view the unit is clearly arched. The platform margins are slightly upturned, and much of the upper surface of the platform has a polygonal ornamentation.
Comparison: The species is similar to Paragondolella bulgarica in its lateral view but differs by the irregular shape of its carina, with denticles increasing and decreasing in height along its length (personal communication of the late Dr. Heinz Kozur).
Material: 12 specimens.
Stratigraphic range and occurrence: Early Bithynian to late Pelsonian of Turkey.
Genus Chiosella Kozur 1989
Type species: Gondolella timorensis Nogami, 1968 (p. 127–128, pl. 10, figs. 17a–17c)
Figs. 5.1, 5.2, 5.5, 5. 6
Remarks: Chiosella is characterized by a very narrow smooth platform that develops from the central-lateral rib (Kozur, 1989, p. 416). The lower surface resembles that of Neospathodus and has an excavated or V-shaped incised keel. The terminal basal cavity is very large and funnel-shaped, flaring. The carina extends over the entire unit and consists of high, equally and moderately inclined, laterally compressed denticles. The similarity between the species C. timorensis and Neogondolella aegaea is so pronounced that several authors proposed them to be synonyms; e.g., Kozur (1989, p. 415).
Chiosella timorensis (Nogami, 1968)
Figs. 5.1, 5.2, 5.5, 5. 6
1968 Gondolella timorensis n. sp.; Nogami, p. 127–128, pl. 10, figs. 17–21.
1970 Spathognathodus gondolelloides; Bender, p. 529– 530, pl. 5, figs. 19, 20.
1970 Neospathodus timorensis; Sweet, p. 256, pl. 2, figs. 22, 23.
timorensis; Nicora, p. 92–98, pl. 1–4, all figs. except fig. 3 of pl. 3.
1988 Kashmirella timorensis; Budurov et al., 107–122, pl. 1, figs. 1–4.
timorensis; Kozur, p. 415–416, pl. 15, figs. 1–3.
Gondolella timorensis timorensis; Mietto et al., p. 311, pl. 1, figs. 1–2.
timorensis; Gaetani et al., p. 195, pl. 17, figs. 8–13.
Chiosella timorensis; Orchard, pl. 1, figs. 1–10, 12–14.
2006 Chiosella timorensis; Grãdinaru et al., pl. 1, figs. 1–7.
2007a Chiosella timorensis; Orchard et al., figs. 5(32–34).
2007b Chiosella timorensis; Orchard et al., figs. 6(32–34, 36–38).
2012 Chiosella timorensis; Goudemand et al., p. 203, figs. 2(1–14), 3(1–8).
2015 Chiosella timorensis; Golding et al., p. 166, figs. 11(4–6).
Original diagnosis (Nogami, 1968; p. 127): Part of Gondolella, characterized by its extremely reduced platform. Kozur (1989, p. 415–416) added that, the segminate to segminiplanate P1 element developed a very narrow or rudimentary platform, commonly asymmetrically, on each side of a high carina. The multielement apparatus of Chiosella has been reconstructed by Orchard (2005).
Description: Elongate P1 element with 12–15 moderately fused denticles becoming increasingly reclined toward the posterior margin. In lateral view, below the base of the denticles, a lateral median ridge develops along most of the length of the unit. The aboral margin is straight, somewhat downturned towards the posterior end. The lower surface is marked by a narrow longitudinal groove that expands posteriorly into a broad basal cavity.
Remarks: This species is very similar to C. gondolelloides, which is distinguished from the latter by its relative length and the development of lateral median ridges. Our specimens have a length/height ratio of about (3.1–3.4) : 1, which is clearly different from the holotype of C. gondolelloides with a length/height ratio of 2.5 : 1 (Bender, 1970) in being much more elongate. The common length/height ratios of C. gondolelloides are about 2.1–2.9 (see specimens by Orchard et al., 2007a, b; Grãdinaru et al., 2006). The development of lateral median ridges is one of the key characters defining this species.
Supporting the inclusion of this species in the genus Neogondolella, Nicora (1977) cited the more or less well- developed platform at all growth-stages, the brim enclosing the posterior end of the carina, as well as width/height/length ratios. Topotypic specimens of Chiosella timorensis have a ratio ranging from 1/2.7/5.5 to 1/2.3/7, Sweet's (1970) species from Pakistan 1/2.8/7 to 1/2.3/5, Nicora's (1977) species from Nevada and Chios 1/2.9/5.8 to 1/2.3/7. These values are in agreement with the ratios calculated by Sweet (1970) for some species of the genus Neogondolella (1/1.5/4 to 1/2/8) and differ from the higher and longer values of the genus Neospathodus (mean value: 1/2/3) (Nicora, 1977).
For Goudemand et al. (2012), the downward deflection of the posterior end varies more as suggested by Sweet: in most specimens the basal margin being straight or sub-straight over most of the unit's length and deflected downwardly posterior of the cusp, the basal margin being sometimes curved along the whole length of the element without distinctive deflection posterior of the cusp (Sweet, 1970).
Stratigraphic range and occurrence: The First Appearance Datum (FAD) of Chiosella timorensis is proposed as the index for the base of the Anisian GSSP (Orchard et al., 2007b). However, Goudemand et al. (2012) found C. timorensis in the late Spathian Haugi Zone of Nevada. The range of this species appears to be late Spathian to Aegean.
Family Incertae Sedis: Taxa that are limited to small numbers (three or four) elements of Ozarkodinid derivation, such as Neostrachanognathus, are difficult to classify within the Family Gondolellidae. The genus Cornudina has had several different multielement reconstructions, and it is not clear which elements may belong to this genus and which do not; it is therefore not possible to place this genus within a known family at this time.
Genus Cornudina Hirschmann, 1959
Type species Ozarkodina breviramulis Tatge, 1956
Figs. 4.20, 4.21, 4.23, 4.24, 4.25
Remarks: Cornudina was described as a form-genus by Hirschmann (1959), and has been reported from Lower to Upper Triassic strata in the Tethyan region. The small-sized ramiform element is characterized by a relatively long cusp, a short anterior process and a posterior process with a small number of denticles. There have been several conflicting multielement reconstructions of this genus. Orchard (2005) considered Cornudina to possess bipennate S3 and S4 elements and a breviform digyrate M element based on his reconstruction of C.? igoi, and referred the genus to the Family Gondolellidae. In contrast, Koike (2016) reconstructed the apparatus of C. breviramulis with extensiform digyrate S1 and S2 elements and placed Cornudina in the Family Ellisonidae. Previously, Koike (1996) had regarded Cornudina as possessing a bimembrate apparatus, prior to his 2016 revision. Kılıç et al. (2018b) pointed out that several elements in the Orchard (2005) reconstruction can be assigned to the form-genera Ketinella and Kamuellerella, which rarely occur with Cornudina. Therefore, the multielement apparatus of Cornudina is still unknown, and as this taxon is limited to a maximum of three elements of Ozarkodinid derivation, it is difficult to classify within the family of Gondolellidae and may well belong to a Family Incertae Cedis.
Cornudina oezdemirae Gedik, 1975
Figs. 4.20, 4.21, 4.24, 4.25
1968 Cornudina breviramulis breviramulis (Tatge); Kozur, pl. 3, fig. 29.
1970 Cornudina?latidentata Kozur and Mostler; Kozur and Mostler, pl. 1, fig. 21.
oezdemirae Gedik; Gedik, pl. 7, figs. 15, 24 (holotype), 29.
breviramulis minor Kozur and Mostler; Koike, pl. 7, fig. 4.
igoi Koike; Koike, figs. 4.1–4.20.
2005 Cornudina igoi; Orchard, Text-fig. 1, P1 element.
2018b Cornudina oezdemirae; Kılıç et al., p. 2, figs. 1.1–1.4, 1.7–1.9.
Diagnosis: Unit composed of a long cusp and an anterior process. The length of anterior processes changes depends on the number of the anterior denticles. Cusp inclined posteriorly 40°–50°. Basal margin of unit ranges in length from 150 to 220 μm and cusp ranges in length from 240 to 340 μm (Koike, 1996). Anterior process possesses 1 to 4 denticles. In some specimens one minute denticle present just behind cusp. Anterior denticles increase in length and inclination toward posterior. These anterior denticles can be fused or separated. Basal cavity laterally expanded and elongated as for the number of the anterior denticles. Basal cavity shows drop shape in cross-section.
Comparisons: Cornudina oezdemirae is distinguished from C. breviramulis in having no distinctive denticle behind the cusp.
Remarks: Cornudina oezdemirae is a unimembrate type composed of the P1 element. C. breviramulis ranges from the Early to Late Triassic but the occurrence of C. oezdemirae is restricted in the Early to Middle Triassic. The denticles on the anterior process of the element of this species vary in number as observed in the P1 element of C. breviramulis.
For reasons of priority, Cornudina oezdemirae Gedik (1975), with one anterior denticle, may include C. igoi Koike (1996), although the latter's anterior process possesses up to 3 discrete denticles. Koike's (2006) comment about up to 4 discrete denticles instead of 1 to 3, modifies the multi-element description of Cornudina in Orchard (2005; text-fig. 1A). Cornudina oezdemirae Gedik shows an evolutionary trend of decreasing total length of the anterior processes and number of discrete denticles, ranging from 3 to 1 minute denticle just behind the cusp. Koike (1999) suggested that C. oezdemirae resembled the S0 element in his reconstruction of Neostrachanognathus tahoensis and therefore would be the senior synonym of N. tahoensis. However, C. oezdemirae commonly occurs separately from other elements of N. tahoensis, and so they are retained as separate species here.
Material: 19 species.
Stratigraphic range and occurrence: Early Spathian– Anisian. C. oezdemirae occurs from the basal part of the Neospathodus triangularis-N. homeri Zone and indicates the early Spathian in the Taho Formation of Japan, whereas in Turkey it indicates the Anisian.
latidentata Kozur and Mostler 1970
1956 Ozarkodina breviramulis; Tatge (pars.), p. 126, pl. 5, fig. 12.
1959 Cornudina breviramulis; Hirschmann, p. 90, pl. 4, figs. 3, 4.
1968 Cornudina breviramulis minor; Kozur, p. 943, pl. 3, figs. 27, 30.
1970 Cornudina latidentata sp. nov.; Kozur and Mostler, p. 457, pl. 1, figs. 11–13, 19.
1972 Cornudina latidentata; Kozur and Mostler, p. 37, pl. 1, figs. 18–23, 26, 27.
1972 Cornudina cf. breviramulis minor; Kozur and Mostler, p. 53, pl. 15, figs. 19, 22, 24, 25.
1972 Cornudina cf. breviramulis breviramulis; Kozur and Mostler, p. 53, pl. 15, fig. 23.
1975 Cornudina latidentata; Trammer, pl. 25, fig. 3.
1975 Cornudina latidentata; Zawidzka, p. 246, pl. 35, figs. 7, 9.
1981 Cornudina?latidentata; Wang and Dai, p. 141, pl. 2, fig. 29.
1992 Cornudina latidentata; Kozur and Mostler, pl. 2, figs. 16, 17.
1999 Cornudina?latidentata; Narkiewicz, pl. 1, fig. 4.
2010 Cornudina latidentata; Götz and Gast, fig. 3, part 6.
2018b Cornudina cf. latidentata, Kılıç et al., figs. 1.5, 1.6, 1.10, 1.11.
2019a Cornudina latidentata; Chen et al., pl. 14, figs. 5A–5D.
Remarks: The single specimen referred to as Cornudina cf. latidentata shares the relatively long anterior denticulated process of this species; however, the fragmentary nature of the specimen and uncertainty about the possible synonymy of C. latidentata with C. oezdemirae (Kılıç et al., 2018b) precludes unambiguous assignment to this species.
Stratigraphic range and occurrence: Anisian of Austria, Poland, Turkey and China.
Genus Neostrachanognathus Koike, 1998
Type species Neostrachanognathus tahoensis Koike, 1998
Original diagnosis: The diagnosis of Neostrachanognathus proposed is based on N. tahoensis by Koike (1998). Neostrachanognathus is characterized by a quadrimembrate apparatus composed of a non-denticulated and three denticulated nongeniculate coniform elements. The base of the element is relatively small and short and the basal cavity shallow. Cusp is proclined and tapered. Denticles of denticulated elements are situated at anterobasal portion (figs. 7 and 9 of Koike, 1998). Coniform elements were referred to as M, S1, S2, and S3 positions by Koike (1998). M element bilaterally subsymmetric adenticulated coniform. S1 element bilaterally subsymmetric denticulated coniform and carries one small proclined anterobasal denticle. S2 element has inwardly bending one or two small to large anterobasal denticles. S3 element possesses long upper basal margin and carries inwardly flexing one or two small to large anterobasal denticles. One posterior denticle may be present on the base. An alternative multielement reconstruction for Neostrachanognathus was proposed by Agematsu et al. (2008) based on natural assemblages of N. tahoensis from Japan, and of isolated specimens of N. sp. from Oman. Agematsu et al. (2008) demonstrated the occurrence of ramiform elements (which could not be differentiated due to poor preservation of the natural assemblages, and thus were all referred to as S elements) with the coniform elements of Neostrachanognathus. These reconstructions lack elements corresponding to S0 and M positions, as occur in typical 15 elements ozarkodinid apparatuses.
Neostrachanognathus tahoensis Koike, 1998
1989 Cratognathodus sp.; Buryi, pl. 2, fig. 6.
1998 Neostrachanognathus tahoensis sp. nov.; Koike, p. 127, 128, figs. 4, 9(1–19, 22).
2008 Neostrachanognathus tahoensis; Agematsu et al., p. 1210, figs. 3A–3C, 4A–4C, 5A–5C, 6A–6B, 6D, 6F.
2017 Neostrachanognathus tahoensis; Kolar-Jurkovšek et al., p. 695, fig. 7.
2019b Neostrachanognathus tahoensis; Chen et al., fig. 6, part 5.
Description: Four elements of this apparatus exhibit common characteristics such as thick wall, relatively small and short base, and proclined and tapered cusp (Agematsu et al., 2008). The element illustrated in Fig. 4.22 is a P2 element in the reconstruction of Agematsu et al. (2008), or an M element in the reconstruction of Koike (1998).
Material: 14 specimens.
Stratigraphic range and occurrence: Lower Spathian?- Aegean of China, Japan, Russia, Turkey, Slovenia and Oman.