Taxonomic scope

The Neogondolella constricta group includes segminiplanate platform-bearing species known or suspected to be characterized by an early growth stage with a narrow, constricted posterior platform, and a relatively low blade–carina. Included among these are species that are only known as later growth stages that resemble other constricta group members. The group is centered on the holotype of N. constricta from the Prida Formation of Nevada, a juvenile element from the upper Anisian Rotelliformis Zone. Very similar juvenile elements occur throughout the upper Anisian and lower Ladinian strata worldwide, but they mature into a variety of different species.

Differentiation of members of the constricta group must generally rely on morphological features of adult elements. The following previously described taxa (with original genus name) are discussed here: Neogondolella aldae Kozur, Krainer and Mostler, Reference Kozur, Krainer and Mostler1994b; Neogondolella bakalovi Budurov and Stefanov, Reference Budurov and Stefanov1972; Neogondolella balkanica Budurov and Stefanov, Reference Budurov and Stefanov1975a; Gondolella constricta Mosher and Clark, Reference Mosher and Clark1965; Neogondolella cornuta Budurov and Stefanov, Reference Budurov and Stefanov1972; Neogondolella excentrica Budurov and Stefanov, Reference Budurov and Stefanov1972; Neogondolella longa Budurov and Stefanov, Reference Budurov and Stefanov1973; Gondolella mesotriassica Kozur and Mostler, Reference Kozur and Mostler1982; Gondolella postcornuta Kovács, Reference Kovács1994; Neogondolella posterolonga Kozur, Krainer and Mostler, Reference Kozur, Krainer and Mostler1994b; Gondolella pseudolonga Kovács, Kozur, and Mietto, Reference Kovács, Kozur and Mietto1980; Neogondolella tardocornuta Budurov and Stefanov, Reference Budurov and Stefanov1984; and Gondolella transita Kozur and Mostler, Reference Kozur and Mostler1971.

Type species

Gondolella mombergensis Tatge, Reference Tatge1956 (Ta 1956/5) from the upper Muschelkalk, Schmidtdiel Quarry, Momberg, near Marburg, Germany.

Neogondolella constricta (Mosher and Clark, Reference Mosher and Clark1965)
 Figures 6, 7

p*  Reference Mosher and Clark1965

Gondolella constricta Mosher and Clark, p. 560, pl. 65, figs. 11, 18, 21, 24, 25 (only).

p  Reference Mosher and Clark1965

Gondolella mombergensis Tatge; Mosher and Clark, p. 560, pl. 65, figs. ?20, 28 (only).

?  Reference Budurov and Stefanov1975a

Neogondolella balkanica Budurov and Stefanov, p. 792–794, pl. 1, figs. 24–31.

p  Reference Rafek1976

Neogondolella constricta; Rafek, pl. IV, fig. 24 (only).

p  Reference Rafek1976

Neogondolella cf. longa Budurov and Stefanov; Rafek, pl. I, figs. 34, 35 (only).

Reference Kovács, Kozur and Mietto1980

Gondolella constricta; Kovács and Kozur, pl. 3, fig. 4a–c.

Reference Kozur, Krainer and Mostler1981

Neogondolella cornuta; Mietto and Petroni, p. 552–553, pl. 57, figs. 5a, b.

Reference Kozur, Krainer and Mostler1981

Neogondolella longa; Mietto and Petroni, p. 553–554, pl. 57, figs. 8a–c.

non  Reference Papšová and Pevný1982

Neogondolella constricta; Papšová and Pevný, Pl. XX, figs. 7–12.

?  Reference Budurov and Stefanov1984

Neogondolella tardocornuta Budurov and Stefanov, p. 605–607, pl. 1, figs. 15–20.

Reference Farabegoli, Levanti, Perri and Veneri1984

Neogondolella cornuta; Farabegoli et al., figs. a1–3, b1–3.

p  Reference Kovács1986

Gondolella constricta; Kovács, pl. IV, fig. 4a–d; pl. V, figs. 5a–d, 6a–d; pl. VI, figs. 3a, b, 4a, b; pl. X, figs. 4a–c, 5a–c, 6a–c (only).

non  Reference Dȕrkoop, Richter and Stritzke1986

Gondolella constricta; Dürkoop et al., pl. 21, figs. 1a–c, 2a, b.

non  Reference Ding and Huang1990

Neogondolella constricta; Ding and Huang, pl. 1, figs. 2, 3.

Reference Koca, Gedik and Balcioğlu1992

Neogondolella constricta; Koca et al., pl. 1, figs. 11a–c, 13.

Reference Koca, Gedik and Balcioğlu1992

Neogondolella cf. cornuta; Koca et al., pl. 1, figs. 15a–c.

Reference Chhabra and Kumar1992

Neogondolella constricta; Chhabra and Kumar, pl. 3, figs. 2, 3a, b, 4a–c, 6a, b.

Reference Kozur, Krainer and Mostler1994b

Neogondolella constricta; Kozur et al., p. 172–174, pl. 4, figs. 12–15, 17, 20.

non  Reference Polak, Havrila, Filo and Pevný1996

Neogondolella constricta; Polak et al., pl. 12, figs. 1–5.

?  Reference Buryi, Baud, Popova, Dickins, Lucas and Zacharov1997

Neogondolella constricta; Buryi, pl. 1, fig. 15.

non  Reference Pevný and Salaj1997

Gondolella constricta; Pevný and Salaj, p. 101–102, pl. IX, figs. 14, 15; pl. X, figs. 1, 2; ?pl. XI, figs. 2, 3.

Reference Pevný and Salaj1997

Gondolella balkanica; Pevný and Salaj, p. 103, pl. XI, figs. 6–8; pl. XV, figs. 5–8.

Reference Orchard and Rieber1999

Neogondolella ex gr. constricta; Orchard and Rieber, pl. 4., figs. 1–12 (multielement).

p  Reference Orchard, Colpron and Nelson2006

Neogondolella ex gr. constricta; Orchard, pl. 5., figs. 9, 10 (only).

non  Reference Sun, Sun, Hao and Jiang2006

Neogondolella constricta; Sun et al., pl. 1, figs. 21–24.

non  Reference Dong and Wang2006

Neogondolella constricta; Dong and Wang, pl. 41, figs. 10, 18.

non  Reference Wu, Yao, Ji and Wang2008

Neogondolella constricta; Wu et al., pl. II, figs. 12–15.

p  Reference Wu, Yao, Ji and Wang2008

Neogondolella mombergensis (Clark and Mosher) (sic); Wu et al., pl. II, figs. 4, 5, 18, 22.

p  Reference Wu, Yao, Ji and Wang2008

Neogondolella navicula (Clark and Mosher) (sic); Wu et al., pl. I, figs. 2, 3, 7, 10.

non  Reference Zhang, Zhou, Lu, Xie, Lou and Liu2009

Neogondolella constricta; Zhang et al., figs. 3. 13, 16.

Reference Orchard2010

Neogondolella constricta; Orchard, fig. 9. 14, 15.

Reference Golding2014

Neogondolella ex gr. constricta morphotype gamma; Golding, fig. 2.24, parts 1–6.

Reference Golding2014

Neogondolella ex gr. constricta morphotype delta; Golding, fig. 2.25, parts 1–6.

non  Reference Sun, Jiang, Sun and Hao2014

Neogondolella constricta; Sun et al., Reference Sun, Jiang, Sun and Hao2014, fig. 3h, k–m.

Reference Lehrmann, Stepchinski, Altiner, Orchard and Montgomery2015

Neogondolella ex gr. constricta; Lehrmann et al., fig. 6.9, 6.10.

non  Reference Bo, Yao, Xiao, Bai and Peng2017

Neogondolella constricta; Bo et al., fig. 4.1–4.4.

Reference Zhang, Orchard, Algeo, Chen, Lyu, Zhao, Kaiho, Ma and Liu2019

Neogondolella constricta; Zhang et al., fig. 7.13a–c.

non  Reference Xie, Liu, Lou, Hu, Zhou, Huang and Wen2019

Neogondolella constricta; Xie et al., fig. 3.2, 3.3, 3.7.

non  Reference Golding2021

Neogondolella ex gr. constricta (Mosher and Clark); Golding, p. 584, pl. 3., figs. 29–35.

non  Reference Qin, Golding, Jiang, Chen, Zhang, Kang, Wang and Yuan2021

Neogondolella constricta; Qin et al., pl. 8., figs. 8–13.

Figure 6.

Neogondolella constricta (Mosher and Clark) from the Rotelliformis Zone, Fossil Hill, Nevada. (1–3) USNM 145189, refigured holotype, clarkei Subzone; (4–6) GSC 141848, FH15, clarkei Subzone; (7) GSC 141849, FH6, burckhardti Subzone; (8–10) GSC 141850, FH24, cricki Subzone; (11, 12) GSC 141851, FH8, burckhardti Subzone.; (13–15) GSC 141852, FH17, vogdesi Subzone; (16, 17) GSC 141853, FH22, cricki Subzone. Scale bar = 200 μm.

Figure 7.

Neogondolella constricta (Mosher and Clark) from B.C. (1–3) GSC 141854, sample 83/205B, Toad Formation, Toad River Canyon, Deleeni Zone; (4, 5) GSC 141855, and (9–11) GSC 141856, both sample 92/AH2, Yellow Bluffs, Alaska Highway, Deleeni Zone; (6–8) GSC 141857, sample 97/WapA6, Llama Formation, Cirque B, Ganoid Ridge, bracketed between the Poseidon and Meginae zones. Scale bar = 200 μm.

Holotype

USNM 145189 (Mosher and Clark, Reference Mosher and Clark1965, pl. 65, figs. 21, 24, 25). Re-illustrated in Figure 6.1–6.3. From sample FH3, clarkei Subzone of the Rotelliformis Zone, upper Anisian, Prida Formation, Fossil Hill, Nevada; deposited in the US National Museum of Natural History.

Diagnosis

The relatively short and arched segminiplanate P₁ element has at first a slender, biconvex platform that broadens with growth and maintains a typical length breadth ratio of ~4:1. A posterior platform constriction present in juveniles disappears during growth as the posterior platform expands and becomes as broad as the median platform. The rounded posterior margin of later growth stages may include a narrow brim around a large, upright to slightly reclined terminal denticle that generally becomes larger than the cusp during growth, which is typically the penultimate carina denticle. During growth, the anterior blade denticles become increasingly elevated relative to those of the median carina, which become increasingly fused. On the underside, a symmetrical basal loop of uniform width surrounds the subterminal pit.

Occurrence

Neogondolella constricta occurs in Nevada throughout the Fossil Hill Member of the Prida Formation at Fossil Hill, where it is abundant (many hundreds of specimens) in the Rotelliformis Zone, but much less so through the Meeki and Occidentalis zones, and into the Subasperum Zone. Elsewhere in Nevada, the species is known from the Wheeler Mine area, and in the Tobin Range. In B.C., it is particularly common in the Deleeni Zone, including in the type locality of that zone and additional locations nearby. Elsewhere in B.C. it ranges through the Poseidon Zone (Table 2).

Neogondolella constricta has a global reach: upper Muschelkalk, northern Germany (Rafek, Reference Rafek1976); Campogrosso section, NE Italy (Mietto and Petroni, Reference Mietto and Petroni1981); middle Bivera Formation, southern Alps, northern Italy (Farabegoli et al., Reference Farabegoli, Levanti, Perri and Veneri1984); Alsóhegy, N. Hungary (Kovács and Kozur, Reference Kozur1980); NE Rudabánya Mountains, Hungary (Kovács, Reference Kovács1986); eastern Turkey (Koca et al., Reference Koca, Gedik and Balcioğlu1992); Kalapani Limestone, northern India (Chhabra and Kumar, Reference Chhabra and Kumar1992); western Carpathians, Slovakia (Pevný and Salaj, Reference Pevný and Salaj1997); chert of Tsentralnaya Mts, Sikhote–Alin (Buryi, Reference Buryi, Baud, Popova, Dickins, Lucas and Zacharov1997); subsurface Polish lowlands (Narkiewicz, Reference Narkiewicz1999); Jones Lake Formation, distal allochthon, south-central Yukon Territory (Orchard, Reference Orchard, Colpron and Nelson2006); Xinyuan Formation, Guandao, Nanpanjiang Basin, South China (Lehrmann et al., Reference Lehrmann, Stepchinski, Altiner, Orchard and Montgomery2015); Kamura Formation, Kamura, central Kyushu Island, Japan (Zhang et al., Reference Zhang, Orchard, Algeo, Chen, Lyu, Zhao, Kaiho, Ma and Liu2019). In China, several reports of N. constricta are discounted (see synonymy) or are insufficiently illustrated (Wang and Wang, Reference Wang and Wang1976; Wang et al., Reference Wang, Wang, Li and Wei2005), but the species does occur in the Qingyan Formation in Guizhou Province (Wu et al., Reference Wu, Yao, Ji and Wang2008). Those reported from Bithynian strata at Deşli Caira, Romania (Golding, Reference Golding2021) are now excluded from this species.

Description

The holotype is a comparatively small, elongate element with a platform that extends throughout the length of the element but narrows at a posterior constriction near the cusp and expands around the final denticle. The axial blade–carina is composed of ~15 relatively low, closely spaced denticles, the anterior ones of which are slightly larger, higher, and more discrete than those of the median carina. The cusp is the penultimate denticle and is twice the size of the adjacent posteriormost denticle. Later growth stages show the posterior platform constriction progressively overgrown to produce at first a tapered posterior outline and then a broad, rounded posterior outline equal in breadth to that of the median part. Concurrently, the anterior denticles of the fixed blade become relatively higher, those of the median carina become increasingly fused into a low ridge, and the posteriormost denticle enlarges and dominates the posterior margin; a narrow posterior platform brim often develops. Very large elements may have some subdued anterior platform crenulation.

Comparisons

Adult specimens of Neogondolella constricta differ from those of N. cornuta in their comparatively shorter and broader platform (4–4.5:1 compared with 5–5.5:1), an outline that is commonly biconvex rather than subrectangular, and in their less conspicuous posterior carina and cusp. Both N. aldae and N. posterolonga differ from N. constricta by possessing platforms that are wider in the anterior and much narrower in the posterior.

Remarks

The distinctive posterior denticle (not the cusp) of Neogondolella cornuta is often larger than that in N. constricta, but later growth stages of both species can display a prominent and reclined terminal denticle, which has been regarded as diagnostic for N. cornuta. Variation in this feature led to separation of both N. balkanica (upright denticle with a platform brim) and N. tardocornuta (upright denticle without a brim) in the Balkans. Nicora and Kovács (Reference Nicora and Kovács1984) previously suggested that N. balkanica was synonymous with N. constricta, while N. tardocornuta was originally combined in N. balkanica by its authors (Budurov and Stefanov, 1975). The holotypes of both Balkan species have platform proportions (~4:1) comparable to that of N. constricta, and variability of the posterior margin in the Nevadan material is such that those species are provisionally included in synonymy here. As pointed out by Kozur et al. (Reference Kozur, Krainer and Mostler1994b), the types of all these species originated in different parts of the late Anisian–early Ladinian, but this does not exclude a long range for N. constricta.

Neogondolella aldae Kozur, Krainer and Mostler, Reference Kozur, Krainer and Mostler1994b
 Figure 8

p  Reference Mosher and Clark1965

Gondolella navicula Huckriede; Mosher and Clark, p. 560–561, pl. 66, figs. 10, 17, 18, 21 (only).

p  Reference Nicora and Kovács1984

Gondolella mombergensis longa (Budurov and Stefanov, Reference Budurov and Stefanov1973); Nicora and Kovács, p. 150, pl. 10, figs. 2, 4, 9 (only).

*  Reference Kozur, Krainer and Mostler1994b

Neogondolella aldae Kozur, Krainer, and Mostler, p. 179–181.

*  Reference Kozur, Krainer and Mostler1994b

Neogondolella aldae aldae Kozur, Krainer, and Mostler, p. 181–182 (see synonymy).

p  Reference Wu, Yao, Ji and Wang2008

Neogondolella mombergensis (Mosher and Clark) (sic); Wu et al., pl. II, fig. 9.

p  Reference Golding2014

Neogondolella ex gr. transita, morphotype beta; Golding, fig. 2.29, parts 1–3.

Reference Golding2014

Neogondolella constricta morphotype beta; Golding, fig. 2.23, parts 1–9.

Figure 8.

Neogondolella aldae Kozur, Krainer, and Mostler from (1–10) the Toad Formation in B.C. and (11–19) the Prida Formation in Nevada. (1–3) GSC 141858, sample 92/AH21, Yellow Bluffs, Alaska Highway, Chischa Zone; (4, 5) GSC 141859, sample 92/AH2, Toad Formation, Yellow Bluffs, Alaska Highway, Deleeni Zone; (6–8) GSC 141860, sample 83/205B, Toad River Canyon, Deleeni Zone; (9, 10) GSC 141861, sample 83/ MJO-Bone. Toad River Canyon, ?Deleeni Zone; (11, 12) GSC 141862, sample FH47, Meeki–Occidentalis zonal boundary; (13–15) GSC 141863, sample SH516, Subasperum Zone; (16, 17) GSC 141864, sample SH531, Occidentalis Zone; (18, 19) GSC 141865, sample SH517, Subasperum Zone. Scale bar = 200 μm.

Holotype

The specimen, figured by Nicora and Kovács (Reference Nicora and Kovács1984, pl. 10, fig. 9) as Gondolella mombergensis longa Budurov and Stefanov. From sample N49 in the middle Prida Formation, Nevadites humboldtensis Smith, Reference Smith1914, beds of the lower Occidentalis Zone at Fossil Hill, Humboldt Range, Nevada.

Occurrence

Mosher and Clark (Reference Mosher and Clark1965) illustrated elements from the Meeki Zone in Nevada, and Kozur et al. (Reference Kozur, Krainer and Mostler1994b) gave a range of lower Meeki and Occidentalis zones. In the present study, the species was found to occur first in the middle of the Rotelliformis Zone and as high as the Subasperum Zone. Neogondolella aldae is one of the more common species of the constricta group in the Humboldt Range, being found in small numbers in about half the studied samples. Elsewhere in Nevada, the species is known from the Tobin Range. In Canada, the species occurs in the Deleeni and Chischa zones at Yellow Bluffs, in the Deleeni Zone of Toad River Canyon, in the Matutinum Zone at Wapiti Lake, and as high as the Meginae Zone along the Alaska Highway at milepost 386 (Table 2).

In Europe, Neogondolella aldae occurs in the Reitzi Zone and Nevadites Zone of the Alps and Bulgaria (Kozur et al., Reference Kozur, Krainer and Mostler1994b) but it has not featured in many studies. A specimen illustrated by Wu et al. (Reference Wu, Yao, Ji and Wang2008) from the Qingyan Formation in Guizhou, China, appears to represent this species.

Description

The platform of this arched segminiplanate element is widest around midlength, with biconvex margins in the anterior two-thirds to three-quarters, and subparallel margins in the remainder of the distinctly narrower posterior platform. Large specimens show increasing lateral growth in the posterior part, which remains narrower than the anterior platform. Overall, the typical length:breadth ratio is about 4:1. The narrowly rounded to subquadrate posterior margin of adult elements may include a very narrow platform brim around a large, moderately reclined terminal denticle, which is separated from the smaller cusp immediately in front of it; both these denticles commonly lie in the posterior constricted part. To the anterior, denticles of the carina of mature elements are largely fused into a low ridge that rises to the posterior and extends to the anterior relatively elevated fixed blade, which is commonly composed of 4 discrete denticles. On the underside of the largest specimens, the pit is anteriorly shifted within the basal scar that extends behind it.

Comparisons

Unlike Neogondolella constricta and N. cornuta, the posterior platform constriction present in juveniles of N. aldae does not disappear during growth as the posterior platform remains narrow. Neogondolella posterolonga differs in its longer and more slender platform. The older N. shoshonensis Nicora, Reference Nicora1976, has a similar platform outline but is narrower with a higher posterior carina.

Remarks

The original description of Neogondolella aldae (Kozur et al., Reference Kozur, Krainer and Mostler1994b) included reference to its distinct terminal “cusp” that was not fused with the posterior platform margin, unlike in N. cornuta. As discussed, previously, such fusion is variable and a more reliable character to distinguish the two species is their platform shape. Note that the true cusp in the adults of the constricta group is the penultimate denticle of the carina and not the larger terminal denticle.

Some Nevadan specimens previously illustrated by Nicora and Kovács (Reference Nicora and Kovács1984) and included in Neogondolella aldae by Kozur et al. (Reference Kozur, Krainer and Mostler1994b) do not exhibit the constricted posterior platform present in the holotype; these are excluded from the present species.

Neogondolella cornuta Budurov and Stefanov, Reference Budurov and Stefanov1972
 Figure 9.1–9.6, 9.9–9.15

p  Reference Mosher and Clark1965

Gondolella constricta Mosher and Clark, p. 560, pl. 65, fig. 22 (only).

p  Reference Mosher and Clark1965

Gondolella mombergensis Tatge; Mosher and Clark, p. 560, pl. 65, fig. 29 (only).

p  Reference Mosher and Clark1965

Gondolella navicula Huckriede; Mosher and Clark, p. 560–561, pl. 66, figs. 19, 20 (only).

*  Reference Budurov and Stefanov1972

Neogondolella cornuta Budurov and Stefanov, p. 839–840, pl. 3, figs. 9–15, 20–22.

Reference Trammer1975

Gondolella cornuta; Trammer, pl. 22, figs. 8a, b, 9a, b.

?  Reference Kovács, Kozur and Mietto1980

Gondolella pseudolonga Kovács, Kozur and Mietto, p. 218–219, pl. 1, figs. 1–4.

Reference Vrielynck1984

Gondolella cornuta; Vrielynck, p. 187–189, pl. 3, figs. 4a–c.

p  Reference Vrielynck1984

Gondolella pseudolonga; Vrielynck, p. 194–195, pl. 3, figs. 2a–c (only)

Reference Orchard and Austin1986

Neogondolella constricta (Mosher and Clark); Orchard, pl. 5.4, fig. 8 (only).

non  Reference Dȕrkoop, Richter and Stritzke1986

Gondolella cornuta; Dürkoop et al., pl. 17, fig. 5a, b; pl. 20, figs. 13a–c, 15a, b.

Reference Uroševič and Sudar1991

Neogondolella cornuta; Uroševič and Sudar, pl. 1, fig. 13.

?  Reference Ramovš and Goričan1995

Neogondolella cornuta; Ramovš and Goričan, pl. 8, fig. 4.

Reference Pevný and Salaj1997

Gondolella cornuta; Pevný and Salaj, pl. XII, figs. 3, 11.

?  Reference Narkiewicz1999

Neogondolella cornuta > N. mesotriassica; Narkiewicz, pl. 3, figs. 3, 4.

Reference Orchard2005

Neogondolella ex gr. constricta; Orchard, fig. 10A–I (multielement).

non  Reference Wang, Wang, Li and Wei2005

Neogondolella cornuta; Wang et al., pl. II, fig. 20.

non  Reference Dong and Wang2006

Neogondolella constricta cornuta; Dong and Wang, pl. 41, fig. 3.

Reference Orchard, Colpron and Nelson2006

Neogondolella ex gr. constricta; Orchard, pl. 4, fig. 15.

Reference Nakrem, Orchard, Weitschat, Hounslow, Beatty and Mørk2008

Neogondolella ex gr. constricta; Nakrem et al., fig. 5.18–5.20.

Reference Orchard2010

Neogondolella cornuta; Orchard, fig. 9.12, 9.13.

non  Reference Sun, Jiang, Sun and Hao2014

Neogondolella constricta cornuta; Sun et al., fig. 3.g?, i, j.

?  Reference Sun, Jiang, Ji and Hao2016

Neogondolella constricta cornuta; Sun et al., fig. 3.5.

non  Reference Xie, Liu, Lou, Hu, Zhou, Huang and Wen2019

Neogondolella constricta cornuta; Xie et al., fig. 3.12.

non  Reference Qin, Golding, Jiang, Chen, Zhang, Kang, Wang and Yuan2021

Neogondolella cornuta; Qin et al., pl. 3, figs. 3, 4; pl. 7, fig. 4; pl. 9, figs. 9–11.

Reference Karádi, Budai, Haas, Vörös, Piros, Dunkl and Tóth2022

Neogondolella cornuta; Karádi et al., fig. 9D.

Reference Karádi, Budai, Haas, Vörös, Piros, Dunkl and Tóth2022

Neogondolella pseudolonga; Karádi et al., fig. 9C.

Figure 9.

(1–6, 9–15) Neogondolella cornuta Budurov and Stefanov from the Prida Formation, Nevada, Occidentalis Zone; (11–15) the Llama Formation, B.C., Chischa Zone. (1, 2) GSC 141866, sample FH56; (3, 4) GSC 141867, sample SH524; (5, 6) GSC 132583, sample SH524; (9, 10) GSC 141869, sample SH527; (11–13) GSC 141870; (14, 15) GSC 141871, both from sample 97/WapB18, Cirque B, Ganoid Ridge. (7, 8) Neogondolella aff. N. cornuta Budurov and Stefanov; GSC 141872, sample SH524. Scale bar = 200 μm (1–13), 250 μm (14, 15).

Holotype

Budurov and Stefanov, Reference Budurov and Stefanov1972, pl. 3, figs. 20–22, Bu 1045/1, from middle–upper Illyrian strata, III γ = cornuta conodont zone; Golo–Bârdo mountains south of Pernik, Bulgaria. Budurov's collections (Bu) are in the Geologisches Institut, Sofia, Bulgaria.

Occurrence

In Nevada, Neogondolella cornuta appears in the Meeki Zone, is abundant (many hundred specimens) through the Occidentalis Zone and is less common in the Subasperum Zone. It also occurs at Wheeler Mine and in the Tobin Range. In B.C., it is known from the Chischa and Poseidon zones at Wapiti Lake, and with the bamberi ammonoid fauna at milepost 386, Alaska Highway (Table 2).

In addition to the Bulgarian types that characterize the Illyrian (Budurov and Trifonova, Reference Budurov and Trifonova1995), Neogondolella cornuta is recorded in southwestern Holy Cross Mountains, Poland (Trammer, Reference Trammer1975); Carnic Alps–Dolomites, NE Italy (Vrielynck, Reference Vrielynck1984); Stuhini Group, Stikine terrane, northern B.C. (Orchard, Reference Orchard and Austin1986, Reference Orchard, Orchard and McCracken1991); eastern Serbia (Uroševič and Sudar, Reference Uroševič and Sudar1991); central Slovenia (Ramovš and Goričan, Reference Ramovš and Goričan1995); western Carpathians, Slovakia (Pevný and Salaj, Reference Pevný and Salaj1997); ?subsurface Polish lowlands (Narkiewicz, Reference Narkiewicz1999); Table Mountain Formation, Sylvester Allochthon, northern B.C. (Orchard, Reference Orchard, Colpron and Nelson2006); Botneheia Formation, Milne Edwardsfjellet, Svalbard (Nakrem et al., Reference Nakrem, Orchard, Weitschat, Hounslow, Beatty and Mørk2008), and the Transdanubian Range, Hungary (Karádi et al., Reference Karádi, Budai, Haas, Vörös, Piros, Dunkl and Tóth2022). In China, a probable juvenile growth stage of this species was illustrated by Sun et al. (Reference Sun, Jiang, Ji and Hao2016) from the Upper Guanling Formation in Guizhou Province, but other reports are suspect (see synonymy).

Description

A relatively elongate, arched segminiplanate P₁ element with a comparatively long and slender subrectangular platform with subparallel platform margins and a typical length:breadth ratio of ~5–5.5:1. Submature elements show the characteristic biconvex anterior platform and constricted posterior that disappears during growth as the posterior platform broadens. A prominent, slightly to strongly reclined and terminal denticle (“horn”) is characteristic of mature elements but this may be overtaken by the cusp in the largest specimens, wherein a narrow platform brim may be developed on the broad, rounded posterior margin. During growth, the anterior blade denticles become slightly elevated relative to those of the median carina, which may be wholly fused. The lowest part of the carina is at element midlength, beyond which the posterior fused carina is often relatively elevated in front of the large posterior denticles. On the underside, the pit is anteriorly shifted within the basal loop, which extends a short distance to the posterior with a rounded or subquadrate outline.

Comparisons

Neogondolella cornuta was regarded as a later growth stage and therefore a synonym of N. constricta by Nicora and Kovács (Reference Nicora and Kovács1984), but in the present work the taxon is interpreted as its direct descendant. Small specimens of the two species may be indistinguishable, but the relatively long platform and longer posterior keel characteristic of N. cornuta become more common features in the constricta group populations through the upper Anisian. Posterior platform configuration may appear similar in the two species, but greater weight is given here to the relative platform length and outline.

Neogondolella aldae differs from N. cornuta in outline, with the relatively broad anterior and reduced posterior platform of the former contrasting with the sub-parallel margins of the latter. The platform of N. posterolonga is like that of early growth stages of N. cornuta and may have arisen through neotony. According to Kovács (Reference Kovács1994), the most diagnostic feature for distinguishing N. cornuta from its successor species N. postcornuta is the penultimate denticle of the carina in specimens of moderate size: this is a conspicuous cusp in the former, ancestral species but is not evident in N. postcornuta, which also has the terminal denticle fused to the platform.

Remarks

Gondolella pseudolonga, which is based on a submature growth stage from Italy (Kovács et al., Reference Kovács, Kozur and Mietto1980), has denticulation characteristic of relatively early growth stages of Neogondolella ex gr. constricta. The character of adult N. pseudolonga is conjectural, but the platform length to breadth ratio of its holotype is 5:1, which corresponds to that of both N. cornuta and N. posterolonga. Neogondolella pseudolonga may be a junior synonym of N. cornuta, although Kozur et al. (Reference Kozur, Krainer and Mostler1994b, p. 174) regarded it as an earlier growth stage and synonym of N. longa from the excentrica Zone of western Bulgaria. However, the holotype of N. longa has a length: breadth ratio of ~6:1 and is narrower than the present species, although its variability is unknown.

Neogondolella cornuta ladinica Kozur et al., Reference Kozur, Krainer and Mostler1994b, is based on elements with an extended basal scar on an upturned basal posterior margin, a feature that becomes more common through the range of the species in Austria, finally replacing the nominate subspecies. Differentiation of such forms has not proven possible in the North American material.

Remarks

These elements resemble Neogondolella cornuta in all respects other than they have a posteriormost denticle that is offset from the main carinal axis. They are distinctly longer than Neogondolella excentrica primitiva n. subsp., but show a similar, generally rudimentary posterior elaboration. In N. e. excentrica, a posterior secondary process and keel are developed, which are absent in N. aff. N. cornuta. About 85 elements with this morphology occur, always with N. cornuta; this is a little more common than N. e. excentrica.

Holotype

Bu 1707/1, from lα = excentrica conodont-Zone 1, Golo–Bârdo mountains south of Pernik, Bulgaria. Budurov's collections (Bu) are in the Geologisches Institut, Sofia, Bulgaria.

Description

The P₁ elements assembled here are elongate segminiplanate conodonts in which a posterior platform constriction occurs in front of a variably inturned posterior platform that may bear up to 3 additional nodes posterior of the inconspicuous cusp. On the underside, one side of an asymmetrical basal scar extends as a posterior keel. In adult specimens, the anterior blade and posterior carina denticles are higher and more discrete than those of the more fused median carina.

Remarks

Many populations of the Neogondolella constricta group from Nevada and B.C. include posteriorly asymmetric elements with a variably inturned posterior platform process above a posterior keel. In a few specimens, there is no deflection of the posterior platform, although an extended keel is well developed, implying transition from other members of the constricta group. Most of these forms previously have been assigned to either N. excentrica or to N. transita. Kozur et al. (Reference Kozur, Krainer and Mostler1994b) regarded these two species as synonyms, with the subsymmetrical platform of N. transita being connected with the strongly curved platforms of N. excentrica through a morphological continuum. Budurov and Stefanov (Reference Budurov and Stefanov1972) included a variety of platform symmetries in their species, but none resembles the holotype of N. transita (platform ratio 4:1; straight axis; large, triangular cusp; upturned, pointed posterior platform tip; aff. N. suhodolica Budurov and Stefanov, Reference Budurov and Stefanov1973). As noted by Kovács (Reference Kovács1984), a comparative study of populations of each taxon in their respective type areas is needed to resolve the scope of each. In this work, we subdivide N. excentrica into three subspecies, two of which are new. These differ in their relative platform dimensions and posterior platform configuration. Records of N. transita in Hungary and Italy are shown as N. ex gr. excentrica in Figure 5.

Neogondolella excentrica excentrica Budurov and Stepanov, Reference Budurov and Stefanov1972
 Figure 10.1?, 10.2–10.7

Reference Mosher and Clark1965

Polygnathus tethydis Huckriede; Mosher and Clark, p. 563, pl. 66, fig. 13.

*  Reference Budurov and Stefanov1972

Neogondolella excentrica Budurov and Stefanov, p. 840–841, pl. 4, figs. 9–28.

p  Reference Trammer1975

Gondolella excentrica (Budurov and Stefanov); Trammer, pl. 25, figs. 4a–c (only).

p  Reference Zawidzka1975

Gondolella excentrica; Zawidzka, pl. 42, figs. 5a, b (only).

Reference Zawidzka1975

Gondolella navicula Huckriede; Zawidzka, pl. 40, figs. 4a, b.

Reference Rafek1976

Neogondolella transita Kozur and Mostler; Rafek, pl. IV, figs. 31–33.

Reference Nicora1976

Neogondolella excentrica; Nicora, p. 639–640, pl. 84, figs. 3–5.

Reference Mietto and Petroni1979

Neogondolella excentrica; Mietto and Petroni, p. 9, pl. I, figs. 5a–c.

Reference Mietto and Petroni1981

Gondolella transita; Mietto and Petroni, p. 555–556, pl. 57, figs. 9a–c.

?  Reference Mietto and Petroni1981

Gondolella basisymmetrica huckriedei (Budurov and Stefanov); Mietto and Petroni, p. 555–556, pl. 57, figs. 2a, b.

Reference Kolar-Jurkovšek1983

Neogondolella excentrica; Kolar-Jurkovšek, p. 341–342, pl. 12, figs. 1a–d.

Reference Vrielynck1984

Gondolella lindstroemi (Budurov and Stefanov); Vrielynck, p. 192–193, pl. 3, figs. 5a–c.

?  Reference Uroševič and Sudar1991

Gondolella transita (Kozur and Mostler, Reference Kozur and Mostler1971); Uroševič and Sudar, pl. 1, fig. 16.

Reference Márquez-Aliaga, Valenzuela-Rios, Calvet and Budurov2000

Neogondolella excentrica; Márquez-Aliaga et al., fig. 6.11.

Figure 10.

(1) Neogondolella excentrica aff. excentrica Budurov and Stefanov from Prida Formation, Fossil Hill, Nevada. GSC 141873, sample FH13, Rotelliformis Zone/clarkei Subzone. (2–7) Neogondolella excentrica excentrica Budurov and Stefanov from (2) Llama Formation, B.C. and (3–7) Prida Formation, Saurian Hill, Nevada. (2) GSC 141874, sample WAP-B17, Cirque B, Ganoid Ridge, Chischa Zone; (3–5) GSC 141875, sample SH529; (6, 7) GSC 141876, sample SH529, both Occidentalis Zone. (8–15) Neogondolella excentrica sigmoidalis n. subsp. from (8–12) sample SH512, Prida Formation, Nevada, Subasperum Zone, and (13–15) Llama Formation, B.C. (8–10) GSC 141877, holotype; (11, 12) GSC 141878; (13–15) GSC 141879, sample 97/WapA6, Cirque B, Ganoid Ridge, Wapiti Lake area, bracketed between Poseidon and Meginae zones. Scale bar = 200 μm.

Diagnosis

As for species plus this slender morphotype has a length:breadth ratio in adult specimens of ≥5:1; commonly subparallel platform margins; an asymmetric posterior margin bearing a tapered, often pointed and offset posterior platform; and a slightly to strongly extended postero-lateral basal keel. The posterior platform is typically inturned and may bear several denticles forming a secondary carina.

Occurrence

About 60 specimens of Neogondolella excentrica excentrica occur in collections from the Occidentalis and lower Subasperum zones in Nevada; the posterior fragment illustrated by Mosher and Clark (Reference Mosher and Clark1965) came from the Protrachyceras Beds. Nicora (Reference Nicora1976) also illustrated the species from Fossil Hill. Rare specimens of N. e. aff. excentrica occur in the Rotelliformis Zone. The subspecies occurs also in Chischa Zone on Ganoid Ridge, and as high as the bamberi Fauna along the Alaska Highway in B.C.

In Europe, Kozur et al. (Reference Kozur, Krainer and Mostler1994b) noted specimens like these occurred in the Nevadites Zone and lower Curionii Zone. Examples of this subspecies are reported from southwestern Holy Cross Mountains, Poland (Trammer, Reference Trammer1975); Upper Muschelkalk, northern Germany (Rafek, Reference Rafek1976); ?San Ulderico section, NE Italy (Mietto and Petroni, Reference Mietto and Petroni1979); Campogrosso section, NE Italy (Mietto and Petroni, Reference Mietto and Petroni1981); Idrske Krnice, Slovenia (Kolar-Jurkovšek, Reference Kolar-Jurkovšek1983); Carnic Alps-Dolomites, NE Italy (Vrielynck, Reference Vrielynck1984); Eastern Serbia (Uroševič and Sudar, Reference Uroševič and Sudar1991); NE Spain (Márquez-Aliaga et al., Reference Márquez-Aliaga, Valenzuela-Rios, Calvet and Budurov2000).

Remarks

This subspecies corresponds to the holotype of the species and other elements originally illustrated by Budurov and Stefanov (Reference Budurov and Stefanov1972), most of which have an inturned posterior platform. The posterior platform is variably developed in Nevadan material, but the presence of a distinct secondary lobe or process is regarded as diagnostic. Although typically offset posteriorly, relatively straight elements occur rarely in the Nevadan material. Mature specimens of N. e. excentrica have long rectangular platforms comparable to that of associated N. cornuta. Two much smaller elements with a strongly deflected posterior platform (Fig. 10.1) occur earlier than typical specimens and are given an aff. designation.

Neogondolella excentrica primitiva new subspecies
 Figure 11.1–11.11, 11.12?–11.14?

?  Reference Rafek1976

Neogondolella prava (Kozur); Rafek, pl. IV, figs. 28, 29.

Reference Nicora1976

Neogondolella excentrica; Nicora, p. 639–640, pl. 84, figs. 1a, b, 2a, b.

?  Reference Kozur and Mirăuța1980

Gondolella transita Kozur and Mirăuța, taf. 1, figs. 1a–d.

?  Reference Bagnoli1982

Gondolella cf. transita Kozur and Mostler; Bagnoli, p. 7, pl. 1, figs. 4a, b.

Reference Uroševič and Sudar1991

Neogondolella excentrica Budurov and Stepanov; Uroševič and Sudar, pl. 1, fig. 15.

?  Reference Kozur, Krainer and Mostler1994b

Neogondolella sp. aff. N. transita (Kozur and Mostler); Kozur et al., pl. 3, figs. 21, 24.

p  Reference Wu, Yao, Ji and Wang2008

Neogondolella constricta (Mosher and Clark); Wu et al., pl. II, figs. 12, 15.

p?  Reference Wu, Yao, Ji and Wang2008

Neogondolella mombergensis (Mosher and Clark) (sic); Wu et al., pl. I, fig. 13.

p?  Reference Golding2014

Neogondolella ex gr. constricta, morphotype epsilon; Golding, fig. 2.26, parts 1–3.

Reference Sun, Jiang, Sun and Hao2014

Neogondolella excentrica; Sun et al., fig. 4e.

Figure 11.

(1–11) Neogondolella excentrica primitiva n. subsp. (1–7, 9–11), from Prida Formation, Nevada. (1, 2) GSC 141880, sample FH8, Rotelliformis Zone/ burckhardti Subzone; (3, 4) GSC 141881, sample FH44, Meeki Zone/dunni Subzone; (5–7) GSC 141882, holotype, FH6, Rotelliformis Zone/burckhardti Subzone; (8) GSC 141883, sample 92AH-2, Deleeni Zone; (9–11) GSC 141884, sample FH48, Occidentalis Zone; (12–14) Neogondolella excentrica aff. primitiva n. subsp. GSC 141885, sample AH26, Minor Zone. Scale bar = 200 μm.

Holotype

GSC 141882 (Fig. 11.5–11.7), from the Fossil Hill Member of the Prida Formation, burckhardti Subzone of the Rotelliformis Zone (sample FH-6), Fossil Hill, Nevada.

Diagnosis

This is a relatively short subspecies of N. excentrica with a length: breadth 4.5: 1; a short, obliquely directed, posteriorly rounded postero-lateral platform lobe; a relatively low blade/carina in front of the 1 or 2 larger offset terminal denticles; and a weakly extended, asymmetrical basal scar posterior of the pit on the lower side.

Occurrence

In Nevada, about 35 elements are assigned to Neogondolella excentrica primitiva n. subsp., mostly from the Rotelliformis Zone, with rare elements occurring in the Meeki Zone at Fossil Hill and in the Tobin Range. Nicora (Reference Nicora1976) also illustrated the subspecies from the Star Canyon section in Nevada. In B.C., the subspecies occurs in the Deleeni Zone at Yellow Bluffs. A specimen assigned to N. e. aff. primitiva n. subsp. occurs in the middle Anisian Minor Zone along the Alaska Highway (Fig. 11.12–11.14).

Kozur et al. (Reference Kozur, Krainer and Mostler1994b) noted forms similar to this subspecies occurring in the Reitziites reitzi Zone in Austria, and additional records may occur in the Upper Muschelkalk of northern Germany (Rafek, Reference Rafek1976); in the northern Apuseni Mountains in Romania; in Punta Bianca, NW Italy (Bagnoli, Reference Bagnoli1982); eastern Serbia (Uroševič and Sudar, Reference Uroševič and Sudar1991); in the Qingyan Formation in Guizhou Province (Wu et al., Reference Wu, Yao, Ji and Wang2008); and the Upper Guanling Formation in Guizhou Province, China (Sun et al., Reference Sun, Jiang, Sun and Hao2014).

Etymology

From Latin prīmitīvus, signifying the first representative of the species.

Comparisons

These elements share relative platform dimensions with Neogondolella constricta but differ from that species in their asymmetric posterior platforms. They differ from the nominate subspecies in its comparatively shorter length, lower posterior carina height, shorter basal scar, and the rounded posterior platform lobe. The latter margin is like that of N. e. sigmoidalis n. subsp., but both the entire element and the posterior basal keel of Neogondolella excentrica primitiva n. subsp. is much shorter. Kozur et al. (Reference Kozur, Krainer and Mostler1994b) assigned similar forms to N. sp. aff. transita, although only a posterior fragment was illustrated. An older specimen assigned to N. e. aff. primitiva n. subsp. differs from these elements in possessing a slightly higher blade–carina.

Remarks

The asymmetric posterior platform of Neogondolella excentrica primitiva n. subsp. is generally manifest as a deflected rounded lobe that may appear as a secondary process, as in the holotype, or simply as a posterior-lateral expansion with an incurved posterior carina. Commonly there is an additional offset posteriormost denticle like that seen in younger and longer elements assigned to Neogondolella aff. N. cornuta.

Holotype

GSC 141877 (Fig. 10.8–10.10), from the upper member of the Prida Formation, upper Subasperum Zone (sample 512), Saurian Hill, Nevada.

Diagnosis

A subspecies of Neogondolella excentrica with an elongate, weakly arched platform of largely uniform width and a length:breadth ratio of 5.5:1. In upper view, the posteriormost one-fifth of the platform has a sigmoidal outline, is inturned, and terminates in a rounded posterior margin. A low blade/carina has relatively discrete anterior denticles, partly fused medial carina, and several large and discrete denticles posterior of the cusp. The basal scar extends beneath the posterior inturned platform far posterior of the pit.

Occurrence

This relatively uncommon subspecies is represented by about 12 elements from the Subasperum Zone at Saurian Hill in Nevada, and the Matutinum and Poseidon zones at Wapiti Lake, Oyster Springs, and Tuchodi Bluffs in B.C. Specimens from the Chischa Zone have less-developed keels, whereas typical specimens occur with ammonoids of the bamberi Fauna.

Other records are those of the ?Upper Muschelkalk of northern Germany (Rafek, Reference Rafek1976); the Curionii Zone at Felsőörs in Hungary (Kovács, Reference Kovács1994) and in the Belvedere section in the Italian Dolomites (Muttoni et al., Reference Muttoni, Nicora, Brack and Kent2004); the Carnic Alps in Austria (Kozur et al., Reference Kozur, Krainer and Lutz1994a); and the Kalapani Limestone, northern India (Chhabra and Kumar, Reference Chhabra and Kumar1992).

Etymology

From the Greek word sigmoeidḗs, shaped like the letter sigma.

Remarks

The long, slender platform and extended keel differentiates Neogondolella e. sigmoidalis n. subsp. from Neogondolella e. primitiva n. subsp., while the rounded posterior platform distinguishes it from N. e. excentrica. The specimen illustrated by Kozur et al. (Reference Kozur, Krainer and Lutz1994a) differs from typical N. aequidentata Kozur, Krainer, and Lutz, Reference Kozur, Krainer and Lutz1994a, in its asymmetry and lower, less arched carina.

In lower Ladinian samples from B.C., elements of Neogondolella posterolonga occur in which posterior platform asymmetry occurs without an extended keel on the underside. These appear to be transitional to Neogondolella e. sigmoidalis n. subsp., which shares the same relative platform dimensions. This mirrors the intermediate nature of N. aff. N. cornuta between N. cornuta and N. e. excentrica, and the variants included in N. e. primitiva n. subsp.

Neogondolella ex gr. mesotriassica (Kozur and Mostler, Reference Kozur and Mostler1982)
Figure 12

p* Reference Kozur and Mostler1982

Gondolella mesotriassica Kozur and Mostler, p. 293–294, pl. 1, figs. ?2, 3, 4.

p Reference Kozur, Krainer and Mostler1994b

Neogondolella mesotriassica; Kozur et al., pl. 4, figs. 7–9 (only).

Reference Budurov and Stefanov2014

Neogonolella constricta longa Budurov and Stafanov; Sun et al., fig. 4d.

Figure 12.

Neogondolella ex gr. mesotriassica (Kozur and Mostler) from (1–3) Toad Formation, B.C. and (4–9) Prida Formation, Nevada. (1–3) GSC 141898, sample 83/215F, Toad River Canyon, Minor–Deleeni zonal boundary; (4–6) GSC 141899, sample FH13, Rotelliformis Zone/clarkei Subzone; (7–9) GSC 141900, sample FH48, Occidentalis Zone/hyatti Subzone. Scale bar = 200 μm.

Holotype

Kozur and Mostler, pl. 1, fig. 4, sample BM 44/75, from within the Reiflingen limestones at Buchberg, near Göstling, lower Austria.

Occurrence

Slightly different specimens assigned to this group occur sporadically through the upper Anisian in Nevada, largely in the Rotelliformis Zone, but also in younger strata. In B.C., the occurrence of this species is bracketed by the Minor and Deleeni zones on Toad River. The characteristic form starts at the base of the Reitzi Zone in Austria (Kozur et al., Reference Kozur, Krainer and Mostler1994b). A similar element was illustrated from the Upper Guanling Formation in Guizhou Province, China (Sun et al., Reference Sun, Jiang, Sun and Hao2014).

Description

The segminiplanate P₁ elements are arched, with a subrectangular platform with a squared-off or truncated posterior margin that may have a v-shaped notch visible in upper view. The platform is slightly broader at midlength and tapers in both directions before often broadening at the posterior end. The blade denticles are more discrete than the largely fused median carina, and together they impart an arcuate upper profile to the element. The carina ends in a small terminal denticle that is separated from those to the anterior, including the cusp. Additional denticles commonly are developed on the posterior platform margin. The basal pit is subterminal in the keel, with an asymmetrical basal loop that reflects the lateral expansion of the posterior end.

Remarks

The holotype of Neogondolella mesotriassica has a v-shaped notch on the posterior margin, but it has neither a truncated posterior platform, nor accessory posterior denticles, as cited in the original diagnosis. The terminal denticle of the holotype is discrete and quite unlike those of the two paratypes, which have the final denticle fused with the posterior platform margin. The latter feature led Kovács et al. (Reference Kovács, Nicora, Szabo and Balini1990, p. 171) to suggest that one paratype was an example of the gamma morphotype of N. constricta, later named N. postcornuta (Kovács, Reference Kovács1994). On the contrary, Kozur et al. (Reference Kozur, Krainer and Mostler1994b) regarded most examples of the latter species as examples of N. mesotriassica, but did not provide a formal taxonomic revision. Both species are retained here based on the divergent morphology of the holotypes.

These elements resemble Paragondolella bifurcata, which is also characterized by a squared-off posterior margin and accessory denticles, but that species differs in its high carina and blade. Neogondolella quasiconstricta n. sp. has similar features but is much broader posteriorly.

Uncommon examples of Neogondolella mesotriassica from both Nevada and B.C. differ in detail but are united by the narrow, truncated, and denticulate character of the posterior margin. The morphology of the holotype and the diversity in the current material suggest further revision will be necessary within this group. The three examples illustrated are from progressively younger horizons and appear to lengthen and show anterior pit migration, which is the same trend recognized in other members of the constricta group.

Neogondolella postcornuta (Kovács, Reference Kovács1994)
 Figure 13

Reference Trammer1975

Gondolella longa; Trammer, pl. 23, figs. 2a, b, 3a, b.

p Reference Kovács, Nicora, Szabo and Balini1990

Gondolella constricta morphotype γ Kovács et al., p. 188, pl. 1, fig. 4a–c; pl. 3, fig. 5.

* Reference Kovács1994

Gondolella constricta postcornuta Kovács, p. 484, pl. 1, figs. 4, 5; pl. 2, figs. 2–5; pl. 6, fig. 4 (see synonymy).

Figure 13.

Neogondolella postcornuta (Kovács) from Prida Formation, Fossil Hill, Nevada. (1, 2) GSC 141901, sample FH35, Meeki Zone/nevadanus Subzone; (3, 4) GSC 141902, sample FH47, Meeki–Occidentalis zonal boundary; (5–7) GSC 141903, sample FH8, Rotelliformis Zone/burckhardti Subzone; (8–10) GSC 141904, sample FH50, Occidentalis Zone. Scale bar = 200 μm.

Holotype

Kovács, p. 484, pl. 1, fig. 4a–d, catalogue number T-6451, sample No. 4, Öskü road-side section, Balaton Highland, Hungary, reposited in the Museum of the Hungarian Geological Survey (Kovács et al., Reference Kovács, Nicora, Szabo and Balini1990, fig. 10). Lower Curionii Zone, Vászoly Limestone Member of the Buchenstein Formation.

Occurrence

Similar specimens occur throughout upper Anisian and into Ladinian strata at Fossil Hill, Nevada, but their occurrence is sporadic because they are not always distinguishable from some growth stages of N. cornuta. The species has not been recognized in B.C.

In the Balaton Highland, Hungary, Neogondolella postcornuta occurs from the uppermost part of the meriani B horizon to the lower Curionii Zone. The first primitive representatives of the species occur in the uppermost bed of the meriani B horizon, whereas common typical ones occur in the liepoldti horizon (Kovács, Reference Kovács1994). A very similar element to the largest illustrated here was figured from southwestern Holy Cross Mountains, Poland (Trammer, Reference Trammer1975). Similar transitional forms occur in the upper Curionii Zone in the Vászoly and Felsőörs sections (Kovács, Reference Kovács1994).

Description

The segminiplanate P₁ element is relatively narrow, elongate, and slightly arched with a posteriorly inclined main denticle completely fused with the posterior platform end. The platform is narrowly biconvex for the most part but narrows in its posterior part. The cusp is inconspicuous in the low carina that passes anteriorly into higher and less fused denticles. The small basal pit is subterminal in position.

Comparisons

According to Kovács (Reference Kovács1994), Neogondolella postcornuta differs from intermediate-sized elements of its forebear Neogondolella cornuta by the lack of a conspicuous cusp anterior of the last denticle of the carina, and by the complete fusion of the terminal denticle to the platform end. However, Kovács (Reference Kovács1994) stressed that these distinguishing characteristics applied chiefly to elements of intermediate size, and both early juvenile and hyperadult growth stages were less distinctive. The Nevadan specimens of this species do not exhibit the subparallel lateral platform margins and truncated posterior margin seen in N. cornuta.

As discussed by Kovács (Reference Kovács1994), N. postcornuta gives rise to N. bakalovi, a Balkan species characterized by a strongly extended posterior platform. Although rare specimens from Nevada show a similar feature, typical elements of the latter species are not recorded in North America. Specimens of N. posterolonga have a similar platform shape but are narrower, longer, and lack the fused posterior denticle.

Remarks

Elements exhibiting the characteristic posterior configuration of Neogondolella postcornuta occur sporadically through the range of N. cornuta, and it is difficult to separate many smaller specimens in which a terminal denticle is located on the platform margin. Chen et al. (Reference Chen, Krystyn, Orchard, Lai and Richoz2015) provided a broad review of Middle Triassic conodonts and suggested N. postcornuta was in fact a junior synonym of N. pseudolonga. As discussed previously, N. pseudolonga is here regarded as a probable synonym of N. cornuta or N. posterolonga. This emphasizes the difficulty of interpreting different growth stages of these allied species.

Neogondolella posterolonga Kozur, Krainer, and Mostler, Reference Kozur, Krainer and Mostler1994b
Figure 14

p Reference Rafek1976

Neogondolella navicula (Huckriede); Rafek, Pl. III, fig. 7 (only).

? Reference Bagnoli1982

Gondolella pseudolonga Kovács, Nicora, and Mietto; Bagnoli, p. 6, pl. 1, figs. 5a, b.

p Reference Nicora and Kovács1984

Gondolella constricta Mosher and Clark; Nicora and Kovács, p. 144–148, pl. 8, fig. 3.

p Reference Nicora and Kovács1984

Gondolella mombergensis mombergensis Huckriede; Nicora and Kovács, p. 149–150, pl. 9, figs. ?7, ?10.

p Reference Nicora and Kovács1984

Gondolella mombergensis longa (Budurov and Stefanov); Nicora and Kovács, p. 150, pl. 10, figs. 6, 7.

* Reference Kozur, Krainer and Mostler1994b

Neogondolella aldae posterolonga Kozur et al., p. 182–183.

Reference Ramovš and Goričan1995

Neogondolella constricta; Ramovš and Goričan, pl. 9, fig. 3a, b.

Reference Pevný and Salaj1997

Gondolella constricta; Pevný and Salaj, pl. XI, figs. 2, 3.

? Reference Orchard, Cordey, Rui, Bamber, Mamet, Struik, Sano and Taylor2001

Neogondolella ex gr. constricta; Orchard et al., pl. 1, fig. 18.

? Reference Sano and Orchard2004

Neogondolella ex gr. constricta; Sano and Orchard, fig. 6.13, 6.14.

p Reference Golding2014

Neogondolella transita morphotype beta; Golding, fig. 2.29, parts 4–6.

p Reference Golding and Orchard2021

Neogondolella ex gr. constricta; Golding and Orchard, p. 29, pl. 15, figs. 1–4.

Figure 14.

Neogondolella posterolonga Kozur, Krainer and Mostler from (1–11) Prida Formation, Nevada, Subasperum Zone, and (12–21) from B.C. (1–3) GSC 141890; (4–6) GSC 141891, both from sample SH520; (7, 8) GSC 141892; (9–11) GSC 141893, both from Subasperum Zone SH534; (12–14) GSC 141894, sample 97/WapA6, Llama Formation, Poseidon? Zone; (15, 16) GSC 141895, sample 82/AH1, Liard Formation, near milepost 386, Alaska Highway, 2 m below Meginae Zone; (17–19) GSC 141896, from sample GSC cur. No. O-83862, Llama Formation, Wapiti Lake, Matutinum Zone; (20, 21) GSC 141897, sample 01/URC9, Toad Formation, Ursula Creek, Williston Lake, 95 m above the Permian–Triassic Boundary. Scale bar = 200 μm.

Holotype

The specimen figured by Nicora and Kovács (Reference Nicora and Kovács1984, pl. 10, fig. 7) as N. mombergensis longa, from sample N50, in the middle Prida Formation, lower Occidentalis Zone age at Fossil Hill, Humboldt Range, Nevada.

Occurrence

Typical representatives of Neogondolella posterolonga appear in the Meeki Zone and form a larger component of the faunas in the Occidentalis and Subasperum zones in Nevada. Kozur et al. (Reference Kozur, Krainer and Mostler1994b) recorded first occurrences in the upper Meeki Zone, but some early growth stages of constricta group elements from the Rotelliformis Zone are similar. Elsewhere in Nevada, the species is known from Wheeler Mine, and in the Tobin Range. In B.C., examples occur in the Chischa Zone at Yellow Bluff; the Matutinum Zone at Wapiti Lake; the Poseidon Zone at Ganoid Ridge and Tuchodi Bluff; and the Meginae Zone along the Alaska Highway at milepost 386 (Table 2).

The specimen from Hungary comes from the Curionii Zone (Kovács, Reference Kovács1994). The species is also recorded in the southwestern Holy Cross Mountains, Poland (Trammer, Reference Trammer1975); Upper Muschelkalk, northern Germany (Rafek, Reference Rafek1976); ?Punta Bianca, NW Italy (Bagnoli, Reference Bagnoli1982); central Slovenia (Ramovš and Goričan, Reference Ramovš and Goričan1995); western Carpathians, Slovakia (Pevný and Salaj, Reference Pevný and Salaj1997); and ?Necoslie Breccia, Cache Creek terrane, Nechako (Orchard et al., Reference Orchard, Cordey, Rui, Bamber, Mamet, Struik, Sano and Taylor2001; Sano and Orchard, Reference Sano and Orchard2004; Golding and Orchard, Reference Golding and Orchard2021).

Description

The platform of this arched segminiplanate element is slender, widest anterior of unit mid-length, and relatively long and narrow for the posterior one-fourth to one-third of its length. Typical length:breadth ratios are 5.0–5.5:1. The blade carina is like other constricta group taxa, with more discrete anterior denticles, a lower, largely fused medial carina in later growth stages, and higher, more discrete posterior denticles, including an often-enlarged cusp and a prominent weakly to strongly inclined terminal denticle around which a very narrow platform brim may be developed. On the underside, there is a short, often irregular posterior extension of the basal loop.

Remarks

Kozur et al. (Reference Kozur, Krainer and Mostler1994b) originally included Neogondolella posterolonga as a subspecies of N. aldae, but it is here elevated to a species. In contrast to the latter, N. posterolonga has a much longer and narrower, less biconvex platform. In N. constricta and N. cornuta, the constriction disappears during later growth whereas in the present species the posterior platform is noticeably narrower. In this sense, the species is considered to retain a juvenile morphology in adult specimens.

Neogondolella posterolonga has been confused with N. longa, but that species is longer, narrower (length:breadth ratio ~6:1), and of more uniform width and with more discrete denticles. Neogondolella pseudolonga was regarded as an early growth stage of N. longa by Kozur et al. (Reference Kozur, Krainer and Mostler1994b) but it may be related to the present species or to N. cornuta. Both N. postcornuta and N. bakalovi may have a similar platform shape, but they have a terminal denticle fused with the platform margin.

Holotype

GSC 141906 (Fig. 15.4–15.6), from the Fossil Hill Member of the Prida Formation, vogdesi Subzone of the Rotelliformis Zone (sample FH17), Fossil Hill, Nevada.

Figure 15.

(1–7) Neogondolella quasiconstricta n. sp. from Prida Formation, Nevada. (1–3) GSC 141905, sample FH6, Rotelliformis Zone/burckhardti Subzone; (4–6) GSC 141906, holotype, sample FH17, Rotelliformis Zone/vogdesi Subzone; (7) GSC 141907, sample FH24, Rotelliformis Zone/cricki Subzone. (8–15) Neogondolella quasicornuta n. sp. from Prida Formation, Nevada and (9, 10) Liard Formation, B.C. (8) GSC 141908, sample SH524, Occidentalis Zone; (9, 10) GSC 141909, sample 92-AH-3, Alaska Highway, bamberi Fauna; (11, 12) GSC 141910, sample SH504, Subasperum Zone; (13–15) GSC 141911, holotype, sample SH519, Occidentalis Zone. Scale bar = 200 μm.

Diagnosis

The platform of the comparatively large P₁ element has a length:breadth ratio of 3.5–4:1, an expanded postero-lateral margin, and commonly accessory denticles on one or both sides of the prominent terminal denticle. The blade carina is low throughout, with more discrete blade denticles passing into a largely fused median carina. The pit is subterminal within the triangular keel.

Occurrence

Neogondolella quasiconstricta n. sp. occurs principally in the Rotelliformis Zone in Nevada, always in association with, but as a minor component of abundant Neogondolella constricta collections. Younger occurrences are far less common. Elsewhere in Nevada, the species is known from the Tobin Range, and in Canada it occurs in the late Anisian of North Tetsa Phosphate section on the Alaska Highway. About 25 elements were assigned to this species.

The species is also recognized from the base of the Reitzi Zone in Felsőörs, Hungary (Szabo et al., Reference Szabo, Kovács, Lelkes and Oravecz-Scheffer1980; Kovács, Reference Kovács and Vörös2003); Han–Bulog Limestone, SE Bosnia (Sudar, Reference Sudar1982); ?middle Bivera Formation, southern Alps, northern Italy (Farabegoli et al., Reference Farabegoli, Levanti, Perri and Veneri1984), and southern Alps (Pisa et al., Reference Pisa, Perri and Veneri1980). Some elements from the the Qingyan Formation in Guizhou Province, China (Wu et al., Reference Wu, Yao, Ji and Wang2008) could be representatives of this species, although crucial lateral views were not provided.

Description

The large segminiplanate P₁ element has a platform with a length:breadth ratio of 3.5–4:1 and an expanded posterior end with a subtriangular outline; the median platform is generally narrower but may be biconvex. The blade–carina is low throughout, with 4–5 discrete anterior blade denticles rising above the downturned anterior platform, and a largely fused median carina. The terminal posterior denticle is prominent and is commonly accompanied by discrete accessory denticles on one or both sides of it. The pit is subterminal within the basal scar that expands progressively to the posterior where the triangular outline mirrors that of the platform.

Etymology

Combination of the Latin quasi- as if, and constricta, the often-co-occurring species that shares the same platform length:breadth ratio.

Comparisons

Neogondolella quasiconstricta n. sp. is similar to, and has often been combined with Paragondolella bifurcata, which differs in having a distinctly high blade and carina, as is characteristic of the genus and the associated Pelsonian species, P. bulgarica and P. hanbulogi. Gondolella fueloepi pseudobifurcata Kovács, Reference Kovács1994, from the costatus horizon of the Reitzi Zone resembles Neogondolella quasiconstricta n. sp. but it has a distinctive posterior platform brim.

Remarks

Typical specimens of P. bifurcata illustrating the diagnostic blade–carina feature were figured by Budurov (Reference Budurov1980) and Qin et al. (Reference Qin, Golding, Jiang, Chen, Zhang, Kang, Wang and Yuan2021). On the contrary, many specimens assigned to P. bifurcata have a low carina (see synonymy), and although this may result from allometric growth (Chen et al., Reference Chen, Neubauer, Krystyn and Richoz2016), there are no high-bladed elements with this platform configuration in the Nevadan material.

Muschelkalk neogondolellins with a single secondary carina developed as nodes posterior of the cusp were assigned to Gondolella mombergensis prava by Kozur (Reference Kozur1968a, Reference Kozurb), and that designation was adopted by Szabo et al. (Reference Szabo, Kovács, Lelkes and Oravecz-Scheffer1980). Subsequently these forms have generally been assigned to Paragondolella bifurcata (e.g., Kovács, Reference Kovács and Vörös2003), however the holotype of Neogondolella prava (Kozur, Reference Kozur1968a, pl. 1, fig. 2a, b) is clearly allied with its Germanic associate N. mombergensis, as shown by similar arched platforms, blade–carina configuration, and geniculate anterior platform (see refigured holotype in Orchard and Rieber, 1998, fig. 1). Rafek (Reference Rafek1976, fig. 4) illustrated a variety of N. prava morphotypes from the upper Muschelkalk, which correspond to neither P. bifurcata nor N. quasiconstricta n. sp.

Neogondolella quasiconstricta n. sp. often occurs with both Neogondolella constricta and N. excentrica primitiva n. subsp., with which it shares relative platform dimensions and a relatively low blade–carina. The three species differ in their posterior configurations, as do the Pelsonian triad of species mentioned above.

Holotype

GSC 141911 (Fig. 15.13–15.15), from the Fossil Hill Member of the Prida Formation, Occidentalis Zone (sample SH519), Saurian Hill, Nevada.

Diagnosis

The P₁ platform element is comparatively long and narrow with a length:breadth ratio of 5–6:1, and subparallel lateral margins that broaden near the posterior end. The blade–carina is low throughout, higher in the anterior, fused in the central part, and slightly elevated in the posterior one-third. A prominent terminal denticle is commonly flanked by accessory denticles. The pit is slightly shifted anterior in the keel, which extends beneath the postero-lateral platform expansion posterior of the pit.

Occurrence

Neogondolella quasicornuta n. sp. is uncommon from high in the Meeki Zone through the low Occidentalis Zone and into the Subasperum Zone at Fossil Hill, and in the latter zone at Wheeler Mine, Nevada. About 50 elements are represented within the abundant collections of Neogondolella cornuta from Nevada. The species is not yet identified in B.C.

In Europe, similar elements are known from ?San Ulderico section, NE Italy (Mietto and Petroni, Reference Mietto and Petroni1979) (no upper view); ?Balaton Highland, Hungary (Szabo et al., Reference Szabo, Kovács, Lelkes and Oravecz-Scheffer1980; Kovács, Reference Kovács and Vörös2003) (no lower view); and Idrske Krnice, Slovenia (Kolar-Jurkovšek, Reference Kolar-Jurkovšek1983).

Description

The elongate segminiplanate P₁ elements of this species have a length:breadth ratio of 5–6:1. The lateral platform margins are subparallel for much of the element length before broadening near the posterior end. The generally low blade–carina resembles that seen in Neogondolella cornuta with more discrete anterior blade denticles, a fused median carina, and slightly elevated, less-fused posterior carinal denticles. A very prominent terminal denticle is accompanied by one or more accessory denticles on the posterior margin. The pit is slightly shifted anterior in the keel, which extends beneath the postero-lateral platform expansion.

Etymology

Combination of the Latin quasi- as if, and cornuta, the species that co-occurs and shares its platform length:breadth ratio.

Remarks

Neogondolella quasicornuta n. sp. shares relative platform dimensions with both N. cornuta and N. excentrica excentrica, with which it often co-occurs. As in the much shorter N. quasiconstricta n. sp., the three associated species differ in their posterior platform configurations. The specimen from Balaton Highland is relatively longer than any others illustrated from Hungary, but no lower view is shown.