Novispathodus, Orchard, 2005

Leu, Marc, Bucher, Hugo, Vennemann, Torsten, Bagherpour, Borhan, Ji, Cheng, Brosse, Morgane & Goudemand, Nicolas, 2022, A Unitary Association-based conodont biozonation of the Smithian-Spathian boundary (Early Triassic) and associated biotic crisis from South China, Swiss Journal of Palaeontology (19) 141 (1), pp. 1-61 : 25-32

publication ID	https://doi.org/ 10.1186/s13358-022-00259-x
persistent identifier	https://treatment.plazi.org/id/038B1D1D-6471-FFAE-FCA1-EB1BD0D0FA07
treatment provided by	Felipe
scientific name	Novispathodus
status

Treatment

Genus NOVISPATHODUS Orchard, 2005

Type species and holotype. Neospathodus abruptus Orchard, 1995, pp. 118–119 , figs. 3.23–24.

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Fig. 16 View Fig Neogondolellinae , Novispathodinae and Mullerinae from Shanggang and Youping cascade. Magnification is ×80.The scale bar is 400 μm. All elements are considered to be P 1 elements if not specifically identified otherwise. A, F, M Discretella ? n. sp. C; A YC16,PIMUZ 39115; F YC16,

PIMUZ 39116; M SHA342C, PIMUZ 39117. B Discretella ? n. sp. B;YC16, PIMUZ 39113. C Discretella aff. discreta (MÜller);YC22,PIMUZ 39106. D, S Discretella discreta (MÜller) ; D YC16, PIMUZ 39110; S SHA304C, PIMUZ 39111. E Novispathodus ex gr. abruptus (Orchard) ;YC41, PIMUZ 39225. G–I, K,

L, N, O Discretella pseudodieneri n. sp.; G YC16, PIMUZ 39124; H YC25, PIMUZ 39125; I SHA304C,PIMUZ 39126; K YC22, PIMUZ 39127, L YC25, PIMUZ 39128; N SHA304C,PIMUZ 39129; O SHA343C,PIMUZ SQL54979. J Guangxidella bransoni (MÜller) ;YC16, PIMUZ 39140. P–R Neospathodus bevelledi n. sp.; P SHA 304C,PIMUZ 39178; Q SHA333C, PIMUZ 39179; R YC12, PIMUZ 39180

Type stratum and locality. Jabral Safra , Oman .

Remarks. Te genus Novispathodus was introduced as a new genus with a 15-element apparatus by Orchard (2005) and revised by Goudemand et al. (2012b) based on the swapping of the S 1 and S 2 positions. Additional to the type species ( Nv. abruptus ) and other species whose multi-element apparatus has been reconstructed (e.g. Nv. pingdingshanensis ( Goudemand et al., 2012b) , Nv. waageni (unpublished)), we tentatively assign also the following species to Novispathodus on the basis that they co-occur with S and M elements that are reminiscent of Novispathodus .

Novispathodus pingdingshanensis ( Zhao & Orchard, 2007)

Fig. 17D–F; M–P, R, S, U, X View Fig , AA, AB, AD, AF

*2007 Neospathodus pingdingshanensis n. sp.; Zhao & Orchard, Zhao et al., p.36, pl. 1, fig. 4A–C.

2012a Novispathodus pingdingshanensis (Zhao & Orchard) ; Goudemand & Orchard in Goudemand et al., p. 1030–1031, figs. 2B, F, G, I–J, M, P, Q, AD, 3T-U, 6.

2013 Neospathodus pingdingshanensis Zhao & Orchard ; Chen et al., p 825, fig. 3.10, 3.12.

2013 Neospathodus waageni subsp. nov. A; Metcalfe et al., p. 1144, figs. 9.1–9.5, 9.7, 9.8, 9.10.

2014 Novispathodus pingdingshanensis (Zhao & Orchard) ; Maekawa & Igo in Shigeta, p. 239–240, figs. 171.13–171.31.

2015 Novispathodus pingdingshanensis (Zhao & Orchard) ; Chen et al., p. 111, 112, figs. 7.1–7.4, 8.5, 8.6.

2016 Novispathodus ex. gr. pingdingshanensis (Zhao & Orchard) ; Komatsu et al., p. 69, figs. 5.4a–5.5c.

2016 Neospathodus robustus Koike ; Chen & Kolar-Jurkovšek in Chen et al., p. 93, fig. 9.5 (only).

2018 Novispathodus pingdingshanensis (Zhao & Orchard) ; Maekawa in Maekawa et al., p. 36–37, figs. 20.2–20.18, 21.1–21.13.

2019 Novispathodus pingdingshanensis (Zhao & Orchard) ; Chen et al., fig. 3, nr. 8.

2019 Novispathodus pingdingshanensis (Zhao & Orchard) ; Liu et al., p. 13, pl. 3 fig. 5 (only).

Material.> 100 specimens.

Diagnosis (Zhao and Orchard, in Zhao et al., 2007; emended by Goudemand, in Goudemand et al., 2012b). Small segminate P 1 elements characterized by a length/ height ratio in the range of 1.32–2.34, and about 4–9 robust, wide, mostly fused, and distinctively posteriorly recurved denticles. In lateral view, the basal margin is usually straight. A large, broadly expanded oval to sub-rounded basal cavity is upturned on the inner margin and flat to downturned on the outer margin.

Remarks. Goudemand (in Goudemand et al., 2012b) revised the original diagnosis by noticing that the basal margin is not necessarily straight and therefore, the most strikingly difference to Nv. waageni is the denticulation: the denticles axes are distinctively recurved posteriorly whereas in Nv. waageni and Nv. abruptus , the denticles are straight, inclined or radiating. Te two or three denticles anterior of the cusp are often clearly asymmetrical and the posterior edge of the element is much shorter than the anterior one. In most sections worldwide, Nv. pingdingshanensis first occurs within the positive δ 13 Ccarb excursion of the latest Smithian and may extend to the earliest Spathian ( Goudemand et al., 2019; Leu et al., 2019; Zhang et al., 2019). Some elements within our rich material resemble Ns. Pingdingshanensis , but appear to have a relatively small basal cavity, more posteriorly recurved denticles or more numerous denticles that what was previously described for this species. Tese elements

( Figs. 17C, J, K, T View Fig ; 18E, I–K View Fig ; 19O, Q View Fig ; 20H, N–O, S View Fig ; here assigned to N. ex gr. pingdingshanensis ) may deserve differentiation in the future. Two elements illustrated by Metcalfe et al., (2013, figs. 9.6, 9.9, p. 1144) as ‘ Neospathodus waageni subsp. nov. A’ display a higher anterior end and more recurved denticles than the other elements they included in ‘ Neospathodus waageni subsp. nov. A’, which we synonymized with Ns. pingdingshanensis : elements like these two elements do not seem to fit in Ns. pingdingshanensis and may deserve assignment to a new species. Some further elements resembling Ns. pingdingshanensis but distinctively shorter than Ns. pingdingshanensis are herein assigned to a new species ( Nv. gryphus n. sp. see below). Occurrence. Worldwide occurrence. China: Jinya/ breakage, it is not excluded that some of these elements Waili area, Guangxi ( Goudemand et al., 2012b) Jiarong, were broken and were subsequently repaired, thus coversouthern Guizhou ( Chen et al., 2013, 2015). Anshun Fm., ing such traces at the surface of the crown. Goudemand Qingyan section, Guizhou ( Ji et al., 2011). Chaohu (Zhao and Orchard (in Goudemand et al., 2012b) differentiated et al., 2007, 2008). Daxiakou, Hubei ( Zhao et al., 2013). similar but longer elements as Nv. aff. pingdingshanensis Vietnam: Bac Tuy Fm., Xenoceltites variocostatus and ( Goudemand et al., 2012b, figs. 2K, 2L) because they are Tirolites beds, ( Komatsu et al., 2016; Shigeta et al., 2014). lower and their denticles more recurved than in elements Canada: Scythogondolella mosheri zone, Wapiti Lake of Nv. pingdingshanensis : their denticles are so recurved ( Orchard & Zonneveld, 2009). Australia: Hovea Member that their upper profile appears straight; as such they are of Kockatea Shale, upper part of Neospathodus waageni reminiscent of the present species, with whom they may Zone, Smithian substage ( Metcalfe et al., 2013). be closely related. In comparison with Nv. soleiformis

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Fig. 17 View Fig Novispathodinae from Qiakong,Laren, Shanggang,and Lilong. Magnification is × 80.The scale bar is 400 μm. All elements are considered

to be P 1 elements if not specifically identified otherwise. A–C, J, T, AA Novispathodus ex gr. pingdingshanensis (Zhao & Orchard) ; A, QIA138,PIMUZ 39226; B QIA138, PIMUZ 39227; C LAR202, PIMUZ 39228; J QIA136, PIMUZ 39229; T LIL504, PIMUZ 39230; AA LIL508,PIMUZ 39231. D–F, M–P, R, S,

U, X, AB, AD Novispathodus pingdingshanensis (Zhao & Orchard) ; D QIA135, PIMUZ 39259; E LIL508,PIMUZ 39260; F LIL506, PIMUZ 39261; M LIL506, PIMUZ 39262; N LIL506,PIMUZ 39263; O LIL507, PIMUZ 39264; P LIL506, PIMUZ 39265; R LIL508,PIMUZ 39266; S LAR204,PIMUZ 39267; U LIL507, PIMUZ 39268; X LIL508, PIMUZ 39269; AB LIL508, PIMUZ 39270; AD LIL508, PIMUZ 39271. G, I, Q Novispathodus cf. ? gryphus n. sp.; G QIA136, PIMUZ 39204; I QIA136, PIMUZ 39205; Q QIA135, PIMUZ 39206. H, K, L Novispathodus gryphus n. sp.; H QIA135, PIMUZ 39243; K LIL506, PIMUZ 39244; L LIL506,PIMUZ 39245. V Novispathodus praebrevissimus n. sp.; LIL507, PIMUZ 39273. W, Y, Z, AE Novispathodus ex gr. abruptus (Orchard) ; W LIL505, PIMUZ 39208; Y SHA346,PIMUZ 39209; Z LIL506, PIMUZ 39210; AE SHA346, PIMUZ 39211. AC sp. indet.; LIL504, PIMUZ 39295. AF Novispathodus pingdingshanensis (P 1 cluster) (Zhao & Orchard); QIA136, PIMUZ 39272

( Zhao & Orchard, 2008) Nv. gryphus n. sp. has a higher Novispathodus gryphus n. sp. carina with less numerous but more posteriorly recurved Fig. 17G–I, K View Fig ?, L denticles. Furthermore, the basal cavity of Nv. gryphus n.

sp. does not extend as a wide deep groove anteriorly like Etymology: from the Greek root ‘gryph’ referring to the in Nv. soleiformis . hooked shape of the P 1 element. Occurrence. Luolou Formation, within the latest Smith- Holotype: specimen illustrated in Fig. 17H View Fig . ian black shales, Guangxi and Guizhou, South China (this Paratype: specimen illustrated in Fig. 17L. View Fig study). Type locality: Qiakong, Luolou formation, Guizhou Province, China. Novispathodus ex gr. abruptus ( Orchard, 1995) Type level: Luolou Formation, within latest Smithian Figs. 16E View Fig ; 17W, Y, Z View Fig , AE; 18H; 20A–G, K, L, M, Q, X black shales. Number of specimens. ca. 10 specimens. 1981 Neospathodus homeri Bender ; Koike, pl. 1, fig. 5.

1984 Neospathodus sp. A ; Hatleberg & Clark, pl. 3, fig. 8, Diagnosis. A species with a short segminate-to-segminis- 21. caphateP 1 element with 3 to 5 highly recurved denticles *1995 Neospathodus abruptus n. sp.; Orchard, p. 118, and a relatively large, sub-rounded basal cavity. 119, figs. 3.16–3.19, 3.23–3.26. Description. Te P 1 element is segminate to segminisca- 2005 Novispathodus abruptus (Orchard) ; Orchard, p. 90, phate, the sub-rounded basal cavity extending to most of text-fig. 16. the length of the element with a tapering at the anterior 2009 Novispathodus abruptus (Orchard) ; Orchard & end. Te denticles are mostly fused and highly recurved Zonneveld, p. 784, fig. 15 parts 34–37. posteriorly: although the denticles get larger from the 2012a Novispathodus sp. nov. A; Goudemand & Orchard anterior to the posterior, except for the posteriormost in Goudemand et al., p. 1031, figs. 2A, R?, Z?. one, the height of the carina looks sub-uniform because 2012a Novispathodus sp. nov. B; Goudemand & Orchard of the way the denticles are recurved. Te length-to- in Goudemand et al., p. 1031, fig. 3 V. height ratio is about 1:1. 2018 Novispathodus abruptus (Orchard) ; Maekawa in Remarks. Tese elements resemble strongly the homolo- Maekawa et al., p. 33, figs., 18.1–18.3, 18.20, 18.23, 18.24, gous ones of Nv. pingdingshanensis , except that they are 18.26, 18.27 (only). much shorter and bear less denticles, which are usually more recurved. Still, some may be confused with broken Material.> 50 specimens. elements of Nv. pingdingshanensis whose anteriormost end is missing ( Fig. 17G View Fig ). Hence the anterior end must be Diagnosis. As in Orchard, 1995. inspected for traces of breakage. Although the specimens Remarks. Here we consider Nv. abruptus in a broad sense, we have seen do not show obvious, surfacial traces of including elements that correspond to Nv. abruptus sensu (See figure on next page.) Fig. 18 View Fig Novispathodinae from Laren and Lilong. Magnification is ×80.The scale bar is 400 μm. All elements are considered to be P 1 elements if not specifically identified otherwise. A, B, D, F, L–N, Q, S, T Novispathodus praebrevissimus n. sp.; A LIL506,PIMUZ 39274; B LAR207, PIMUZ 39275; D LIL508, PIMUZ 39276; F LIL507, PIMUZ 39277; L LIL507, PIMUZ 39278; M LIL509, PIMUZ 39279; N LIL507,PIMUZ 39280; Q LIL507, PIMUZ 39281; S LIL507, PIMUZ 39282; T LIL507, PIMUZ 39283. C, O, R Novispathodus? praebrevissimus n. sp.; C LAR204,PIMUZ 39284; O LIL507, PIMUZ 39285; R LIL507, PIMUZ 39286. E, I Novispathodus ex gr. pingdingshanensis (Zhao & Orchard) ; E LIL507,PIMUZ 39232, I LIL507, PIMUZ 39233. G Triassospathodus aff. symmetricus (Orchard) ;LIL507, PIMUZ 39309. H Novispathodus ex gr. abruptus (Orchard) ; LIL507, PIMUZ 39212. J, K Novispathodus praebrevissimus (juvenile) n. sp.; J LIL507, PIMUZ 39287; K LIL507,PIMUZ 39288. P Novispathodus robustispinus (Zhao & Orchard) ; LIL507,PIMUZ 39290 stricto together with forms that have been suggested to deserve assignment to separate species, such as Nv. sp. nov. A and Nv. sp. nov. B Goudemand and Orchard (in Goudemand et al., 2012b). We consider the most diagnostic feature of Nv. ex gr. abruptus is the terminal 1–3 progressively smaller denticles at the posterior end.

Te P 1 element of this species is less robust, more rectangular with more fused denticles than that of Ic. crassatus . Its morphology also recalls that of the homologous element in Nv. pingdingshanensis , but its basal cavity is relatively smaller and its denticles are not as posteriorly recurved nor usually as broad as in the latter. Te P 1 element of Tr. symmetricus has more posteriorly reclined denticles, it may have a small terminal denticle but not several of increasingly smaller size. Yet, the distinction between Tr. symmetricus and Nv. abruptus may be confusing. Te P 1 element of Tr. homeri has a more elongated basal cavity and a more developed, posteriorly reclined and laterally deflected process than in both Tr. symmetricus and Nv. abruptus . Goudemand and Orchard (in Goudemand et al., 2012b) implicitly suggested that elements like those of their Nv. sp. nov. B, where the small terminal denticles are not increasingly smaller but of equal (small) height instead, may be transitional between Nv. abruptus and Tr. homeri (compare the elements illustrated in figs. 2.9 and 3.17 of Orchard 1995, assigned to Tr. homeri and Nv. Abruptus , respectively). Note further that, based on the material from Tsoteng, it is likely that suchP 1 elements were still associated with a Novispathodus apparatus. Tis suggests that P 1 elements like that of Tr. homeri may have evolved before the more substantial modifications of the rest of the apparatus implied by the difference between Novispathodus and Triassospathodus . Occurrence. Worldwide occurrence in latest Smithian and early Spathian rocks. Xenoceltites View in CoL and Tirolites View in CoL beds within Nv. pingdingshanensis and Nv. brevissimus zones, Japan ( Koike, 1981; Maekawa et al., 2018). Oman ( Orchard, 1995). British Columbia ( Orchard & Zonneveld, 2009), South China; Goudemand et al., 2012b; this study).