North of Calama, Antofagasta Region

One gladius from a locality south of Quebrada Quinchamale, ~5 km northwest of Cerro Jaspe, was collected by Andrew Tomlison in 2005 and deposited at the Servicio Nacional de Geología y Minería (SNGM–1849), Santiago, Chile - SERNAGEOMIN. It was figured (Rubilar and Pérez d'A., Reference Rubilar and Pérez d'A.2006, fig. 2C) as ‘Trachyteuthis sp.’ and is here described and refigured as Pseudoteudopsis perezi n. gen. n. sp. (see Fig. 4.1–4.3 in Systematic paleontology). Ammonites identified as Rehmannia (R.) cf. paucicostata (Tornquist, Reference Tornquist1898) and Neuqueniceras antipodum (Tornquist, Reference Tornquist1898) were recovered from the same level (Rubilar and Pérez d'A., Reference Rubilar and Pérez d'A.2006, p. 113) indicating reference to the lower Callovian uppermost bodenbenderi to lowermost proximum zones of the Andean ammonite zonation (cf. Riccardi and Westermann, Reference Riccardi and Westermann1991; Riccardi, Reference Riccardi2008).

Cordillera de Domeyko, northeast of Taltal

The material consists of four specimens, three of which are included in T. covacevichi and one in T. chilensis n. sp. (see the following). They were collected by Vladimir Covacevich † (then at SERNAGEOMIN) and/or Hans-Peter Schultze (Kansas University) in March 1994, and deposited in the paleontological collection of the SERNAGEOMIN (SNGM–8245 to 8248).

The material of T. covacevichi came from Quebrada del Profeta, H. P. Schultze collection (Holotype, original MNHN, SGO.PI.6437a, b, figured by Fuchs and Schultze, Reference Fuchs and Schultze2008, p. 43, figs. 3, 4; plaster cast SNGM–8245, here figured in Fig. 2.1); Cerro Islote, V. Covacevich collection (SNGM–8248; figured by Rubilar and Pérez d'A., Reference Rubilar and Pérez d'A.2006, fig. 2D; here figured in Fig. 2.3); and Quebrada Sandón, H.P. Schultze, and V. Covacevich collection (original MNHN, SGO.PI.6438, figured by Fuchs and Schultze, Reference Fuchs and Schultze2008, fig. 5; plaster cast SNGM–8245, here figured in Fig. 2.5).

Figure 2

Trachyteuthis covacevichi Fuchs and Schultze, Reference Fuchs and Schultze2008, middle Oxfordian (transversarium Zone), Antofagasta Region, Chile. (1, 2) Holotype (plaster cast, SNGM-8245), Quebrada del Profeta, H.P. Schultze collection (original SGO.PI 6437): (1) dorsal view of plaster cast (coated with ammonium chloride); (2) same as (1) (drawing with gladius reconstructed outline). (3, 4) Nearly complete gladius, Cerro Islote, V. Covacevich collection (SNGM-8248): (3) dorsal view (coated with ammonium chloride); (4) same as (3) (drawing with gladius reconstructed outline). (5) Juvenile gladius (rubber cast of internal mold, SNGM-8247; original, SGO.PI 6438), Quebrada Sandón, H.P. Schultze and V. Covacevich collection. All figures 0.90x.

The only specimen of T. chilensis n. sp. came from Quebrada del Profeta, V. Covacevich collection (SNGM–8246; figured by Rubilar and Pérez d'A., Reference Rubilar and Pérez d'A.2006, fig. 2A, 2B, as ‘Trachyteuthis sp.’ and refigured here in Fig. 3.1).

Figure 3

Trachyteuthis chilensis n. sp., Holotype (SNGM-8246, V. Covacevich collection), middle Oxfordian (transversarium Zone), Quebrada del Profeta, Antofagasta Region, Chile. (1) Gladius dorsal view (coated with ammonium chloride), 1x; (2) same as (1) (drawing with gladius reconstructed outline), 1x; (3) dorsum counterpart (rubber cast) showing median field granulated area, 0.90x. X–X´=lateral ridges along median field.

Ammonites referred by V. Covacevich (SERNAGEOMIN internal report, 1995) to the genera Mirosphinctes Schindewolf, Reference Schindewolf1926, Euaspidoceras Spath, Reference Spath1931, Cubaspidoceras Myczyński, Reference Myczyński1976, and ‘Perisphinctes’ Waagen, Reference Waagen1869 were found at the same levels, indicating an Oxfordian age (see Rubilar and Perez, d'A., Reference Rubilar and Pérez d'A.2006). Fuchs and Schultze (Reference Fuchs and Schultze2008, p. 44) labeled the type horizon of T. covacevichi as “uncertain (probably cordatum zone), early Oxfordian” without much elaboration although, from the stratigraphic accounts of the Quebrada del Profeta section given by Chong and Förster (Reference Chong and Förster1976, p. 147), Schultze (Reference Schultze1989, p. 186), Gygi and Hillebrandt (Reference Gygi and Hillebrandt1991, p. 142) and Hillebrandt et al. (Reference Hillebrandt, von, Kossler and Gröschke2012, p. 67), the specimens of T. covacevichi from Quebrada del Profeta level ‘10’ (cf. Schultze, Reference Schultze1989, p. 186, pl. 1, fig. 2; Fuchs, Reference Fuchs2006a, pl. 14, fig. F; Fuchs and Schultze, Reference Fuchs and Schultze2008, figs. 3, 4; here Fig. 2.1, 2.2) were found together with ammonites belonging to the subgenera Antilloceras Wierzbowski, Reference Wierzbowski1976, and Gregoryceras Spath, Reference Spath1924, which characterize the middle Oxfordian transversarium Zone, and ~6 m above a level with Caracoliceras dunkeri (Steinmann, Reference Steinmann1881), ascribed to the lower–middle Oxfordian uppermost Cordatum to Plicatilis Zones.

This fauna occurs in concretions, which also contain fishes, crustaceans, bivalves, and Lingula (cf. Schultze, Reference Schultze1989), within a succession of black sandy shales interbedded with fine-grained calcareous sandstones and thin limestone layers. Paleoenvironmental interpretations indicate dysaerobic shallow waters similar to those of the Posidonien Schiefer of southern Germany (cf. Chong and Förster, Reference Chong and Förster1976; Schultze, Reference Schultze1989).

The specimens from Cerro Islote (Rubilar and Pérez d'A., Reference Rubilar and Pérez d'A.2006, fig. 2D; here Fig. 2.3, 2.4) and Quebrada Sandon (Fuchs and Schultze, Reference Fuchs and Schultze2008, fig. 5; here Fig. 2.5) are most likely from the same level, i.e., equivalent to the middle Oxfordian transversarium Zone (cf. Gygi and Hillebrandt, Reference Gygi and Hillebrandt1991).

Repository and institutional abbreviations

Sgo–PI, Paleontología Invertebrados, Museo Nacional de Historia Natural, Santiago de Chile, Chile. SNGM, Colección Paleontológica Servicio Nacional de Geología y Minería (SERNAGEOMIN), Santiago de Chile, Chile.

Abbreviations for morphologic terms and dimensions (cf. Fuchs and Larson, Reference Fuchs and Larson2011, fig. 1)

GL=gladius length; GW=maximum gladius width; GW_{1/2 GL}=gladius width at half GL; GW_HZ=gladius width at anterior end of hyperbolar zone (HZ); GW_LF=gladius width at anterior end of lateral fields (LF); HZL=hyperbolar zone length; LFL=lateral fields length; A_ia=angle of diverging inner asymptotes; A_ga=angle of diverging granulate area.

Subclass Coleoidea Bather, Reference Bather1888

Order ?Teuthida Naef, Reference Naef1916

Family Trachyteuthididae Naef, Reference Naef1921

Remarks

Evolutionary history and classification at the ordinal level of coleoids is still highly debated (see Lindgren et al., Reference Lindgren, Pankey, Hochberg and Oakley2012 and references therein). Thus, as no new information on this issue is added by the material here described and a discussion on higher-level taxonomy of gladius-bearing coleoids is beyond the scope of this paper, the Trachyteutididae are here tentatively placed in the order Teuthida, following traditional views on the group systematics (cf. Naef, Reference Naef1922; Jeletzky, Reference Jeletzky1966).

It should be mentioned, however, that a three-part gladius with broad median field and wings (lateral fields), which was originally used to link fossils with modern teuthids, has been considered a plesiomorphic structure, which probably evolved at least four times in extant lineages and, therefore, is not a reliable character for determining relationships (see Young et al., Reference Young, Vecchione and Donovan1998; Vecchione et al., Reference Vecchione, Young, Donovan and Rodhouse1999). The one feature that would allow correct systematic assignment of teuthids is the arm crown (presence or absence of tentacles or filaments), but it is rarely preserved in the fossil record, except for a handful of exceptional (‘Lagerstatten’) localities (see Donovan and Fuchs, Reference Donovan and Fuchs2016), and when they are preserved (as in Mastigophora Owen, Reference Owen1856 and Plesioteuthis Wagner, Reference Wagner1859), they are subject to different interpretations (e.g., Naef, Reference Naef1922; Jeletzky, Reference Jeletzky1966; Bandel and Leich, Reference Bandel and Leich1986; Young et al., Reference Young, Vecchione and Donovan1998; Vecchione et al., Reference Vecchione, Young, Donovan and Rodhouse1999; Fuchs, Reference Fuchs2006a, Reference Fuchsb, Reference Fuchs2014; Fuchs et al., Reference Fuchs, Engeser and Keupp2007; Fuchs and Larson, Reference Fuchs and Larson2011; Klug et al., Reference Klug, Schweigert, Dietl and Fuchs2005, Reference Klug, Fuchs, Schweigert, Röper and Tischlinger2015; Donovan and Fuchs, Reference Donovan and Fuchs2016). Thus, relationships of families based only on gladii remain uncertain at present (Donovan et al., Reference Donovan, Doguzhaeva and Mutvei2003).

Genus Trachyteuthis Meyer, Reference Meyer1846

(=Coccoteuthis Owen, Reference Owen1855; ?Voltzia Schevill, Reference Schevill1950)

Type species

Trachyteuthis ensiformis Meyer, Reference Meyer1846, Tithonian, Solnhofen, Germany, by monotypy (see Donovan, Reference Donovan1995) [= T. hastiformis (Rüppell, Reference Rüppell1829)]

Other species

Trachyteuthis latipinnis (Owen, Reference Owen1855) from the Kimmeridge Clay of England and Tithonian of Germany; T. zhuravlevi Hecker and Hecker (Reference Hecker and Hecker1955) from the Tithonian of the Lower Volga region, Russia; T. willisi Wade (Reference Wade1993) from the Albian of Australia; T. nusplingensis Fuchs, Engeser, and Keupp (Reference Fuchs, Engeser and Keupp2007) from the upper Kimmeridgian of southern Germany; T. teudopsiformis Fuchs, Engeser, and Keupp (Reference Fuchs, Engeser and Keupp2007) from the lower Tithonian of southern Germany; T. covacevichi Fuchs and Schultze (Reference Fuchs and Schultze2008) from the Oxfordian of Chile; T. bacchiai Fuchs and Larson (Reference Fuchs and Larson2011) from the Cenomanian of Lebanon; and possibly ?T. palmeri (Schevill, Reference Schevill1950) from the Oxfordian of Cuba. Another Cretaceous record has been reported, but not figured, from the Aptian of Germany (Engeser and Reitner, Reference Engeser and Reitner1985).

Diagnosis

Trachyteuthid gladius with a hyperbolar zone length/gladius length ratio of 0.34–0.52. Anterior gladius end rounded or weakly pointed (modified from Fuchs and Larson, Reference Fuchs and Larson2011, p. 816; see Remarks)

Occurrence

Most records of Trachyteuthis are from the lower Tithonian lithographic limestones of Solnhofen and Nusplingen, southern Germany, from which at least 50 specimens have been preserved in different museums (Fuchs et al., Reference Fuchs, Engeser and Keupp2007, p. 576; according to D. Fuchs [personal communication, March 28, 2016], this number refers to the specimens used for morphometric comparisons, and there are ~10 times more accumulated in museums and private collections); ~23 of these were figured between 1825 and 2006, and until 2007 were considered (but see the following) to belong in a single species, T. hastiformis. A few additional specimens were also mentioned and/or figured from other regions in the Northern Hemisphere: three from the Tithonian of Russia (Trautschold, Reference Trautschold1866; Hecker and Hecker, Reference Hecker and Hecker1955), nine from the Kimmeridgian–Tithonian of England (Owen, Reference Owen1855; Donovan, Reference Donovan1977; Hewitt and Wignall, Reference Hewitt and Wignall1988), (?) one from the Aptian of Germany (Engeser and Reitner, Reference Engeser and Reitner1985), four from the Cenomanian of Lebanon (Fuchs, Reference Fuchs2006b, Reference Fuchs2007; Fuchs and Larson, Reference Fuchs and Larson2011), and (?) two from the Oxfordian of Cuba (Schevill, Reference Schevill1950). Eleven Trachyteuthis specimens are known from the Southern Hemisphere: five from the Oxfordian of Chile (Schultze, Reference Schultze1989; Rubilar and Perez d'A., Reference Rubilar and Pérez d'A.2006; Fuchs and Schultze, Reference Fuchs and Schultze2008), one from the lower Tithonian of Antarctica (Doyle, Reference Doyle1991), and five from the Albian of Australia (Wade, Reference Wade1993).

Remarks

As noted by Doyle et al. (Reference Doyle, Donovan and Nixon1994, p. 11) and Donovan (Reference Donovan1995, p. 1), Trachyteuthis was proposed by Meyer (Reference Meyer1846, p. 598) on the basis of at least two specimens for which he introduced two specific names, of which only one, T. ensiformis, is valid (International Commission on Zoological Nomenclature [ICZN], 1999, Art. 12.2.7) as the name was accompanied by reference to an unnamed specimen figured by Münster (Reference Münster1846, pl. 9, fig. 3). T. ensiformis is, therefore, the type species of Trachyteuthis Meyer, and T. hastiformis Rüppell, Reference Rüppell1829 is a senior synonym. As Meyer (Reference Meyer1846, p. 598) identified his own material with Münster’s (Reference Münster1846) figure, it is clear that the type series contained more than one specimen, and Münster’s specimen was, therefore, designated lectotype of the species (cf. Doyle et al., Reference Doyle, Donovan and Nixon1994, p. 11; Donovan, Reference Donovan1995, p. 2). Meyer (Reference Meyer1846) did not mention, and was probably unaware of, the fact that Münster’s (Reference Münster1846, pl. 9, fig. 3) specimen had been figured, together with another three specimens, under the name ‘Sepia linguata’ by d’Orbigny (1839, in Férussac and d’Orbigny, 1835–Reference Férussac and d’Orbigny1848, p. 292, pl. 16, fig. 3, a slightly reduced mirror image of Münster, Reference Münster1846, pl. 9, fig. 3). Since then, most subsequent authors have accepted that T. ensiformis Meyer, Reference Meyer1846, is a (junior) subjective synonym of T. hastiformis, a conclusion that does not invalidate the availability of Meyer’s (Reference Meyer1846) specific name (ICZN, 1999, Art. 10.6).

Although the holotype—by monotypy—of T. hastiformis was figured by Rüppell (Reference Rüppell1829, pl. 3, fig. 2), it is worth mentioning that Rüppell (Reference Rüppell1829, p. 9) indicated that the first figured specimen of the species (cf. also d’Orbigny, 1839, in Férussac and d’Orbigny 1835–Reference Férussac and d’Orbigny1848, p. 290) is probably represented by a drawing by the eighteenth-century copper engraver G.W. Knorr, which was included without identification in a monograph by Walch (1768–Reference Walch1773, pl. 22, fig. 2), who considered it a fish remain, an identification that was upheld by Germar (Reference Germar1826, p. 109). Another specimen of this species (see Fuchs et al., Reference Fuchs, Engeser and Keupp2007, p. 579) was figured by König (Reference König1825, pl. 17, fig. 201) as ‘Sepia prisca,’ and although this name constitutes the senior available name (ICZN, 1999, Art. 12.2.7), it should be considered a nomen oblitum and Rüppell’s (Reference Rüppell1829) younger name, T. hastiformis, a nomen protectum (ICZN, 1999, Art. 23.9.1 and 23.9.2).

Münster (Reference Münster1837) introduced another seven new specific names—S. obscura, S. linguata, S. regularis, S. gracilis, S. venusta, S. antiqua, and S. caudata—but they have been considered nomina nuda as they lacked descriptions and/or illustrations or any other indication that could validate them (cf. ICZN, 1999, Art. 12). d’Orbigny (1839, in Férussac and d’Orbigny, 1835–Reference Férussac and d’Orbigny1848), however, on the basis of Münster’s unpublished notes and drawings (cf. Wagner, Reference Wagner1860, p. 759; Fuchs et al., Reference Fuchs, Engeser and Keupp2007, p. 575), described and figured ‘S. antiqua,’ ‘S. caudata,’ and ‘S. linguata,’ the last one including the material referred by Münster (Reference Münster1837) to the three remaining specific names. Only ‘Sepia venusta’ was regarded an ‘anomalous form’ of a different species. Thus, d’Orbigny (1839, in Férussac and d’Orbigny 1835–Reference Férussac and d’Orbigny1848; cf. also d’Orbigny, Reference d’Orbigny1845), validated the three mentioned names, and became their author, as he was the first to publish them in a way that satisfies the criteria of availability (ICZN, 1999, Art. 50), although he clearly stated that he thought that all the specimens included under these specific names were in fact different growth stages and/or damaged representatives of a single species, i.e., Sepia hastiformis Rüppell.

It is worth mentioning that other specimens introduced by Münster (Reference Münster1837) as nomina nuda, i.e., S. obscura, S. regularis, S. gracilis, were placed by d’Orbigny (1839, in Férussac and d’Orbigny, 1835–Reference Férussac and d’Orbigny1848, p. 292; Reference Férussac and d’Orbigny1845, p. 162) in S. linguata. As the name of this species, even if regarded as a synonym of T. hastiformis, also complies with the criteria of availability (ICZN, 1999, Art. 11), one of the syntypes should be selected as lectotype whenever the original collection is restudied.

Almost all specimens included in T. hastiformis are from the lower Tithonian lithographic limestones of Solnhofen and Nusplingen, southern Germany. However, as a result of a morphological comparison by Fuchs et al. (Reference Fuchs, Engeser and Keupp2007, p. 576), based on at least 50 specimens from those localities deposited in different museums, Trachyteuthis hastiformis was restricted to specimens with coarse and irregular dorsal granulation and a median field with a spindle-shaped elevation, a feature first noted by Wagner (Reference Wagner1860, p. 755). Two new species were also recognized, i.e., T. nusplingensis Fuchs, Engeser, and Keupp, Reference Fuchs, Engeser and Keupp2007, and T. teudopsiformis Fuchs, Engeser, and Keupp, Reference Fuchs, Engeser and Keupp2007. T. nusplingensis was characterized by a flat dorsal gladius and regular and fine granulation, whereas T. teudopsiformis was said to have a gladius median field with a pronounced median keel, narrow granulation restricted to the posterior keel, and sharply pointed anterior gladius. According to Fuchs et al. (Reference Fuchs, Engeser and Keupp2007, p. 588), reliable comparisons with Trachyteuthis representatives from other localities and stratigraphic levels are difficult or impossible, as in most cases morphological knowledge is antiquated or based on only a few poorly preserved specimens. In the opinion of Fuchs et al. (Reference Fuchs, Engeser and Keupp2007), all those specimens should be considered as Trachyteuthis sp. until additional records and morphological features justify specific distinctions.

On the basis of the mentioned diagnostic features, especially the presence or absence of spindle-shaped elevation on the median field of the gladius, besides the lectotype only ~10 of the ~23 figured specimens assigned to T. hastiformis between 1829 and 2006 should remain in this species (i.e., d’Orbigny, 1839, in Férussac and d’Orbigny, 1835–Reference Férussac and d’Orbigny1848, pl. 14, fig. 2, pl. 16, fig. 1; Meyer, Reference Meyer1855, pl. 19, figs. 1, 2—var. media and minor of Wagner, Reference Wagner1859—; Rietschel, Reference Rietschel1977, fig. 6; Bandel and Leich, Reference Bandel and Leich1986, figs. 14–16; Donovan, Reference Donovan1995, fig. 1; Fuchs, Reference Fuchs2006a, pl. 14, figs. B, C), while ~13 should be excluded (i.e., d’Orbigny, 1839, in Férussac and d’Orbigny, 1835–Reference Férussac and d’Orbigny1848, p. 290, pl. 14, fig. 1; pl. 15, figs. 1, 3; d’Orbigny, Reference d’Orbigny1845, pl. 5, fig. 4, pl. 6, fig. 1; Quenstedt, Reference Quenstedt1849, pl. 31, fig. 25; Pictet, Reference Pictet1854, pl. 48, fig. 3; Zittel, Reference Zittel1885, fig. 710; Crick, Reference Crick1896, pl. 14; Naef, Reference Naef1922, fig. 51; Donovan, Reference Donovan1977, figs. 8, 9; Doyle, Reference Doyle1991, text-figs. 2A, B). For the latter there are at least two additional names available besides T. linguatus: T. latipinnis (Owen, Reference Owen1855) and T. zhuravlevi (Hecker and Hecker, Reference Hecker and Hecker1955). Resolution of this issue must await future studies of the European material.

Coccoteuthis, with C. latipinnis as type species, was introduced by Owen (Reference Owen1855, p. 125, pl. 7) for two specimens from the Kimmeridge clay of England. Owen (Reference Owen1855, p. 125) compared his material with several specimens from the Tithonian of Solnhofen, figured by König (Reference König1825, pl. 17, fig. 201) as ‘Sepia prisca,’ by Rüppell (Reference Rüppell1829, p. 9, pl. 3, fig. 2) as ‘Sepia hastiformis,’ by d’Orbigny (1839, in Ferussac and d’Orbigny, 1835–Reference Férussac and d’Orbigny1848, pl. 14, figs. 1, 2) as ‘Sepia antiqua Münster,’ and undesignated by Münster (Reference Münster1846, pl. 9, fig. 1), although he mentioned that C. latipinnis was broader in proportion to its length, a feature apparently shared by material from southern Germany, as exemplified by a specimen figured by Quenstedt (Reference Quenstedt1849, p. 493, pl. 32, fig. 1) as T. hastiformis. Owen (Reference Owen1855) was apparently unaware of Meyer’s (Reference Meyer1846) proposal of Trachyteuthis, with T. ensiformis as type species, for part of Münster’s (Reference Münster1846) material. Wagner (Reference Wagner1860) proposed the synonymy of Trachyteuthis Meyer, Reference Meyer1846 and Coccoteuthis Owen, Reference Owen1855, although he retained the latter as a senior synonym mentioning that figures and description were only given by Meyer in 1855. A similar view was taken by Keferstein (Reference Keferstein1866, p. 1441), Trautschold (Reference Trautschold1866, p. 15), and Crick (Reference Crick1896, p. 440), but Zittel (Reference Zittel1885, Reference Zittel1895) took the opposite view, as did Naef (Reference Naef1922, p. 138) and all subsequent authors.

Trachyteuthus zhuravlevi (Hecker and Hecker, Reference Hecker and Hecker1955) was introduced for one specimen from the Tithonian of Ulyanovsk (= Simbirsk) province, originally described as “Coccoteuthis hastiformis Rüpp.” by Trautschold (Reference Trautschold1866, p. 15, pl. 4), and two other specimens (Hecker and Hecker, Reference Hecker and Hecker1955, p. 40, pl. 2, figs. 2-3, text-figs. 4, ?5 [reproduced in part by Krymholts, Reference Krymholts1958, pl. 71, fig. 3] + ?1) from the Tithonian of Pugachyov, Saratov province, also in the lower Volga River region. T. zhuravlevi is very similar to T. hastiformis, but according to Hecker and Hecker’s (Reference Hecker and Hecker1955) interpretation of a specimen of this last species figured by Quenstedt (Reference Quenstedt1849, pl. 31, fig. 25), it would differ in a narrow median band having granulae and keel, whereas in T. hastiformis they would cover the total median field of the gladius. In fact, both species appear to differ in the presence or absence of a spindle-shaped elevation on the gladius median field, a feature regarded by Fuchs et al. (Reference Fuchs, Engeser and Keupp2007, p. 579) as diagnostic for T. hastiformis (see the preceding).

Voltzia, with V. palmeri as type species, was introduced by Schevill (Reference Schevill1950, p. 99, pl. 23, figs. 1–3) for two specimens from the Oxfordian of Cuba. According to Shevill (1950, p. 99) differences with Trachyteuthis were in the more prominent and chevron-shaped growth lines of the central field and the presence of a ventral “convex axial deposit smaller than but resembling the phragmocone of Sepia.” Schevill (Reference Schevill1950, p. 100) considered that “there is no close relationship between Voltzia and Trachyteuthis,” noting differences in proportions and stressing the presence of a “modified phragmocone” as an essential contrast with regard to representatives of the latter as well as to “a series of Solnhofen-Eichstätt teuthoids” he had the opportunity to examine. The paratype, which was not figured, was said to have only two-thirds of the anterior part preserved, while the holotype, as described and figured by Schevill (Reference Schevill1950, pl. 23, fig. 1) lacks the posterior margin and the right lateral field, whereas the left lateral field is not clearly visible. Moreover, Schevill’s (Reference Schevill1950, p. 99–100) description mentions, “dorsal surface tuberculate, more finely toward edges,” a feature that is not clearly visible in the figure, especially with regard to arrangement and regularity. However, as all dorsal features of the anterior part and median field described and figured by Schevill (Reference Schevill1950, pl. 23, figs. 1–3) do not differ from those known in Trachyteuthis, and even if the existence of a ventral phragmocone-like ‘deposit’ remains doubtful (cf. Waage, Reference Waage1965, p. 17) and cannot be checked because the type material is missing from the collections of the Museum of Comparative Zoology, Cambridge, USA (cf. Jeletzky, Reference Jeletzky1966, p. 107), Voltzia has been considered, tentatively (Jeletzky, Reference Jeletzky1966, p. 45, 107; Donovan, Reference Donovan1977, p. 31) or definitely (Riegraf, Reference Riegraf1995, p. 151; Fuchs et al., Reference Fuchs, Engeser and Keupp2007, p. 579), a synonym of Trachyteuthis. It could also be considered a nomen dubium (ICZN, 75.5, G).

Although the hyperbolar zone length/gladius length ratios of these species has been considered diagnostic for Trachyteuthis (Fuchs et al., Reference Fuchs, Engeser and Keupp2007), i.e., HZL/GL=0.40–0.45, they differ from those shown by T. covacevichi (HZL/GL =0.45–0.52) and the new species T. chilensis n. sp. (HZL/GL=0.34), recognized in the following. Thus, an emendation to the diagnosis of Trachyteuthis seems to be necessary (see the preceding).

Trachyteuthis covacevichi Fuchs and Schultze, Reference Fuchs and Schultze2008

Figure 2.1–2.5

1989 Plesioteuthis sp.; Reference SchultzeSchultze, p. 186, pl. 1, fig. 2

2006 Trachyteuthis sp.; Fuchs, Reference Fuchs2006a, pl. 14, fig. F

pars 2006 Trachyteuthis sp.; Reference Rubilar and Pérez d'A.Rubilar and Pérez d'A., fig. 2D, non figs. 2 A, 2B (= Trachyteuthis chilensis n. sp., see below), non fig. 2C (= Pseudoteudopsis perezi n. gen. n. sp., see the following).

2008 Trachyteuthis covacevichi Reference Fuchs and SchultzeFuchs and Schultze, p. 43, figs. 3A–3C, 4 (Fuchs, Reference Fuchs2006a, pl. 14, fig. F refigured), 5.

Holotype

Specimen consisting of an almost complete gladius, SGO.PI. 6437a–b, here refigured in Figure 2.1.

Diagnosis

Gladius with a hyperbolar length/gladius length ratio of 0.40–0.52. Anterior gladius end more or less rounded (modified from Fuchs and Schultze, Reference Fuchs and Schultze2008, p. 42; see Remarks).

Occurrence

Middle Oxfordian transversarium Zone of Quebrada del Profeta, Antofagasta Region, Chile. As explained in the preceding section on Material and geological setting, the type horizon was referred to the Oxfordian by Rubilar and Pérez, d'A. (Reference Rubilar and Pérez d'A.2006) due to the presence of the ammonite genera Mirosphinctes Schindewolf, Euaspidoceras Spath, Cubaspidoceras Myczynski, and ‘Perisphinctes’ Waagen, and labeled as “uncertain (probably cordatum zone), early Oxfordian,” without other considerations, by Fuchs and Schultze (Reference Fuchs and Schultze2008, p. 44). However, as mentioned in the preceding Material and geological setting section, the specimens of T. covacevichi from Quebrada del Profeta level ‘10’ were found together with ammonites belonging to the subgenera Antilloceras Wierzbowski and Gregoryceras Spath, which characterize the middle Oxfordian transversarium Zone.

Description

The holotype (Fig. 2.1, 2.2) is a three-dimensional uncompressed gladius with an elongate outline in which the posterior margin and the right lateral field have not been preserved. It has an inferred length (GL) of ~124.2 mm. Maximum gladius width (GW)=58.2 mm at maximum expansion of the wing’s lateral fields, about 76.6 mm of gladius length measured from the anterior end. Gladius width decreases steadily toward the anterior margin, amounting to 45.6 mm at half of gladius length and 25.8 mm at 14.5 mm of the anterior margin. From there, the margin converges into a weakly rounded frontal tip. Median field is longitudinally and transversally arcuate, with maximum convexity in the gladius posterior end. Inner asymptotes separating the median field from the hyperbolar zones are clearly visible and diverge at an angle of 40°, whereas limit between the hyperbolar zone and lateral field is transitional. The gladius surface shows three successive layers partially exposed; the granulated dorsal (outer) layer is poorly represented and barely visible in a few patchy remnants on the gladius anterior right side and on the median field left slope at 65 mm of the anterior margin. The dorsal surface of the relatively thick intermediate layer shows ~18–20 growth lines of alternating strength per cm; on the median field they are strongly concave and projected toward the posterior margin to become almost parallel to the lateral margins on both sides of the median field. At about 83 mm of gladius length, measured from the anterior end, growth lines bend first outward and then backward, following the expansion of the lateral wings, forming the hyperbolar zone (HZ) and the related (inner) asymptote and merging gradually with the lateral field outer margin. Growth lines are weakly visible on the inner wall layer. Longitudinal ridges are also present on the outer surface of the intermediate layer along the median field, four of which (two, 0.11 mm apart, on each side of the median plane) are well visible in the posterior half of the gladius, coinciding with the point where the growth lines start to be parallel to the lateral margins; ridges of each pair are ~0.11 mm apart and diverge from the posterior margin end forming an angle of 15°. The area between both pairs of ridges and the axis of the median field is weakly depressed.

The second specimen (SNGM–8248; Fig. 2.3, 2.4) included in this species is also a three-dimensional uncompressed gladius with an elongated outline in which the posterior margin and anterior part of the right lateral field have not been preserved. It is smaller than the holotype and has an inferred length (GL) of 102 mm. Maximum gladius width (GW=48 mm) corresponds to the maximum expansion of the lateral fields of the wings at about 59.2 mm of gladius length measured from the anterior end. Gladius width decreases steadily toward the anterior margin, amounting to 33.8 mm at half of gladius length and 27.6 mm at 10.2 mm of the anterior margin. From there, the margin converges into a weakly rounded frontal tip. Median field is longitudinally and transversally arcuate, with maximum transversal convexity in the gladius posterior end. Inner asymptotes separating the median field from the hyperbolar zones are clearly visible at the angle of convergence of the arcuate median field and the flat lateral wings and diverge with an angle of 40°, whereas limit between the hyperbolar zone and lateral field is less pronounced and only visible on the gladius posterior right side. The gladius surface shows successive layers partially exposed but not clearly differentiated due to preservation; the granulated dorsal (outer) layer is poorly represented and barely visible in a few patchy remnants on the median field. The dorsal surface of what appears to be the intermediate layer shows some barely visible growth lines on some parts of the median field and on the lateral fields. Longitudinal protuberances like ridges are also present on both sides of the median field and are well visible in the gladius middle region. The areas between both ridges and the axis of the median field are slightly depressed.

A small juvenile gladius (SGO.PI.6438, SNGM–8247; Fig. 2.5) is preserved as a three-dimensional external mold exposing half of the gladius anterior part, left wing lateral field, and posterior margin, whereas other parts of the mold are filled with sediment. It has a length (GL) of 54.3 mm. The inferred maximum gladius width (GW=24.8 mm) is on the maximum expansion of the wings’ lateral fields, at about 37 mm of gladius length measured from the anterior end. Gladius width decreases steadily toward the anterior margin, amounting to 18 mm at half of gladius length and 10.8 mm at 6 mm of the anterior margin. From there the margin converges into a triangular frontal tip. The gladius anterior half is longitudinally and transversally arcuate. The gladius surface shows barely visible growth lines parallel to the margins in the left wing and close to the anterior, lateral and posterior margins (see Table 1).

Table 1

Measurements in millimeters for Trachyteuthis covacevichi Fuchs and Schultze, Reference Fuchs and Schultze2008. GL=gladius length; GW=maximum gladius width; GW_{1/2 GL}=gladius width at half GL; GW_HZ=gladius width at anterior end of hyperbolar zone (HZ); GW_LF=gladius width at anterior end of lateral fields (LF); HZL=hyperbolar zone length; LFL=lateral fields length; A_ia=angle of diverging inner asymptotes; A_ga=angle of diverging granulate area. Values in parentheses=relation of each dimension to GL.

Material

The holotype, one plaster cast of a small (juvenile) gladius (SGO. –PI.6438) from tributaries north of Quebrada del Profeta, and three nearly complete gladii from a concretionary level, Quebrada del Profeta (SNGM–8245), Cerro Islote (SNGM–8248), and Quebrada Sandon (SNGM–8247), middle Oxfordian transversarium Zone, Antofagasta Region, Chile.

Remarks

Some rather small differences between holotype dimensions reported by Fuchs and Schultze (Reference Fuchs and Schultze2008, p. 43) and those recorded here could be because preservation prevents accurate measurements. The difference in recorded GW_1/2GL (38 vs. 45.6 mm) could be due to reconstruction of missing parts. Difference in LFL values (38 vs. 54.3) is probably related to different interpretations of structure limits. The new specimen figured here (Fig. 2.3, 2.4) indicates that the 0.40–0.45 hyperbolar zone length/gladius length ratio given by Fuchs and Schultze (Reference Fuchs and Schultze2008, p. 42) in the species diagnosis should be enlarged to 0.40–0.52.

As indicated by Fuchs and Schultze (Reference Fuchs and Schultze2008, p. 45), the gladius of T. covacevichi differs from those of T. hastiformis and T. nusplingensis, from the Tithonian of southern Germany, by being relatively wider (GW/GL=0.46–0.47 vs. 0.35–0.39) and more compact (LFL/GL=0.39–0.48 vs. 0.32–0.35; GW_HZ/GL=0.30–0.38 vs. 0.27–0.28). It also lacks the spindle-shaped elevation in the median field of T. hastiformis. As mentioned in the preceding, its gladius is also wider and more compact than that of T. chilensis n. sp. Comparison with T. cf. hastiformis from the Tithonian of Antarctica (Doyle, Reference Doyle1991, p. 172, text-fig. 2A, 2B) is hindered by the poor preservation of the Antarctic material.

Trachyteuthis chilensis new species

Figure 3.1–3.3

pars 2006 Trachyteuthis sp.; Reference Rubilar and Pérez d'A.Rubilar and Pérez d'A., fig. 2A, 2B, non fig. 2C (= Pseudoteudopsis perezi n. gen. n. sp., see below); non Fig. 2D (= Trachyteuthis covacevichi Fuchs and Schultze, Reference Fuchs and Schultze2008; see the preceding).

Holotype

The almost complete gladius with counterpart of anterior region, SNGM–8246, here figured in Figure 2.1–2.3, middle Oxfordian, Quebrada del Profeta, Antofagasta, Chile.

Diagnosis

Compressed gladius of medium size with elongated outline, weakly rounded frontal tip, and pointed posterior margin, median field laterally limited by two weakly raised blunt ridges that diverge from the posterior to the anterior margin at an angle of about 20°, passing inward into two shallow depressions bordering on both sides a rounded weak elevation along the median field, granulae on dorsal surface regularly arranged following the anterior margin.

Occurrence

Thus far species only known from the middle Oxfordian transversarium Zone of Quebrada del Profeta, Antofagasta Region, Chile.

Description

The holotype is a compressed gladius with an elongated outline where the posterior margin and the left and partly the right lateral fields have not been preserved. It has an inferred length (GL) of ~221 mm. Maximum gladius width (GW=~75.3 mm) corresponds to the maximum expansion of the wings’ lateral fields at about 150.8 mm of the gladius length measured from the anterior end. Gladius width decreases steadily toward the anterior margin, amounting to 55.8 mm at half of gladius length and 38 mm at 17.6 mm of the anterior margin. From there, the margin converges into a weakly rounded frontal tip. On both sides along the median field are two weakly raised blunt ridges (indicated with X and X' in Fig. 3.2) that diverge from the posterior to the anterior margin at an angle of about 20° and are clearly visible at about half of gladius length; outward they are limited by a flat marginal area, and inward they pass into two shallow depressions bordering on both sides a rounded weak elevation along the median field; depressions fade away anteriorly, where the median field becomes flat, and posteriorly where it is bluntly arched. An inner asymptote separating the median field from the hyperbolar zone is barely visible on the posterior right side, and the limit between the hyperbolar zone and the lateral field is marked by a very weak ridge. The estimated inner asymptote angle amounts to about 40°. The gladius surface shows three successive layers partially exposed; the granulated dorsal (outer) layer is poorly represented and barely visible in a few patchy remnants, e.g., on the gladius anterior right side and on the median field right depression at about half of the gladius length. The dorsal surface of the intermediate layer shows growth lines near the anterior margin, where they are strongly concave on the median field and projected toward the posterior margin to become almost parallel to the lateral margins; they are of alternating strength, the stronger amounting to about four per centimeter, while the weaker are barely visible. Very weak longitudinal ridges are also present on the outer surface of the intermediate layer along the lateral margins. A counterpart of the gladius anterior region (Fig. 2.3), with a length of 116.4 mm, shows the imprint of the dorsal gladius surface, where granulae are clearly exposed; they are regularly arranged following the growth lines of the median field and the shape of the gladius anterior margin; they reach their maximum size on the median line between 20 and ~60 mm of the anterior margin to become smaller toward the anterior, posterior, and lateral margins. The rounded weak elevation along the median field is clearly visible along the total gladius length, whereas the weak left ridge and concave depression are restricted to an area at about half of the gladius length (see Table 2).

Table 2

Measurements in millimeters for Trachyteuthis chilensis n. sp. GL=gladius length; GW=maximum gladius width; GW_{1/2 GL}=gladius width at half GL; GW_HZ=gladius width at anterior end of hyperbolar zone (HZ); GW_LF=gladius width at anterior end of lateral fields (LF); HZL=hyperbolar zone length; LFL=lateral fields length; A_ia=angle of diverging inner asymptotes; A_ga=angle of diverging granulate area. Values in parentheses=relation of each dimension to GL.

Etymology

From Chile, country of provenience of the holotype.

Material

Holotype, SNGM–8246.

Remarks

Trachyteuthis chilensis resembles T. hastiformis in the general outline, but differs in the absence of a spindle-shaped elevation in the gladius median field, in the regular arrangement and size variation of granulae, and perhaps, as reconstructed here, in the more elongated gladius with more pointed posterior and rounded anterior margins. It should be noted, however, that the presence in the middle part of the gladius of T. chilensis of a weak rounded elevation with two shallow depressions on either side could represent an incipient development of the spindle-shaped elevation that characterizes the younger species. Comparison with T. cf. hastiformis from the Tithonian of Antarctica (Doyle, Reference Doyle1991, p. 172, text-fig. 2A–2B) is hindered by the poor preservation of the Antarctic material.

Trachyteuthis chilensis differs from T. latipinnis (Owen, Reference Owen1855) by being smaller (GL=~221 mm vs. ~308 mm) and by the more elongated outline of the gladius and median field, the more rounded and flat gladius anterior margin, and the more regular granule arrangement.

Comparison with T. zhuravlevi Hecker and Hecker, Reference Hecker and Hecker1955 is made difficult by the incomplete and poor preservation of the Russian specimens, although one of them (Hecker and Hecker, Reference Hecker and Hecker1955, pl. 2, fig. 2, text-fig. 4; Krymholts, Reference Krymholts1958, pl. 71, fig. 3) appears to have a broader gladius posterior margin and a narrower median field.

Trachyteuthis covacevichi has a smaller (GL=54.3–124.2 mm vs. ~221 mm), relatively wider (GW/GL=0.46–0.47 vs. 0.34), and more compact gladius (LFL/GL=0.39–0.48 vs. 0.29; W_HZ/_GL=0.32–0.38 vs. 0.28).

Comparison with T. (?) palmeri (Schevill, Reference Schevill1950; p. 99, pl. 23, figs. 1, 2), from the Oxfordian of Cuba, is hindered by the incomplete preservation of the Cuban specimens—which are lost—where the lateral fields, hyperbolar zones, and tubercles are missing or are not clearly visible in the figures. Some proportions look rather similar, although the Cuban species appears to differ from T. chilensis in being smaller (GL=197 mm vs. 221 mm) and broader (GW_1/2GL=0.28 vs. 0.25) and having a more anteriorly pointed gladius.

Genus Pseudoteudopsis new genus

Type species

Pseudoteudopsis perezi new species, by monotypy.

Diagnosis

As for the species.

Occurrence

Thus far only known from the Callovian of the Antofagasta Region, Chile.

Etymology

Pseudo (Gr.), meaning false, and Teudopsis (Teuthis, Gr., calamar; øps, Gr., appearance), referring to the false similarity with Teudopsis Deslongchamps, Reference Deslongchamps1835.

Remarks

Pseudoteudopsis n. gen. resembles Teudopsis Deslongchamps, Reference Deslongchamps1835 in the pronounced median keel, but the latter is characterized by gladius margins converging rather rapidly in a pointed blade-like or spike-like anterior margin or a free rhachis (cf. Fuchs and Weis, Reference Fuchs and Weiss2010). Moreover, Pseudoteudopsis n. gen. differs in its elongated outline with rounded posterior and anterior margins, flat and poorly developed wings, presence of granulae on the dorsal surface, and an entire median keel of almost the same width from the posterior to the anterior margin.

Pseudoteudopsis perezi new species

Figure 4.1–4.3

Figure 4

Pseudoteudopsis perezi n. gen. n. sp. Holotype, SNGM-1849, A. Tomlinson collection, lower Callovian (uppermost bodenbenderi to lowermost proximum Zones), Quebrada Quinchamale, northwest of Cerro Jaspe, Antofagasta province, Chile. (1) Rubber cast of external mold (coated with ammonium chloride); (2) same as (1) (drawing with gladius reconstructed outline); (3) detail of (1), external mold showing granulate area. (1, 2) 1x; (3) 4x.

pars 2006 Trachyteuthis sp.; Reference Rubilar and Pérez d'A.Rubilar and Pérez d'A., fig. 2C; non fig. 2A, 2B (=Trachyteuthis chilensis n. sp., see the preceding), non fig. 2D (=Trachyteuthis covacevichi Fuchs and Schultze, Reference Fuchs and Schultze2008; see the preceding).

Holotype

A three-dimensional external mold of a gladius of which a rubber cast is figured in Figure 4.1, 4.2, collected by Andrew Tomlison in 2005 and deposited at the SERNAGEOMIN (SNGM–1849), lower Callovian.

Diagnosis

Elongated outline with rounded posterior and anterior margins, flat and poorly developed wings, presence of granulae on the dorsal surface, and a pronounced median ridge of almost the same width from the posterior to the anterior margin.

Type horizon

Lower Callovian uppermost bodenbenderi to lowermost proximum zones, south of Quebrada Quinchamale, ~5 km northwest of Cerro Jaspe.

Description

The holotype is represented by a three-dimensional external mold of a gladius with an elongated outline where the posterior margin and the left lateral wing are not preserved. It has an inferred length (GL) of 169 mm. Maximum gladius width (GW=71 mm) on maximum expansion of the wings’ lateral fields, at about 114.5 mm of gladius length measured from the anterior end. Gladius width decreases steadily toward the posterior margin with 51.5 mm at the beginning of the hyperbolar zones, from which the inner asymptotes continue bordering a relatively prominent median field, which reaches 22.4 mm in width just before forming a rounded posterior tip. Inner asymptotes diverge with an angle of 50° and separate the median field from the hyperbolar zone, which is not differentiated from the lateral fields. The gladius margins converge gradually toward the anterior margin reaching 32.9 mm in width just before forming a blunt anterior margin with a very small radius of curvature. Median field is flat in the gladius anterior half and becomes progressively more arcuate toward the posterior margin. The internal mold shows some patchy shell remnants and areas with irregular granulae and longitudinal fine ridges subparallel to gladius margin. Along the median line there is a prominent rounded ridge with a rather uniform width of ~9.3 mm from the posterior to anterior margin (see Table 3).

Table 3

Measurements in millimeters for Pseudoteudopsis perezi n. gen. n. sp. GL=gladius length; GW=maximum gladius width; GW_{1/2 GL}=gladius width at half GL; GW_HZ=gladius width at anterior end of hyperbolar zone (HZ); GW_LF=gladius width at anterior end of lateral fields (LF); HZL=hyperbolar zone length; LFL=lateral fields length; A_ia=angle of diverging inner asymptotes; A_ga=angle of diverging granulate area. Values in parentheses=relation of each dimension to GL.

Etymology

After my colleague and friend the late Ernesto Pérez d’Angelo (1932–2013), who promoted the initiation of this study, in recognition of his life-long contributions to the paleontology of Chile.

Material

The holotype.

Remarks

Pseudoteudopsis perezi n. sp. resembles Teudopsis sp. described by Fischer and Riou (Reference Fischer and Riou1982, p. 15, pl. 3, fig. 7) from the lower Callovian of Voulte-sur-Rhône, France, in the presence of a prominent median keel having a similar width from the posterior to the anterior margin. The French specimen—a doubtful Teudopsis according to Fuchs and Weis (Reference Fuchs and Weiss2010, p. 355)—is poorly preserved and consists of the posterior half of the gladius median field, in which the lateral wings are not visible. Its inclusion in Teudopsis was assumed (Fischer and Riou, Reference Fischer and Riou1982, p. 16) because the strong obliquity of the growth lines suggested a pointed rhachis. This specimen appears to differ from Pseudoteudopsis perezi n. gen. n. sp. in the pointed rhachis and the fact that the median keel is formed by 4–5 narrower ridges, but poor preservation prevents a close comparison.

A species of Trachyteuthis with a median keel, i.e., Trachyteuthis teudopsiformis Fuchs, Engeser, and Keupp (Reference Fuchs, Engeser and Keupp2007, p. 584, fig. 6) from the lower Tithonian of southern Germany, differs in the relatively broader gladius, more developed wings, and less prominent median keel that becomes wider and flattened toward the anterior margin.