New late Smithian (Early Triassic) ammonoids from the Lusitaniadalen Member, Vikinghogda Formation, Svalbard

: A single carbonate concretion from the uppermost Lusitaniadalen Member (‘Fish Niveau’, Vikinghøgda Formation) at Stensiöfjellet, Spitsbergen, contains a fairly rich assemblage of late Smithian (Early Triassic, Olenekian) ammonoids. The main species identified are Wasatchites tridentinus Spath, 1934, Wasatchites cf. distractus (Waagen, 1895), Anasibirites kingianus (Waagen, 1895), Arctoprionites nodosus (Frebold, 1930), Arctoprionites resseri (Mathews, 1929) and Xenoceltites subevolutus Spath, 1930. Two taxa are here discussed: Prionitid sp. and Arctoceras erebori sp. nov. The assemblage provides new information about the Early Triassic biostratigraphy and palaeogeography of the Boreal Realm. The composition of the Spitsbergen fauna is in good agreement with other assemblages within and outside the Boreal Realm and essentially comprises cosmopolitan taxa. Only Arctoceras erebori sp. nov. and Prionitid sp. are as yet restricted to Spitsbergen. The newly reported occurrence of low paleolatitude taxa from Spitsbergen strengthens the cosmopolitan distribution of ammonoids during late Smithian time, thus improving existing correlation. A single carbonate concretion from the uppermost Lusitaniadalen Member (‘Fish Niveau’, Vikinghøgda Formation) at Stensiöfjellet, Spitsbergen, contains a fairly rich assemblage of late Smithian (Early Triassic, Olenekian) ammonoids. The main species identified are Wasatchites tridentinus Spath, 1934, Wasatchites cf. distractus (Waagen, 1895), Anasibirites kingianus (Waagen, 1895), Arctoprionites nodosus (Frebold, 1930), Arctoprionites resseri (Mathews, 1929) and Xenoceltites subevolutus Spath, 1930. Two taxa are here discussed: Prionitid sp. and Arctoceras erebori sp. nov. The assemblage provides new information about the Early Triassic biostratigraphy and palaeogeography of the Boreal Realm. The composition of the Spitsbergen fauna is in good agreement with other assemblages within and outside the Boreal Realm and essentially comprises cosmopolitan taxa. Only Arctoceras erebori sp. nov. and Prionitid sp. are as yet restricted to Spitsbergen. The newly reported occurrence of low paleolatitude taxa from Spitsbergen strengthens the cosmopolitan distribution of ammonoids during late Smithian time, thus improving existing correlation


Introduction
A growing interest is focused on the usefulness of Early to Middle Triassic ammonoids of the Boreal Realm as high-resolution biostratigraphic markers, which in part benefit the increased petroleum exploration activity in the region and its need for accurate biochronological correlation. A revision of ammonoid taxonomy, biostratigraphy and palaeobiogeography of the Boreal Triassic is therefore required. The present paper provides new interpretations of a late Smithian fauna from Spitsbergen, and a new taxon is presented.
Boreal Early Triassic ammonoid faunas were characterised mainly by low alpha (= local) diversity, significant intraspecific variation (e.g., Hammer & Bucher, 2005;Weitschat, 2008) and high endemism (Tozer, 1982;Brayard et al., 2006). The Early Triassic Boreal Realm can be divided into western (Sverdrup Basin) and eastern (Siberian) provinces according to taxonomic differences of the ammonoid assemblages. Svalbard was located somewhat in between, presenting affinities to both of these provinces, although such affinities were not constant. Based on these intermediate affiliations, the establishment of a Svalbardian province was proposed by Weitschat & Dagys (1989).
The Tardus Zone defines the late Smithian in the Boreal Realm, and is characterised by cosmopolitan prionitids and xenoceltitids (e.g., Xenoceltites Spath, 1930); (Brühwiler et al., 2010). The late Smithian faunas of Svalbard are closely related to the faunas of the Sverdrup
The occurrence of the arctoceratid genus Arctoceras Hyatt, 1900 in Svalbard has long been controversial in the literature. According to various authors (e.g., Kummel, 1961;Tozer & Parker, 1968), it ranges through the whole of the Smithian in Svalbard. It was designated as an index taxon by Korčinskaja (e.g., 1986) to define a new zonation, which was, however, not confirmed by Weitschat & Dagys (1989). Instead, they retained the classic two-zone scheme (Romunderi and Tardus) for the Smithian in Svalbard. Nevertheless, Weitschat & Lehman (1978) stated that Arctoceras does range up into the Tardus Zone, whereas Weitschat & Dagys (1989) did not confirm the presence of arctoceratids in the late Smithian.

Material, location and stratigraphy
The ammonoid fauna discussed here was collected by the late Wolfgang Weitschat (University of Hamburg) from a single carbonate concretion in a horizon informally known as the 'Fish Niveau' (Wiman, 1910) in the uppermost Lusitaniadalen Member of the Vikinghøgda Formation at Stensiöfjellet (Fig. 2). It corresponds to the ammonoid biostratigraphic level of the Tardus Zone.
The material was originally stored in Hamburg, and then Basin (Weitschat & Lehmann, 1978 Brayard et al., 2013;Jattiot et al., in press). In Svalbard (as in the Queen Elizabeth Islands, Arctic Canada), the genus Xenoceltites is associated with Anasibirites and Wasatchites, unlike other Boreal localities (Tozer, 1961;Weitschat & Lehmann, 1978). At low and middle palaeolatitudes (e.g., China, western USA) it also occurs with the prionitid fauna, but the latest Smithian is represented by an additional zone characterised by the xenoceltitid genus Glyptophiceras (Waagen, 1895), above the prionitid beds. Therefore, while the Boreal Tardus Zone can be easily correlated worldwide, the subdivision of the uppermost Smithian into an Anasibirites / Wasatchites fauna followed by a Glyptophiceras / Xenoceltites fauna cannot be extended to higher palaeolatitudes (Brühwiler et al., 2010). The separation between Xenoceltites and Glyptophiceras is still unclear -whether the latter is an evolutionary transitional form or an example of intraspecific variation is still a matter of debate (H. Bucher, pers. comm. to V.P., 2017). transferred to the Natural History Museum (NHM) in Oslo. Original descriptions of sections and faunas are given by Weitschat & Lehmann (1978). The Lusitaniadalen Member (Fig. 3) is composed mainly of dark-grey shales with sandstones, calcareous siltstones and fossiliferous limestone concretions. The sand-/ siltstone content increases in an upward-shallowing trend (Mørk et al., 1999;Lundschien et al., 2014). Carbonate concretions are more prominent in the middle and upper parts, and are most likely of early diagenetic origin (Mørk et al., 1999;Hounslow et al., 2008). The depositional setting is interpreted as a moderately deep shelf environment (Mørk et al., 1999). The red square represents the concretion horizon from which the studied fauna most likely came (modified from Vigran et al., 2014, figs. 18 & 38a).

Discussion
The studied fauna is in good agreement with other late Smithian assemblages worldwide, and is most closely related to the Sverdrup Basin among all Boreal regions.
Xenoceltites is associated with Wasatchites, confirming previous such reports (e.g., Weitschat & Lehmann, 1978;Weitschat & Dagys, 1989). However, the fauna is also characterised by species that have not been previously recorded from Svalbard.
Arctoprionites resseri, for instance, has been recorded in British Columbia and in the USA (Tozer, 1994;Brayard et al., 2013;Jattiot et al., in press). This find suggests a broader palaeogeographic range for this taxon than previously thought, and confirms the faunal similarities between the Svalbard Early Triassic and the Canadian regions, as also highlighted by the presence of Arctoprionites nodosus, which is only known from Arctic Canada outside Svalbard, its type locality (Frebold, 1930;Tozer, 1994).
Similarly, the occurrence of specimens similar to Wasatchites distractus, common in the Tethyan Realm but unknown at higher latitudes, strengthens the cosmopolitan composition of the late Smithian ammonoid faunas. This species is generally recorded from regions such as Salt Range, Spiti and Tibet, where the lower part of the Boreal Tardus Zone is represented by the Wasatchites distractus beds (Brühwiler et al., 2012b;Brayard et al., 2013).
The occurrence of arctoceratids in this level is also worth noting. Findings of Arctoceras erebori sp. nov. at Stensiöfjellet confirm the range of this genus up to the uppermost Smithian, as suggested by e.g., Kummel (1961) and Weitschat & Lehmann (1978), with Arctoceras erebori sp. nov. possibly restricted to the Tardus Zone.
The lack of additional well-preserved specimens of the taxon Prionitid sp. hinders a more definite taxonomic classification as well as a discussion of its potential palaeogeographic significance.
As Wasatchites cf. distractus and specimens comparable to Prionitid sp. have not been recorded in the Canadian regions, two questions arise: whether the Svalbard fauna may be distinctive, and therefore the erection of a distinct Svalbardian Province (Weitschat & Dagys, 1989) should be reconsidered, or whether a sampling bias should be taken into account. Further evidence is therefore needed.

Methods
The classical parameters for ammonoid biometry (see e.g., Korn, 2010) were measured: diameter (D), whorl width (W), whorl height (H) and umbilical diameter (U). Ratios of each parameter against corresponding diameter D (W/D, H/D and U/D) were plotted, in addition to the ratio W/H to describe the shape of the whorl section.
Both absolute values and ratios have been plotted against D when at least four measurements could be obtained. The total number of specimens assigned to a genus/ species is labelled as N, and the number of measured specimens as n.
One specimen described herein (PMO 227.988) was imaged with a Nikon Metrology XT H 225 ST microfocus CT scanner at the Natural History Museum, University of Oslo. The scanning was carried out at 120 kV, 120 uA, 1 s exposure time, 0.5 mm tin filter, 3000 rotations.

Results
The specimens were assigned to nine species. The Xenoceltidae family is represented by a few specimens of Xenoceltites subevolutus (Fig. 4A-B The specimens are generally well preserved, with rare evidence of breakage/collapse and diagenetic compression. With some exceptions, no information about the orientation in the concretion is available. The ceratitic suture lines are visible on only a few specimens. The assemblage is composed mainly of immature individuals, as indicated by morphological observations and statistical analyses, with a very similar conch shape but large variation in size, shell coiling and ornamentation among specimens. Ornamentation is thus the key character for taxonomical identification. The assemblage also provides good material for studying intraspecific variation, as exemplified by Wasatchites (Fig. 5). As intraspecific variation in Triassic ammonoids has been discussed widely in the literature (e.g., Dagys & Weitschat, 1993b;Weitschat, 2008), we will focus on the implications of the studied assemblage for palaeogeography and biostratigraphy.

Conclusions
The studied fauna, most of which is characterised by a few cosmopolitan prionitid genera (Wasatchites and Anasibirites being the most common), is in good agreement with other faunas from the Tardus Zone worldwide. These results also strengthen correlations between Svalbard and the Canadian regions, as well as between palaeolatitudes. The concretion described here also provided new insights that are of importance for Early Triassic ammonoid taxonomy, biostratigraphy and palaeogeography: • Low-latitude ammonoid taxa such as Wasatchites cf. distractus, which until now has not been reported from any Boreal locality, and Arctoprionites resseri, recorded from British Columbia and USA but new in Svalbard, suggest faunal exchange between Svalbard and the Tethys to a higher degree than previously thought. These occurrences improve the correlation between palaeolatitudes.
• The presence of arctoceratids at this level confirms the range of the genus Arctoceras up to the Tardus Zone in Spitsbergen, as already suggested in some previous reports. This is not the first late Smithian occurrence of taxa known from middle Smithian strata. Jattiot et al. (2015) recorded two specimens of Galfettites omani Brühwiler & Bucher (2012a) in a late Smithian fauna from Noe Tobe, Timor, representing the first and youngest known occurrence of this taxon. Another arctoceratid, Churkites Okuneva, 1990, occurs in the latest middle Smithian in the western USA, but also in the latest Smithian in South Primorye (Shigeta & Kumagae, 2015). This may indicate that the late Smithian extinction was more gradual than was initially thought.
• The problematic Prionitid sp., which could not be assigned with certainty to any known taxon of the same family, may be evidence of an even higher early late Smithian diversification of the prionitids. On this matter, further investigation might show whether the niches vacated by other families stimulated the radiation of the prionitids.

Taxonomical descriptions
By Veronica Piazza.
All taxonomical work was conducted with reference to the manual 'Procedure in Taxonomy' (Schenk et al., 1956). The scale bar is 1 cm, unless indicated. The repository of the labelled specimens is abbreviated as PMO (Palaeontological collections of the Natural History Museum, University of Oslo, Norway). Background data are provided in a thesis by the first author (Piazza, 2015).
Occurrence. Rare in the whole assemblage, N = 6.
Description. Involute discoidal shell with oval whorl section and convex flanks. Venter commonly arched, smooth or presenting slightly arched to straight ventral ribs. Moderately deep and narrow umbilicus with rounded umbilical shoulder and fairly high-angled umbilical wall. Fairly smooth shell; ornamentation consisting of faint sinuous and regularly spaced ribs most visible on the flanks, as observed on Fig. 6B. Possible constriction observed on the body chamber of specimen PMO 227.988 (Figs. 6C-D, 7). Smooth/slightly ribbed inner whorls. Fairly straight growth lines, although somewhat arched on the venter (Fig. 6A). Poorly preserved ceratitic suture lines, with elongated lobes, broad rounded saddles and denticulated ventral lobes (cf., Fig. 7).
Discussion. This taxon is assigned to the family Prionitidae due to the striking, shared characters: compressed shell, narrow umbilicus with rounded umbilical shoulder and high-angled oblique umbilical wall becoming steeper towards the body chamber. The specimens differ from Arctoprionites (Frebold, 1930) and Hemiprionites Spath, 1929 by their lack of tabulate venter, and from Anasibirites Mojsisovics, 1896 andWasatchites Mathews, 1929 by the ornamentation style. Specimen PMO 227.988 is the best preserved and likely the most mature specimen. Its internal features were observed through CT scanning, and a very high-quality preservation of the internal septa was revealed (Fig. 7), but no further information on the possible constriction Genus Arctoceras Hyatt, 1900 Type species. Ceratites polaris Mojsisovics, 1896 Arctoceras erebori sp. nov. Fig. 6E Description. Subinvolute and extremely discoidal shell with trapezoidal whorl section. Narrow and fairly deep umbilicus with high and very steep (c. 90°) umbilical could be obtained. Therefore, this feature cannot be interpreted as a constriction or healed wound with any degree of certainty. If it is a constriction, it may be unique to this prionitid, as such features have not been reported in other prionitids. The CT scanning analysis revealed that the inner whorls are filled with high X-ray density material (probably barite), and the septa are not visible in this region. The other collected specimens of Prionitid sp. do not show such unusual X-ray preservation.
The most difficult comparison has been with the genus Anasibirites. As discussed in the recent revision by Jattiot et al. (2015), this genus can be identified by its distinctive megastriae, which are ridge-like structures generated by discontinuities in secretion of the shell. Megastriae are asymmetric, superficial features, while ribs on the contrary are more symmetric and their formation also involves the deeper mantle layers. Specimens here described do not present megastriae, and possess smooth to slightly ribbed inner whorls. Thus, Prionitid sp. is not comparable to A. kingianus.
Because of the uncertainty of the assignment of our specimens to known prionitid taxa, the possibility of a new taxon has been considered. However, the lack of additional well-preserved material with new diagnostic features has led to an open classification. morphology and to the shape of the venter and whorls. The presence of strigation is not a new feature in the genus, as seen e.g., in A. strigatus Brayard & Bucher, 2008, where it is most prominent on the flanks, and A. tuberculatum (Smith, 1932); however, both species exhibit clearly different ornamentation. The studied specimens, moreover, bear weaker ornamentation and more irregular and less prominent distance between the folds than in A. blomstrandi (Lindström, 1865), which is the main arctoceratid recorded from the Smithian beds of Spitsbergen. This species (and its synonyms) has never been reported to exhibit a keel at any growth stage. It is possible that this keel-like structure on the holotype's body chamber is simply an artefact of preservation, e.g., compaction fracture.
Arctoceratids are usually a more typical component of middle Smithian faunas. At one locality on Ellesmere Island (Arctic Canada), Arctoceras is associated with Wasatchites (Tozer, 1961). The presence of arctoceratids in this level confirms that the vertical range of Arctoceras extends up into the Tardus Zone, as suggested by previous biostratigraphic research on Smithian faunas in Svalbard, as discussed above.
wall and sharply angled umbilical shoulder. Venter ranges from arched to broadly rounded. Specimen PMO 210.489 has what appears to be a keeled venter on the terminal part of the preserved body chamber but no ornamentation is visible on the venter other than strigation. Flatter flanks on the more compressed specimens. Further comments on ornamentation are prevented by the poor preservation of the specimens other than holotype. However, radial/gently sinuous folds on the flanks are observed on the best preserved specimens (e.g., PMO 227.985, Fig. 6E-F, which shows finer striae in between the folds). The folds become more distant towards the body chamber. Very well preserved suture lines were found only on specimen PMO 227.985 (Fig. 6G), with two elongated, prominently denticulated lateral lobes and rounded saddles. Denticulation very pronounced with bifurcation on the ventral lobe.
Discussion. The specimens here discussed are assigned to the genus Arctoceras mainly due to the umbilical