[The published version of this article is available online: Zverkov, N. G. and Efimov, V.M. 2019. Revision of Undorosaurus, a mysterious Late Jurassic ichthyosaur of the Boreal Realm. Journal of Systematic Palaeontology, http://dx.doi.org/10.1080/14772019.2018.1515793] Revision of Undorosaurus, a mysterious Late Jurassic ichthyosaur of the Boreal Realm Nikolay G. Zverkov1, 2, 3 & Vladimir M. Efimov4 1Geological Faculty, Lomonosov Moscow State University, Leninskie Gory 1, Moscow 119991, Russia 2Geological Institute of the Russian Academy of Sciences, Pyzhevsky lane 7, Moscow 119017, Russia 3Borissiak Paleontological Institute of the Russian Academy of Sciences, Profsoyuznaya st., 123, Moscow 117997, Russia; e-mail: [email protected] 4Undory Paleontological Museum, village of Undory, Ulyanovsk Region, 433312, Russia Recent study of ophthalmosaurid ichthyosaurs has brought us a number of new taxa, however, the validity of several ophthalmosaurid taxa from the Volgian (Tithonian) of European Russia still remains unclear, complicating the comparisons and in some cases affecting taxonomic decisions of new contributions. A revision of the type series of all three species of Undorosaurus, erected by Efimov in 1999, reveals the potential validity of two of them. This contradicts previous research, which concluded that only the type species, U. gorodischensis, is valid. Furthermore, examination of the holotype of Cryopterygius kristiansenae from the coeval strata of Svalbard show that it is synonymous with Undorosaurus gorodischensis sharing all diagnostic features of the species, especially those related to forelimb morphology: humerus with extensive anteroposteriorly elongate proximal end, poorly pronounced trochanter dorsalis and reduced deltopectoral crest; and ulna proximodistally elongate and not involved in perichondral ossification on its whole posterior edge. This supports the idea of intensive exchange of ichthyosaurs between the Middle Russian Sea and other Boreal seas in the Late Jurassic. In order to resolve the phylogenetic position of Undorosaurus within Ophthalmosauridae as well as relations of other ophthalmosaurids, a new dataset including 33 taxa and 106 characters, 23 of which are new, was compiled. The results of this analysis challenge all previous phylogenetic hypotheses for Ophthalmosauridae in a number of aspects including Undorosaurus spp. recovered deeply nested within Platypterygiinae as a sister group to derived platypterygiines. Keywords. Ichthyosaurs, Ophthalmosauridae, Undorosaurus, Upper Jurassic, Volgian, Tithonian Introduction Ophthalmosauridae is a highly derived clade of ichthyosaurs that appeared in the Middle Jurassic (Fernández 1999, 2003; Druckenmiller & Maxwell 2014) and dominated until the extinction of ichthyosaurs in the early Late Cretaceous (Fischer et al. 2016). In the last decade, new ophthalmosaurid taxa have been erected from the Arctic (Maxwell 2010; Druckenmiller et al. 2012; Roberts et al. 2014; Delsett et al. 2017) and Europe (Paparella et al. 2017; Tyborowski 2016), that have illuminated the previously underestimated taxonomic diversity of Late Jurassic ichthyosaurs. This ‘renaissance’ in the study of Late Jurassic ichthyosaurs was preceded by a conservative period when their diversity was considered to be ‘on the low side’ (Maisch & Matzke 2000, p. 90), and when Late Jurassic ichthyosaurs of the Northern Hemisphere were considered to be represented by only three genera: Ophthalmosaurus, and the incomparably rarer Brachypterygius and Nannopterygius (Maisch & Matzke 2000). Since 1997 several ophthalmosaurid taxa (three genera and twelve species) have been erected, based on materials from the Upper Jurassic (Volgian/Tithonian) of European Russia (Arkhangelsky 1997, 1998, 2001b; Efimov 1998, 1999 a, b). These genera were soon regarded as subjective junior synonyms of Ophthalmosaurus and Brachypterygius (Maisch & Matzke 2000; McGowan & Motani 2003), although the validity of most of Russian ichthyosaurs was questioned without first-hand examination (Storrs et al. 2000; Maisch & Matzke 2000; McGowan & Motani 2003; Maisch 2010). The first attempts to reassess some of these taxa have been undertaken only recently (Arkhangelsky & Zverkov 2014; Zverkov et al. 2015a). However, the type series of Undorosaurus has been awaiting a thorough required revision. Despite all the debates on its validity and poorly understood osteology (see e. g. Maisch & Matzke 2000; McGowan & Motani 2003; Maisch 2010) Undorosaurus is frequently used for comparative purposes when other Upper Jurassic ophthalmosaurids are discussed (e.g. Maxwell 2010; Druckenmiller et al. 2012; Roberts et al. 2014; Paparella et al. 2017; Delsett et al. 2017). The phylogenetic position of Undorosaurus is no less controversial. First attempts to include Undorosaurus in a phylogenetic context were undertaken only 15 years after the taxon was described (Arkhangelsky & Zverkov 2014; Roberts et al. 2014). The position of Undorosaurus sufficiently varied sufficiently in recovered phylogenies that the need of redescription of the type materials is clear. Therefore, some researchers preferred to avoid including Undorosaurus into phylogenetic analyses (Ji et al. 2016; Fisher et al. 2016), whereas others continued to consider this taxon, despite the absence of adequate data on its morphology (Maxwell et al. 2016; Paparella et al. 2017; Delsett et al. 2017; Moon 2017). This paper is a part of the ongoing project on taxonomic and phylogenetic revision of the Upper Jurassic ichthyosaurs of the Boreal Realm. The goals of this contribution are to redescribe the type materials of Undorosaurus, and to assess the position of this genus in a modern taxonomic and phylogenetic framework. Materials and methods The principal focus of this paper is the type materials of Undorosaurus gorodischensis, U. nessovi and U. khorlovensis, which are stored in Undory Palaeontological museum (UPM). Additionally NGZ has personally examined the holotype of Cryopterygius kristiansenae PMO 214.578, which is on display in the Natural History Museum, University of Oslo, Norway. • UPM EP-II-20(572), the holotype of Undorosaurus gorodischensis, is a relatively complete, disarticulated skeleton, including most of the skull, 27 vertebrae, ribs, pectoral and pelvic girdles and limbs. It was collected from Ulyanovsk Province, bank of the Volga River near Gorodischi village; Upper Jurassic, middle Volgian (Tithonian), Epivirgatites nikitini Ammonite Biozone. • UPM EP-II-23(744) is a fragmentary skeleton, with incomplete left mandibular ramus, stapes, several teeth, 36 vertebral centra including atlas-axis complex, rib fragments; fragmentary left coracoid and scapula, incomplete left forelimb. It was collected from thre bank of the Volga River near Gorodischi village, Ulyanovsk Province; Upper Jurassic, upper Volgian (Tithonian), Garniericeras catenulatum Ammonite Biozone. • UPM EP-II-21(1075) is an incomplete skeleton comprising fragments of the premaxilla and nasal, basioccipital, mandibular fragments, atlas-axis and 17 presacral vertebrae, fragments of the coracoids and scapulae, incomplete left forelimb and fragments of right forelimb. It was collected from the bank of the Volga River near Gorodischi village, Ulyanovsk Province; Upper Jurassic, middle Volgian (Tithonian), Epivirgatites nikitini Ammonite Biozone. • UPM EP-II-22(1073), an incomplete skeleton, comprises: fragmentary left nasal, left prefrontal, basioccipital, opisthotic, and stapes; complete left and partial right quadrates; mandibular fragments, hyoids, atlas-axis and 17 vertebrae, rib fragments, right coracoid and scapular fragments, incomplete humerus, radius, ulna, and autopodial elements. It was collected from the bank of the Volga River near Gorodischi village, Ulyanovsk Province; Upper Jurassic, middle Volgian (Tithonian), Epivirgatites nikitini Ammonite Biozone. • UPM EP-II-24(785), the holotype of U. nessovi, comprises a disarticulated skeleton, including basisphenoid, basioccipital (severely eroded), right stapes, quadrates, articular and surangular, hyoid, atlas-axis and 15 vertebrae, rib fragments, coracoid, scapulae, nearly complete right and fragmentary left forelimbs. It was collected from the bank of the Volga River near Slantsevy Rudnik village, Ulyanovsk Province; Upper Jurassic, middle Volgian (Tithonian), Virgatites virgatus Ammonite Biozone. • UPM EP-II-27(870), the holotype of U. khorlovensis, comprises fragments of the snout, 3 vertebrae, incomplete scapulae, humeri, radius, femur fragment. It was collected at a phosphate mine near Khorlovo village, Voskresensk District, Moscow Province; Upper Jurassic, middle Volgian (Tithonian), Virgatites virgatus Ammonite Biozone. • YKM 44028-7 is an articulated left forelimb in matrix collected at the bank of the Volga River near ‘Detskiy sanatorium’, Ulyanovsk province; Upper Jurassic, middle Volgian (Tithonian), Epivirgatites nikitini Ammonite Biozone. • PMO 214.578, the holotype of Cryopterygius kristiansenae, is a nearly complete skeleton, from Svalbard, Norway; Upper Jurassic, middle Volgian (middle Tithonian), Crendonites anguinus Ammonite Biozone. Comments on the geological setting Because of high faunal provincialism among marine invertebrates during the latest Jurassic and earliest Cretaceous, independent regional marine stages are often used. In the so-called Pan-Boreal Superrealm, the Volgian and Ryazanian stages are used instead of the Tithonian and Berriasian international units. Although the bases of the Tithonian and Volgian stages