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New Osteostracans from the Lower Devonian Terrigenous Deposits of Podolia, Ukraine

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New Osteostracans from the Lower Devonian Terrigenous Deposits of Podolia, Ukraine New osteostracans from the Lower Devonian terrigenous deposits of Podolia, Ukraine VICTOR VOICHYSHYN Voichyshyn, V. 2006. New osteostracans from the Lower Devonian terrigenous deposits of Podolia, Ukraine. Acta Palaeontologica Polonica 51 (1): 131–142. New osteostracans (Agnatha, Osteostraci) Zenaspis dzieduszyckii sp. nov. and Wladysagitta janvieri gen. et sp. nov., as well as new form of Diademaspis are described from the Lower Devonian of Podolia (Ukraine). Among them Zenaspis dzieduszyckii sp. nov. differs from all other representatives of the genus by its smaller overall size and relatively larger or− bits. Wladysagitta janvieri is remarkable by having a rostral process. This new species together with “Cephalaspis” acutirostris Stensiö, 1932 from Old Red deposits of Shropshire (Great Britain) are proposed to be united under common generic name Wladysagitta. Since Diademaspis sp. is completely unknown in outline of its shield, it could not be named as new species, although it should be a really new form. New diagnostic features, such as relative width of the orbits, rela− tive width of the dorsal field, and the position of the posterior end of the lateral fields relatively to the margins of the headshield, are proposed to differentiate genera within the Zenaspididae. Probably, the rostral process of osteostracans should be considered as a multifunctional device having at least a sensory function for food−search, possibly hydrody− namic functions, or may play a role in scaring predators. Key words: Agnatha, Osteostraci, Devonian, Podolia, taxonomy, morphology. Victor Voichyshyn [[email protected]], Department of the Taxonomy of Modern and Fossil Biota, State Museum of Natural History NASU, Teatralna Str. 18, 79008, Lviv, Ukraine. Introduction leva member, is notable for the most diversity of agnathan species. The second zone (corresponding to the rest of the Podolia (Fig. 1A, B) is the only region in Ukraine where the Khmeleva member) has been still characterized insufficien− Lower Devonian deposits with remains of ichthyofauna tly because of not numerous and poorly preserved vertebrate come to the surface. During about 150 years of collecting fossils. And the third zone (apparently corresponding to the here the vertebrate fossils have been found in more than 90 Strypa member) contains several species which never have localities situated in outcrops along banks of the Dniester been recorded for the first zone (Voichyshyn 2001b). River and its northern tributaries, and in sandstone quarries. Findings of Podolian cephalaspids (Agnatha, Osteostraci) At present faunal list of Early Devonian agnathans and fishes were already mentioned more than 100 years ago (Alth 1874). from Podolia number 72 species, including 8 Thelodonti, 39 Despite that, they had been the subject of only a few studies Heterostraci, 19 Osteostraci, 4 Placodermi, 1 Acanthodii, until recently (Zych 1937; Pauča 1941; Balabai 1962). During and 1 Holocephali (Voichyshyn 2001a, modified). the last two decades regional Early Devonian faunas including In Podolia, Lower Devonian redbeds strata (the Old Red cephalaspids have been studied more intensively (Belles−Isles Formation or Dniester Series) are up to 1800 m thick and and Janvier 1984; Janvier 1985b, 1988; Afanassieva 1989, range from Lochkovian to Eifelian in age (Narbutas 1984; 1990, 1991; Afanassieva and Voichyshyn 1991; Voichyshyn Drygant 2000, 2003). In their lower part (Ustechko and 1994, 1998; Voichyshyn and Solodkyi 2004). These investi− Khmeleva members of the Dniester Series) they consist of gations revealed some depositional peculiarities of the Podo− multi−coloured, mainly red, fine−grained cross−bedding mas− lian representatives of this group. Specifically, Podolian ce− sive quartz sandstones and siltstones with seams of argillites phalaspids, except for one specimen of Parameteoraspis (Drygant 2000). (Afanassieva 1991), are linked to terrigenous sediments. Most Zych (1927), Brotzen (1933), and Balabai (1960) tried to of the 19 species known from Podolia are derived from the divide the Podolian section of Lower Devonian terrigenous first faunal zone of the Podolian Old Red, according to the deposits into biostratigraphical (faunal) zones. Among these subdivisions proposed by Brotzen (1933). This also applies to division schemes the Brotzen’s one should be regarded as the the material described below. In contrast, only a few speci− most suitable. It has three zones. The first of them, according mens of three species of the Benneviaspididae (Belles−Isles to Narbutas (1984), including the stratigraphical interval and Janvier 1984; Afanassieva 1989, 1990, 1991) have so far from the Ustechko member to the lowest part of the Khme− been reported from deposits referred to the third faunal zone. Acta Palaeontol. Pol. 51 (1): 131–142, 2006 http://app.pan.pl/acta51/app51−131.pdf 132 ACTA PALAEONTOLOGICA POLONICA 51 (1), 2006 sons and conclusions are exclusively based on the external RUSSIA Ternopil' morphology of the headshield. POLAND UKRAINE Measurements (Fig. 2).—A, distance between the pineal fo− ramen and the posterior margin of the median dorsal field; B, PODOLIA pre−pineal length of shield: distance between the anterior ROMANIA margin of the shield and the pineal foramen; C, post−pineal length of shield: distance between the pineal foramen and the BLACK SEA posterior margin of the shield; G, maximum length of lateral field: distance between the anterior and posterior ends of the Zalishchyky lateral field; L, maximum length of the shield; Lc, length of cornual processes: distance between the middle of the short− 48°50' 3km est line connecting the tip of the pectoral sinus to the lateral Dzhuryn margin of the shield and the tip of the cornual process; Lrc, distance between the anterior margin of the shield (without rostral process) and the tip of the cornual process; Ol, orbit length: distance between anterior and posterior margins of Ustechko the orbit; Omin, minimum distance between orbits; Os, orbit width: distance between lateral margins of the orbits; Q, dis− 25°30' 25°45' tance between the anterior margin of the shield and the nasohypophysial opening; S, maximum width of shield; Sd, maximum width of median dorsal field; Si, width of the ab− Dniester dominal part of the shield. Institutional abbreviations.—BM(NH), British Museum of Natural History; The Natural History Museum (London); Horodnytsia GMLSU, Geological Museum of Lviv State University, De− 48°40' partment of Geology (Lviv); SMNH, State Museum of Natu− ral History, National Academy of Sciences of Ukraine (Lviv). Fig. 1. Contours of Ukraine and the neighbouring countries (A), simplified map of Podolia with main cities and major tributaries of Dniester (B), local− ity map, showing position of the points of finds of the studied material (indi− Systematic palaeontology cated by an asterisk) (C). Class Osteostraci Lankester, 1868 Here, I describe three cephalaspid taxa from Podolia, two of Subclass Cornuata Janvier, 1985a which belong to the Zenaspidida. However, the third taxon, Order Zenaspidida Stensiö, 1958 which is referred here to a new genus, is represented by material that is not sufficiently well−preserved for allowing its systematic Family Zenaspididae Stensiö, 1958 assignment within the “horseshoe−shielded” osteostracans. Type genus: Zenaspis Lankester, 1870, see below. Diagnosis.—Zenaspidida of average to large size. The head− shield is dome−shaped and massive. The abdominal part of Material and methods the shield has a low, more or less developed, median dorsal crest. The hypophysial division of the nasohypophysial The material examined for this study was mainly collected opening is, as a rule, larger than the nasal division. The pi− by the author in 1992 and 1994 at the Ustechko outcrops, and neal plate is feebly developed or absent. The median dorsal in 1992 in quarries of Horodnytsia (Fig. 1). In addition, some field is broad. The lateral fields have a conspicuous dilata− specimens kept in the Museum of Lviv State University, De− tion (or widening) in their posterior parts reaching back− partment of Geology were also studied. No information wards no further than the proximal part of the dorsal surface about the collector(s) and date of collecting is available for of the cornual processes. The ornamentation includes either these specimens. single large tubercles, or groups of tubercles of different All the material was mechanically prepared, if necessary, size on tesserae. and cleaned, then measured, photographed with an Olympus Genera included.—Diademaspis Janvier, 1985, and, possi− C−2100 camera and studied under optical zoom. bly, Tegaspis Wängsjö, 1952. Podolian agnathans from terrigenous deposits typically Remarks.—The genus Tegaspis has several features in which do not exhibit traces of endoskeleton. Therefore, all compari− it appreciably differs from other zenaspidids, such as the VOICHYSHYN—NEW OSTEOSTRACANS FROM PODOLIA 133 shape of the headshield (its anterior margin and cornual pro− cesses), approximately same size of the two divisions of the 10 mm nasohypophysial opening, a comparatively narrow median Q B dorsal field, and the position of the posterior parts of the lat− Omin eral fields, which does not extend on the dorsal surface of Os cornual processes at all. G Ol L Genus Zenaspis Lankester, 1870 sensu Janvier A 1985b Lrc Type species: Cephalaspis salweyi Egerton, 1857, Lower Devonian of C Great Britain. Diagnosis.—Zenaspididae of moderate size (L = 90–130 mm). The width of the shield is greater than its length (L/S = Lc 0.6–0.7). The prepineal part of the shield is shorter than the postpineal one (B/C = 0.8–0.9). The cornual processes are Sd comparatively short. The abdominal part of the shield is short Si with a massive median dorsal crest. The hypophysial division of the nasohypophysial opening is somewhat larger than the S nasal division. The orbital openings are small (S/Os = 32–42) with a short distance between them (Omin/Os = 2.5–3.3). The median dorsal field is comparatively broad (index S/Sd is Fig. 2. Zenaspis salweyi (Egerton, 1857) (after Janvier 1985a: fig. 63B) with a designation of the shield measurements (after Afanassieva 1991, 9–11).
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