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УЧЕНЫЕ ЗАПИСКИ КАЗАНСКОГО УНИВЕРСИТЕТА. СЕРИЯ ЕСТЕСТВЕННЫЕ НАУКИ
2022, Т. 164, кн. 4 С.567-576
ISSN 2542-064X (Print) ISSN 2500-218X (Online)
ORIGINAL ARTICLE
UDC 567.31:551.734
doi: 10.26907/2542-064X.2022.4.567-576
FISH ASSEMBLAGES FROM THE MIDDLE-UPPER DEVONIAN OF THE MIDDLE URALS, RUSSIA
A.O. Ivanov
St. Petersburg State University, St. Petersburg, 199034 Russia Kazan Federal University, Kazan, 420008 Russia
Abstract
New assemblages of diverse fishes from the Givetian-Famennian were described in six sections in the Middle Urals: Pokrovskoe, Baronskaya, Sulem, Pershino, Yokva, and Vilva. Diademodus, Wellerodus, and Mimipiscis genera were discovered in the Urals for the first time. Data on the occurrence of Phoebodus species in the Pokrovskoe section were used to specify the boundary between the Phoebodus sophiae and Ph. latus phoebodontid zones. Four phoebodontid zones can be recognized in the sections of the Middle Urals.
Keywords: fishes, Devonian, Middle Urals
In the Urals region, fish assemblages have been recorded earlier in the Late Devonian of the South Urals [1-3] and the Famennian of the Polar Urals [4], but fishes from the Middle and Late Devonian of the Middle Urals are still poorly known, with only one Phoebodus species recently described [5].
A new collection of diverse and abundant fishes from the Givetian-Famennian deposits of the Middle Urals is studied in this paper. The fish microremains were obtained from the processed conodont samples (coll. by A.Z. Bikbaev and M.P. Snigireva, Institute of Geology and Geochemistry, Ekaterinburg) and micrographed using Hitachi S-3400N, Cambridge CamScan-4, and Tescan VEGA-II XMU scanning electron microscopes. All specimens described here are housed at the Paleontological Museum of St. Petersburg State University (PMSPU).
The new assemblages of fishes under study come from six sections (localities) of the Middle Urals: Pokrovskoe, Baronskaya, Sulem, Pershino, Yokva, and Vilva. Five sections (Pokrovskoe, Baronskaya, Sulem, Pershino, and Yokva) are located in the Sverdlovsk Region, and one (Vilva) in the Perm Region.
The Pokrovskoe section is in the vicinity of the village of Pokrovskoe on the Bob-rovka River (left tributary of the Irbit River), 10 km west of the town of Artemovsky. The Givetian and Frasnian deposits of the Vysotinkian and Brodovkian regional stages (RS) are exposed on both banks of the river for over 2 km. The Givetian deposits of
Introduction
Geological Setting
the Vysotinkian RS include sandy limestones with clay interbeds, while the deposits of the Brodovkian RS, corresponding to the Lower Frasnian and possibly Upper Givetian, contain unbedded and massive reef limestones [6-8].
The Baronskaya section is situated on the right bank of the Mezhevaya Utka River (right tributary of the Chusovaya River), 6.5 km from the river mouth. The Upper Fras-nian deposits of the Askynian RS and the Lower Famennian deposits of the Makarovian RS are represented by limestones, dolomitized limestones with intercalations of marls, mudstones, clayey and calcareous-siliceous shales [9, 10].
The Sulem section is on the left bank of the Sulem River, near the river mouth, 15 km south of the Baronskaya section. The Upper Frasnian deposits of the Askynian RS and the Lower Famennian deposits of the Makarovian RS are formed by micritic and dolomitized limestones with intercalations of marls and clayey limestones [11].
The Yokva section is on the right bank of the Chusovaya River, 3 km downstream of the village of Yokva. The section comprises the Upper Frasnian-Tournaisian deposits. The Upper Frasnian deposits of the Askynian RS and the Famennian deposits of the Makarovian-Lytvian RS of the section consist of limestones and dolomites with diverse fossils [12].
The Pershino section is located near the village of Pershino on the Rezh River, 10 km north of the town of Rezh. The massive limestones of the Upper Frasnian deposits of the Gubinskian RS and the detrital limestones and siltstones of the Lower Famennian deposits of the Shameyskian RS are exposed in the section [10, 13].
The Vilva (Krivoy Rog) section is located on the left bank of the Vilva River, 2 km southeast of the town of Gremyachinsk. The deposits of the section belong to the Fras-nian Sargaevian, Domanikian, Mendymian, Askynian RS and Lower Famennian Makarovian RS. The Upper Frasnian and Lower Famennian deposits are composed of limestones, dolomites, and marls with various invertebrate fossils [14].
Fish Assemblages
Pokrovskoe section. The diverse and abundant fish remains (Fig. 1) of the Pok-rovskoe section were found at four levels: in the interval of the Givetian of the Upper Polygnathus varcus, Upper Klapperina disparilis, Skeletognathus norrisi, and Frasnian of the Upper Mesotaxis falsiovalis conodont zones (CZ). The fish assemblage of the Upper Po. varcus CZ is made up of the teeth of the chondrichthyans Phoebodus fastigatus and Ph. sophiae, as well as ptyctodontid plates (Fig. 1.25), acanthodiform scales, teeth of struniiform sarcopterygians, and actinopterygian scales. The fish remains from the Upper K. disparilis CZ are as follows: teeth of Phoebodus curvatus, Ph. fastigatus, Ph. Sophiae (Fig. 1.2-8), and Ph. sp.; chondrichthyan scales of "Ohiolepis", protacrodontid, ctena-canthid, and hybodontid types (Fig. 1.14, 16, 20, 21); uncompleted plates of ptyctodon-tids (Fig. 1.24) and undetermined placoderms; acanthodiform scales (Fig. 1.23); jaw fragments of osteolepiforms; teeth and jaw fragments of struniiforms; teeth, jaw fragments (Fig. 1.30), and scales of actinopterygians, including Moythomasia sp. The assemblage of the S. norrisi CZ includes: the teeth of Phoebodus curvatus, Ph. fastigatus, Ph. latus (Fig. 1.9, 10, 12, 13), and Ph. sp.; chondrichthyan scales of "Ohiolepis", protacrodontid, ctenacanthid, and hybodontid types (Fig. 1.15, 19, 22); plate fragments of undetermined placoderms; osteolepiform scales; struniiform teeth, fragments of jaws, and scales (Fig. 1.26, 28, 29); teeth, jaw fragments, and scales of actinopterygians,
Fig. 1. Fish remains from the Givetian-Frasnian of the Pokrovskoe section. 1, 11, 17, 18, 27, 33 - Upper M. falsiovalis CZ; 2-8, 14, 16, 20, 21, 30 - Upper K. disparilis CZ; 9, 10, 12, 13, 15, 19, 22-24, 26, 28, 29, 31, 32, 34-39 - S. norrisi CZ; 25 - Upper Po. varcus CZ. 1-4 -Phoebodus fastigatus Ginter et Ivanov, 1992, teeth, occlusal views: 1 - PMSPU 97-1, 2 -PMSPU 97-2, 3 - PMSPU 97-3, 4 - PMSPU 97-4. 5-7 - Ph. sophiae St. John et Worthen, 1875, teeth, occlusal views: 5 - PMSPU 97-5, 6 - PMSPU 97-6, 7 - PMSPU 97-7. 8-10 -Ph. curvatus Ivanov, 2021, teeth, occlusal (8a, 9, 10) and labial (8b) views: 8 - PMSPU 74-2, 9 - PMSPU 74-4, 10 - PMSPU 74-5. 11-13 - Ph. latus Ginter et Ivanov, 1995, teeth, occlusal
views: 11 - PMSPU 97-8, 12 - PMSPU 97-9, 13 - PMSPU 97-10. 14-22 - Chondrichthyan scales, crown views: 14, 15 - Ohiolepis type, 14 - PMSPU 97-11, 15 - PMSPU 97-12; 16-21 -Ctenacanth type, 16 - PMSPU 97-13, 17 - PMSPU 97-14, 18 - PMSPU 97-15, 19 - PMSPU 97-16, 20 - PMSPU 97-17, 21 - PMSPU 97-18; 22 - Tessera-like scale, PMSPU 97-19. 23 -Acanthodiformes indet., scale, oblique crown view, PMSPU 97-20. 24, 25 - Ptyctodontida indet.: 24 - upper tooth plate, lingual view, PMSPU 97-21; 25 - dermal clasping element, ventral view, PMSPU 97-22. 26-29 - Struniiformes indet.: 26 - jaw fragment, lingual view, PMSPU 97-23; 27 - parasymphysial whorl, lateral view, PMSPU 97-24; 28, 29 - teeth, 28 -lateral view, PMSPU 97-25, 29 - lingual view, PMSPU 97-26. 30-32, 39 - Actinopterygii indet.: 30 - jaw fragment, labial view, PMSPU 97-27; 31 - skull bone, external view, PMSPU 97-28; 32 - fulcral scale, external view, PMSPU 97-29; 39 - flank scale, external view, PMSPU 97-30. 33-36 - Moythomasia sp., flank scales, external views: 33 - PMSPU 97-31, 34 - PMSPU 97-32, 35 - PMSPU 97-33, 36 - PMSPU 97-34. 37, 38 -Mimipiscis sp., flank scales, external views: 37 - PMSPU 97-35, 38 - PMSPU 97-36. Scale bars: 1-23 - 100 ^m; 24-29 - 500 ^m; 30-39 - 300 ^m
including Moythomasia sp. and Mimipiscis sp. (Fig. 1.31, 32, 34-39). A number of fish remains were recovered from the Upper M. falsiovalis CZ: the teeth of Phoebodus fas-tigatus, Ph. latus (Fig. 1.1, 11), and Ph. sp.; chondrichthyan scales of protacrodontid and ctenacanthid types (Fig. 1.17, 18); osteolepiform scale fragments; struniiform teeth, fragments of jaws, and scales (Fig. 1.27); teeth, jaw fragments, and scales of actinoptery-gians, including Moythomasia sp. and Mimipiscis sp. (Fig. 1.33). The record of Omalo-dus teeth in the Pokrovskoe section is erroneous [8] - there are no known findings of omalodontids in the Urals.
Baronskaya section. An assemblage of diverse fishes from the Frasnian of the Lower Palmatolepis rhenana CZ of the Baronskaya section comprises the following remains: the teeth of the chondrichthyans Phoebodus bifurcatus (Fig. 2.1), Ph. curvatus, Ph. fastigatus (Fig. 2.2), Ph. latus, Ph. sp., Diademodus sp. (Fig. 2.3), and Protacodus sp. (Fig. 2.4); chondrichthyan scales of the ctenacanthid type; acanthodiform scales; struniiform teeth; scales of the actinopterygians Moythomasia sp. and Mimipiscis sp. The teeth of Phoebodus bifurcatus and the fragments of actinop-terygian scales were found in the Palmatolepis linguiformis CZ of the section, and a tooth of Cladodoides sp. was obtained from the Famennian of the Middle Palmatolepis triangularis CZ (Fig. 3).
Sulem section. The fish remains from the Sulem section were discovered in four levels: Frasnian of the Lower Pa. rhenana, Pa. linguiformis, Famennian of the Lower and Upper triangularis CZ. The teeth of Phoebodus bifurcates and Ph. latus (Fig. 2.5, 6) and the struniiform teeth occurred in the Lower Pa. rhenana CZ. The chondrichthyan scales of the ctenacanthid type, placoderm scale, acanthodiform and actinopterygian scales were collected in the Pa. linguiformis CZ. The chondrichthyan teeth of Cladodoides sp. and the unidentified cladodontomorph, actinopterygian bone fragments were found in the Lower Pa. triangularis CZ. The teeth of Wellerodus sp. (Fig. 2.7) and Cladodoides sp. (Fig. 2.8), struniiform teeth, and actinopterygian scales were obtained from the Upper Pa. triangularis CZ.
Pershino section. A tooth of Phoebodus curvatus was recorded in the Lower Pa. rhenana CZ of the Pershino section. The teeth of Protacodus sp., scales of the ctenacanthid type, and placoderm fragments occurred in the Lower Pa. triangularis CZ of same section. The assemblage of the Middle-Upper Pa. triangularis CZ of the section
Fig. 2. The chondrichthyan teeth (1-15, 17-22, 25-28) and scales (16, 23, 24) from the Late Devonian of the Baronskaya section (1-4), Sulem section (5-8), Vilva section (9-17), Pershino section (18-24), and Yokva section (25-28). 1-6, 9-15 - Lower Pa. rhenana CZ; 7, 8, 16 - Upper Pa. triangularis CZ; 17 - Uppermost Pa. crepida CZ; 18-24 - Middle-Upper Pa. triangularis CZ; 25 - Pa. linguiformis CZ; 26, 27 - Pa. postera-Middle Pa. expansa CZ; 28 - Middle Pa. expansa-Middle S. praesulcata CZ. 1, 9 - Phoebodus fastigatus Ginter et Ivanov, 1992, occlusal views: 1 - PMSPU 97-37, 9 - PMSPU 97-45. 2, 6, 10 - Phoebodus bifurcatus Ginter et Ivanov, 1992, occlusal views: 2 - PMSPU 97-38, 6 - PMSPU 97-42, 10 - PMSPU 97-46. 3, 13 -Diademodus sp.: 3 - oblique occlusal view, PMSPU 97-39; 13 - occlusal (13a) and oblique lateral (13b) views, PMSPU 97-49. 4, 15, 21, 22, 25 - Protacodus sp., occlusal (4, 15, 21) and labial (22, 25) views: 4 - PMSPU 97-40, 15 - PMSPU 97-51, 21 - PMSPU 97-56, 22 - PMSPU 97-57, 25 -PMSPU 97-60. 5, 11 - Phoebodus latus Ginter et Ivanov, 1995, occlusal views: 5 - PMSPU 97-41, 11 - PMSPU 97-47. 7 - Wellerodus sp., occlusal (7a) and labial (7b) views, PMSPU 97-43. 8 -Cladodoides sp., occlusal view, PMSPU 97-44. 12 - Phoebodus curvatus Ivanov, 2021, occlusal view, PMSPU 97-48. 14 - Protacrodontidae indet., occlusal view, PMSPU 97-50. 16, 23 - Ctena-canth type scales, crown views: 16 - PMSPU 97-52, 23 - PMSPU 97-58. 17 - cf. Cladodoides sp., oblique labial view, PMSPU 97-53. 18 - Symmoriiformes indet., labial view, PMSPU 97-54. 19 -Phoebodus typicus Ginter et Ivanov, 1995, occlusal view, PMSPU 97-54. 20 - Squatinactis sp., occlusal view, PMSPU 97-55. 24 - Protacrodontid type scale, crown view, PMSPU 97-59. 26 -Thrinacodus tranquillus Ginter, 2000, oblique occlusal view, PMSPU 97-61. 27 - Squatinactis caudispinatus Lund et Zangerl, 1974, labial (27a) and occlusal (27b) views, PMSPU 97-62. 28 -Thrinacodus ferox (Turner, 1982), oblique occlusal view, PMSPU 97-63. Scale bars: 300 цт
Fig. 3. Distribution of fish taxa in the sections of the Middle Urals. List of conodont zones:
Polygnathus timorensis, Polygnatus rhenanus, Polygnathus varcus, Polygnathus ansatus, "Ozarkodina" semialternans, Schmidtognathus hermanni, Polygnathus cristatus, Polygnathus cristatus ectypus, Klapperina disparilis, Polygnathus dengleri sagitta, Polygnathus dengleri dengleri, Skeletognathus norrisi, Ancyrodella rotundiloba pristina, Ancyrodella rotundiloba soluta, Ancyrodella rotundiloba rotundiloba, Ancyrodella rugosa, Mesotaxis falsiovalis, Pal-matolepis transitans, Palmatolepis punctata, Ancyognathus primus, "Ozarkodina" nonaginta, Palmatolepis housei, Palmatolepis proversa, Palmatolepis plana, Palmatolepis hassi, Palmat-olepis feisti, Palmatolepis jamieae, Palmatolepis winchelli, Palmatolepis bogartensis, Palmat-olepis rhenana, Palmatolepis linguiformis, Palmatolepis ultima, Palmatolepis subperlobata, Palmatolepis triangularis, Palmatolepis delicatula platys, Palmatolepis minuta minuta, Pal-matolepis crepida crepida, Palmatolepis crepida, Palmatolepis termini, Palmatolepis glabra prima, Palmatolepis glabra pectinata
includes: the teeth of Phoebodus typicus (Fig. 2.19), Squatinactis sp. (Fig. 2.20), Protacodus sp. (Fig. 2.21, 22), and Symmoriiformes indet. (Fig. 2.18); scales of the cte-nacanthid and protacrodontid types (Fig. 2.23, 24); fragments of ptyctodontid plates; scales ofMimipiscis sp. and bone fragments of actinopterygians.
Yokva section. The fish remains in the Yokva section were found in two levels of the Upper Frasnian and two levels of Upper and Uppermost Famennian. A tooth of Phoebodus sp. was collected from the Upper Pa. rhenana CZ. The teeth of Protacodus sp. (Fig. 2.24) were found in the Pa. linguiformis CZ. The teeth of Thrinacodus ferox, Th. tranquillus (Fig. 2.26), Squatinactis caudispinatus (Fig. 2.27), along with scales of
the ctenacanthid type, placoderm fragments, actinopterygian teeth were recorded in the Palmatolepis postera-Middle Palmatolepis expansa CZ interval. The teeth of Thrinacodus ferox (Fig. 2.28), Squatinactis caudispinatus, and actinopterygian teeth were reported from in the Middle Palmatolepis expansa-Middle Siphonodella praesul-cata CZ interval.
Vilva section. Diverse fish fossils were recorded in the Lower and Upper Fras-nian, Lower Famennian of the Vilva section (Fig. 3).
The anterior supragnathal plate of Plourdosteus sp., plate fragments of arthrodires and bothriolepids, osteolepiform scales were found in the Palmatolepis transitans CZ. The assemblage of the Lower Pa. rhenana CZ comprises the teeth of Phoebodus bifurcatus (Fig. 2.10), Ph. curvatus (Fig. 2.12), Ph. fastigatus (Fig. 2.9), Ph. latus (Fig. 2.11), Wellerodus sp., Diademodus sp. (Fig. 2.13), Protacodus sp. (Fig. 2.15), Protacrodontidae indet. (Fig. 2.14), and stethacanthid, as well as chondrichthyan scales of the ctenacanthid type, fragments of ptyctodontid and bothriolepid plates, struniiform teeth, osteolepiform scales, and actinopterygian teeth and scales. The fragments of placoderm plates, acanthodiform scales, fragments of struniiform scales, and scales of Moythomasia sp. were found in the Lower Palmatolepis triangularis CZ. A tooth of Protacodus sp., scales of the ctenacanthid type (Fig. 2.16), acanthodiform scales, fragments of struniiform scales, scales of Mimipiscis sp. were discovered in the Upper Pa. triangularis CZ. The fish remains from the Lower Palmatolepis crepida CZ include the chondrichthyan scales of the ctenacanthid and hybodontid types, acanthodiform scales, struniiform teeth, osteolepiform scales, and actinopterygian toothed plates. The tooth of cf. Cladodoides sp. (Fig. 2.17) was collected in the Uppermost Pa. crepida CZ.
Conclusions
The fish assemblages from the Givetian-Famennian of the Middle Urals are taxo-nomically diverse and represented by widely distributed taxa. Among chondrichthy-ans, phoebodontids are dominant. Some taxa, such as Diademodus, Wellerodus, and Mimipiscis, were reported for the first time from the Urals. Data on the distribution of Phoebodus species in the Givetian-Frasnian of the Pokrovskoe section allow updating the boundary between the Phoebodus sophiae and Ph. latus phoebodontid zones [15]. The Phoebodus sophiae Zone corresponds to the Lower Po. varcus-K. disparilis CZ and the Ph. latus Zone to the Lower M. falsiovalis-Palmatolepis jamieae CZ. The following phoebodontid zones can be recognized in the sections of the Middle Urals: Phoebodus sophiae, Ph. latus, Ph. bifurcatus, and Ph. typicus.
Acknowledgments. My sincere thanks to A.Z. Bikbaev (Leading Engineer) and M.P. Snigireva (PhD in Geology and Mineralogy, Leading Engineer) from Zavaritsky Institute of Geology and Geochemistry, Ural Branch, Russian Academy of Sciences for providing the collection and the stratigraphic data, as well as to N.S. Vlasenko (Engineer, St. Petersburg State University) and R.A. Rakitov (PhD in Biology, Senior Scientist, Borissiak Paleontological Institute, Russian Academy of Sciences) for their assistance with SEM imaging.
The scientific research was performed at the Center for Geo-Environmental Research and Modeling (GEOMODEL) and the Centre for Microscopy and Microanaly-sis of Research Park of St. Petersburg State University.
This study was partly funded by the Russian Foundation for Basic Research (project
no. 20-05-00445a). This paper was supported by the Kazan Federal University Strategic
Academic Leadership Program (PRI0RITY-2030).
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Received August 29, 2022
Ivanov Alexander Olegovich, PhD in Geology and Mineralogy, Associated Professor, Department of Sedimentary Geology, Institute of Earth Sciences; Senior Researcher, Institute of Geology and Petroleum Technologies
St. Petersburg State University
Universitetskaya Emb., 7/9, St. Petersburg, 199034 Russia Kazan Federal University
ul. Kremlevskaya, 18, Kazan, 420008 Russia E-mail: IvanovA-Paleo@yandex.ru
ОРИГИНАЛЬНАЯ СТАТЬЯ
УДК 567.31:551.734 doi: 10.26907/2542-064X.2022.4.567-576
Комплексы рыб из среднего - верхнего девона Среднего Урала, Россия
А.О. Иванов
Санкт-Петербургский государственный университет, г. Санкт-Петербург, 199034, Россия Казанский (Приволжский) федеральный университет, г. Казань, 420008, Россия
Аннотация
Новые комплексы разнообразных рыб из живета - фамена описаны из шести разрезов Среднего Урала: Покровское, Баронская, Сулем, Першино, Йоква и Вильва. Представители Diademodus, Wellerodus и Mimipiscis впервые отмечены на Урале. Распространение видов Phoebodus в разрезе Покровское позволяет уточнить положение границы фебодонтидных зон Phoebodus sophiae и Ph. latus. Полученные результаты свидетельствуют о том, что в разрезах Среднего Урала могут быть прослежены четыре фебодонтидные зоны.
Ключевые слова: рыбы, девон, Средней Урал
Благодарности. Автор выражает благодарность сотрудникам ИГГ Уро РАН А.З. Бикбаеву (ведущий инженер) и М.П. Снигиревой (канд. геол.-минерал. наук, ведущий инженер) за предоставленную коллекцию и стратиграфические данные, а также Н.С. Власенко (инженер, СПбГУ)
и Р.А. Ракитову (канд. биол. наук, с.н.с., ПИН РАН) за помощь при работе на сканирующих электронных микроскопах.
Исследования проведены с использованием оборудования ресурсных центров «Геомодель» и «Микроскопии и микроанализа» Научного парка СПбГУ.
Работа выполнена за счет средств Программы стратегического академического лидерства Казанского (Приволжского) федерального университета (ПРИОРИТЕТ-2030) и при частичной поддержке РФФИ (проект № 20-05-00445а).
Поступила в редакцию 29.08.2022
Иванов Александр Олегович, кандидат геолого-минералогических наук, доцент кафедры осадочной геологии Института наук о Земле; старший научный сотрудник Института геологии и нефтегазовых технологий
Санкт-Петербургский государственный университет
Университетская наб., д. 7/9, г. Санкт-Петербург, 199034, Россия Казанский (Приволжский) федеральный университет ул. Кремлевская, д. 18, г. Казань, 420008, Россия E-mail: IvanovA-Paleo@yandex.ru
<For citation: Ivanov A.O. Fish assemblages from the Middle-Upper Devonian of the Middle Urals, Russia. Uchenye Zapiski Kazanskogo Universiteta. Seriya Estestvennye Nauki, 2022, vol. 164, no. 4, pp. 567-576. doi: 10.26907/2542-064X.2022.4.567-576.
<Для цитирования: Ivanov A.O. Fish assemblages from the Middle-Upper Devonian of the Middle Urals, Russia // Учен. зап. Казан. ун-та. Сер. Естеств. науки. - 2022. - Т. 164, кн. 4. - С. 567-576. - doi: 10.26907/2542-064X.2022.4.567-576.
