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УЧЕНЫЕ ЗАПИСКИ КАЗАНСКОГО УНИВЕРСИТЕТА. СЕРИЯ ЕСТЕСТВЕННЫЕ НАУКИ
2024, Т. 166, кн. 3 С.548-556
ISSN 2542-064X (Print) ISSN 2500-218X (Online)
ORIGINAL ARTICLE
UDC 567:551.734
doi: 10.26907/2542-064X.2024.3.548-556
Fish Assemblage from the Givetian (Middle Devonian) of the Kuznetsk Basin, Russia
A.O. Ivanov ab, N.G. Izokh c
aSt. Petersburg State University, St. Petersburg, 199034 Russia bKazan Federal University, Kazan, 420008 Russia cTrofimuk Institute of Petroleum Geology and Geophysics, Siberian Branch, Russian Academy of Sciences, Novosibirsk, 630090 Russia
A diverse fish assemblage was reported from the Mazalovskiy Kitat and Alchedat beds (Givetian, Middle Devonian) of the Kuznetsk Basin (Siberia, Russia). Ptyctodontid and acan-thothoracid placoderms; chondrichthyans such as the phoedodontiforms Phoebodus fastigatus and Ph. sophiae, the omalodontiform Omalodus grabaui, the cladodontomorph Cladodoides, euselachian Protacrodus, "Ohiolepis, and "Cladolepis"; Cheiracanthoides and acanthodiform acanthodians; actinopterygian Moythomasia; and struniiform sarcopterygians occur in the assemblage. The teeth of phoebodontiforms and omalodontiform are dominant among the fish mi-croremains. The assemblage contains widely distributed taxa. The occurrences of Cladodoides and Protacrodus in the Givetian of Kuznetsk Basin are the oldest in the world.
Keywords: fishes, Middle Devonian, Kuznetsk Basin, Siberia
The fish microremains earlier have been recorded in the Givetian Mazalovskiy Kitat and Alchedat formations of Kuznetsk Basin, Siberia [1]. The placoderm "Ptyctodus" sp.; the chondrichthyans Phoebodus floweri Wells, 1944, Ph.? bryanti Wells, 1944, Protacrodus sp., Cladodus sp., and Ohiolepis sp.; the acanthodians Devononchus cf. laevis Gross and Nostolepis? sp. A.; the sarcopterygians Laccognathus sp. and Glyptolepis sp.; actinopterygians Moythomasia durgaringa Gardiner et Bartram, 1977, "M." antiqua (Williams, 1886), and Kentuckia hlavini Dunkle, 1964 were reported from three sections of these formations. Subsequently, Phoebodus floweri was redescribed as Ph. fastigatus Ginter et Ivanov, 1992, and Ph.? bryanti - as Omalodus bryanti Wells, 1944 [2], but O. bryanti now is a junior synonym of O. grabaui (Hussakof et Bryant, 1918) [3].
In this article, the collection of new and known specimens from the Mazalovskiy Kitat (Icriodus brevis conodont Zone (CZ) or Polygnathus timorensis standard CZ) and Alchedat (Icriodus difficiis CZ or Polygnathus varcus - "Ozarkodina" semialternans standard CZ) beds (Mazalovskokitatian Regional Stage, Givetian) was studied.
Abstract
Introduction
1. Material and Methods
Fish microremains were collected from the terrigenous-carbonate deposits of three sections (1/89, E8211, and E8212) near Anzhero-Sudzhensk, in the (Mazalovskiy) Kitat River basin (Kemerovo Region, Western Siberia) (Fig. 1).
Fig. 1. Maps showing the localities studied for the Givetian fishes
Section 1/89, located about 3 km upstream from the village of Lebedyanka, is an outcrop on the left bank of the Kitat River. Here, the fish microremains were found in one sample (1/89-1-1) from the limestones of the Mazalovskiy Kitat beds, which also contained abundant stromatoporoids, corals, trilobites, ostracods, brachiopods, crinoids, and conodonts [1]. Sections E8211 (3/89) and E8212 (2/89) are situated on the right bank of the Alchedat River, at the southern margin of the village of Leb-edyanka [1, 4]. Section E8211 is a small abandoned quarry exposing the deposits of the Mazalovskiy Kitat beds and the lower part of the Alchedat beds (Fig. 2). The fish microremains were collected from five samples of the Alchedat beds. Section E8212 exposes the deposits of the upper part of the Alchedat beds, with the fish microremains found in four samples (Fig. 2).
The fish microremains were obtained from the processed conodont samples and photomicrographed using scanning electron microscopy (S-3400N (Hitachi, Japan), Cambridge CamScan-4 (Cambridge Instruments, UK), and VEGA-II XMU (Tescan, Czech Republic)). The described specimens are housed at the Paleontological Museum of St. Petersburg State University (PMSPU).
2. Fish Assemblage
The fish microremains under study include placoderm plate fragments; chondrich-thyan teeth and scales; acanthodian jaw fragment and scales; actinopterygian teeth, tooth plates, jaw fragments, and scales; and sarcopterygian teeth and scale fragments.
Fig. 2. Distribution of fish taxa in sections E8211 (3/89) (a) and E8212 (2/89) (b). Stratigraphic columns according to [9]. Abbreviations: Bed no. - bed number, Con. zone - conodont zone, R. Stage - regional stage. Legend: 1 - massive limestones, 2 - bedded limestones, 3 - clayey limestones 4 - sandstones, 5 - mudstones, 6 - tabulat corals, 7 - correlation lines
The placoderm remains are represented by the plate fragments of Ptyctodon-tida indet. and the tesserae of Acanthothoraci indet. (Fig. 3.1), which resemble the tesserae of Ohioaspis [5]. The chondrichthyan microremains are the teeth of the phoebodontiforms Phoebodus fastigatus (Figs. 4.5-7) and Ph. sophiae (Figs. 4.1-4); the omalodontiform Omalodus grabaui (Figs. 4.8-12); the cladodontomorph Cladodoides sp. (Fig. 4.13); and the euselachian Protacodus sp. (Fig. 4.14). They are also the scales of "Ohiolepis" (Figs. 4.18, 19), "Cladolepis", ctenacanthid (Figs. 4.15, 16), and protacrodontid types (Fig. 4.17), as well as unidentified tessera-like scales (Fig. 4.20). The acanthodian scales belong to Acanthodiformes indet. (Fig. 3.3) and Cheiracanthoides sp. (Fig. 3.4), resembling the scales of Cheiracanthoides comptus Wells, 1944 [6]. The acanthodian jaw fragment (Fig. 3.2) is similar to the dentigerous jaws of some ischnacanthiforms [7, 8]. The actinop-terygian scales (Figs. 3.5, 6) were attributed to Moythomasia sp., other remains (Figs. 3.7, 8) are undetermined. The sarcopterygian microremains are represented by struniiform teeth and unidentified scale fragments. Among fish microremains, the teeth of phoebodontiforms and omalodontiform are dominant.
Fig. 3. Fish microremains from the Givetian (Alchedat beds, Icriodus difficilis CZ) of the Kuznetsk Basin: 1 - sample 3/89-3-21, 2-5- sample 3/89-3-191, 6 - sample 3/89-3-17, 7, 8 -sample 2/89-5-12. 1 - Placoderm Acanthothoraci indet. (cf. Ohioaspis), tessera, crown view, PMSPU 112-18. 2 - 4 - acanthodian remains: 2 - dentigerous jaw fragment, occlusal view, PMSPU 112-19; 3 - acanthodiform scale, crown view, PMSPU 112-20; 4 - Cheiracanthoides sp., scale, oblique crown view, PMSPU 112-21. Actinopterygian remains: 5, 6 - Moythomasia sp., flank scales, external views, 5 - PMSPU 112-22, 6 - PMSPU 112-23; 7 - jaw fragment, labial view, PMSPU 112-24; 8 - tooth plate, occlusal view, PMSPU 112-25. Scale bars: 200 ^m
The results obtained demonstrate that the fish assemblage from section 1/89 of the Lower Givetian Mazalovskiy Kitat beds comprises Phoebodus fastigatus, Ph. sp., "Ohiolepis", "Cladolepis", Acanthodiform indet., and Moythomasia sp. The fishes from the lower part of the Middle Givetian Alchedat beds of section E8211 were assigned to Ptyctodontidae indet., Acanthothoraci indet. (cf. Ohioaspis), Placodermi indet., Phoebodus fastigatus, Ph. sophiae, Omalodus grabaui, Cladodoides sp., Protacrodus sp., "Ohiolepis", "Cladolepis", Chondrichthyes indet., Cheiracanthoides sp., Acanthodiformes indet., Acanthodii indet., Moythomasia sp., Actinopterygii indet., and Sarcopterygii indet. (Fig. 2, a). The upper part of the Alchedat beds of section E8212 contains Placodermi indet., Phoebodus sp., "Ohiolepis", Chondrichthyes indet., Acanthodii indet., Actinopterygii indet., and Struniiformes indet. (Fig. 2, b).
Considering the above, the diverse fish assemblage from the Givetian of the Kuznetsk Basin includes widely distributed taxa. Besides the described assemblage, the Givetian species Phoebodus sophiae is known from the Middle varcus-disparilis CZ of northeastern Australia; central Iran; Southern Mauritania; the Holy Cross Mountains of Poland; the Rhenish Slate Mountains of Germany; Spain; Portugal; the US states of Iowa, Indiana, and New York [10]; as well as from the Upper varcus-Upper disparilis CZ of the Middle Urals in Russia. Ph. fastigatus has been reported from the Givetian-Frasnian (interval of varcus-rhenana CZ) of the Kuznetsk Basin and the Middle and Southern Urals of Russia; Poland; Spain; the US states of Iowa, Indiana, New York, and Utah; Western Australia; China; Mauritania; Morocco [10]; and possibly from the hemiansatus CZ of the Lower Givetian [11, 12]. The phoedo-dontiform chondrichthyans first appeared in the Givetian and became very diverse in the Late Devonian.
Fig. 4. Chondrichthyan microremains (1-14 - teeth and 15-20 - scales) from the Givetian of the Kuznetsk Basin: 1-4, 6-15, 17-20 - Alchedat beds, Icriodus difficilis CZ: 1, 4, 6, 12 - sample E8211-13/3; 2, 3, 7-9, 11, 13, 14, 17, 19, 20 - sample 3/89-3-19; 10, 15 - sample 3/89-3-25;
18 - sample 2/89-5-12; 5, 16 - Mazalovskiy Kitat beds, Icriodus brevis CZ, sample 1/89-1-1. 1-4 - Phoebodus sophiae St. John et Worthen, 1875, occlusal (1, 2a, 3, 4) and oblique labial (2b) views, 1 - PMSPU 112-1, 2 - PMSPU 112-2, 3 - PMSPU 112-3, 4 - PMSPU 112-4. 5-7 - Ph. fastigatus Ginteret Ivanov, 1992, occlusal views, 5 - holotype, PMSPU 5-1, 6 -PMSPU 112-5, 7 - PMSPU 112-6. 8-12 - Omalodus grabaui (Hussakof et Btyant, 1918), labial (8-10) and lingual (11, 12) views, 8 - PMSPU 7-4, 9 - PMSPU 112-7, 10 - PMSPU 112-8, 11 - PMSPU 7-3, 12 - PMSPU 112-9. 13 - Cladodoides sp., occlusal (13a), lingual (13b), and oblique labial (13c) views, PMSPU 112-10. 14 - Protacrodus sp., lingual (14a), occlusal (14b), oblique labial (14c), and oblique lateral (14d) views, PMSPU 112-11. 15, 16 - ctenacanthid type, crown views, 15 - PMSPU 112-12, 16 - PMSPU 112-13. 17 - protacrodontid type, crown view, PMSPU 112-14. 18, 19 - "Ohiolepis" type, crown views, 18 - PMSPU 112-15,
19 - PMSPU 112-16. 20 - tessera-like scale, crown view, PMSPU 112-17. Scale bars: 200 ^m
Omalodus grabaui was recorded in the Middle Givetian-Lower Frasnian of Poland; Spain; the Kuznetsk Basin of Russia; Morocco; the US states of New York,
Iowa, Indiana, Kentucky, and Ohio [3]. Cladodoides has been recorded from the Upper Frasnian of Germany; the Upper Frasnian-Middle Famennian of Poland, Moravia, Morocco, and the South Urals of Russia; the Lower Famennian of France and Armenia; the Upper Frasnian-Middle Famennian of Iran; the Lower Famennian and Tournaisian of Western Australia; possibly from the Lower Carboniferous of Yakutia in Russia; and New Mexico in the USA [3]. This taxon has been also found in the Frasnian-Famennian of the Kuznetsk Basin. Protacodus is known from the Upper Devonian-Pennsylvanian of various regions. The occurrences of the cladodontomorph Cladodoides and the euselachian Protacodus in the Givetian of the Kuznetsk Basin are the oldest in the world.
Conclusions
The fish assemblage from the Givetian of the Kuznetsk Basin contains the mi-croremains of ptyctodontid and acanthothoracid placoderms; chondrichthyans such as the phoedodontiforms Phoebodus fastigatus and Ph. Sophiae, the omalodontiform Omalodus grabaui, the cladodontomorph Cladodoides, the euselachian Protacrodus, "Ohiolepis, and "Cladolepis"; Cheiracanthoides and acanthodiform acanthodians; the actinopterygian Moythomasia; and struniiform sarcopterygians. This fish assemblage includes widely distributed taxa such as Phoebodus fastigatus, Ph. sophiae, Omalodus grabaui, Cladodoides, Protacrodus, and Moythomasia. The occurrences of the clado-dontomorph Cladodoides and the euselachian Protacrodus in the Middle Givetian of the Kuznetsk Basin are the earliest records in the world.
Acknowledgments. We are grateful to N.S. Vlasenko (Engineer, St. Petersburg State University) and R.A. Rakitov (PhD in Biology, Senior Research Fellow, Bo-rissiak Paleontological Institute, Russian Academy of Sciences) for their assistance with SEM imaging. The scientific research was performed at the Center for Geo-Envi-ronmental Research and Modelling (GEOMODEL), the Center for X-ray Diffraction Studies, and the Centre for Microscopy and Microanalysis (Research Park of St. Petersburg State University).
This study was supported by the Kazan Federal University Strategic Academic Leadership Program (PRI0RITY-2030) and performed as part of the state assignment to the Russian Academy of Sciences (project no. FWZZ-2022-0003).
Conflicts of Interest. The authors declare no conflicts of interest.
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9. Izokh N.G. New Givetian conodonts of the genus Icriodus from the Kolyvan-Tom Folded Area (South of West Siberia). Paleontol. J., 2023, vol. 57, no. 5, pp. 553-559. https://doi.org/10.1134/S0031030123050027.
10. Ivanov A.O. A new phoebodontid shark from the Devonian of the Urals and the distribution of Phoebodus species. Paleontol. J., 2021, vol. 55, no. 3, pp. 301-310. https://doi.org/10.1134/S0031030121030096.
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Received February 28, 2024 Accepted April 11, 2024
Ivanov Alexandr Olegovich, PhD in Geology and Mineralogy, Associated Professor, Department of Sedimentary Geology, Institute of Earth Sciences; Senior Research Fellow, Laboratory of Stratigraphy of Oil and Gas Bearing Reservoirs, Institute of Geology and Petroleum Technologies St. Petersburg University
Universitetskaya nab., 7/9, St. Petersburg, 199034 Russia Kazan Federal University
ul. Kremlevskaya, 18, Kazan, 420008 Russia E-mail: IvanovA-Paleo@yandex.ru Izokh Nadezhda Georgievna, PhD in Geology and Mineralogy, Senior Research Fellow, Laboratory of Paleozoic Paleontology and Stratigraphy
Trofimuk Institute of Petroleum Geology and Geophysics, Siberian Branch, Russian Academy of Sciences
pr. Akademika Koptyugina, 3, Novosibirsk, 630090 Russia E-mail: IzokhNG@ipgg.sbras.ru
ОРИГИНАЛЬНАЯ СТАТЬЯ
УДК 567:551.734 doi: 10.26907/2542-064X.2024.3.548-556
Комплекс рыб из живетских отложений (средний девон) Кузнецкого бассейна
А.О. Иванов12, Н.Г. Изох3 'Санкт-Петербургский государственный университет, г. Санкт-Петербург, 199034, Россия 2Казанский (Приволжский) федеральный университет, г. Казань, 420008, Россия 3Институт нефтегазовой геологии и геофизики им. А.А. Трофимука СО РАН, г. Новосибирск, 630090, Россия
Аннотация
В статье рассмотрен комплекс разнообразных рыб из живетских мазаловско-китатских и ал-чедатских слоев Кузнецкого бассейна Сибири. В комплексе встречены тиктодонтидные и акан-тоторацидные плакодермы, хрящевые такие как фебодонтиды Phoebodus fastigatus и Ph. sophiae, омалодонтид Omalodus grabaui, кладодонтоморф Cladodoides, эвселяхий Protacrodus, "Ohiolepis, "Cladolepis"; акантоды Cheiracanthoides и акантодиформ; лучеперая Moythomasia и струниифор-мный лопастеперый. Зубы фебодонтид и омалодонтид преобладают в комплексе среди других микроостатков рыб. Комплекс содержит широко распространенные таксоны. Находки Cladodoides и Protacrodus в живете Кузбасса - древнейшие в мире.
Ключевые слова: рыбы, средний девон, Кузнецкий бассейн, Сибирь.
Благодарности. Авторы выражают благодарность 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», «Рентгенодифракционные методы исследования» и «Микроскопии и микроанализа» Научного парка СПбГУ.
Работа выполнена за счет средств Программы стратегического академического лидерства Казанского (Приволжского) федерального университета (ПРИОРИТЕТ-2030) и в рамках госзадания РАН (проект FWZZ-2022-0003).
Конфликт интересов. Авторы заявляют об отсутствии конфликта интересов.
Поступила в редакцию 28.02.2024 Принята к публикации 11.04.2024
Иванов Александр Олегович, кандидат геолого-минералогических наук, доцент, доцент кафедры осадочной геологии Института наук о Земле; старший научный сотрудник лаборатории стратиграфии нефтегазоносных резервуаров Института геологии и нефтегазовых технологий Санкт-Петербургский государственный университет
Университетская наб., д. 7/9, г. Санкт-Петербург, 199034, Россия Казанский федеральный университет
ул. Кремлевская, д. 18, г. Казань, 420008, Россия E-mail: IvanovA-Paleo@yandex.ru Изох Надежда Георгиевна, кандидат геолого-минералогических наук, старший научный сотрудник лаборатории палеонтологии и стратиграфии палеозоя
Институт нефтегазовой геологии и геофизики им. А.А. Трофимука СО РАН
просп. Академика Коптюга, д. 3, г. Новосибирск, 630090, Россия E-mail: izokhng@ipgg.sbras.ru
For citation: Ivanov A.O., Izokh N.G. Fish assemblage from the Givetian (Middle Devonian) of the Kuznetsk Basin, Russia. Uchenye Zapiski Kazanskogo Universiteta. Seriya Estestvennye Nauki, 2024, vol. 166, no. 3, pp. 548-556. https://doi.org/10.26907/2542-064X.2024.3.548-556.
Для цитирования: Ivanov A.O., Izokh N.G. Fish assemblage from the Givetian (Middle Devonian) of the Kuznetsk Basin, Russia // Учен. зап. Казан. ун-та. Сер. Естеств. науки. 2024. Т. 166, кн. 3. С. 548-556. https://doi.org/10.26907/2542-064X.2024.3.548-556.
