First record of malformed trilobites from the Silurian of Ukraine

This article presents the first documented case of malformed trilobite exoskeletons from the Silurian of Ukraine. Fossils of animals with injuries and pathologies are of significant palaeobiological value, as they provide essential insights into the evolution of predator-prey interactions, trophic n...

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Published in:Доповіді НАН України
Date:2025
Main Author: Dernov, V.S.
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Language:English
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Cite this:First record of malformed trilobites from the Silurian of Ukraine / V.S. Dernov // Доповіді Національної академії наук України. — 2025. — № 4. — С. 3-9. — Бібліогр.: 37 назв. — англ.

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First record of malformed trilobites from the Silurian of Ukraine / V.S. Dernov // Доповіді Національної академії наук України. — 2025. — № 4. — С. 3-9. — Бібліогр.: 37 назв. — англ.
1025-6415
https://nasplib.isofts.kiev.ua/handle/123456789/206611
565.393:551.733.3(477.43.)44
https://doi.org/10.15407/dopovidi2025.04.003
This article presents the first documented case of malformed trilobite exoskeletons from the Silurian of Ukraine. Fossils of animals with injuries and pathologies are of significant palaeobiological value, as they provide essential insights into the evolution of predator-prey interactions, trophic networks, animal behavior, and genetic anomalies. The material under study consists of two pygidia of Bumastus restevensis from the Rykhta Formation (Ludlow, Upper Silurian) in Khmelnytskyi Oblast (western Ukraine, the Volyn-Podillia Monocline). A comparative analysis of these malformed exoskeletons with similar specimens from other global localities reveals that they are unique, with no direct analogues. The injuries observed are interpreted as “predation scars,” indicative of unsuccessful predatory attacks. The distinctive morphology of these malformations suggests that they were most likely inflicted by cephalopods, arthropods, such as phyllocarid crustaceans and/or aquatic chelicerates, or fishes.
Досліджено перший задокументований випадок прижиттєво пошкоджених екзоскелетів трилобітів із силурійських відкладів України. Скам’янілості тварин з пошкодженнями та патологіями мають значну палеобіологічну цінність, оскільки дають змогу зрозуміти еволюцію взаємодій хижака і жертви, трофічних мереж, поведінки тварин та генетичних аномалій. Досліджений матеріал складається з двох пігідіїв Bumastus restevensis з рихтівської світи (лудлов, верхній силур) Хмельницької області (Волино-Подільська монокліналь). Порівняльний аналіз цих екзоскелетів з аналогічними зразками з інших місцезнаходжень світу показує, що вони є унікальними і не мають прямих аналогів. Виявлені пошкодження інтерпретуються як «шрами хижацтва», що свідчать про невдалі напади хижаків. Характерна морфологія цих ушкоджень дає підставу припустити, що вони, найімовірніше, були завдані головоногими молюсками, членистоногими, такими як філокариди та/або водні хеліцерові, або рибами.
I extend my sincere thanks to Drs Volodymyr Grytsenko and Halyna Anfimova (National Museum of Natural History of the NAS of Ukraine, Kyiv) for the opportunity to examine the trilobite specimens collected by the late Dr. Leonid Konstantynenko. I am also grateful to Dr. Tamara Nemyrovska (Institute of Geological Sciences of the NAS of Ukraine, Kyiv) for her valuable advice on the article. The author is sincerely grateful to anonymous reviewers whose comments and suggestions improved the quality of the final version of the manuscript.
en
Видавничий дім "Академперіодика" НАН України
Доповіді НАН України
Науки про Землю
First record of malformed trilobites from the Silurian of Ukraine
Перша знахідка прижиттєво пошкоджених трилобітів у силурійських відкладах України
Article
published earlier
institution Digital Library of Periodicals of National Academy of Sciences of Ukraine
collection DSpace DC
title First record of malformed trilobites from the Silurian of Ukraine
spellingShingle First record of malformed trilobites from the Silurian of Ukraine
Dernov, V.S.
Науки про Землю
title_short First record of malformed trilobites from the Silurian of Ukraine
title_full First record of malformed trilobites from the Silurian of Ukraine
title_fullStr First record of malformed trilobites from the Silurian of Ukraine
title_full_unstemmed First record of malformed trilobites from the Silurian of Ukraine
title_sort first record of malformed trilobites from the silurian of ukraine
author Dernov, V.S.
author_facet Dernov, V.S.
topic Науки про Землю
topic_facet Науки про Землю
publishDate 2025
language English
container_title Доповіді НАН України
publisher Видавничий дім "Академперіодика" НАН України
format Article
title_alt Перша знахідка прижиттєво пошкоджених трилобітів у силурійських відкладах України
description This article presents the first documented case of malformed trilobite exoskeletons from the Silurian of Ukraine. Fossils of animals with injuries and pathologies are of significant palaeobiological value, as they provide essential insights into the evolution of predator-prey interactions, trophic networks, animal behavior, and genetic anomalies. The material under study consists of two pygidia of Bumastus restevensis from the Rykhta Formation (Ludlow, Upper Silurian) in Khmelnytskyi Oblast (western Ukraine, the Volyn-Podillia Monocline). A comparative analysis of these malformed exoskeletons with similar specimens from other global localities reveals that they are unique, with no direct analogues. The injuries observed are interpreted as “predation scars,” indicative of unsuccessful predatory attacks. The distinctive morphology of these malformations suggests that they were most likely inflicted by cephalopods, arthropods, such as phyllocarid crustaceans and/or aquatic chelicerates, or fishes. Досліджено перший задокументований випадок прижиттєво пошкоджених екзоскелетів трилобітів із силурійських відкладів України. Скам’янілості тварин з пошкодженнями та патологіями мають значну палеобіологічну цінність, оскільки дають змогу зрозуміти еволюцію взаємодій хижака і жертви, трофічних мереж, поведінки тварин та генетичних аномалій. Досліджений матеріал складається з двох пігідіїв Bumastus restevensis з рихтівської світи (лудлов, верхній силур) Хмельницької області (Волино-Подільська монокліналь). Порівняльний аналіз цих екзоскелетів з аналогічними зразками з інших місцезнаходжень світу показує, що вони є унікальними і не мають прямих аналогів. Виявлені пошкодження інтерпретуються як «шрами хижацтва», що свідчать про невдалі напади хижаків. Характерна морфологія цих ушкоджень дає підставу припустити, що вони, найімовірніше, були завдані головоногими молюсками, членистоногими, такими як філокариди та/або водні хеліцерові, або рибами.
issn 1025-6415
url https://nasplib.isofts.kiev.ua/handle/123456789/206611
citation_txt First record of malformed trilobites from the Silurian of Ukraine / V.S. Dernov // Доповіді Національної академії наук України. — 2025. — № 4. — С. 3-9. — Бібліогр.: 37 назв. — англ.
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fulltext 3 ОПОВІДІ НАЦІОНАЛЬНОЇ АКАДЕМІЇ НАУК УКРАЇНИ НАУКИ ПРО ЗЕМЛЮ GEOSCIENCES ISSN 1025-6415. Допов. Нац. акад. наук Укр. 2025. № 4: 3—9 C i t a t i o n: Dernov V.S. First record of malformed trilobites from the Silurian of Ukraine. Dopov. Nac. akad. nauk Ukr. 2025. No. 4. P. 3—9. https://doi.org/10.15407/dopovidi2025.04.003 © Publisher PH «Akademperiodyka» of the NAS of Ukraine, 2025. Th is is an open access article under the CC BY-NC-ND license (https://creativecommons.org/licenses/by-nc-nd/4.0/) https://doi.org/10.15407/dopovidi2025.04.003 UDC 565.393:551.733.3(477.43.)44 V.S. Dernov, https://orcid.org/0000-0002-5873-394X Institute of Geological Sciences of the NAS of Ukraine, Kyiv, Ukraine National Museum of Natural History of the NAS of Ukraine, Kyiv, Ukraine E-mail: vitalydernov@gmail.com First record of malformed trilobites from the Silurian of Ukraine Presented by Academician of the NAS of Ukraine S.B. Shekhunova Th is article presents the fi rst documented case of malformed trilobite exoskeletons from the Silurian of Ukraine. Fossils of animals with injuries and pathologies are of signifi cant palaeobiological value, as they provide essential insights into the evolution of predator-prey interactions, trophic networks, animal behavior, and genetic anomalies. Th e material under study consists of two pygidia of Bumastus restevensis from the Rykhta Formation (Ludlow, Upper Silurian) in Khmelnytskyi Oblast (western Ukraine, the Volyn-Podillia Monocline). A comparative analysis of these malformed exoskeletons with similar specimens from other global localities reveals that they are unique, with no direct analogues. Th e injuries observed are interpreted as “predation scars,” indicative of unsuccessful predatory attacks. Th e distinctive morphology of these malformations suggests that they were most likely infl icted by cephalopods, arthropods, such as phyllocarid crustaceans and/or aquatic chelicerates, or fi shes. Key words: exoskeleton malformation, Trilobita, Ludlow, Volyn-Podillia Monocline, “predation scars”. Introduction. Th e fossilized remains of extinct animals exhibiting injuries and pathologies are essential for understanding the evolution of predator-prey interactions, trophic networks, animal behavior, congenital anomalies, growth defects, and related phenomena [1, 2]. Trilobites, due to their biomineralised exoskeletons, possess a high preservation potential and therefore serve as a signifi cant source of data on abnormalities in fossil arthropods [3—7 and references therein]. In Ukraine, trilobites have been reported from the Cambrian to Devonian strata of the Volyn- Podillia Monocline [8, 9], the Carboniferous deposits of the Lviv Palaeozoic Trough [10], the Do- nets Basin, and the Dnipro-Donets Depression [11, 12], as well as Permian strata of the Crimean Geosyncline [13, 14]. To date, only a single case of a malformed trilobite has been documented from Ukraine [10: Fig. 4J]. Typically, abnormal trilobites comprise around 0.05 % of a given popu- lation; however, their frequency can locally reach up to 2 %, or even 5 % in exceptional cases [15]. 4 ISSN 1025-6415. Dopov. Nac. akad. nauk Ukr. 2025. No. 4 V.S. Dernov Th e aim of this article is to investigate the palaeoecological signifi cance of the fi rst document- ed occurrence of malformed trilobite exoskeletons from the Silurian of Ukraine. Material and methods. Th e material under investigation consists of two moderately preserved pygidia of Bumastus restevensis Balashova, 1975 (specimens NMNHU-G 9763/02 and NMNHU- G 9763/09). Th ese specimens were collected by the late Dr. Leonid Konstantynenko (1943–2014) from the Rykhta Formation (Ludlow, Late Silurian) at outcrop No. 21 (outcrop numbering as per [16: Fig. 2]), located near the village of Hrynchuk in Khmelnytskyi Oblast. Th is locality lies within the Volyn-Podillia Monocline on the southwestern margin of the East European Craton (Fig. 1). Specimens NMNH-G 9763/02 and NMNH-G 9763/09 are housed in the Department of Geology of the National Museum of Natural History, National Academy of Sciences of Ukraine (Kyiv; acronym: NMNHU). Prior to photography, the fossils were coated with ammonium chlo- ride (NH4Cl). Th e terminology used in this article follows the traditional classifi cation of trilobite exoskeleton malformations [3–7]. Geological setting. Th e trilobite specimens examined in this study originate from the Late Silurian Rykhta Formation (Fig. 2). Th is formation extends across the Volyn-Podillia Monocline and the western part of the eastern slope of the Lviv Palaeozoic Trough. It comprises a 22—48 m thick sequence of argillaceous limestones interbedded with marlstone layers and thin-bedded dolomites. In the Dniester River region, the upper part of the formation consists of a succession of domerites and dolomites, up to 7.0 m thick. North of the reference section near the village of Rykhta (Khmelnytskyi Oblast), the formation is laterally replaced by dolomites of the Toky For- mation and domerites of the Velytsi Formation. Based on lithological characteristics, the forma- tion is subdivided into the Hrynchuk Subformation (argillaceous limestones and marlstones) and the Isakovytsi Subformation (thin-bedded dolomites) [16, 17]. Th e Rykhta formation contains a diverse and abundant marine biota, including calcareous algae, chitinozoans, corals, stromatopo- roids, brachiopods, bivalves, tentaculitids, and ostracods [16, 17 and references therein]. UKRAINE Neogene Palaeogene Lutsk Rivne Ternopil Khmelnytskyi Ivano-Frankivsk Uzhhorod Chernivtsi 50 km Lviv Sluch R. Prut R. Dniester R. Tysa R. Cretaceous Upper Palaeozoic Devonian Silurian Proterozoic Locality N Fig. 1. Schematic geological map of western Ukraine showing the geographical location of the fossil site with malformed trilobites 5ISSN 1025-6415. Допов. Нац. акад. наук Укр. 2025. № 4 First record of malformed trilobites from the Silurian of Ukraine Description of the malformed Bumastus restevensis. Specimen NMNHU-G 9763/02 (Fig. 3, a, c) is an almost completely preserved pygidium, measuring 24 mm in width and 19 mm in length. It displays two elongated pits located on the left side of the pygidium: one near the posterior mar- gin (the larger pit), and one near the mid-length of the pygidium (the smaller pit). Th e larger pit is wedge-shaped, slightly curved towards the lateral margin of the pygidium, and measures 5 mm in Series Pridoli Ludlow SI LU RI AN Sk ala M aly ni vt si Ba go vy tsy a Ky ta yh or od Bagovytsya Ternava Konivka Tsviklivtsi Rykhta Varnytsya Trubchyn Dzvenyhorod Pryhorodok Furmanivka Gorstian Ludfordian Homerian Telychian Sheinwoodian Wenlock Liandovery Stage Group Formation a b c d Fig. 2. Silurian stratigraphy of Podillia in western Ukraine and the position of the Rykhta Formation. Stratigraphic scheme modified after [17] Fig. 3. Malformed pygidia of Bumastus restevensis Balashova, 1975 from the Late Silurian Rykhta For- mation of western Ukraine. a, c — specimen NMN- HU-G 9763/02 (a — dorsal view of the pygidium, c — enlarged malformation); b, d — specimen NMNHU- G 9763/09 (b  — dorsal view of the pygidium, d — enlarged malformation). Scale bars — 5 mm▶ ▶ 6 ISSN 1025-6415. Dopov. Nac. akad. nauk Ukr. 2025. No. 4 V.S. Dernov length and 0.15—0.70 mm in width. Its depth and width gradually decrease from anterior (near the anterior margin of the pygidium) to posterior, reaching a maximum depth of approximately 0.5 mm Th e smaller pit is ellipsoidal or teardrop-shaped, located approximately 4 mm from the larger pit. It measures 2 mm in length and 1 mm in width at its widest point and is partially fi lled with matrix. Specimen NMNHU-G 9763/09 (see Fig. 3, b, d) is a nearly complete, though poorly preserved, pygidium measuring 17 mm in length and approximately 20 mm in width. Th e malformation con- sists of a crescent-shaped pit in the posterior part of the pygidium, near the sagittal line. Th e pit’s width along the anterior-posterior axis is 2 mm, with a length of approximately 2 mm and a depth of about 1 mm. Th e greatest depth is at the posterior end of the pit (near the posterior margin of the pygidium), with the depth gradually decreasing towards the edges. Th e transverse profi le of the pit is V-shaped, with steep walls and a narrow, sharply concave bottom with angular margins. Discussion and concluding remarks. Th e Palaeozoic deposits of Podillia have experienced minimal tectonic deformation or metamorphism, supporting the interpretation that the trilobite exoskeleton injuries described here occurred during the animals’ lifetime. Th e malformations observed in the studied specimens are interpreted as the result of unsuc- cessful predatory attacks (“predation scars” as described in [2—4]), likely by cephalopods, arthro- pods or fi shes. Th is interpretation is supported by the characteristic morphology of the injuries, which is distinct from those typically caused by other factors [cf. 2—7]. Th e elongated pits on pygidium NMNHU-G 9763/02 are nearly symmetrically placed. How- ever, the force responsible for these damages was insuffi cient to produce a smaller pit that fully corresponded morphologically to the larger one. Th is discrepancy could be due to the direction of the predator’s attack on the trilobite and the considerable bulging of the pygidium, which may have prevented the formation of perfectly symmetrical damage. Additionally, uneven regenera- tion of the injury across the pygidium may have further contributed to the asymmetry. Th e malformation in specimen NMNHU-G 9763/02 bears a strong resemblance to a similar injury observed on the pygidium of the Ordovician trilobite Parabarrandia bohemica (Novák, 1884) from the Czech Republic [18]. Th is injury was interpreted as a partially healed wound from a failed predatory attack by a cephalopod or arthropod during the trilobite’s life [18]. Th e injury in specimen NMNHU-G 9763/02 is also similar to injury types 4 and 5 observed on the shells of the brachiopod Ivdelinia pulchra Franchi, Schemm-Gregory et Klug, 2013 from the Givetian (Middle Devonian) of Morocco [19]. Cephalopods are likely the agents responsible for these injuries, as they are known to produce paired fractures with their beaks [20—24]. Th erefore, the paired injuries in the Bumastus specimen can reasonably be attributed to cephalopods, albeit with some caution [19]. Silurian cephalopods of Podillia are relatively well-studied (see review in [25]). Fossils of Apty- chopsis, calcite structures interpreted as the aperture operculum of orthocerid nautiloids [26], parts of the jaw apparatus of orthocerids [27], or protective shields used by orthocerids for defense [28], have been found in Silurian deposits of Podillia [29: pl. 2, fi g. 4]. However, there is no evidence to suggest that Aptychopsis was part of the jaw apparatus of nautiloids. Moreover, the morphology of the malformation on specimen NMNHU-G 9763/09 indicates that the predator’s tool was slightly pointed and semi-circular in plan, whereas Aptychopsis are typically fl at or only slightly convex. Arthropods capable of causing injuries, as observed in the trilobite specimens, include phylloca- rids, eurypterids, and other aquatic chelicerates [1, 30]. Although phyllocarids — known as scaven- gers and predators [31] — have not yet been recorded from the Silurian of Ukraine, eurypterids have. 7ISSN 1025-6415. Допов. Нац. акад. наук Укр. 2025. № 4 First record of malformed trilobites from the Silurian of Ukraine Baltoeurypterus tetragonophthalmus (Fischer de Waldheim, 1839) [32, 33] and Pterygotus sp. [34], as well as the synziphosurine Pasternakevia podolica Selden et Drygant, 1987 [32, 33], are known from the Ludlow deposits in Podillia. However, since these chelicerates are found in la- goonal facies that lack trilobites [32, 35], the likelihood that they caused the injuries observed in the trilobites is considered negligible. Th e Silurian Period marked the emergence of the fi rst jawed fi shes, including placoderms and acanthodians [36]. Rare and poorly preserved fi sh remains occur in the Ludlow strata of Ukraine [37]. Th e shape of the malformation on the pygidia of Bumastus restevensis resembles the shape of the upper jaw morphology of some placoderms, suggesting that these early fi shes might also have been responsible for the damage investigated in this study. Acknowledgments. I extend my sincere thanks to Drs Volodymyr Grytsenko and Halyna An- fi mova (National Museum of Natural History of the NAS of Ukraine, Kyiv) for the opportunity to examine the trilobite specimens collected by the late Dr. Leonid Konstantynenko. I am also grateful to Dr. Tamara Nemyrovska (Institute of Geological Sciences of the NAS of Ukraine, Kyiv) for her valu- able advice on the article. Th e author is sincerely grateful to anonymous reviewers whose comments and suggestions improved the quality of the fi nal version of the manuscript. REFERENCES 1. Mundy, D . J. C. (1982). A note on the predation of brachiopods from the Dinantian reef limestones of Cracoe, North Yorkshire. Trans. Leeds Geol. Ass., 9, No. 6, pp. 73-83. 2. Babcock, L. E. (2003). Trilobites in Paleozoic predator-prey systems, and their role in reorganization of early Paleozoic ecosystems. In: Kelley, P. H., Kowalewski, M. & Hansen, T. A. (Eds.). Predator-prey interactions in the fossil record (pp. 55-92). Springer. https://doi.org/10.1007/978-1-4615-0161-9_4 3. Owen, A. W. (1985). Trilobite abnormalities. Trans. R. Soc. Edinb: Earth Sci., 76, No. 2-3, pp. 255-272. https:// doi.org/10.1017/S0263593300010488 4. Babcock, L. E. (1993). Trilobite malformations and the fossil record of behavioral asymmetry. J. Paleontol., 67, No. 2, pp. 217-229. https://doi.org/10.1017/s0022336000032145 5. Bicknell, R. D. C., Smith, P. M. & Miller-Camp, J. (2024). An atlas of malformed trilobites from North American repositories Part 1. The Indiana University Paleontological Collection. Am. Mus. Novit., 4026, pp. 1-16. https:// doi.org/10.1206/4026.1 6. Bicknell, R. D. C., Smith, P. M. & Hopkins, M. J. (2024). An atlas of malformed trilobites from North American repositories Part 2. The American Museum of Natural History. Am. Mus. Novit., 4027, pp. 1-36. https://doi. org/10.1206/4027.1 7. Bicknell, R. D. C., Smith, P. M., Amati, L., & Hopkins, M. J. (2025). 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Дернов, https://orcid.org/0000-0002-5873-394X Інститут геологічних наук НАН України, Київ, Україна Національний науково-природничий музей НАН України, Київ, Україна E-mail: vitalydernov@gmail.com ПЕРША ЗНАХІДКА ПРИЖИТТЄВО ПОШКОДЖЕНИХ ТРИЛОБІТІВ У СИЛУРІЙСЬКИХ ВІДКЛАДАХ УКРАЇНИ Досліджено перший задокументований випадок прижиттєво пошкоджених екзоскелетів трилобітів із си- лурійських відкладів України. Скам’янілості тварин з пошкодженнями та патологіями мають значну па- леобіологічну цінність, оскільки дають змогу зрозуміти еволюцію взаємодій хижака і жертви, трофічних мереж, поведінки тварин та генетичних аномалій. Досліджений матеріал складається з двох пігідіїв Bumastus restevensis з рихтівської світи (лудлов, верхній силур) Хмельницької області (Волино-Подільська монокліналь). Порівняльний аналіз цих екзоскелетів з аналогічними зразками з інших місцезнаходжень світу показує, що вони є унікальними і не мають прямих аналогів. Виявлені пошкодження інтерпретують- ся як «шрами хижацтва», що свідчать про невдалі напади хижаків. Характерна морфологія цих ушкоджень дає підставу припустити, що вони, найімовірніше, були завдані головоногими молюсками, членистоноги- ми, такими як філокариди та/або водні хеліцерові, або рибами. Ключові слова: прижиттєві пошкодження екзоскелета, Trilobita, лудлов, Волино-Подільська монокліналь, “шрами хижацтва”.