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Article

Vertebrate Coprolites Reveal Diversity of Prey Fishes in the Oligocene Carpathian Basin of the Paratethys

by
Malgorzata Bienkowska-Wasiluk
1,*,
Piotr Bajdek
2 and
Mateusz Granica
1
1
Faculty of Geology, University of Warsaw, Żwirki i Wigury 93, 02-089 Warsaw, Poland
2
Independent Researcher, 42-202 Częstochowa, Poland
*
Author to whom correspondence should be addressed.
Diversity 2025, 17(8), 507; https://doi.org/10.3390/d17080507
Submission received: 27 June 2025 / Revised: 17 July 2025 / Accepted: 18 July 2025 / Published: 24 July 2025
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Versions Notes

Abstract

Coprolites from the Oligocene Menilite Formation of the Outer Carpathians in southeastern Poland were investigated to reveal the diversity of prey fishes consumed by coprolite producers. The material comprises 186 coprolites from seven localities. The coprolites are either sub-spherical, or elongate, and although classified into eight shape categories, display a morphological continuum. The phosphatic matrix is preserved in 28% of the specimens. Fish remains, including bones and scales, are preserved in 94% of the coprolite specimens. In 31% of specimens, these remains belong to the orders Perciformes, Gadiformes, Clupeiformes, and Aulopiformes. Prey sizes were estimated and compared to the sizes of fishes preserved as articulated skeletons from the same formation, that inhabited the Carpathian Basin of the Paratethys. The results demonstrate that coprolite analysis provides a significant paleontological data, which can be applied to infer fish diversity in other regions of the Paratethys, as well as in other sedimentary basins.

1. Introduction

The diversity of extinct actinopterygian fishes can be assessed through multiple approaches, including morphological analysis of skeletons, otoliths, teeth, bones and scales, coprolites, trace fossils, and molecular genetics. Coprolites, or fossil faeces, provide significant paleontological insights into diet and predation (e.g., [1,2,3,4,5,6]).
In this study, we analysed a large number of coprolites to demonstrate how they provide insights into fish diversity, particularly regarding the prey consumed by coprolite producers and predator–prey relationships. Our study integrated new materials with a re-examination of previously published [5] non-spiral coprolites supposedly produced by teleost fishes, to reconstruct coprolite succession during Oligocene in the Carpathian Basin of the Paratethys.

Geological Setting

The materials studied were collected from the Oligocene deposits of the Menilite Formation of the Outer Carpathians in southeastern Poland. The Oligocene deposits occur in the upper part of an Upper Jurassic–Lower Miocene sequence. Deposits of the Outer Carpathians were detached from their basement and trust northward mainly in the Early and Middle Miocene [7,8]. The new materials studied were collected from deposits of the Silesian and Skole units (nappes) of the Outer Carpathians in seven localities (Figure 1). Three of these are in the Skole Unit: Hermanowa, Jamna Dolna, and Krępak. Four localities are in the Silesian Unit: Jasienica Rosielna, Rudawka Rymanowska, Rogi, and Winnica. Our analyses included specimens from two localities in the Skole Unit—Futoma and Wola Czudecka—that were studied by Bajdek and Bienkowska-Wasiluk [5].
In the Hermanowa locality (near Rzeszów city), specimens were collected from the Rudawka Tractionite Member, ichthyofaunal zone IPM2 (abbreviation HE in [11]).
The Jamna Dolna locality (near the city of Przemyśl) has two outcrops [12]. In the first outcrop, the specimens were recovered from the Rudawka Tractionite Member, ichthyofaunal zone IPM2, in the second outcrop from the Korzeniówka Member, ichthyofaunal zone IPM6. The second outcrop had been correlated previously with the Krępak Member, ichthyofaunal zone IPM3 [12], but since collection of the index taxon Argyropelecus cosmovici, it has been correlated with the ichthyofaunal zone IPM6.
In the Krępak locality (near the city of Przemyśl), specimens were collected from the Korzeniówka Member, ichthyofaunal zone IPM6.
In the Jasienica Rosielna, Rogi (RO1 in [11]), and Winnica (W in [11]) localities (near Krosno town), specimens were derived from the Upper Menilite Beds, ichthyofaunal zone IPM2.
In the Rudawka Rymanowska (RR in [11,12,13]) locality (in area of Sanok town), the specimens were collected from horizon of the Tylawa Limestones, ichthyofaunal zone IPM2.
The Rudawka Tractionite Member, the Tylawa Limestones, and the Upper Menilite Beds are correlated to the nannoplankton zone NP23 of Martini [14] and are dated to the early Oligocene, ca. 31 Ma (Figure 2). The part of the Korzeniówka Member with ichthyofaunal zone IPM6 recognised is correlated to the zone NP25 and is dated to the late Oligocene, ca. 26 Ma.
In the Futoma and Wola Czudecka localities, specimens were collected from the Dynów Marl Member, ichthyofaunal zone IPM1, correlated to the nannoplankton zone NP23 and dated to the early Oligocene, ca. 32 Ma [5].
Around the Eocene/Oligocene boundary, intense tectonic activity, global cooling, and a sea-level fall resulted in the formation of the Paratethys through its separation from the Tethys Ocean. Basin isolation resulted in oxygen-depleted conditions during the Oligocene and caused deposition of organic-rich sediments. The Menilite Formation of the Outer Carpathians was deposited in the Central Paratethys (Figure 1C). Interpretations of the depositional system of the Menilite Formation are widely variable. Based on sedimentological and paleontological analyses including foraminifera and fishes, alongside other considerations, deposition on the continental slope, submarine ridges, and basin floor (e.g., [7,11,15,16,17]), and locally on the shelf is assumed (e.g., [15,18,19,20]).
Climatic change and basin isolation led to salinity and oxygen concentration stratification of the water column. The alternating deposition of turbidites, organic C-rich mudstones, diatomites, and coccolith limestones in the Paratethys has been interpreted as the result of fluctuations in sea level, salinity, temperature, and nutrient supply [21,22,23,24]. Formation of organic C-rich deposits is related to increased productivity. During early Rupelian (about the onset of the nannoplankton zone NP23), connection between Paratethys and the world ocean was closed (see e.g., [21,25]).
Deposits of the Dynów Marl Member were formed during intensive pelagic carbonate sedimentation induced by phytoplankton blooms of low-diversity coccolithophorid assemblages [26,27]. Blooms of calcareous nannoplankton are interpreted as the results of low salinity in surface water related to the inflow of fresh water into the basin and high nutrient supply [26,27]. Low salinity was suggested by Studencka et al. [28] on the basis of bivalve assemblage analysis. For the mesopelagic zone, the normal salinity has been inferred based on fish fauna [29].
Coprolites occur in laminated marls or marl laminae interbedding beds of marls.
The fine-grained nature, presence of primary structures, and abundant nannoplankton content of the marl laminae and laminated marls suggest that they originated via pelagic settling through the water column and/or deposition from low-concentration turbidity currents (see [30]). The presence of wavy-anastomosing carbonaceous kerogenous laminae in similar and synchronous marls of the bituminous marl formation from Romania suggests a benthic microbial mat origin [31]. The marl laminae and laminated marls are intercalated with slabby marls, mostly fine-grained turbidites (see [32]).
The Rudawka Tractionite Member consists of organic C-rich shales and sandstone-shale couplets that were deposited by low-concentration turbidity currents and hemipelagic settling from suspension. The disappearance of mesopelagic fishes such as Stomiiformes and Myctophiformes that have been recorded in Dynów Marl Member and absence of bioturbations suggests that the chemocline was positioned relatively high in the water column (see [33]).
The Tylawa Limestone Horizon include limestones intercalated with shales and sandstones. Limestones were formed during intensive blooms of calcareous nannoplankton (see e.g., [34,35]) that were monospecific [27] and are related to low salinity in surface water. In laminated limestones and calcareous grey shales with well-preserved fishes, microbially induced sedimentary structures interpreted as benthic microbial mats occur [36]. The presence of microbial mats could be responsible for low δ15Norg values recorded by Bojanowski et al. [37].
Coprolites occur in laminated limestones and calcareous grey shales. The fine-grained nature, presence of primary structures, presence of microbially induced sedimentary structures, and abundant nanoplankton content in these deposits reveal that they originated via pelagic settling from suspension. Other organic C-rich shales and sandstone–shale couplets were deposited by low-concentration turbidity currents and hemipelagic settling from suspension. Absence of mesopelagic fishes such as Stomiiformes and Myctophiformes suggests that the chemocline was positioned relatively high in the water column.
The Upper Menilite Beds in the Silesian unit consists of organic C-rich shales, sandstones and cherts, which were deposited by low-concentration turbidity currents and hemipelagic settling from suspension. Fish assemblages, similar to those in the Tylawa Limestone Horizon, suggest a relatively high position of the chemocline.
During the later Rupelian (upper NP23, NP24), seaways with the open ocean were partly restored as sea-level rose (e.g., [25,27]). In the Chattian (NP24-25), the entire Paratethys was inhabited by a fully marine fauna (e.g., Stomiiformes and Myctophiformes).
Deposits of the Korzeniówka Member include black shales, sandstones, and grey or green shales. The black shales dominate in the Member and are either allochthonous sediments deposited by low-concentration turbidity currents, or autochthonous (background) hemipelagic sediments. Sandstone–shale couplets were deposited by low-concentration turbidity currents and hemipelagic settling from suspension. Grey or green shales are most likely background hemipelagic sediments deposited slowly from suspension. These shales, which show bioturbation, can be attributed to improved oxygenation in the sediment [12,33]. The presence of mesopelagic fishes such as Stomiiformes and Myctophiformes suggests that the chemocline was positioned lower in the water column [33] than during formation of deposits of Rudawka Tractionite Member, Tylawa Limestone Horizon, and Upper Menilite Beds.

2. Materials and Methods

The material comprises 186 coprolite specimens, mostly preserved on 88 slabs and their 77 counterslabs, giving in total 274 catalogue numbers (Table S1). Slabs contain from one to thirteen coprolites. The material is housed at the Muzeum Geologiczne im. Stanisława Józefa Thugutta, Faculty of Geology, University of Warsaw, Poland (MWGUW). The material has been collected from seven sites: Hermanowa, Jamna Dolna, Jasienica Rosielna, Krępak, Rudawka Rymanowska, Rogi, and Winnica. Photographs were taken with digital microscope Keyence VHX-7000 (Keyence, Itasca, IL, USA) and camera Nikon DSLR (Nikon, Tokyo, Japan) at the Faculty of Geology, University of Warsaw.

2.1. Morphological Analyses of Coprolites

The collection was subdivided into eight shape categories based on different shapes: (A) sub-spherical, (B) oval or short elongate, (C) tear-shaped or conical, (D) elongate straight, (E) linear curved, (F) linear S-shaped, (G) linear sinuous, (H) unclassified (irregular, incomplete or poorly preserved) (Figure 3). The shape was observed on slab surface of rock sample. Most of the specimens were strongly flattened during compaction processes, but we assume that their shapes on the slab surface are well preserved, as it was observed in fishes [12].
To visualize shape categories, we plotted measurements of specimens, their width, and length. To assess the presence of distinguished groups in the fossil record, we plotted specimen measurements of width and length against ichthyofaunal zones. The relationships between specimen widths and lengths were evaluated after calculation of Spearman and Pearson correlation coefficients for each of the distinguished shape category in order to determine monotonic or linear relationships (Table S2).
In addition to the 186 coprolites newly described herein, our statistical analyses included 16 specimens from the underlying Rupelian Dynów Marl Member of the Menilite Formation collected in two localities (Futoma and Wola Czudecka, Figure 1B), that were studied by Bajdek and Bienkowska-Wasiluk [5].

2.2. Inclusions in Coprolites

Inclusions were identified and their taxonomy was recognised when possible. To visualize the presence of inclusions, we plotted measurements of coprolite specimens (width and length) against recognised prey.
We collected measurements of skeletal elements of well-preserved complete fish skeletons and recorded the sizes of these fishes from the same deposits (Menilite Formation) as well as recent fishes (Table S3–S6). Therefore, we obtained data on the proportions of body length (standard length—SL) to particular remains. Such proportions were used to estimate the size of fish prey consumed by predators (see [38,39,40,41,42]).
From the measurements data, a linear regression equation was established for each prey order between the measurement of skeletal element and SL (Table S6).

3. Results

3.1. Morphological Diversity of Coprolites

Although the collection initially displays considerable size and morphological variation, the cross-plot of length versus width for the eight shape categories of coprolites (Figure 4A) reveals that these groups are not distinctly separated, rather they form a morphological continuum. Most specimens are elongate, with various tear-shaped and curved forms representing intermediate stages within a complex succession of progressively more elongate and typically more curved morphologies. Sub-spherical and linear sinuous specimens represent the extremes of this continuum.
For both diameter and length of specimens, the mode (4 mm, 18 mm; n = 202) is typically smaller than the median (5.5 mm, 18.5 mm; n = 202), which is smaller than the arithmetic mean (7.47 mm, 22.56 mm; n = 202). This indicates that the collection is dominated by small coprolite specimens, while larger specimens are progressively less abundant. However, the diameter of coprolites classified as elongate straight (n = 15) displays a reverse pattern with most specimens being wide (mode = 6 mm, median = 5.5 mm, arithmetic mean = 5.18 mm), suggesting that wider faeces had a more solid consistency on average, while narrower faecal strings were more easily bent. The length of the “linear sinuous” specimens (n = 19) also shows a reverse pattern, with the mode (45 mm) being larger than the median (36 mm), which is larger than the arithmetic mean (32.26 mm). This reflects the fact that specimens classified as “linear sinuous” tend to be particularly long, with a mean length/diameter ratio of 9.43 (Table S2).
The diameter of the specimens varies from 0.5 mm in ZI/75/039/1/a (Figure 3K) classified as “linear sinuous” to 28 mm in the unclassified specimen ZI/75/085/2 (Table S1). Coprolites classified as “sub-spherical” show on average the largest but also the most variable diameter (arithmetic mean = 12.08 mm, standard deviation = 8.11 mm). Average diameter and diameter variability tend to decrease with increasing length/diameter ratio (Table S2). This indicates that whereas more or less spherical forms are encountered among all sizes, clearly elongate specimens are mostly very narrow in width (Figure 4A). Conversely, the length varies from 3.75 mm in ZI/75/043/3 classified as “sub-spherical” to 95 mm in ZI/75/033/a (Figure 3I) classified as “linear S-shaped”.
For most shape categories of coprolites, with the exception of those classified as “linear curved”, Spearman and Pearson correlation coefficients are comparable (Table S2). Linear curved coprolites show a moderate linear but a weak monotonic positive relationship between their widths and lengths (Spearman’s ρ = 0.12, Pearson’s r = 0.43). Correlation coefficients of other groups tend to decrease with increasing coprolite length, being highest in the “sub-spherical” (Spearman’s ρ = 0.98, Pearson’s r = 0.99) and the lowest in the “linear S-shaped” specimens (Spearman’s ρ = 0.17, Pearson’s r = 0.14).

3.2. Stratigraphic Distribution of Coprolite Shape Categories

Coprolites from the IPM 1 ichthyofaunal zone are “linear curved”, “linear S-shaped”, and “linear sinuous”. Coprolites from IPM 2 show large variability, with all shape categories present, dominated by “oval or short elongate”, “tear-shaped or conical”, and “linear curved”. Coprolites from IPM 6 are dominated by “sub-spherical” category.
Our statistical analyses included the arithmetic and geometric means of the specimen diameters and lengths within the eight distinguished shape categories (Table S2). The typically similar values of arithmetic and geometric means revealed by our statistical results are reflected by the relatively even spatial distribution of specimens on the scatter plot (Figure 4A). This is particularly evident for coprolites classified as “oval or short elongate” and “elongate straight” (Table S2). In contrast, coprolites belonging to the shape categories “sub-spherical”, “linear S-shaped”, and “linear sinuous” show a slightly less even distribution in the scatter plot (Figure 4A), which may be explained by their mixed stratigraphic provenance (Figure 4B). “Sub-spherical” coprolites are clearly subdivided into two groups comprising specimens from either the IPM2 or IPM6 ichthyofaunal zones (Figure 4A,B). Meanwhile, the “linear S-shaped” and “linear sinuous” groups include a mixture of specimens in the zones IPM1 and IPM2 (Figure 4A,B).

3.3. Matrix Preservation

Preservation of the matrix is partly dependent on lithology. All coprolites (four specimens) from the black shales of Jamna Dolna lack the matrix. Specimens from other contexts either lack the matrix or show the matrix of a relatively stronger diagenetic alteration. In Rudawka Rymanowska, the matrix is preserved (Figure 5) in 84% (16/19) of specimens from the laminated limestone, 27% (17/62) of specimens from the calcareous grey shale, and 0% (0/4) of specimens from the grey shale. None of the coprolite specimens from the brown shales of Jasienica Rosielna (0/77), Jamna Dolna (0/5), Rogi (0/3), Winnica (0/3), Hermanowa (0/2), and Krępak (0/2) has a preserved matrix.

3.4. Inclusions

Coprolite inclusions are most useful for identifying prey affinity. Inclusions are abundant in the studied coprolites and consist of bones and scales of actinopterygian fish. Most coprolites, 94% (174/186) contain inclusions and 31% (58/186) have remains recognisable to order, family or species (Figure 4C and Figure 6). Recognised remains include Perciformes, Gadiformes, Clupeiformes, and Aulopiformes (Table 1). The most common are remains of Perciformes, which constitute 66% of coprolites with inclusions (38/58), and nearly half of these coprolites (17/38) contain Oligoserranoides budensis (see [43]), while the remainder contain remains that may belong to O. budensis, but they are indeterminant. Remains of Gadiformes constitute 22% of coprolites with inclusions (13/58), and among them are mostly scales similar to Palaeogadus (see [44]). The Clupeiformes and Aulopiformes (Holosteus) comprise 9% and 3%, respectively, of coprolites with inclusions.

3.5. Sizes of Fishes in the Menilite Formation

The assemblage of fishes of the Menilite Formation, preserved as skeletons (Figure 7), comprises fishes with various body sizes, ranging from about 10 mm to approximately 1350 mm SL (Figure 8, Table S7).
The fishes of the Dynów Marl Member and the lower members of the Menilite Formation comprise assemblage of the IPM 1 zone. The assemblage contains Aeoliscus, Anenchelum glarisianum, Beksinskiella longimana, ‘Glossanodon’ musceli, Oligophus moravicus, Palimphyes sp., Palaeorhynchus sp., Scopeloides glarisianus, and Vinciguerria obscura. The most abundant fishes in this assemblage are small and medium-sized fishes; they have standard lengths ranging from 20 to 140 mm (Figure 8). A single genus (Anenchelum) of medium- to large-sized fish with SL from 50 to 1350 mm comprise about 10% of the assemblage.
The fishes of the Upper Menilite Beds, the Rudawka Tractionite Member, and the Tylawa Limestone Horizon comprise the assemblage of IPM2 zone. The assemblage contains Anenchelum glarisianum, Beksinskiella longimana, ‘Glossanodon’ musceli, Oligoserranoides budensis, Palaeogadus simionescui, and Trachinus minutus. Among the most abundant fishes in this assemblage are three small-sized fishes ranging from 10 to 54 mm (Figure 8, ‘G.’ musceli, O. budensis, T. minutus), one small- to medium-sized fish (SL 21–96 mm, B. longimana) and two small- to large-sized fishes (SL 13–1350 mm, P. simionescui and A. glarisianum).
The fishes of the Korzeniówka Member belong to the assemblage of the IPM6 zone. Among the most abundant fishes of this assemblage are three small-sized fishes ranging from 13 to 56 mm (Figure 8, Idrissia sp., Eomyctophum sp., Argyropelecus sp.), two small- to medium-sized fishes (SL 21–96 mm, Clupeiformes indet., Syngnathus sp.) and one small- to large-sized fish (SL 50–1350 mm, Anenchelum glarisianum).
The sizes of fishes in these three assemblages appear to be similar, with each assemblage dominated by small- to medium-sized fishes.

3.6. Sizes of Preys from Coprolites

Taxonomically recognisable remains in coprolites include scales, vertebrae. and bones of the head such as opercle, preopercle, maxilla, premaxilla, and mandible. These remains were used to estimate the size of prey using the linear regression for each order between the measurement of skeletal element and SL (Figure 9, Table S6).
The relation of particular measurement of skeletal element to body length is variable between taxa (Figure 9). Therefore, we applied for interpretation of taxa of the same order only taxa with similar morphology and no taxa with distinctly different morphology.
We estimated the size of prey for all Perciformes, Gadiformes, Clupeiformes, and Aulopiformes (Table 1, Figure 10) recognised in investigated coprolites. Each interpreted size of fish, shown as standard length, has a range of values. This is due to a fact that the analysed skeletal elements may be of different sizes for the same size of an individual or achieved measurements of these elements have variability for similar sized fish. For example, sizes of scales are variable (see [45,46]) in a single individual. Vertebrae are larger in the abdominal region than the ones near the caudal fin. The achieved measurements of skeletal elements have variability because of two reasons: (1) measurements of bones and scales from coprolites may be slightly smaller in result of contact or cover by another skeletal element or matrix; (2) measurements of bones and scales from well-preserved skeletons may be slightly reduced in result of contact or cover of another skeletal element.
The achieved results show that prey fishes (Table 1, Figure 10) assigned to Perciformes were small-sized fishes from 22 to 51 mm of SL. The Gadiformes were medium-sized fishes from 61 to 271 mm of SL. The Clupeiformes were medium-sized fishes from 34 to 112 mm of SL. The Aulopiformes were medium-sized fishes from about 86 to about 220 mm of SL. Three size results for Perciformes that exceeded the known data for complete fish skeletons from the Menilite Formation were rejected (see Figure 10), as implausible. 3.7. The prey sizes compared to sizes of fishes preserved as skeletons in the Menilite Formation
From the skeletal record of fishes, we can reconstruct sizes of fishes during the formation of deposits of the Menilite Formation (see Table S7). The Perciformes represented by Oligoserranoides budensis were small-sized fishes from 19 to 51 mm of SL [43]. The Gadiformes (Palaeogadus) were medium-sized fishes from 13 to 278 mm of SL [47]. The Clupeiformes (Beksinskiella, Sanalosa) were small- to medium-sized fishes from 19 to 96 mm of SL [48,49,50]. The Aulopiformes (Holosteus) were medium- to large-sized fishes from 150 mm to about 500 mm of SL.
Sizes of prey fishes assigned to Perciformes are the same as sizes of Perciformes recorded as well-preserved skeletons. Sizes of prey fishes assigned to Gadiformes are nearly the same as sizes of fishes recorded as well-preserved skeletons. Our results reveal the presence of only medium-sized Clupeiformes that have been prey fishes, nearly the same, slightly larger as sizes of fishes from the skeletal record. Fishes also preyed on medium sized Aulopiformes, but we have not recognised large Aulopiformes in coprolites.
Perciformes and among them Oligoserranoides budensis were the main food sources for predatory fishes.

4. Discussion

4.1. Succession of Coprolite Shape Categories Throughout the Menilite Formation

The coprolite shape categories from the Menilite Formation, Western Carpathian Mountains, form a morphological continuum from sub-spherical, thorough more or less elongate to distinctly elongate, than distinct groups of morphotyphes. Investigated coprolites have been collected from five outcrops of middle Rupelian deposits and two outcrops of the middle Chattian deposits of the Formation. Most specimens from the Rupelian Rudawka Tractionite Member, Upper Menilite Beds, and Tylawa Limestone resemble in morphology those previously described by Bajdek and Bienkowska-Wasiluk [5] from the underlying Rupelian Dynów Marl Member of the Menilite Formation (Figure 2), but show larger variability of shape categories. Coprolites from Rudawka Tractionite Member, Upper Menilite Beds, and Tylawa Limestone are mostly strongly elongate, often sinuous forms (e.g., MWGUW ZI/75/033, MWGUW ZI/75/046/a, MWGUW ZI/75/039/1/a, Figure 3), while sub-spherical shapes are far less abundant (e.g., MWGUW ZI/75/028/2, Figure 3B). Some of the sub-spherical forms likely represent incompletely preserved elongate or tear-shaped coprolites. In contrast, in the Chattian Korzeniówka Member (Figure 2) only sub-spherical coprolite forms have been found (e.g., MWGUW ZI/75/027/a, MWGUW ZI/75/030/b, MWGUW ZI/75/031, MWGUW ZI/75/032/b, Figure 3A). These have large diameters, ranging from 18.25 to 26 mm. Sub-spherical coprolites from Rupelian have various diameters, ranging from 3 to 19 mm, with most being smaller than those from Chattian. The diameter of faeces correlates with the total body length of the animal, as it was employed for some groups such as crocodiles [51]. Therefore, the large diameters of sub-spherical coprolites from Chattian indicate larger producers than those producing similar shaped coprolites during the Rupelian. This contrasts with skeletal record, as fish sizes from Rupelian and Chattian are similar. Fish sizes from the Rupelian IPM 1 and IPM 2 zones range from 10 to 1350 mm SL, while those from Chattian IPM 6 zone range from 13 to 1350 mm (Figure 8). Most common are small to medium-sized fishes preserved as skeletons in both Rupelian and Chattian deposits.
The diameter of all coprolites from Rupelian ranges from 0.5 to 28 mm and indicates variable sizes of producers. Although the correlation of coprolite diameter with fish length for investigated fishes is not possible to establish and requires further research, the variability of coprolite diameters indicates large variability in the length of coprolite producers in the Menilite Formation. However, interpretations of differences in estimated predator sizes between Rupelian and Chattian based on coprolite diameter needs additional research.
Predators are capable of feeding on preys of smaller size than the predator size [52,53]. The size variability of prey fishes recognised in the Menilite Formation is from 22 to 271 mm of SL, and reveals the presence of predators larger than these sizes. Although the results do not reveal many large predators, some of them are known from skeletal records. They belong to scombriform Anenchelum glarisianum, and gadiform Palaeogadus simionescui. Large predators are usually numerically rarer in the food web than smaller species [54], and therefore rare in the fossil record. The prey length generally ranges from 10% to 46% of predator length [52] for freshwater fishes. Such relation reveals possible predator lengths ranging from 47 to 2710 mm in the Menilite Formation.

4.2. Preservation of Coprolites in the Menilite Formation

Although rocks of the Menilite Formation are compacted, possibly resulting in flattening and increase of diameters and lengths of coprolites, their shapes, visible on rock surfaces (e.g., linear curved, linear S-shaped) and bedding planes, are well preserved and not deformed (e.g., bended, cracked) in result of other diagenetic processes. The coprolites from the Menilite Formation contain organic remains that were swallowed and excreted by fishes in oxygenated higher parts of water column. The faeces sank and were deposited on the sea bottom under low-oxygen conditions [5].
Assemblages from Rupelian limestones, marls, and calcareous grey shales (Dynów Marl Member, Tylawa Limestones) contain many well-preserved coprolites with matrix and/or inclusions represented by fish remains. Assemblages from Rupelian brown shales, and Chattian black shales do not contain a preserved matrix, but many contain inclusions of fish remains. This variability in matrix preservation reflects different depositional conditions, including the types of sediment components and diagenetic processes. Sediment rich in calcite micrite protected faeces better than sediment without calcite. The presence of microbial mats seems to be an important factor for coprolite preservation, as it was demonstrated for fish preservation in limestones and calcareous grey shales of Tylawa Limestones by Leonowicz et al. [36] and inferred for marls of the bituminous marl formation from Romania [31], which are synchronous with and similar to marls of the Dynów Marl Member.

4.3. Ecology of Fishes from the Menilite Formation

The Menilite Formation contains various assemblages of fishes. The changes in the taxonomic composition of the assemblages thorough the section are significant. They reflect changes in environment, e.g., the water column oxygenation in the basin [11], that is supported by ichnological record [33]. Fishes of the Menilite Formation have been included in several ecologic groups, e.g., epipelagic, meso-bathypelagic, benthopelagic, demersal, bathydemersal, neritic, and reef [11,33]. The epipelagic fishes (e.g., Clupeiformes, some Scombriformes) live in pelagic waters from the surface to 150–200 m. The meso-bathypelagic fishes (e.g., Myctophiformes, Stomiiformes) live in pelagic waters from 150–200 m to 4000 m. The benthopelagic fishes (e.g., Gadiformes, some Scombriformes) live near the bottom (up to 100 m above bottom), mainly at 200 to 1000 m water depth. The demersal and bathydemersal fishes live on the sea floor. The neritic and reef fishes live in shallow waters from the surface to 200 m.
Fishes of the Stomiiformes (Gonostomatidae, Phosichthyidae) and Myctophiformes (Myctophidae) constitute a significant part (42%, [29]) of the assemblage of the Dynów Marl Member. Their recent relatives inhabit the deep pelagic zones. The assemblage contains epipelagic fishes (e.g., Scombriformes: Euzaphlegidae) and benthopelagic fishes (e.g., Scombriformes: Trichiuridae, Gadiformes, Argentiniformes).
The assemblage of the Dynów Marl Member and the lower members of the Menilite Formation, that belongs to IPM 1, contain 48% of meso-bathypelagic taxa, 30% of epipelagic taxa, 17% of benthopelagic taxa, and 5% of neritic and reef taxa [11].
The assemblage of the Upper Menilite Beds, the Rudawka Tractionite Member and the Tylawa Limestone Horizon belongs to IPM2. The assemblage of IPM2 includes 71% of benthopelagic, demersal, and bathydemersal fishes, 24% of epipelagic taxa, 3% of neritic and reef taxa, but only 2% of meso-bathypelagic taxa [33]. The most common fishes of the assemblage are representatives of Clupeiformes, Argentiniformes, Gadiformes, Perciformes, and Scombriformes [11,12,55].
The assemblage of the Korzeniówka Member represents IPM6. The assemblage of IPM6 is dominated by meso-bathypelagic taxa (45%) and epipelagic taxa (35%). Benthopelagic, demersal, and bathydemersal fishes constitute 18%. Neritic and reef fishes are rare (2%, [33]). The contribution of ecological groups of fishes in this assemblage is similar to the assemblage recognised in the Dynów Marl Member, but taxonomic content differs. Deep-water fishes are represented by Stomiiformes (Gonostomatidae, Phosichthyidae) and Myctophiformes (Myctophidae) similar to the assemblage of the Dynów Marl Member, but mostly represented by other genera. The most common fishes of the assemblage of IPM 6 are representatives of Clupeiformes, Myctophiformes, Syngnathiformes, Stomiiformes, Gadiformes, Argentiniformes, and Scombriformes [11,12].
Prey fishes of IPM 1 include epipelagic Scombriformes, meso-bathypelagic Myctophiformes, and benthopelagic Gadiformes.
Prey fishes of IPM 2 include epipelagic Clupeiformes, benthopelagic Gadiformes, possibly demersal Perciformes, possibly mesopelagic Aulopiformes.
Prey fishes of IPM 6 belong to epipelagic Clupeiformes.
It shows that predators feed on different ecological groups, living in different depths of the pelagic zone, as well as close to bottom.

4.4. Relation of Prey Sizes to Skeletal Record of Fishes in the Menilite Formation

The estimated sizes of prey fishes belonging to Perciformes and Gadiformes are nearly the same as sizes of individuals of these orders from skeletal record from the Menilite Formation. Therefore, coprolites can be successfully applied in size estimations of fishes living in the basins. The estimated sizes of some Clupeiformes found in coprolites slightly larger than from skeletal record from the formation can reveal the presence of larger individuals than previously known. The rich record of Perciformes, Gadiformes, perciform Oligoserranoides budensis, and presence of Clupeiformes in coprolites reveals the presence of most common fishes in the Menilite Formation known from the skeletal record. Therefore, coprolites can be successfully applied in identifying the most common fishes living in the basins.
In the investigated material, 94% of coprolites contain fish inclusions, 31% contain fishes recognizable to order, and 9% contain fishes recognizable to species.

5. Conclusions

The morphological succession of coprolites in the Menilite Formation, Western Carpathian Mountains, begins with assemblages composed solely of strongly elongate, linear forms which were described from the lower Rupelian Dynów Marl Member [5], through assemblages dominated by comparable elongate forms from the middle Rupelian Rudawka Tractionite Member, Upper Menilite Beds, and Tylawa Limestone, to the assemblage of sub-spherical coprolites from the middle Chattian Korzeniówka Member. The morphological variability observed in coprolites produced by teleost fishes is greater than previously reported [5] and demonstrate a morphologic continuum rather than distinct morphotypes.
Analysis of fish remains, preserved within the investigated coprolites, revealed the presence of common fishes in the Carpathian Basin. Measurements of these fish remains allowed the estimation of both prey and predator sizes in the basin. The results indicate the presence of preyed fishes of variable lengths, all of them small- to medium-sized fishes within size range of fishes observed in the skeletal record.
Our method provides significant potential for inferring fish diversity in other Cenozoic and Mesozoic deposits. We propose that in instances where the skeletal record is absent, the close relatives (extant or fossil) could be used in estimation of size of fishes recognised in coprolites.

Supplementary Materials

The following supporting information can be downloaded at https://zenodo.org/records/16027949 (accessed on 30 June 2025), DOI: 10.5281/zenodo.16027948. Figure S1: Measurements of the bones of fishes. Table S1: Catalogue of coprolite specimens. Table S2: Coprolite descriptive statistics. Table S3: Selected measurements of Perciformes. Table S4: Selected measurements of Clupeiformes. Table S5: Selected measurements (in mm) of Gadiformes and Aulopiformes. Table S6: Linear regression equations of standard length (SL; mm) on bone and scale dimensions (mm) for the four prey orders and percent relative errors of each skeletal remain (see Figure S1 for measure details). Coefficient of determination (r2), number of data pairs in regression (n) and SL range are indicated, x—dimension of bone or scale. Table S7: The size and abundance of the most abundant fishes preserved as complete skeletons of the Menilite Formation in assemblages IPM1, IPM2, and IPM 6 [56,57,58].

Author Contributions

Conceptualization, M.B.-W. and P.B.; methodology, M.B.-W., P.B., and M.G.; analyses and conclusions M.B.-W. and P.B.; resources, M.B.-W.; writing—original draft preparation, M.B.-W. and P.B.; writing—review and editing, M.B.-W., P.B. and M.G. All authors have read and agreed to the published version of the manuscript.

Funding

The research was funded by the Faculty of Geology, University of Warsaw (grant WG 501-D113-01-1130202).

Institutional Review Board Statement

Not applicable.

Data Availability Statement

The original contributions presented in the study are included in the article and in Supplementary Materials; further inquiries can be directed to the corresponding author. The following supporting information can be downloaded at https://zenodo.org/records/16027949 (accessed on 30 June 2025), DOI: 10.5281/zenodo.16027948.

Acknowledgments

We express our gratitude to Radosław Wasiluk, Marcin Pałdyna, Damian Smoleń, Rafał Nawrot, the Spirifer Geological Society for collecting some of the specimens, and help during fieldwork; to Robert Szybiak, Albin Jamróz, and Grzegorz Salwa for observations of diversity of coprolites in the Menilite Formation. We acknowledge two anonymous reviewers for their valuable comments and reviews that improved the quality of the manuscript.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. Location maps. (A) Study area within Central Europe. (B) Localities where the specimens were collected (white dots) with two localities described by Bajdek and Bienkowska-Wasiluk [5] within simplified geological map of the Polish part of the Outer Carpathians (modified from [9]). (C) Carpathian Basin of Central Paratethys in paleogeography of the Oligocene (adopted from [10]).
Figure 1. Location maps. (A) Study area within Central Europe. (B) Localities where the specimens were collected (white dots) with two localities described by Bajdek and Bienkowska-Wasiluk [5] within simplified geological map of the Polish part of the Outer Carpathians (modified from [9]). (C) Carpathian Basin of Central Paratethys in paleogeography of the Oligocene (adopted from [10]).
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Figure 2. General stratigraphic relationships of the Oligocene deposits of the Polish Outer Carpathians (after [11]) and position of the studied and discussed sections designated by white arrows and text.
Figure 2. General stratigraphic relationships of the Oligocene deposits of the Polish Outer Carpathians (after [11]) and position of the studied and discussed sections designated by white arrows and text.
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Figure 3. Coprolites from the Menilite Formation, examples of eight shape categories. (A,B) sub-spherical, (A) MWGUW ZI/75/027/a; (B) MWGUW ZI/75/028/2; (C,D) oval or short elongate, (C) MWGUW ZI/75/066, (D) MWGUW ZI/75/102/a; (E,F) tear-shaped or conical, (E) MWGUW ZI/75/048; (F) MWGUW ZI/75/112/1/a; (G) elongate straight, ZI/75/040/3; (H) linear curved MWGUW ZI/75/040/1; (I,J) linear S-shaped, (I) ZI/75/033/a, (J) MWGUW ZI/75/046/a; (K) linear sinuous, ZI/75/039/1/a; (L,M) unclassified (irregular, incomplete or poorly preserved), (L) MWGUW ZI/75/069/2, (M) MWGUW ZI/75/104/a. Scale bar: 10 mm.
Figure 3. Coprolites from the Menilite Formation, examples of eight shape categories. (A,B) sub-spherical, (A) MWGUW ZI/75/027/a; (B) MWGUW ZI/75/028/2; (C,D) oval or short elongate, (C) MWGUW ZI/75/066, (D) MWGUW ZI/75/102/a; (E,F) tear-shaped or conical, (E) MWGUW ZI/75/048; (F) MWGUW ZI/75/112/1/a; (G) elongate straight, ZI/75/040/3; (H) linear curved MWGUW ZI/75/040/1; (I,J) linear S-shaped, (I) ZI/75/033/a, (J) MWGUW ZI/75/046/a; (K) linear sinuous, ZI/75/039/1/a; (L,M) unclassified (irregular, incomplete or poorly preserved), (L) MWGUW ZI/75/069/2, (M) MWGUW ZI/75/104/a. Scale bar: 10 mm.
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Figure 4. Relationship between width and length of coprolites and (A) shape categories, (B) ichthyofaunal zones, and (C) the taxonomy of included prey remains.
Figure 4. Relationship between width and length of coprolites and (A) shape categories, (B) ichthyofaunal zones, and (C) the taxonomy of included prey remains.
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Figure 5. Preservation of matrix in coprolites. (A–D) Matrix well-preserved in laminated limestone, (A,B) MWGUW ZI/75/035/a and MWGUW ZI/75/035/b; (C,D) MWGUW ZI/75/041/2. (E,F) Matrix well-preserved in the calcareous grey shale, (E) MWGUW ZI/75/073, (F) MWGUW ZI/75/075/a. (G,H) Imprint of matrix in brown shale, (G) MWGUW ZI/75/109/a, (H) MWGUW ZI/75/105/1/a. Scale bar: 1 mm (D), 5 mm (C,H), 10 mm (A,B,E–G).
Figure 5. Preservation of matrix in coprolites. (A–D) Matrix well-preserved in laminated limestone, (A,B) MWGUW ZI/75/035/a and MWGUW ZI/75/035/b; (C,D) MWGUW ZI/75/041/2. (E,F) Matrix well-preserved in the calcareous grey shale, (E) MWGUW ZI/75/073, (F) MWGUW ZI/75/075/a. (G,H) Imprint of matrix in brown shale, (G) MWGUW ZI/75/109/a, (H) MWGUW ZI/75/105/1/a. Scale bar: 1 mm (D), 5 mm (C,H), 10 mm (A,B,E–G).
Diversity 17 00507 g005
Diversity 17 00507 g006
Figure 6. Remains of fishes in coprolites, photos, and superimposed interpretative drawings. (A–J) perciform Oligoserranoides budensis, (A,B) opercle, MWGUW ZI/75/065; (C,D) preopercle, MWGUW ZI/75/047/4/a; (E,F) scales, MWGUW ZI/75/029/2/b; (G,H) scales, MWGUW ZI/75/037; (I,J) vertebrae, MWGUW ZI/75/037. (K,N) clupeiform, (K–L) preopercle, MWGUW ZI/75/031, (M,N) scale, MWGUW ZI/75/038. (O–T) gadiform, (O,P) vertebrae, MWGUW ZI/75/060/3/b; (Q,R) scale, MWGUW ZI/75/039/2/b; (S,T) scales, MWGUW ZI/75/039/1/a. Scale bar: 1 mm.
Figure 6. Remains of fishes in coprolites, photos, and superimposed interpretative drawings. (A–J) perciform Oligoserranoides budensis, (A,B) opercle, MWGUW ZI/75/065; (C,D) preopercle, MWGUW ZI/75/047/4/a; (E,F) scales, MWGUW ZI/75/029/2/b; (G,H) scales, MWGUW ZI/75/037; (I,J) vertebrae, MWGUW ZI/75/037. (K,N) clupeiform, (K–L) preopercle, MWGUW ZI/75/031, (M,N) scale, MWGUW ZI/75/038. (O–T) gadiform, (O,P) vertebrae, MWGUW ZI/75/060/3/b; (Q,R) scale, MWGUW ZI/75/039/2/b; (S,T) scales, MWGUW ZI/75/039/1/a. Scale bar: 1 mm.
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Diversity 17 00507 g007
Figure 7. Fish skeleton Oligoserranoides budensis, MWGUW ZI/57/014/a (a) from the Menilite Formation associated by coprolites, MWGUW ZI/75/039/1/b (b), MWGUW ZI/75/039/2/b (c), MWGUW ZI/75/039/3/b (d), brown shale. Scale bar: 10 mm.
Figure 7. Fish skeleton Oligoserranoides budensis, MWGUW ZI/57/014/a (a) from the Menilite Formation associated by coprolites, MWGUW ZI/75/039/1/b (b), MWGUW ZI/75/039/2/b (c), MWGUW ZI/75/039/3/b (d), brown shale. Scale bar: 10 mm.
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Diversity 17 00507 g008
Figure 8. Standard lengths (in mm) of the most abundant fishes preserved as skeletons in the Menilite Formation of assemblages of ichthyofaunal zones IPM 1, IPM 2, and IPM 6.
Figure 8. Standard lengths (in mm) of the most abundant fishes preserved as skeletons in the Menilite Formation of assemblages of ichthyofaunal zones IPM 1, IPM 2, and IPM 6.
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Diversity 17 00507 g009
Figure 9. Location of maxilla, opercle and preopercle, and relations of selected remains measurements to standard length for Perciformes, Clupeiformes, and Gadiformes.
Figure 9. Location of maxilla, opercle and preopercle, and relations of selected remains measurements to standard length for Perciformes, Clupeiformes, and Gadiformes.
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Diversity 17 00507 g010
Figure 10. Estimated prey sizes of fishes recognised in coprolites from the Menilite Formation.
Figure 10. Estimated prey sizes of fishes recognised in coprolites from the Menilite Formation.
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Table 1. Coprolite specimens with remains of Perciformes, Gadiformes, Clupeiformes, and Aulopiformes, size of inclusion analysed in estimation of prey size, and estimated prey size.
Table 1. Coprolite specimens with remains of Perciformes, Gadiformes, Clupeiformes, and Aulopiformes, size of inclusion analysed in estimation of prey size, and estimated prey size.
Specimen
Number		InclusionsTaxonomy of InclusionsSize (mm)Estimated Prey Size (SL in mm)
ZI/75/029/2/a, ZI/75/029/2/bPerciform-like scales similar to Oligoserranoides budensisPerciformesMaximum diameter of scale 1–1.540–46
ZI/75/031Preopercle of ClupeiformesClupeiformesLength of preopercle l’pop 448
ZI/75/037Perciform-like scales similar to Oligoserranoides budensis,
vertebrae		PerciformesHeight of vertebra 1, maximum diameter of scale 1.536–40
ZI/75/038Vertebrae and scales of
Clupeiformes 		ClupeiformesHeight of vertebra 1, maximum diameter of scale 2.535–48
ZI/75/039/1/a, ZI/75/039/1/b, Scales of Gadiformes, family Merlucciidae, similar to
Palaeogadus, bones		GadiformesMaximum diameter of scale 1.287
ZI/75/039/2/a, ZI/75/039/2/b, Scales of Gadiformes, family Merlucciidae, similar to
Palaeogadus, bones		GadiformesMaximum diameter of scale 1.598–102
ZI/75/039/3/a, ZI/75/039/3/b, Scales of Gadiformes, family Merlucciidae, similar to
Palaeogadus, bones		GadiformesMaximum diameter of scale 1, height of vertebra 182–88
ZI/75/040/1 Perciform-like scales similar to Oligoserranoides budensisPerciformesMaximum diameter of scale 1–1.540–46
ZI/75/042Perciform-like scales similar to Oligoserranoides budensisPerciformesMaximum diameter of scale 1.5–2>48
ZI/75/043/4Scales of Clupeiformes ClupeiformesMaximum diameter of scale 484–112
ZI/75/044/2/a, ZI/75/044/2/b, Bones of perciform
Oligoserranoides budensis		PerciformesDepth of preopercle h’pop 4.534
ZI/75/047/4/a, ZI/75/047/4/bBones of perciform
Oligoserranoides budensis		PerciformesDepth of preopercle h’pop 431
ZI/75/052Perciform-like scales similar to Oligoserranoides budensis, bones, vertebraePerciformesHeight of vertebra 1.2, maximum diameter of scale 132–41
ZI/75/056Scales of Gadiformes, family Merlucciidae, similar to
Palaeogadus		GadiformesMaximum diameter of scale 1.8104–116
ZI/75/060/1Scales of Gadiformes, family Merlucciidae, similar to
Palaeogadus		GadiformesMaximum diameter of scale 177–88
ZI/75/060/3/a, ZI/75/060/3/bVertebrae of Gadiformes, family Merlucciidae, similar to
Palaeogadus		GadiformesHeight of vertebra 0.872
ZI/75/060/4/a, ZI/75/060/4/bPerciform-like scales similar to Oligoserranoides budensis, bonesPerciformesMaximum diameter of scale 0.829–37
ZI/75/060/11Scales of Gadiformes, family Merlucciidae, similar to
Palaeogadus		GadiformesMaximum diameter of scale 177
ZI/75/060/12Perciform-like scales similar to Oligoserranoides budensisPerciformesMaximum diameter of scale 132–46
ZI/75/064/a, ZI/75/064/bPerciform-like scales similar to Oligoserranoides budensisPerciformesMaximum diameter of scale 132–46
ZI/75/065Bones of perciform
Oligoserranoides budensis		PerciformesDepth of preopercle h’pop 7, length of maxilla Lp 539–47
ZI/75/068/1Bones and scales of perciform Oligoserranoides budensisPerciformesLength of maxilla Lp 539
ZI/75/076/a, ZI/75/076/bBones and scales of perciform Oligoserranoides budensisPerciformesDepth of preopercle h’pop 536
ZI/75/077/4Perciform-like scales similar to Oligoserranoides budensis, bonesPerciformesHeight of vertebra 1, maximum diameter of scale 132–36
ZI/75/078/2Bones (including vertebrae) and scales of Gadiformes, family Merlucciidae, similar to PalaeogadusGadiformesHeight of vertebra 0.661–87
ZI/75/081Perciform-like scales similar to Oligoserranoides budensis, bonesPerciformesMaximum diameter of scale 1.540–51
ZI/75/083/1Scales of Clupeiformes ClupeiformesMaximum diameter of scale 484–112
ZI/75/083/2Bones of Clupeiformes ClupeiformesDepth of preopercle h’pop 1063
ZI/75/085/1Bones of perciform
Oligoserranoides budensis		PerciformesDepth of preopercle h’pop 536
ZI/75/085/2Bones of perciform
Oligoserranoides budensis		PerciformesDepth of preopercle h’pop 642
ZI/75/086/1/a, ZI/75/086/1/b Bones and scales of perciform Oligoserranoides budensisPerciformesDepth of opercle 5, scales 1–1.832–41
ZI/75/086/3Bones and scales of perciform Oligoserranoides budensisPerciformesDepth of opercle 4, scales 1–1.532–34
ZI/75/087/4Perciform-like scales similar to Oligoserranoides budensis, bonesPerciformesLength of maxilla Lp 539
ZI/75/087/5Bones and scales of perciform Oligoserranoides budensisPerciformesLength of preopercle l’pop 225
ZI/75/088/2/a, ZI/75/088/2/bPerciform-like scales similar to Oligoserranoides budensis, bonesPerciformesDepth of preopercle h’pop 5.539
ZI/75/088/3/a, ZI/75/088/3/bBones of perciform
Oligoserranoides budensis		PerciformesDepth of preopercle h’pop 326
ZI/75/088/3/a, ZI/75/088/3/bBones of HolosteusAulopiformesHeight of vertebra 2153–220
ZI/75/088/4/a, ZI/75/088/4/bBones of HolosteusAulopiformesHeight of vertebra 186–136
ZI/75/088/5/a, ZI/75/088/5/bPerciform-like scales similar to Oligoserranoides budensis, bonesPerciformesHeight of vertebra 0.830
ZI/75/092/1/a, ZI/75/092/1/b, Bones of perciform
Oligoserranoides budensis		PerciformesHeight of vertebra 0.830
ZI/75/092/4/a, ZI/75/092/4/b, Bones of perciform
Oligoserranoides budensis		PerciformesDepth of preopercle h’pop 431
ZI/75/095/a, ZI/75/095/b Scales of GadiformesGadiformesMaximum diameter of scale 5175–272
ZI/75/096Scales of Gadiformes, family Merlucciidae, similar to
Palaeogadus		GadiformesMaximum diameter of scale 2109–126
ZI/75/099/a, ZI/75/099/bPerciform-like scales similar to Oligoserranoides budensis, bonesPerciformesHeight of vertebra 136
ZI/75/100/3Perciform-like scales similar to Oligoserranoides budensisPerciformesMaximum diameter of scale 1.235
ZI/75/102/a, ZI/75/102/bPerciform-like scales similar to Oligoserranoides budensisPerciformesMaximum diameter of scale 1.540–51
ZI/75/103/1/a, ZI/75/103/1/bPerciform-like scales similar to Oligoserranoides budensis, bonesPerciformesLength of maxilla Lp 4, maximum diameter of scale 131–46
ZI/75/103/3Perciform-like scales similar to Oligoserranoides budensisPerciformesMaximum diameter of scale 1–1.540–46
ZI/75/104/a, ZI/75/104/bScales of Gadiformes, family Merlucciidae, similar to
Palaeogadus		GadiformesMaximum diameter of scale 2.5120–150
ZI/75/105/2Perciform-like scales similar to Oligoserranoides budensis, bonesPerciformesHeight of vertebra 1, maximum diameter of scale about 132–36
ZI/75/107/1/a, ZI/75/107/1/bPerciform-like scales similar to Oligoserranoides budensis, bonesPerciformesHeight of vertebra 136
ZI/75/110/a, ZI/75/110/bBones of perciform
Oligoserranoides budensis		PerciformesDepth of preopercle h’pop 4, length of preopercle l’pop 2.531
ZI/75/111/a, ZI/75/111/bScales of Gadiformes, family Merlucciidae, similar to
Palaeogadus, bones		GadiformesMaximum diameter of scale 1.598–102
ZI/75/112/1/a, ZI/75/112/1/bScales of Gadiformes, family Merlucciidae, similar to
Palaeogadus, bones		GadiformesMaximum diameter of scale 1.5, height of vertebra 182–98
ZI/75/112/2/a, ZI/75/112/2/bBones of perciform
Oligoserranoides budensis		PerciformesHeight of vertebra 0.7, length of preopercle l’pop 225–28
ZI/75/112/3Bones of perciform
Oligoserranoides budensis		PerciformesHeight of vertebra 0.728
ZI/75/113/1/a, ZI/75/113/1/bBones of perciform
Oligoserranoides budensis		PerciformesDepth of preopercle h’pop 642
ZI/75/113/2/a, ZI/75/113/2/bPerciform-like scales similar to Oligoserranoides budensis, bonesPerciformesLength of maxilla Lp 323
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Bienkowska-Wasiluk, M.; Bajdek, P.; Granica, M. Vertebrate Coprolites Reveal Diversity of Prey Fishes in the Oligocene Carpathian Basin of the Paratethys. Diversity 2025, 17, 507. https://doi.org/10.3390/d17080507

AMA Style

Bienkowska-Wasiluk M, Bajdek P, Granica M. Vertebrate Coprolites Reveal Diversity of Prey Fishes in the Oligocene Carpathian Basin of the Paratethys. Diversity. 2025; 17(8):507. https://doi.org/10.3390/d17080507

Chicago/Turabian Style

Bienkowska-Wasiluk, Malgorzata, Piotr Bajdek, and Mateusz Granica. 2025. "Vertebrate Coprolites Reveal Diversity of Prey Fishes in the Oligocene Carpathian Basin of the Paratethys" Diversity 17, no. 8: 507. https://doi.org/10.3390/d17080507

APA Style

Bienkowska-Wasiluk, M., Bajdek, P., & Granica, M. (2025). Vertebrate Coprolites Reveal Diversity of Prey Fishes in the Oligocene Carpathian Basin of the Paratethys. Diversity, 17(8), 507. https://doi.org/10.3390/d17080507

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