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A large coprolite of a carnivorous dinosaur found in Harding County, South Dakota, US
A large Miocene coprolite from South Carolina, US
Coprolites found on the Blahnita riverbed, Romania, showing a seed inclusion (right specimen)
A large coprolite from South Carolina, US
Age: White River Oligocene; Location: Northwest Nebraska; Dimensions: Varies (25 mm × 20 mm); Weight: 8-10 g; Features: Many small inclusions and one has a complete toe bone from a small deer called a leptomeryx.

A coprolite (also known as a coprolith) is fossilized feces. Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour (in this case, diet) rather than morphology. The name derives from Ancient Greek κόπρος (kópros), meaning "dung", and λίθος (líthos), meaning "stone". They were first described by William Buckland in 1829. Before this, they were known as "fossil fir cones" and "bezoar stones". They serve a valuable purpose in paleontology because they provide direct evidence of the predation and diet of extinct organisms.[1] Coprolites may range in size from a few millimetres to over 60 centimetres.

Coprolites, distinct from paleofeces, are fossilized animal dung. Like other fossils, coprolites have had much of their original composition replaced by mineral deposits such as silicates and calcium carbonates. Paleofeces, on the other hand, retain much of their original organic composition and can be reconstituted to determine their original chemical properties, though in practice the term coprolite is also used for ancient human fecal material in archaeological contexts.[2][3][4] The study of coprolites in Japan was pioneered by Michiko Chiura.[5][6]

Initial discovery

[edit]

British fossil hunter Mary Anning noticed as early as 1824 that "bezoar stones" were often found in the abdominal region of ichthyosaur skeletons found in the Lias formation at Lyme Regis.[7] She also noted that if such stones were broken open they often contained fossilized fish bones and scales as well as sometimes bones from smaller ichthyosaurs. These observations by Anning led the geologist William Buckland to propose in 1829 that the stones were fossilized feces and to name them coprolites. Buckland also suspected that the spiral markings on the fossils indicated that ichthyosaurs had spiral ridges in their intestines similar to those of modern sharks and that some of these coprolites were black with ink from swallowed belemnites.[8]

Research value

[edit]

By examining coprolites, paleontologists are able to find information about the diet of the animal (if bones or other food remains are present), such as whether it was a herbivore or a carnivore, and the taphonomy of the coprolites, although the producer is rarely identified unambiguously,[9] especially with more ancient examples.[10] In some instances, knowledge about the anatomy of animals' digestive tracts can be helpful in assigning a coprolite to the animal that produced it, one example being the finding that the Triassic dinosauriform Silesaurus may have been an insectivore, a suggestion which was based on the beak-like jaws of the animal and the high density of beetle remains found in associated coprolites.[11] Further, coprolites can be analyzed for certain minerals that are known to exist in trace amounts in certain species of plant that can still be detected millions of years later.[12] In rare cases, coprolites have even been found to contain well-preserved insect remains.[13] There is also a documented case of a coprolite containing an ichnofossil in the form of footprints of a crocodilian, created when a crocodilian stepped on the faecal matter before it became fossilised.[14]

Recognizing coprolites

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A Miocene pseudocoprolite from Washington state. They are commonly mistaken for coprolites because of their appearance and shape; they are actually of inorganic origin. Scale in mm. See Spencer (1993).

The recognition of coprolites is aided by their structural patterns, such as spiral or annular markings, content, undigested food fragments, and associated fossil remains. The smallest coprolites are often difficult to distinguish from inorganic pellets or from eggs. Most coprolites are composed chiefly of calcium phosphate, along with minor quantities of organic matter. By analyzing coprolites, it is possible to infer the diet of the animal which produced them.

Coprolites have been recorded in deposits ranging in age from the Cambrian period[15] to recent times and are found worldwide. Some of them are useful as index fossils, such as Favreina from the Jurassic period of Haute-Savoie in France.

Some marine deposits contain a high proportion of fecal remains. However, animal excrement is easily fragmented and destroyed, so usually has little chance of becoming fossilized.

Coprolite mining

[edit]

In 1842 the Rev John Stevens Henslow, a professor of botany at St John's College, Cambridge, discovered coprolites just outside Felixstowe in Suffolk in the villages of Trimley St Martin,[16] Falkenham and Kirton[17] and investigated their composition. Realizing their potential as a source of available phosphate once they had been treated with sulfuric acid, he patented an extraction process and set about finding new sources.[18]

Very soon, coprolites were being mined on an industrial scale for use as fertilizer due to their high phosphate content. The major area of extraction occurred over the east of England, centered on Cambridgeshire and the Isle of Ely[19][20] with its refining being carried out in Ipswich by the Fison Company.[20] There is a Coprolite Street near Ipswich docks where the Fisons works once stood.[21]

The industry declined in the 1880s[20][16] but was revived briefly during the First World War to provide phosphates for munitions.[19] A renewed interest in coprolite mining in the First World War extended the area of interest into parts of Buckinghamshire as far west as Woburn Sands.[18]

See also

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Look up coprolite in Wiktionary, the free dictionary.
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Notes

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References

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Coprolite

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A coprolite is the fossilized or mineralized excrement of ancient organisms, preserving traces of their intestinal contents and serving as a valuable in . These structures typically form when fecal matter, rich in indigestible remnants such as bones, scales, shells, or plant material, undergoes rapid and mineralization, often through phosphatization, before can occur. Coprolites are relatively rare due to the perishable nature of organic waste, but they are more commonly preserved in marine environments from organisms like , reptiles, and , appearing as nodular, contorted, or spiral-shaped masses. The recognition of coprolites as fossil feces dates to the early 19th century, when fossil collector discovered specimens in the strata of , , prior to 1824, initially describing them as "bezoar stones." In 1829, geologist coined the term "coprolite" from Greek roots meaning "dung stone" and confirmed their identity as fossilized excrement through detailed analysis, revolutionizing the understanding of prehistoric biology. Buckland's work, including his identification of coprolites from ichthyosaurs and plesiosaurs containing undigested prey remains, established coprolites as key evidence for reconstructing ancient ecosystems. Coprolites hold significant paleontological value by revealing the diets, health, behaviors, and environmental conditions of extinct , often providing unavailable from body fossils. For instance, they can contain identifiable fragments like fish scales, shards, or , allowing scientists to infer trophic relationships, such as carnivorous reptiles preying on belemnites or herbivorous dinosaurs consuming wood during seasonal shortages. Modern analyses employ techniques like computed tomography (CT) scanning and chemical studies to extract detailed information on , , and even gut microbiomes from these fossils. Despite challenges in attributing coprolites to specific producers, ongoing research continues to uncover their role across the eon, from invertebrates to mammals.

Overview

Definition and Etymology

A coprolite is a fossilized specimen of or dung produced by ancient animals, serving as a that records behavioral evidence such as diet and ecology rather than the physical structure of the organism itself. Unlike body fossils, which preserve skeletal or remains, coprolites fall under the broader category of ichnofossils, including bromalites—fossilized digestive residues—that provide indirect insights into prehistoric life without representing the animal's morphology. The term "coprolite" was coined in 1829 by the English geologist and paleontologist William Buckland, who introduced it in his seminal work on fossil discoveries near Oxford, England, to describe mineralized dung concretions initially linked to extinct reptiles like Megalosaurus. Buckland derived the name from the Ancient Greek words kopros (κόπρος), meaning "dung" or "excrement," and lithos (λίθος), meaning "stone," aptly capturing the petrified nature of these specimens. This etymology reflected Buckland's recognition of coprolites as biogenic structures formed through diagenetic processes, distinguishing them from inorganic concretions. While coprolites are biogenic and primarily associated with animal producers, they are not limited to vertebrates; examples include those from such as , like opalized coprolites from deposits, and even early feces containing material from Siluro-Devonian strata.

Physical Characteristics

Coprolites exhibit a wide array of shapes influenced by the digestive systems of their producers, including elongated rod-like or cylindrical forms, spirals with coiled internal structures, discoidal or kidney-shaped variants, and more amorphous or irregular masses. These morphologies aid in preliminary recognition; for instance, spiral coprolites often derive from animals like possessing spiral valves in their intestines. Sizes vary dramatically based on the organism, ranging from a few millimeters in length for coprolites produced by small such as to over 40 centimeters for those from large carnivorous dinosaurs like Tyrannosaurus rex. In terms of composition, coprolites are predominantly mineralized remnants of organic fecal matter, with calcium phosphate (in the form of ) as the most common , though silica or replacements also occur depending on the . Embedded inclusions frequently preserve undigested dietary elements, such as fragments in coprolites, seeds or material in specimens, and grains in various types, reflecting the original meal contents. These inclusions are often pulverized and integrated into the matrix, distinguishing coprolites from similar . Color variations in coprolites typically span to hues, resulting from organic carbon residues and iron oxides, while lighter shades like yellowish-, pinkish, or whitish tones arise from enrichment or specific mineralization processes. Textures externally appear nodular, contorted, or polished smooth, with internal cross-sections revealing concentric layering from successive depositions in the gut or voids created by escaping gases during early decay. These features contribute to their distinctive, often lumpy appearance compared to other fossilized concretions.

Formation

Biological Origin

Coprolites originate from the produced by animals through their digestive systems, which process ingested , extract nutrients, and expel undigested residues as . In vertebrates, the digestive tract—comprising the , , intestines, and or —breaks down via enzymatic and mechanical actions, resulting in that vary in composition based on diet. Carnivorous vertebrates, such as ancient or dinosaurs, produce rich in undigested fragments, scales, or feathers due to incomplete of protein-rich prey, while herbivorous vertebrates like titanosaurs yield containing fibers, , , or bark remnants from cellulose-heavy diets. Invertebrates also contribute to coprolite formation through analogous digestive processes, though their systems are simpler and often segmented. For instance, annelids and arthropods like process food in a , , and , excreting compact fecal pellets or castings that preserve undigested , such as wood particles in coprolites from the Eocene-Oligocene. Coprolites are categorized by their producers, including those from (e.g., spiral forms from spiral valve intestines in ), dinosaurs (e.g., bone shards indicating carnivory in theropods), mammals (e.g., inclusions in therapsid feces), , and even ancient humans, with dietary indicators like bone fragments in examples highlighting trophic levels. Immediately after excretion, several pre-fossilization factors influence the potential for to persist long enough for fossilization. High content promotes rapid bacterial of organic components, accelerating breakdown, while drier conditions slow this process and enhance preservation chances. Bacterial activity, particularly in phosphate-rich , can create microenvironments that inhibit further decay through early mineralization, and exposure to environmental elements like oxygen and temperature fluctuations determines initial stability before burial.

Fossilization Process

The fossilization of coprolites begins with taphonomic pathways that prioritize rapid burial in fine-grained sediments to shield the fecal material from , , and aerobic decay. This initial entombment, often occurring within hours to days after deposition, limits bacterial and oxidation, allowing organic components to persist long enough for mineralization to commence. In many cases, follows, where minerals such as () infiltrate and replace the original organic matrix, transforming the soft, amorphous feces into a durable phosphatic . This process replicates the internal textures and inclusions, preserving evidence of undigested remains while gradually recrystallizing the material over geological timescales. Environmental conditions play a crucial role in favoring coprolite preservation, particularly anaerobic settings that inhibit oxidative breakdown. Low-oxygen environments, such as stagnant lagoons, lake bottoms, or floors, promote the accumulation of phosphate-rich pore waters derived from decaying , creating ideal conditions for phosphatization—the precipitation of minerals like directly onto or within the coprolite. These settings often feature microbial mats or biofilms that accelerate burial and provide a geochemical barrier, with low (below 6.38) and high availability driving the carbonate-phosphate switch essential for mineral . In lacustrine or systems, for instance, seasonal anoxia enhances this process, as seen in deposits where microbial coverage ensured rapid encapsulation. The overall fossilization of coprolites typically unfolds over millions of years through diagenetic and stages, with early mineralization stabilizing the structure within weeks to months before sedimentary compaction and cementation complete the transformation. However, rare instances of accelerated preservation occur in exceptional media like , where coprolites entomb with amber-like fidelity over thousands to millions of years, or in tar pits, where asphaltic impregnation halts decay almost immediately, as evidenced by 50,000-year-old coprolites retaining cellulosic material.00674-6)

Identification

Morphological Features

Coprolites display a range of diagnostic shapes and patterns that facilitate preliminary field identification of their producers, often reflecting the anatomy of the digestive tract. Reptilian coprolites frequently exhibit segmented forms characterized by circumferential constrictions, as seen in crocodilian specimens with rounded ends and periodic narrowing along their length. Spiral patterns are common in coprolites from reptiles and certain , resulting from the in their intestines; these heteropolar structures feature tight coils at one end tapering to an exposed edge, with coil widths varying from 3-4 mm in larger examples. Cross-sections of such coprolites reveal internal layering, where longitudinal and transverse views show stacked spirals or continuous wrappings of material, providing clues to the producer's gut morphology. Surface features further aid in distinguishing coprolites during initial assessment. Desiccation cracks often appear as fine, irregular fissures on the exterior, particularly in exposed specimens, while longitudinal zones of rough texture may indicate areas where fecal material adhered to intestinal walls before expulsion. Impressions from surrounding tissues or ingested items, such as shell fragments, can be visible on the surface, and embedded fossils like parasite eggs—such as cestode ova with distinct morphological traits resembling modern forms—may protrude or be discernible without dissection. Size and weight of coprolites correlate with the body size of the producer, offering proportional insights for identification. For instance, sauropod coprolites can reach up to 40 cm in diameter, reflecting the massive scale of their herbivores, whereas smaller reptiles produce specimens under 10 cm. These dimensions, combined with density variations, help estimate the original animal's mass without invasive analysis.

Analytical Techniques

Non-destructive methods, such as computed (CT) scanning, allow researchers to visualize the internal structures of coprolites without causing damage, enabling the identification of inclusions like bone fragments, plant material, or entire organisms. For instance, synchrotron phase-contrast micro has revealed articulated skeletons and remains within coprolites, providing insights into predator-prey relationships while preserving the specimen for further study. This technique produces high-resolution 3D reconstructions, distinguishing coprolites from similar by highlighting voids and textures indicative of digestive processes. Destructive analytical approaches, including thin-section microscopy and chemical assays like Fourier transform (FTIR) spectroscopy, provide detailed examinations of coprolite microstructure and composition. Thin-section preparation involves slicing and polishing coprolites to 30 μm thickness for petrographic analysis under transmitted light, revealing microcrystalline groundmasses and inclusions such as fish scales or decapod cuticles that indicate dietary habits and diagenetic history. FTIR spectroscopy, applied to powdered samples, identifies components by measuring absorption spectra; for example, the presence of (absorption at 580–600 cm⁻¹) in coprolites suggests carnivorous producers, while carbonates and silicates point to herbivores. Biomolecular techniques further enhance coprolite studies by targeting ancient genetic and isotopic signatures. (aDNA) extraction, often using phenol-chloroform methods followed by PCR amplification, has successfully recovered from coprolites up to 14,000 years old, confirming origin and detecting parasites like Enterobius vermicularis even in microscopically negative samples. measures ratios such as δ¹³C in coprolite or to distinguish dietary contributions from C₃ (e.g., temperate grasses, δ¹³C ≈ -27‰) versus C₄ (e.g., , δ¹³C ≈ -13‰), offering quantitative insights into plant-based diets without relying on macroscopic remains. , via of aDNA, reconstructs gut s by classifying bacterial taxa; analyses of coprolites from show reduced diversity compared to modern feces, with elevated Actinobacteria and diminished Firmicutes, reflecting preservation biases and host-specific communities. Tools like CoproID leverage these metagenomic data to predict coprolite sources (e.g., vs. canine) based on microbiome composition and host DNA proportions.

History

Early Discovery and Studies

The initial scientific recognition of coprolites as fossilized feces occurred in the early , primarily through the work of English . In 1829, Buckland published a seminal paper describing spiral-shaped fossils from the Lias formations near , , which he identified as the excrement of ichthyosaurs based on their contents, including undigested fish bones and scales. This marked the first systematic study of coprolites, distinguishing them from previously misinterpreted objects like stones or plant cones, and he coined the term "coprolite" from the Greek words for dung and stone. Mary Anning, a renowned fossil collector from , played a crucial role in these early discoveries by excavating numerous coprolites and supplying them to Buckland for analysis. Her observations of inclusions such as remains within the s provided key evidence supporting Buckland's interpretations, and her correspondence with him in included detailed descriptions that advanced the understanding of coprolite morphology and association with marine reptiles. Anning's contributions extended the scope of coprolite studies beyond mere collection, emphasizing their potential for revealing dietary habits of extinct animals. During the 19th and early 20th centuries, coprolite research evolved amid debates over their origins, with some paleontologists proposing they were stones or plant cones rather than fecal matter. Buckland's findings countered these views by highlighting organic inclusions incompatible with such interpretations, but the controversy persisted. Classifications based on shape, composition, and stratigraphic context to differentiate them from other concretions helped standardize identification, shifting focus from curiosity to . Key milestones in the included Otto Abel's studies on coprolite inclusions, which integrated them into paleobiological reconstructions of ancient ecosystems and behaviors. Abel examined feces from various formations to infer predator-prey relationships and digestive processes, promoting coprolites as vital for holistic rather than isolated oddities. This period marked a transition to systematic research, laying groundwork for mid-20th-century advancements in analysis.

Commercial Mining

The commercial mining of coprolites emerged in the mid-19th century as a significant industry driven by the demand for phosphate-rich fertilizers to boost agricultural productivity. In , particularly in , , and , extraction began in the late following the recognition of coprolites—fossilized nodules high in —as a viable alternative to and . The boom intensified during the 1840s to 1860s, coinciding with post-Napoleonic War food shortages and advancements in soil chemistry. Similarly, in the United States, coprolite took off in around 1867, centered in the Charleston Basin along rivers such as the Ashley and Cooper, where phosphate deposits including coprolitic nodules were abundant and linked to post-Civil War agricultural recovery. Mining operations typically involved open-pit excavation, with workers using picks, shovels, and later steam-powered dredgers to remove shallow deposits from , , or riverbed layers. Extracted nodules were processed at nearby washmills—often horse- or steam-operated—where they were crushed, washed to remove impurities, and treated with to produce "coprolite manure," a soluble . The scale was substantial: in the UK, annual production peaked at over 258,000 tons by , generating economic value exceeding £1 million yearly and royalties averaging £100 per acre for landowners. In , the industry yielded $300,000 to $400,000 annually in the early 1880s, with state royalties reaching $250,000 per year by 1890, employing thousands and spurring investments of millions in equipment and land. The coprolite trade declined sharply by the 1890s and had largely ended by 1900, supplanted by cheaper imports of (until its exhaustion around 1872), vast rock discoveries in and , and the rise of synthetic fertilizers like those developed by John Bennet Lawes. This shift was accelerated by regulatory measures, such as the UK's 1894 Quarries Act, and economic pressures from weather-related farm failures. The legacy includes profound environmental alterations, with extensive pits, ponds, and lowered fields scarring landscapes in eastern and South Carolina's Lowcountry, some of which persist as wetlands or archaeological sites today.

Applications in Paleontology

Dietary Reconstruction

Coprolites serve as direct evidence for reconstructing ancient animal diets by preserving undigested inclusions that reflect the consumed food sources and trophic habits. These inclusions often include plant microfossils such as grains, granules, and phytoliths, which indicate herbivorous or omnivorous diets dominated by vegetation; fragments and tooth pieces, signaling carnivory through predation or scavenging; and fish scales or spines, pointing to piscivorous feeding in aquatic or semi-aquatic environments. Such macroscopic and microscopic remains provide high-resolution snapshots of individual meals, allowing paleontologists to infer seasonal variations and resource availability without relying solely on skeletal evidence. Notable examples illustrate the diversity of dietary insights from coprolites. In tyrannosaurid coprolites from , crushed fragments constitute 30–50% of the volume, demonstrating a carnivorous diet that included large vertebrates, with the fragmentation patterns suggesting bone-crushing behavior consistent with scavenging or active predation. Similarly, Pleistocene coprolites attributed to herbivores like in contain evidence indicating a diet focused on C3 and C4 grasses in open grasslands, which aligns with dental wear patterns observed in associated Equus fossils. These cases highlight how coprolites reveal specific strategies, such as opportunistic scavenging in apex predators or specialized in ungulates. Quantitative methods enhance dietary reconstructions by estimating proportions of food types and trophic positions. Pollen concentration analysis, for instance, quantifies plant intake by counting grains per gram of coprolite material, enabling estimates of dietary breadth—such as high concentrations of herb pollen (114,000–138,000 grains/ml) indicating dominant herbivory in moa coprolites from New Zealand. Stable isotope ratios, particularly δ15N values, further delineate trophic levels, with enrichments of 3–5‰ per level signaling shifts from primary producers to carnivores, as seen in coprolite associations from Miocene ecosystems where higher δ15N confirmed top-predator status. These approaches, often integrated with techniques like mass spectrometry from the identification section, provide robust, proportional models of ancient diets. A 2024 analysis of over 500 coprolites from the early Jurassic in Poland revealed dietary flexibility in early dinosaurs, contributing to understanding their ecological dominance.

Parasite and Pathogen Studies

Coprolites serve as exceptional repositories for preserved parasite remains, particularly the eggs and cysts of helminths such as nematodes and trematodes, which can endure mineralization or processes over millennia. These structures are often identified through microscopic examination, revealing intact or partially degraded forms that indicate infection intensity and host-parasite dynamics in ancient populations. For instance, in human coprolites from Hinds Cave in , dating to the Archaic period (approximately 4,000–8,000 years ago), pinworm eggs (Enterobius vermicularis) have been documented, suggesting endemic among prehistoric hunter-gatherers in arid environments where dry conditions favored preservation. Such findings highlight how coprolites capture snapshots of parasitic burdens without relying on skeletal evidence. Pathogen detection in coprolites has advanced through () analysis, enabling the recovery of bacterial spores and viral genetic material that illuminate infectious disease histories. Metagenomic sequencing of coprolites has identified diverse microbial communities, including potential zoonotic agents like those from or viruses, providing evidence of disease transmission between humans and animals in prehistoric settings. A notable example is the extraction of viral DNA from a 14th-century coprolite in , which revealed a diverse virome including bacteriophages and giant viruses such as Mimiviridae, offering insights into ancient microbial diversity in human guts. These studies, often employing (PCR) and next-generation sequencing on rehydrated samples, underscore coprolites' utility in tracing evolution and challenges in early human societies. The evolutionary implications of parasite findings in coprolites extend to tracking co-evolution between hosts and parasites across geological timescales, particularly in non-human contexts. In coprolites, helminth eggs, including those of acanthocephalans (thorny-headed worms), have been preserved, indicating ancient infections in reptilian hosts and shedding light on parasite diversification during the age of dinosaurs. For example, putative acanthocephalan eggs recovered from coprolites in , likely from crocodyliform producers, represent some of the earliest direct evidence of these parasites, suggesting their adaptation to vertebrate intermediate hosts predated the K-Pg extinction event. Such discoveries facilitate reconstructions of parasite-host phylogenies and ecological roles in prehistoric ecosystems.

Environmental Insights

Coprolites serve as valuable archives for reconstructing past vegetation and climates through and analysis, which reveals the floral composition of ancient environments ingested by their producers. grains preserved within coprolites can indicate dominant communities, such as grasslands or forests, and shifts between arid and humid conditions based on the ratios of drought-tolerant versus moisture-dependent species. For instance, analysis of Late-Glacial coprolites from the giant Mylodon darwinii in Mylodon Cave, southern , identified from grasses, shrubs, and herbaceous like Fragaria and Azorella, suggesting a transitional landscape with both open, dry steppes and wetter, forested patches during the Pleistocene-Holocene boundary around 10,530 BP. This approach complements stable techniques by providing direct botanical evidence of seasonal or regional climate variability. Faunal interactions within ecosystems are illuminated by coprolites through traces of coprophagy, the consumption of , which highlights nutrient recycling and trophic dynamics. Invertebrate borings, such as tunnels made by insect larvae, observed in approximately one-third of coprolites from the in , , demonstrate rapid post-depositional exploitation by coprophagous organisms, fostering energy transfer across trophic levels in swampy habitats. Similarly, packrat middens—accumulations of fecal pellets from Neotoma —preserve macrofossils and that track fluctuations and succession in arid environments, such as expansions of woodlands during Marine Isotope Stage 3 (~50,000 years BP) in lowlands. The spatial distribution of coprolites aids in habitat reconstruction, particularly by inferring migration routes and resource availability like sources. Concentrations of coprolites in Permian deposits, such as the "Coproland" site in the Rio do Rasto Formation of the Paraná Basin, , with over a thousand specimens featuring flat undersurfaces indicative of submersion, point to persistent aquatic environments that attracted aggregations of and reptiles, suggesting reliable freshwater availability in a Middle to Upper Permian . In Early Permian sites like Arroyo del Agua, , clustered coprolites from diverse vertebrates imply seasonal gatherings around water bodies, informing on migration patterns driven by hydrological features in semi-arid landscapes.

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