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Paleoburrow
Paleoburrow
Paleoburrow
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A paleoburrow is an underground shelter excavated by extinct paleo-vertebrate megafauna that lived in the prehistoric era.[1][2][3] Most paleoburrows are likely made by giant armadillos and large ground sloths, depending on their size.[4] Thousands of examples have been identified across South America, mostly in the Brazilian states of Santa Catarina and Rio Grande do Sul.[4]

The first paleoburrow was discovered in Rondonia by Amilcar Adamy in 2010.[5] Paleoburrows, often exposed by development, are threatened by construction and degradation from rain and the elements.[4] Researchers from several Brazilian universities have formed the Paleoburrows Project to raise awareness about their existence, and fight misinformation.[4][6]

Description

[edit]

Thousands of sites characterized as paleoburrows have been found across South America,[4] such as in Ponta do Abunã, in Rondônia, within the Amazon region,[7] in the Serra do Gandarela National Park, in Minas Gerais,[8] in Monte Bonito, the southern Rio Grande,[9] as well as the Toca do Tatu in Santa Catarina.[10]

After the extinction of the megafauna about 10,000 years ago, some paleoburrows were reused by indigenous human populations. Recent searches indicate that these structures were used as temporary shelters as well as for ritual purposes. In the interior of some paleoburrows, researchers discovered stone tools, ceramic artifacts, human burials, and inscriptions engraved in the walls.[citation needed]

Paleoburrows are classified as paleontological sites; however, if the remains of ancient populations are also found, the site may become the object of research for both paleontologists and archaeologists.[11][12]

Paleoburrows dug by giant ground sloths often have large claw marks along the walls.[4] Many paleoburrows are exposed by local development, but subsequently eventually destroyed by rain, the elements, or construction.[4]

Paleoburrows are sometimes enlarged by erosion.[5] One of the largest reaches 2,000 feet in length over all of its branching tunnels, originally over 6 feet tall and 3 to 5 feet wide.[5] It was made by many animals over generations.[5]

The exact animals which dug the largest burrows, up to 5 feet in diameter, are debated.[5] One group of scientists, including Heinrich Frank, believe that they were dug by giant ground sloths, possibly Catonyx, Glossotherium, or Lestodon.[5] Another group believes that even the largest burrows are attributable to extinct armadillos and/or pampatheres like Pampatherium, Holmesina or Propraopus, even though they were smaller than the sloths.[5]

It is unclear why the burrows were dug to their large sizes.[5] Whether the burrows were dug by giant armadillos or giant sloths, they are much larger than would be necessary.[5]

Crotovina

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Paleoburrows filled with sediments, deposited over the centuries through rainfall and accumulated due to the porosity of the terrain, are called crotovina. Generally, fossils found in crotovinas exhibit large proportions similar to known megafauna of their geological period.[13][14]

See also

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References

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Further reading

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Paleoburrow

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A paleoburrow is an ancient underground tunnel or shelter excavated by extinct megafaunal vertebrates during the Pleistocene epoch, serving as evidence of their burrowing behavior preserved in . These structures, recognized as the world's largest ichnofossils—trace fossils such as burrows or tracks left by prehistoric animals—are primarily found in southern and attributed to giant like Lestodon armatus, which could weigh over four tons and measure up to 15 feet in length. Paleoburrows vary in size but can extend up to 600 meters in length and reach heights of 2 meters or more, with some chambers spanning over 12 feet high and widths sufficient for multiple large animals. Formed more than 10,000 years ago during the , these tunnels were likely dug for shelter from predators, extreme weather, or as seasonal refuges, and their discovery has revolutionized by providing direct insights into the and locomotion of extinct South American . Smaller paleoburrows may have been created by other , such as giant armadillos, highlighting a diverse range of burrowing activities among Ice Age herbivores. The study of paleoburrows, often termed "paleotocas" in , has grown since their systematic identification in the early 2000s, revealing over 1,500 such structures in regions like the state of . These sites preserve trace fossils such as claw marks and burrow structures, and contribute to geoheritage conservation efforts, as they offer unique windows into the and environmental dynamics of prehistoric before human arrival. Ongoing research emphasizes their role in understanding megafaunal extinction patterns and the paleoenvironmental conditions of the continent.

Definition and Characteristics

Definition

A paleoburrow is an underground tunnel or chamber system excavated by extinct vertebrate megafauna during the Pleistocene epoch, preserved as trace fossils, or ichnofossils, in sedimentary rock. These structures represent the largest known ichnofossils globally, providing evidence of subterranean behaviors in ancient ecosystems. Key attributes of paleoburrows include their typically horizontal or gently sloping configuration, often straight or slightly sinuous, with subcircular to subelliptical cross-sections that lack complex biogenic structures such as nests but exhibit clear excavation traces like grooves and scratches from digging. Dimensions vary widely, with lengths up to 600 meters, heights typically 2 meters but up to 4 meters in chambers, and widths up to 4 meters or more, reflecting the scale of the extinct excavators. The term "paleoburrow" derives from the prefix "paleo-" (ancient) and "," entering formal ichnological literature in the with the description of the ichnogenus Megaichnus. In South American contexts, the equivalent "paleotoca" is also used, emphasizing their ancient origins. Paleoburrows are distinguished from caves by their biogenic origin through animal excavation, featuring smooth internal walls and tool marks, in contrast to the abiotic dissolution processes of natural or anthropogenic features. In softer sediments, they often occur as infilled variants known as krotovinas, where the tunnels are backfilled with surrounding material.

Physical Features

Paleoburrows exhibit subhorizontal tunnels that are typically straight or slightly sinuous, with subcircular to subelliptical cross-sections that allow for the passage of large-bodied animals. The walls and ceilings often display smooth surfaces resulting from sediment compaction during excavation and subsequent animal movement, while floors may appear flattened due to infilling sediments. These structures are primarily excavated in consolidated sediments or soft rocks such as weathered volcanic materials and sandstones, contributing to their preserved morphology. Trace evidence within paleoburrows includes long, shallow grooves interpreted as digging marks, which are commonly parallel to one another and grouped in sets suggestive of impressions from two or three claws. These scratches appear on walls and ceilings, with occasional impressions of osteoderms or possible on upper surfaces, indicating direct contact during . In some cases, enlarged portions or intersecting tunnels show sediment displacement patterns aligned with the direction of excavation. Paleoburrows vary significantly in size, with smaller forms classified as Megaichnus minor featuring widths of 0.6–0.9 meters and heights of 0.5–0.7 meters, or slightly larger variants up to 1.5 meters wide and 0.9 meters high. Larger examples, known as Megaichnus major, reach heights of approximately 2 meters and widths up to 4 meters, often narrowing toward the ends, and these megatunnels can extend over 50 meters in length. Total burrow lengths range from several meters to over 600 meters in some systems. Chambers can reach up to 10 meters wide and 4 meters high. As of 2023, a 340-meter-long paleoburrow was documented in , . Internal structures are generally simple, consisting of a main with occasional branching into smaller ramifications or chambers, though complex networks of interconnected burrows occur in densely populated sites. About 70% of larger paleoburrows are partially filled with sediments or rock fragments, and smooth elliptical areas on floors or walls may represent resting zones, but no features indicative of prolonged habitation, such as , are present. These morphological traits provide a scale comparable to burrows constructed by modern large xenarthrans, such as giant armadillos or two-toed sloths, though adapted to extinct megafaunal dimensions.

Geological and Taphonomic Context

Formation Processes

Paleoburrows, known ichnogenus Megaichnus, were excavated by extinct megafaunal xenarthrans, primarily mylodontid such as Glossotherium and Scelidotherium, using their specialized forelimbs equipped with large, curved claws on digits II and IV. These animals used their forelimbs equipped with large, curved claws, where robust humeri and strong radial s provided the mechanical leverage to loosen and displace , creating parallel grooves and scratches visible on burrow walls. Sediment removal occurred through pushing or dragging loosened material outward with the forelimbs, as inferred from the limb proportions that favored force over speed in digging activities. The behavioral drivers for excavation centered on creating temporary or semi-permanent shelters, or domichnia, to evade predators, mitigate , or aestivate during seasonal droughts in the subtropical Pleistocene landscapes of . Uniform wall textures and smooth surfaces in many paleoburrows suggest rapid digging sessions, possibly completed in days or weeks, followed by maintenance through body or intentional rubbing to enlarge passages. Evidence from associated ichnofabrics indicates reoccupation by multiple individuals or species, underscoring the burrows' utility as refuges during periods of environmental stress. Formation required specific environmental conditions, including consolidated but friable substrates such as weathered aeolian sandstones, paleosols, and volcanic rocks in subtropical regions, which could be penetrated and excavated using claws despite their firmness. These sediments, often found on moderate slopes near water sources, provided the ideal friable consistency for claw-based loosening, with harder consolidated layers like basaltic or granitic rocks occasionally worked in later stages. Paleoburrows were predominantly formed during the (approximately 126,000 to 11,700 years ago), aligning with the peak abundance and distribution of South American before their .

Preservation and Identification

Paleoburrows are primarily preserved through taphonomic processes that involve rapid post-abandonment infilling and subsequent , particularly in subtropical environments with aeolian deposits, such as those in southern . Following abandonment, these structures often experience quick with wind-blown sands or debris, which stabilizes the tunnels and prevents significant or erosion. In regions like southern , where many paleoburrows occur in Pleistocene aeolianites and sandstones, cementation occurs via in the porous sediments, leading to the hardening of walls and infill materials over time. This process has allowed traces to endure for more than 10,000 years, spanning the to transition, as evidenced by dated sites in consolidated coastal barrier systems. Recent studies (as of 2024) in regions like the Caminhos dos Cânions do Sul Global Geopark have documented additional paleoburrows, providing further insights into their taphonomic preservation in subtropical paleoenvironments. Identification of paleoburrows relies on distinct criteria that highlight their biogenic origins, such as the presence of diagnostic traces like shallow grooves and scratches on walls and ceilings, which indicate mechanical excavation rather than abiotic formation. Cross-sectional analysis often reveals layered infill with laminations, textural variations, or color differences from surrounding host rock, reflecting episodic deposition from multiple sediment horizons and distinguishing these from non-biogenic features lacking such patterns, such as water-eroded cavities without smooth, friction-polished surfaces. Morphological features, including subhorizontal tunnels with diameters ranging from 0.6 to 4 meters and slightly sinuous paths, further support identification when combined with the absence of typical abiotic indicators like irregular fracturing. Diagnostic challenges arise in differentiating paleoburrows from pseudofossils such as root casts, which typically exhibit vertical branching and fibrous textures without linear scratches, or from surface animal trails that lack subsurface continuity. To confirm biogenic origins, researchers employ 3D mapping techniques, such as or scanning, alongside sedimentological analysis to assess , composition, and depositional fabrics within the infill. These methods help verify the structural discordance of paleoburrows with host strata and rule out natural cavities like lava tubes, which show irregular walls and no biogenic markings. Crotovinas represent a subset of paleoburrows, specifically those excavated in softer substrates like or that undergo complete backfilling with shortly after abandonment. In such environments, processes cause the filled burrows to be transposed downward through bioturbation and pedogenesis, resulting in structures that are fully sediment-infilled without persistent open voids, unlike many exposed paleoburrows in lithified aeolian deposits. This taphonomic pathway preserves crotovinas as subtle traces in soil profiles, often identifiable only through their oblique orientation and infill contrasts relative to the surrounding matrix.

Biological Attribution

Candidate Megafauna

The primary candidates for excavating paleoburrows are giant of the family , such as Scelidotherium leptocephalum, Glossotherium robustum, and Lestodon armatus, with anatomical adaptations suited for extensive digging activities. Attribution to is supported by matching claw marks and biomechanical features, though larger paleoburrows (Megaichnus major) have been proposed for megatheriids like and . These extinct xenarthrans reached lengths of up to 6 meters and weights of approximately 4 tons, with robust forelimbs and massive claws measuring 20-30 centimeters in length, as evidenced by skeletal fossils that indicate powerful leverage for soil displacement. The exact family remains debated, with some studies favoring based on ichnological evidence. Secondary candidates include giant cingulates such as pampatheres (Pampatherium spp.) and glyptodonts (Panochthus fraterculus), which measured about 1.5 meters in length and featured heavily armored forelimbs with enlarged claws optimized for excavation. These xenarthrans, related to modern armadillos but scaled up significantly, exhibited osteoderm-covered limbs that could reinforce digging efficiency against hard soils, as inferred from fossilized forelimb morphology. Species with glyptodont-like armor, including robust humeri and short, powerful metacarpals, further support their capability for creating narrower paleoburrows. These inhabited Pleistocene , spanning open grasslands, savannas, and forested regions where burrowing likely served as protection from predators and environmental stressors. Their occurred between 12,000 and 10,000 years ago, coinciding with rapid climate shifts at the end of the and the arrival of human populations, which together exerted pressures on these large herbivores. Proboscideans, such as gomphotheres present in South America, are excluded as candidates due to the incompatibility of burrow dimensions and internal traces with their trunk-based locomotion and lack of digging adaptations, as burrow morphologies instead match xenarthran skeletal proportions.

Evidence Linking Animals to Burrows

Paleoburrows exhibit distinctive trace fossils on their internal surfaces, including numerous parallel scratches and grooves interpreted as marks from large claws during excavation. These scratches, often 20-25 cm long and oriented in patterns consistent with forelimb strokes, align with the anatomy of extinct ground sloths, such as the three-toed manus of mylodontids like Scelidotherium, which possessed robust claws up to 30 cm in length. Such bioglyphs are absent in burrows attributed to non-mammalian origins, providing ichnological evidence for megafaunal digging activity, particularly Mylodontidae. The dimensions of paleoburrows further support attribution to giant . Tunnels typically measure 1-2 m in height and up to 4 m in width, with some chambers reaching 1.8 m high, correlating closely with the shoulder height of species like americanum (approximately 1.8-2 m) and allowing passage for animals weighing 2-4 tons. Smaller variants, classified as Megaichnus minor, have diameters of 0.9-1.5 m, suitable for mid-sized s such as Scelidotherium, while larger Megaichnus major forms exceed 2 m, potentially accommodating larger megatheriids or mylodontids like Lestodon. These measurements preclude or smaller vertebrate origins and match skeletal reconstructions of sloth forelimbs adapted for powerful . Comparative ichnology reinforces the link to xenarthrans, particularly . Modern giant s (Priodontes maximus) construct burrows with entrances around 0.4-0.6 m wide and tunnels 2-5 m long, showing similar subhorizontal morphology but scaled down proportionally to their 1.5 m body length; this analogy extends to extinct sloths, whose burrows exhibit expanded chambers suggestive of resting or sheltering behavior observed in extant burrowers. The ichnogenus Megaichnus, established for these structures, encompasses species like M. major for sloth-attributed paleoburrows, distinguished by their size and scratch patterns from smaller armadillo traces. Rare associations of fossils with paleoburrows provide direct evidence of occupancy. For instance, a complete Scelidotherium skeleton was discovered within a burrow filled with near , , indicating use as a shelter. Isotopic dating of infill sediments places many paleoburrows in the , around 12,000-10,000 years BP, coinciding with the extinction window of South American megafauna including .

Discovery and Distribution

History of Research

The earliest observations of what are now recognized as paleoburrows date back to the early in southern , where they were initially noted during geological surveys and misattributed to activity or natural erosion processes. In 1933, German-Brazilian geologist Friedrich Kurt Padberg-Drenkpol documented the first known example in the state of Santa Catarina, interpreting the tunnels as potential archaeological structures possibly linked to ancient indigenous or colonial excavations. Similar misattributions persisted, with some locals and early explorers associating the features with Jesuit treasure hoards or artificial mining, leading to anecdotal reports rather than systematic study. A shift toward recognizing their biogenic origins began in the late 20th century. In 1994, Brazilian paleontologists Lúcia P. Bergqvist and Noelia B. Maciel identified burrows in middle Pleistocene sediments as animal-made, challenging prior human-centric interpretations. This laid groundwork for modern ichnology, though widespread acknowledgment remained limited until the early , when over 1,500 paleoburrows were documented across southern and southeastern through targeted fieldwork. Key contributions came from Heinrich T. Frank, who, starting around 2000, led expeditions that uncovered major examples and published initial analyses in 2008, 2010, 2012, and 2013, emphasizing scratch marks and tunnel morphology as evidence of megafaunal excavation. These efforts addressed early gaps in understanding, moving from isolated reports to evidence-based attribution to extinct mammals like giant . Formal ichnological classification advanced in 2016 with the establishment of the genus Megaichnus by Rafael A. Lopez, Heinrich T. Frank, and colleagues, who described the largest paleoburrows (M. major) as traces of Cenozoic megafauna based on biometric and sedimentological data from Brazilian sites. This seminal work, published in Ichnos, integrated comparative anatomy and trace fossil analysis, solidifying paleoburrows as a distinct category of vertebrate ichnofossils. Pre-2010 underestimation of their scale and biogenic nature was further rectified by multidisciplinary teams involving paleontologists, geologists, and sedimentologists. Methodological progress accelerated in the and , transitioning from manual surveys to advanced techniques like scanning and for tunnel volumetrics and . Researchers such as Luiz Carlos Weinschutz applied these tools to map internal structures and analyze sediment infill, enhancing preservation insights without destructive excavation. By the , studies expanded into geoheritage, with paleoburrows integrated into Global Geopark initiatives like Caminhos dos Cânions do Sul, promoting conservation and public education on their Pleistocene origins. This era marked a collaborative phase, with Brazilian universities forming the Paleoburrows Project to combat misinformation and advocate for site protection.

Major Sites and Locations

Paleoburrows are predominantly distributed across , with the highest concentration in southern , where over 1,500 sites have been documented in the states of , Santa Catarina, and within the sediments of the Paraná Basin. These structures occur primarily in paleosols developed on aeolian sandstones and related deposits, often preserved 10-20 meters below the modern surface due to sedimentary infilling and overburden. The geological setting reflects a Pleistocene environment of stabilized dunes and soil horizons suitable for excavation by large vertebrates. Key localities in southern include the Southern Canyons Paths Global , spanning and Santa Catarina, where accessible paleoburrows such as Toca do Tatu in Timbé do Sul—featuring associated —and Engenho Velho in Jacinto Machado exemplify well-preserved networks in weathered outcrops. In the metropolitan area of , a dense cluster of giant paleoburrows has been identified in granitic and substrates, highlighting urban-proximate discoveries. Further north, in , a notable 340-meter-long paleoburrow and a complex of six interconnected tunnels—each around 40 meters long with chambers up to 10 meters wide and 4 meters high—represent some of the largest documented examples, recently designated for protection. Beyond , paleoburrows are rarer but emerging in neighboring countries, aligning with the historical range of Pleistocene . In , large burrows attributed to extinct have been reported. In , similar structures occur in loessoid deposits near San Pedro and along the coast of . Potential sites in include underground tunnels in the Chumbivilcas Province of , interpreted as Pleistocene megafaunal paleoburrows in local geological contexts. Overall, these occurrences are confined to regions with South American Pleistocene distributions and are absent in , where differing faunal assemblages prevailed.

Scientific and Cultural Significance

Paleobiological Insights

Paleoburrows provide critical evidence of complex behavioral patterns among extinct xenarthran , particularly such as Lestodon armatus and americanum. The branching and interconnected chamber structures of these burrows, often extending up to 600 meters in total length with multiple galleries, suggest shelter-seeking in social groups, where branching indicates use by multiple individuals or successive generations rather than solitary excavation. Claw marks and resting hollows within chambers further imply communal occupation, potentially including adults with juveniles, as reoccupation patterns show extended use over time. These features point to burrows serving as refuges from environmental stressors like arid conditions during glacial phases of the Pleistocene, and possibly from predators, adapting to the harsher climates of southern . Ecologically, paleoburrows reveal the role of xenarthran in modification, acting as ecosystem engineers through large-scale excavation that aerated and created stable subterranean microenvironments. These structures, preserved in sediments, facilitated reoccupation by diverse taxa, enhancing in transitional landscapes. Integration with regional records from the terminal Pleistocene indicates that such burrowing occurred at grassland-forest interfaces, where likely influenced vegetation dynamics through sediment displacement and potential in tracked materials, though direct evidence for the latter remains indirect. This underscores their contribution to turnover and landscape heterogeneity in pre-extinction . Recent 2024 research has utilized paleoburrows as indicators of seasonality in environments, while a new of was found inhabiting them, highlighting their ongoing role in modern . The dating of paleoburrows to the terminal Pleistocene, with associated fossils like Catonyx cuvieri calibrated to approximately 9,960 ± 40 BP (mean 11,315 cal BP), demonstrates persistence of burrowing xenarthrans until around 10,000 BP, aligning with the Late Quaternary extinction event in . This temporal overlap with human arrival in implies potential interactions, though direct evidence such as co-occurring human artifacts with remains is limited and requires further research. Over 80% of South American >44 kg, including these burrowers, vanished by this period, highlighting the burrows' value in tracing extinction dynamics. As the first direct evidence of burrowing in xenarthrans, paleoburrows challenge prior assumptions of predominantly arboreal or semi-arboreal lifestyles for many , revealing instead a that emerged or intensified in the Pleistocene. This shifts understanding of their locomotor and ecological capabilities, emphasizing terrestrial behaviors in response to environmental pressures.

Geoheritage and Conservation

Paleoburrows have gained recognition as key elements of geological heritage in , particularly within the Caminhos dos Cânions do Sul Global in the states of Santa Catarina and , where 24 such structures have been documented and four designated as geosites of regional and international significance due to their ichnofossil value. This geopark status underscores their role in preserving paleoenvironmental records from the Pliocene-Pleistocene era. In 2023, the largest known paleoburrow in was officially protected under state legislation to shield it from human impacts, marking a step toward formalized conservation. Despite these advancements, paleoburrows remain vulnerable to multiple threats, including from unguided visitors, exposure and destruction during urban expansion and construction projects, and natural degradation from , water flow, and rockfalls. Over 400 paleoburrows in the metropolitan area alone have been uncovered through large-scale anthropogenic excavations, highlighting the risks posed by development. Mitigation strategies include guided tours and sign-posted access in protected areas, 3D mapping and for , and the establishment of research-oriented reserves within the framework to limit direct access while supporting scientific study. The Paleoburrows Project, a collaborative effort by researchers from Brazilian universities, promotes awareness and mapping to bolster these protections. These sites hold substantial educational value, fostering public engagement through initiatives in the Caminhos dos Cânions do Sul Global , where visitors learn about megafaunal legacies via interpretive trails and geoeducational programs. They also connect to , such as indigenous and settler folklore interpreting the burrows as mystical formations, enhancing outreach in museums and paleontological centers across southern . On a global scale, paleoburrows stand out as the largest ichnofossils ever recorded, analogous to renowned sites like trackways in their ability to reveal behavioral insights, though they uniquely document subterranean megafaunal activity. This distinctiveness has spurred recommendations for international surveys to explore similar structures beyond , emphasizing their potential for broader geoheritage networks.

References

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