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Burrow

A burrow is a hole or tunnel excavated into the ground or other substrate by an animal to serve as a space for habitation, temporary refuge, or protection from predators and environmental conditions.^[1] These structures vary widely in complexity, from simple shafts dug by insects to extensive networks created by mammals, and they play crucial roles in the animals' survival strategies.^[2] Burrows are constructed by a diverse array of species across terrestrial and aquatic environments, including mammals like rabbits, groundhogs, and prairie dogs; reptiles such as tortoises; birds like burrowing owls; and invertebrates including ants, worms, clams, and crustaceans.^[1] Fossorial animals, specialized for burrowing, often possess adaptations such as strong claws, powerful limbs, or streamlined bodies to facilitate excavation, enabling them to rework soil and create habitats that support their lifestyles.^[3] Burrow types can be classified into primary burrows dug directly by the occupant, secondary burrows modified by other species, or tertiary ones reused by yet another animal, reflecting complex interactions within ecosystems.^[4] Ecologically, burrows enhance soil aeration, nutrient cycling, and water infiltration, while providing shelter that boosts biodiversity by hosting multiple species in a single system.^[5] For instance, abandoned burrows from species like aardvarks serve as homes for hyenas, warthogs, and porcupines, fostering habitat diversity and supporting food webs in savannas and grasslands.^[6] These structures also influence landscape patterns, with burrow mounds attracting unique arthropod communities and aiding in seed dispersal or predator avoidance for cohabitants.^[7]

Definition and Characteristics

Definition

A burrow is a hole, tunnel, or complex underground structure excavated into the ground or other substrate by an animal to serve as a space for shelter, nesting, or temporary refuge.^[1] The term "burrow" originates from Middle English borowe or burh, a variant of burgh meaning a fortified dwelling or refuge, derived from Old English burh (stronghold), ultimately tracing back to Proto-Germanic *burgs denoting a protected enclosure or hill-fort.^[8] This etymological root reflects the burrow's role as a secure retreat, evolving by the 14th century to specifically describe animal-dug subterranean habitats. Burrows differ from other animal shelters such as above-ground dens (e.g., bear dens in hollow logs), rock shelters, or hollows used by larger mammals for resting or rearing offspring, though some dens, like those of foxes, are themselves excavated burrow systems.^[9] Nests, by contrast, are usually built above ground from gathered organic materials like twigs, leaves, or silk, primarily for egg-laying or young-rearing in birds, insects, and some mammals.^[10] Caves represent natural geological formations—such as limestone hollows—rather than excavations created by animals, though some species may occupy or modify them.^[11] In terms of scale and complexity, burrows range from simple vertical shafts, exemplified by basic rabbit holes that provide quick access and minimal elaboration, to elaborate multi-chambered networks, such as the interconnected systems in prairie dog towns that support communal living across vast areas.^[12] This variation in design accommodates diverse environmental needs while maintaining the core subterranean nature of the structure.^[13]

Physical Features

Burrows exhibit a variety of structural elements that form their basic architecture, including entrance shafts that provide access from the surface, interconnected tunnels for navigation, enlarged chambers for resting or storage, ventilation holes to facilitate air exchange, and secondary escape routes for alternative exits.^[14]^[1] These components create a network that can range from simple linear passages to complex, multi-level systems, depending on the environmental context.^[15] The physical properties of burrows are profoundly influenced by soil type and material interactions, with loose sandy soils often requiring deeper tunnels for stability to prevent collapse, while cohesive clay soils support more stable excavations at varied depths.^[16] Stability is enhanced in many cases through natural reinforcement of burrow walls, such as the application of mucus or saliva linings that bind soil particles and prevent erosion or caving.^[17]^[18] Size variations in burrows are extensive, with diameters typically ranging from a few millimeters in small invertebrate systems to over a meter in large vertebrate complexes, and depths extending from near-surface levels to several meters below ground.^[19]^[20] These dimensions are constrained by soil mechanics, where sandier substrates necessitate deeper constructions for structural integrity.^[21] Certain burrow designs incorporate environmental adaptations, such as spiral-shaped entrances that twist downward and water-filled sections in proximity to aquatic zones, which help maintain internal conditions amid varying moisture levels.^[18]^[22]

Animal Groups and Burrows

Vertebrate Burrows

Vertebrate burrows are subterranean structures excavated or occupied by animals with backbones, serving as shelters, nesting sites, or dormancy chambers, and varying widely in complexity based on the species' ecology and behavior. These burrows often reflect adaptations to specific environments, such as arid soils or aquatic sediments, and can range from simple tunnels to elaborate networks supporting social groups. Among mammals, rabbits construct extensive warrens that function as multi-generational complexes, with interconnected tunnels and chambers supporting breeding colonies over successive years. These structures can span significant areas, with some systems extending up to hundreds of meters in total tunnel length, allowing for escape routes and communal living.^[23] Prairie dogs form social networks known as towns, which are vast burrow systems covering up to 400 hectares or more, divided into wards that facilitate group vigilance and foraging. Badger setts consist of deep underground chambers designed for family units, often reaching depths of several meters with multiple entrances and sleeping areas that accommodate social groups of adults and young. Birds also utilize burrows for nesting, with burrowing owls creating ground-level nests in abandoned mammal burrows or self-dug tunnels up to 3 meters long, often decorated with dung to attract insects. Kingfishers excavate riverbank tunnels for nesting, typically 1 to 2 meters in length and slightly upward-sloping to prevent flooding, where both parents share incubation duties in a terminal chamber. Reptiles and amphibians employ burrows for survival in harsh conditions; desert tortoises dig burrows for aestivation during extreme summer heat, retreating into these refuges to conserve water and regulate temperature. In arid regions, certain frogs construct burrows with plugged entrances using soil or mucus, sealing themselves in to prevent desiccation during dry periods and emerging after rains. Fish burrows occur in sedimentary environments, as seen in jawfish, which maintain vertical burrows in marine rubble, reinforced with stones or coral fragments up to 30-50 cm deep for shelter and mouthbrooding eggs. Lungfish in freshwater systems create aestivation burrows during seasonal droughts, forming flask-shaped chambers lined with a mucus cocoon to survive months of estivation until water returns. Vertebrates excavate burrows using specialized traits, such as claws for scratch-digging in species like badgers and prairie dogs, incisor teeth for chisel-digging in rodents, or body pushing with the head to compact soil. Burrow use varies from solitary, as in territorial blind mole rats that aggressively defend individual tunnels, to highly social systems like prairie dog towns, where communal digging enhances group protection and resource sharing.

Invertebrate Burrows

Invertebrate burrows exhibit remarkable diversity in form and function, often characterized by their micro-scale dimensions, typically with diameters under 1 cm, which facilitate high-density networks in soil, sediment, or wood substrates. These structures contrast with the larger, more singular burrows of vertebrates by enabling intricate, interconnected systems that support colonial or solitary lifestyles tied to specific ecological niches. Invertebrates, lacking rigid endoskeletons, rely on flexible body plans to excavate and inhabit these spaces, producing burrows that range from simple tunnels to complex architectures adapted for protection, feeding, and reproduction. Among insects, ant colonies form extensive subterranean networks with chambers dedicated to brood rearing, food storage, and fungal cultivation, often extending meters deep with branched tunnels reinforced by soil particles. Termite mounds represent elaborate above-ground extensions of underground burrows, featuring ventilated tunnels and chambers that regulate internal climate through passive airflow.^[24] Solitary bees, such as mining species, construct individual burrows in soil or decayed wood, provisioning them with pollen for larval development before sealing them temporarily.^[25] Other arthropods demonstrate specialized burrow designs; for instance, trapdoor spiders line their vertical shafts with silk secreted via spinnerets and chelicerae, capping entrances with hinged, camouflaged doors for ambush predation.^[26] Crayfish, particularly burrowing species in aquatic margins, build chimney-like structures atop tunnels in stream banks, using these protrusions to enhance burrow ventilation and access surface water.^[27] In mollusks and annelids, clams embed in marine or estuarine sediments, extending siphons through U-shaped burrows to filter feed while minimizing exposure. Earthworms create semi-permanent vertical channels that promote soil aeration by facilitating water and air percolation.^[28] Invertebrate burrowing often employs appendages like mandibles in insects for scraping and excavating soil, chelicerae in arachnids for manipulating silk and debris, or peristaltic body contractions in worms for propulsion through sediment. These burrows are frequently temporary or seasonal, aligned with life cycles such as larval stages or reproductive periods, allowing invertebrates to abandon and re-excavate as environmental conditions shift.^[29]

Construction Processes

Excavation Methods

Animals employ a variety of biomechanical and behavioral techniques to excavate burrows, primarily relying on mechanical forces to displace soil or sediment. These methods are adapted to the animal's anatomy and the substrate's properties, such as hardness, cohesion, and moisture content. In subterranean mammals, excavation often involves creating cylindrical tunnels that match the body diameter to minimize energy expenditure and ensure structural stability.^[30] Mechanical digging is the predominant technique, utilizing specialized appendages or body movements to loosen and remove material. Moles (Talpidae) exemplify limb-based excavation, employing oversized front paws with broad palms and sharp claws in a breast-stroke motion to scrape soil forward and push it aside. This allows them to dig at rates sufficient to construct extensive tunnel networks, with forelimbs rotating outward for efficient displacement. Pocket gophers (Geomyidae), in contrast, combine forelimbs with powerful incisor teeth protruding from the jaws to chisel through roots and compact soils; their lower incisors grind against upper ones to maintain sharpness during digging, enabling them to excavate in harder substrates where limbs alone would be insufficient.^[31]^[32]^[33] Body undulation serves as a primary excavation method for limbless burrowers like certain snakes, particularly in loose granular media such as sand. Species like sand boas (e.g., Eryx spp.) propagate lateral waves along their body to generate thrust against the burrow walls, fluidizing the substrate and propelling forward while minimizing slippage; this undulatory locomotion reduces penetration resistance by fracturing or compressing sediment ahead. In soft-bodied invertebrates, similar peristaltic undulations—rhythmic contractions expanding the anterior body—create space by compacting walls and allowing backward expulsion of material.^[19] Chemical assistance is even less common, with no well-documented cases of acid secretion specifically for softening soil in burrowing ants; formic acid in species like Formica is primarily defensive rather than excavatory.^[34] Energy efficiency in excavation is influenced by substrate resistance and body size, with burrowing costs ranging from 360 to 3,400 times higher than surface locomotion due to soil compaction forces. Smaller animals like tuco-tucos (Ctenomys talarum) exhibit elevated metabolic rates in harder soils, while tunnel diameter optimization—typically 3–15 cm—balances excavation effort against collapse risk. Burrowing speeds vary widely. Soil displacement techniques further enhance efficiency: moles push loosened earth backward with hindlimbs before surfacing it via molehills, gophers perform in-tunnel somersaults to carry soil in cheek pouches or kick it rearward, and undulators like snakes compact material against burrow walls for later expulsion.^[35]^[30]^[36] Environmental constraints, particularly hard or rocky substrates, drive specialized adaptations. Aardvarks (Orycteropus afer) feature reinforced snouts with keratinized tips and powerful foreclaws to pry open termite mounds and dig in compacted soils, allowing penetration where softer-nosed burrowers cannot; their nostrils seal during excavation to exclude dust. These adaptations ensure survival in diverse soils, from loose sands to clay-rich earth, by distributing forces to prevent tool or limb damage.^[37]^[38]

Modification and Reuse

Animals frequently modify the internal structure of burrows after initial excavation to enhance functionality and comfort. For instance, red foxes (Vulpes vulpes) line their den chambers with dry grass, leaves, feathers, and fur to provide insulation and a soft bedding for pups.^[39] Similarly, pocket gophers (Thomomys spp.) construct specialized chambers within their burrow systems for storing plant roots and tubers, allowing them to cache food for later consumption without repeated surface foraging.^[40] To regulate access and protect against intruders or environmental threats, many burrowers add earthen plugs or seals at entrances; pocket gophers, for example, routinely block tunnel openings with soil to maintain burrow integrity and prevent flooding or predation.^[40] Burrows are often repurposed by different species, leveraging existing structures to reduce excavation energy costs. Burrowing owls (Athene cunicularia) commonly occupy abandoned prairie dog (Cynomys ludovicianus) burrows, which provide ready-made shelter in open grasslands; this reuse is particularly prevalent in the Great Plains, where up to 90% of owl nests in some areas are sited in former prairie dog towns.^[41] Red foxes similarly adopt and modify burrows originally dug by prey species, such as woodchucks (Marmota monax) or rabbits (Oryctolagus cuniculus), expanding entrances or adding linings to suit their needs.^[39] These modifications are driven by behavioral adaptations to life stage or environmental changes. Expansion of burrow systems often occurs to accommodate growing family sizes; subterranean mammals like pocket gophers enlarge tunnels and chambers during breeding seasons to house litters and increased food stores.^[16] Seasonal shifts may prompt shortening or sealing of tunnels for energy conservation. Over generations, burrow systems exhibit succession patterns where primary excavators yield to secondary occupants, fostering complex ecological interactions. In prairie dog colonies, initial diggers create extensive networks that, after abandonment due to predation or disease, are successively inhabited by burrowing owls, snakes, and insects, enhancing habitat heterogeneity and supporting biodiversity in grassland ecosystems.^[42] This multi-species occupancy can persist for years, with structures modified incrementally by each user, demonstrating the long-term durability and adaptability of burrow architectures.^[43]

Functions and Adaptations

Protection Mechanisms

Burrows function as primary defensive structures for many burrowing animals, incorporating architectural features and behavioral adaptations that deter predators and enhance survival. These mechanisms exploit the burrow's subterranean nature to provide refuge from aerial, terrestrial, and burrowing threats, allowing inhabitants to evade detection or direct confrontation.^[44] Predator deterrence often relies on burrow design elements such as multiple entrances and hidden locations, which facilitate rapid escape and confuse pursuing predators. For instance, California ground squirrels construct burrow systems with numerous interconnected entrances, enabling quick evasion of non-burrowing predators like coyotes or hawks by providing alternative exit routes.^[45] Similarly, meerkats (Suricata suricatta) utilize complex burrow networks with multiple bolt-holes during foraging, while incorporating sentinel behavior where group members perch atop mounds or near entrances to scan for threats and emit alarm calls, alerting others to retreat underground. This vigilance is particularly effective in open savannas, where sentinels reduce group predation risk through coordinated warnings. Physical barriers within burrows further impede predator intrusion, including collapsible tunnels and blocking mechanisms tailored to specific threats. Pocket gophers (Thomomys spp.), for example, rapidly construct earthen plugs to seal tunnels against invading snakes, effectively creating temporary barriers that prevent deeper access.^[40] In some rodent species, such as certain desert-dwelling gerbils, burrows feature reinforced chambers with loose soil that can collapse under pressure from digging predators, buying time for escape.^[45] Although less common in mammals, flooded chambers observed in semi-aquatic burrowers like muskrats (Ondatra zibethicus) deter terrestrial predators by inundating tunnels during high water, a strategy that parallels defensive flooding in some arid-zone rodents during rare floods.^[46] Camouflage and detection avoidance enhance burrow security by minimizing predator awareness of the refuge. Many burrowing species, including prairie dogs (Cynomys spp.), construct surface mounds that mimic natural soil elevations, blending entrances into the landscape to avoid visual cues.^[44] Scent masking provides an additional layer; California ground squirrels chew and rub snakeskins on their fur and burrow entrances to overlay their odor with reptilian scents, reducing detectability by mammalian predators reliant on olfaction.^[47] A notable case study is the burrow systems of wombats (Vombatus ursinus and Lasiorhinus spp.), which evade dingoes (Canis dingo) through exceptional depth and complexity. These burrows extend up to 30 meters in length and 3.5 meters deep, featuring multiple chambers and entrances interconnected by narrow, twisting tunnels that exhaust pursuing predators and limit maneuverability.^[48] Wombats reinforce defense by positioning their cartilaginous, rock-hard rumps at entrances to block access, capable of crushing intruders against the tunnel roof.^[49] This combination allows wombats to spend approximately 70% of their time underground, surviving in predator-rich environments where surface activity would be lethal.^[48]

Environmental Regulation

Burrows play a crucial role in thermoregulation by providing insulation against extreme external temperatures, creating stable microclimates that help occupants maintain optimal body temperatures. In arid environments, the greater bilby (Macrotis lagotis) utilizes spiral burrows up to 2 meters deep, which maintain a relatively constant internal temperature of approximately 23–24°C, buffering against surface highs exceeding 40°C in summer.^[50] This stability reduces thermal stress, with burrow temperatures showing significantly less variability than surface conditions, often 5–10°C cooler during peak heat.^[51] Similarly, in temperate regions, burrowing mammals like the European badger (Meles meles) experience burrow temperatures that are more moderate year-round, minimizing energy costs for heating or cooling the body.^[52] Humidity control and ventilation are facilitated by burrow architecture, which promotes airflow and water retention to prevent desiccation or excessive moisture. Tunnel designs in species such as prairie dogs (Cynomys spp.) incorporate multiple entrances and chambers that drive convective airflow through buoyancy effects, exchanging stale air for fresher, oxygen-rich air while moderating humidity levels.^[53] In amphibians like the burrowing sandhill frog (Arenophryne rotunda), burrows in sandy soils retain up to 28% moisture at saturation, allowing the frogs to absorb water osmotically through their skin during estivation, thus maintaining hydration in dry conditions.^[54] These features ensure higher relative humidity inside burrows compared to arid surface conditions.^[55] Beyond temperature and humidity, burrows regulate other environmental factors essential for survival, such as protection from desiccation in arid-adapted species and enhanced oxygenation in aquatic sediments. Arid-dwelling reptiles, including the desert tortoise (Gopherus agassizii), retreat into burrows that shield them from evaporative water loss, with internal conditions sustaining body water balance during prolonged droughts.^[56] In marine sediments, infaunal bivalves like the soft-shell clam (Mya arenaria) construct U- or Y-shaped burrows that pump oxygenated water through siphons, increasing local oxygen penetration by up to 5–10 cm into otherwise hypoxic layers.^[57] This ventilation prevents sulfide toxicity and supports respiration in low-oxygen environments.^[58] The stable microclimates of burrows also influence physiological processes, notably by lowering metabolic rates and conserving energy. In desert tortoises, burrow temperatures of 25–32°C during the day allow for reduced oxygen consumption rates compared to surface exposure, enabling metabolic depression that can halve energy expenditure during inactivity.^[56] Similarly, dormant African hinge-back tortoises (Kinixys spekii) in burrows exhibit oxygen uptake rates 50–70% lower than active individuals at equivalent temperatures (10–25°C), facilitating survival through seasonal dry periods.^[59] These adaptations underscore how burrows optimize physiological efficiency for long-term occupancy.

Ecological and Historical Significance

Modern Ecological Roles

Burrows constructed by earthworms and rodents play a crucial role in soil aeration and nutrient cycling within modern ecosystems. Earthworm burrows enhance soil porosity, facilitating greater water infiltration rates—up to four times higher in tilled fields compared to unburrowed soil—and promoting root growth by improving oxygen availability to plant roots.^[60] These activities also accelerate organic matter decomposition and nutrient recycling, incorporating residues into deeper soil layers and enriching fertility through casts high in organic carbon and nitrogen.^[61] Similarly, rodent burrows, such as those of pocket gophers and ground squirrels, increase soil porosity and aeration, leading to higher water infiltration and reduced soil compaction around burrow entrances, which supports enhanced root penetration and plant establishment in grasslands and agricultural fields.^[62] Abandoned rodent tunnels further contribute to long-term drainage and nutrient distribution by creating persistent channels.^[63] Beyond soil processes, burrows serve as vital habitats and refuges for non-burrowing species, boosting biodiversity in contemporary landscapes. Abandoned mammal burrows, including those of rodents and tortoises, provide stable microenvironments with moderated temperatures and humidity, attracting insects such as beetles, crickets, and bumble bees that nest in these cavities.^[64] For instance, gopher tortoise burrows host over 300 invertebrate species, many of which rely on the burrows for shelter and are absent from surrounding areas, while abandoned rodent tunnels offer nesting sites for bumble bees and carpenter bees in grasslands and forests.^[65] These structures can persist for up to 10 years post-abandonment, extending their role as refugia for diverse taxa during environmental stresses like drought or fire.^[64] Burrows also mediate trophic interactions, influencing predator-prey dynamics and seed dispersal in ecosystems. By offering refugia from surface predators and stressors, burrows alter encounter rates and survival, as seen in microbial communities protected within predator burrows or chipmunks selecting burrow sites with features that enhance predator detection.^[66] In vertebrate systems, burrows facilitate prey protection—such as prairie dog towns serving as hunting grounds and escapes for black-footed ferrets—while thwarting predators through complex architectures that limit access.^[67] Additionally, burrowing rodents like kangaroo rats promote seed dispersal via mound caches and scatter-hoarding, where stored seeds germinate after pilferage or abandonment, enhancing plant diversity and recruitment in arid and grassland habitats.^[68] In human-modified landscapes, burrows present both challenges and opportunities in agriculture and conservation. Pocket gopher burrows damage crops by feeding on roots and stems, disrupting irrigation through tunnels that divert water, and contaminating hay with soil mounds, leading to equipment wear and yield losses exceeding $10 million annually in major Nebraska field crops.^[69] Conversely, in conservation efforts, burrows of keystone species like gopher tortoises support endangered taxa—providing habitat for over 360 co-occurring species, including the indigo snake and burrowing owl—while prairie dog colonies sustain black-footed ferrets through burrow networks mapped for habitat restoration.^[70] These roles underscore the need for integrated management to balance ecological benefits against agricultural impacts.^[71]

Fossil Record

Burrows preserved as trace fossils, known as ichnofossils, provide critical evidence of ancient animal behaviors without relying on body fossils. These structures are classified into ichnogenera based on morphology, such as Skolithos, which represents simple vertical burrows typically attributed to suspension-feeding annelids or phoronids in firm substrates, and Thalassinoides, featuring complex branching networks formed by deposit-feeding crustaceans in softgrounds.^[72] Other notable examples include Treptichnus pedum, a serpentine burrow indicating probing behavior, and Planolites, unlined horizontal tunnels suggesting temporary dwellings.^[73] This classification system, developed through ichnotaxonomy, emphasizes architectural patterns like branching, lining, and spreite (leaf-like infills) to distinguish behaviors such as dwelling, feeding, or locomotion.^[74] The geological record of burrows begins in the Ediacaran Period around 565 million years ago with simple, shallow traces like Helminthoidichnites, narrow horizontal trails less than a few millimeters wide and rarely exceeding the sediment-water interface, reflecting minimal bioturbation.^[75] By the terminal Ediacaran (~555–541 Ma), traces show slight increases in complexity, including short vertical shafts and rare three-dimensional forms like Treptichnus. Recent discoveries as of 2025, including three-dimensional burrow systems from approximately 550-million-year-old strata in China, indicate that Ediacaran animals began exploring sediments in three dimensions, suggesting more complex behaviors than previously thought.^[76]^[73] The Cambrian Explosion, starting ~541 Ma, marks a dramatic diversification, with the Fortunian Stage introducing deeper burrows (up to 4–5 cm) and diverse ichnogenera such as Arenicolites (U-shaped ventilation structures) and early Skolithos, alongside intense bioturbation indices rising to 6 in offshore settings.^[75] Thalassinoides networks proliferate in the Cambrian Age 2 (~529–521 Ma), indicating expanded infaunal ecospace.^[72] This timeline is delineated by formations like the Chapel Island (Newfoundland) and Wood Canyon (California), where Treptichnus pedum defines the Ediacaran-Cambrian boundary.^[73] Interpretation of these trace fossils involves analyzing cross-sections to reveal internal architecture, such as marginal tubes in Phycosiphon or boxwork patterns in Thalassinoides, often using computed tomography (CT) scans for three-dimensional reconstruction in core samples.^[77] Filling materials provide behavioral clues: passive infills of host sediment suggest open burrows abandoned post-construction, while active backfilling with lined walls (e.g., meniscate structures) indicates sediment processing for feeding or stability.^[78] Associations with body fossils, though rare, enhance inferences; for instance, arthropod traces like Rusophycus near trilobite remains link traces to specific taxa, while ichnofabrics (overlapping burrow densities) gauge community interactions and substrate conditions.^[73] These methods, combining field slabs, X-radiography, and density contrasts, allow reconstruction of ancient seafloor ecosystems.^[77] Burrow trace fossils offer profound evolutionary insights, evidencing the emergence of motile bilaterian animals in the Ediacaran (~560–540 Ma) capable of horizontal grazing and shallow penetration, predating the Cambrian radiation.^[75] These early traces suggest bilaterians like annelid-like worms adapted to dysoxic conditions, with behaviors such as biofilm scraping in Helminthoidichnites indicating coelomate body plans for efficient burrowing.^[75] The Cambrian surge in burrow depth and complexity reflects ecological innovations, including suspension feeding in vertical Skolithos and network dwelling in Thalassinoides, which deepened the mixed layer and enhanced sediment oxygenation by ventilating sulfidic zones.^[72] This bioturbation likely amplified global oxygenation events around 521 Ma, stabilizing phosphorus cycles and facilitating metazoan diversification during the Cambrian Explosion.^[75]

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Biomechanical adaptations for burrowing in the incisor enamel ...
Jun 16, 2021 · Geomyidae (pocket gophers) are small‐ to medium‐bodied rodents with anatomies highly adapted to a subterranean‐burrowing (fossorial) lifestyle ...
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Morphological constraints in the digging apparatus of pocket ...
Aug 7, 2025 · We present a discussion of the structural and functional constraints on jaws and forelimbs in gophers as well as an analysis of historical ...
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(PDF) Wood ants produce a potent antimicrobial agent by applying ...
Wood ants fight pathogens by incorporating tree resin with antimicrobial properties into their nests. They also produce large quantities of formic acid in ...
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The Energy Cost of Burrowing by the Pocket Gopher Thomomys bottae
Burrowing can require 360–3,400 times more energy than moving the same distance on the surface, depending on soil type.
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Rabbits burrowing
Mar 29, 2008 · We found our pair of rabbits taking it in turn to dig the same burrow went a distance of about 4ft-5ft in a day. Our yardstick was when the ...Missing: speed | Show results with:speed
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Aardvark | African Wildlife Foundation
They have an elongated head, with a long, protruding, narrow snout and nostrils that can be sealed. ... Life in an aardvark burrow. Aardvarks love to dig, and ...
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The biomechanics of burrowing and boring
Jan 15, 2015 · Burrowers and borers are ecosystem engineers that alter their physical environments through bioturbation, bioirrigation and bioerosion.
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[PDF] Red Fox - Amazon S3
Their basic structure consists of the main channel, with a chamber or a widening of the main channel, lined with grasses and other ... Gray fox have been reported ...
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Pocket Gophers - UC IPM
Gophers seal the openings to the burrow system with earthen plugs. Short, sloping lateral tunnels connect the main burrow system to the surface; gophers ...
[43]
Nest-Site Selection In A High-Density Colony of Burrowing Owls
Dec 1, 2018 · Higher Burrowing Owl nest success in prairie dog towns may reflect benefits gained from alarm calling and the predator dilution effect of ...
[44]
Seasonal temperature acclimatization in a semi-fossorial mammal ...
Mar 16, 2018 · Our objective was to explore the role of burrows as thermal refuges for a small endotherm, the pygmy rabbit (Brachylagus idahoensis), during the summer and ...
[45]
Burrowing Owls Require Mutualist Species and Ample Interior ...
Sep 19, 2024 · Our study serves as an example of how an ecosystem approach, including conservation of the fossorial mammal species burrowing owls depend upon ...Missing: foxes rabbits
[46]
11 Evolutionary Ecology of Burrow Construction and Social Life
In general, burrows are central to the life history of many species, functioning as protective dwellings against predators and environmental extremes. Within ...<|separator|>
[47]
The Evolutionary Advantage of Burrowing Underground
Many animals dig dens for protection from weather, disasters, or predators, and this behavior may have allowed some species to survive when others have died ...Missing: zoology | Show results with:zoology
[48]
[PDF] Predator avoidance: Mechanisms - eScholarship
Escape to burrows affords rodents immediate refuge from nonburrowing predators, but burrows are also places of danger for digging predators and especially ...
[49]
Burrowing Invasive Species: An Unquantified Erosion Risk at the ...
Jul 31, 2019 · We demonstrate how burrowing invasive species represent a potentially significant but unquantified erosion risk at aquatic margins.
[50]
Ground Squirrels Chew Snakeskins to Mask Their Scent
Feb 5, 2008 · Squirrel pups, which are not resistant, do the chewing, as do their mothers, which are resistant but look after the pups in the burrows. Ms ...
[51]
Burrow use and ranging behaviour of the southern hairy‐nosed ...
Aug 7, 2025 · With over 75% of their time spent below ground, wombats depend on burrows to survive the harsh Australian environment, including protection from ...
[52]
Wombats' deadly bums: how they use their 'skull-crushing' rumps to ...
Nov 3, 2020 · The rump of the wombat is hard as rock, used for defence, burrowing, bonding, mating and possibly violently crushing the skulls of its enemies against the roof ...
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https://www.mdpi.com/2313-7673/10/11/754
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Relic bilby (Macrotis lagotis) refuge burrows - CSIRO Publishing
Jun 7, 2013 · Both temperature and humidity were also significantly less variable in the burrows.
[55]
Under the weather?—The direct effects of climate warming on a ...
Feb 15, 2018 · The burrow system provides an exceptional buffer against maximum temperature extremes (Figure 6). Hourly temperatures at mid and deep depths ...
[56]
Learning from Nature: Bio-Inspired Designs and Strategies ... - MDPI
This mechanism plays a crucial role in regulating the airflow in many animal dwellings, such as in prairie dog burrows, where the pressure-gradient-driven ...
[57]
[PDF] Water relations of the burrowing sandhill frog, Arenophryne rotunda ...
The water retention curve for sand taken from A. rotunda burrows (Fig. 2) shows that sand held 28.4% moisture at saturation point (0 kPa) but quickly lost.
[58]
Greater bilby burrows: Important structures for a range of species in ...
Oct 23, 2025 · Bilby burrows provide a refuge from environmental extremes, and have less variable temperature and humidity fluctuations than control ...
[59]
How temperature, humidity, and burrow selection affect evaporative ...
Aug 6, 2025 · Desert tortoises conserve water and lower metabolic expenditures by retreating into burrows (Bulova 2002; Wilson et al. 2001). Therefore, one ...
[60]
Behavioural patterns of the soft-shell clam Mya arenaria
Jul 18, 2019 · It follows that advective processes must have contributed to the sediment oxygenation and oxidation around the siphon. During the V stage ...
[61]
[PDF] Toxic sulfide concentrations in the sediments and water column of ...
Many sediment infauna circulate oxygenated seawater through their body cavities or burrows, thus maintaining sulfide concentrations near their body tissues ...<|control11|><|separator|>
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https://pmc.ncbi.nlm.nih.gov/articles/PMC6920383/
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Soil Properties, Part 2 of 3: Biological Characteristics | Extension
Earthworms. Earthworm burrows enhance water infiltration and soil aeration. Fields that are “tilled” by earthworm tunneling can absorb water at a rate four ...
[64]
[PDF] Earthworms - Natural Resources Conservation Service - USDA
They improve soil porosity by burrowing and mixing soil. As they feed, earthworms participate in plant residue decomposition, nutrient cycling, and ...
[65]
Testing for consistency in the impacts of a burrowing ecosystem ...
Dec 18, 2019 · Less compact soils at burrow entrances will be more aerated28 and may have higher water infiltration rates63, which may also explain the drier ...
[66]
[PDF] The bioturbation transport of chemicals in surface soils
Burrows or tunnels made by animals increase soil porosity, promote aeration of the soil, and increase infiltration of water. The effects of these burrow systems ...
[67]
Gopherus polyphemus - USDA Forest Service
Because gopher tortoise burrows provide habitat for other species, gopher tortoise presence may increase local animal diversity. Many other animals seek ...<|separator|>
[68]
Rusty Patched Bumble Bee (Bombus affinis) | U.S. Fish & Wildlife ...
Rusty patched bumble bee nests are typically 1 to 4 feet underground in abandoned rodent nests or other mammal burrows and occasionally at the soil surface or ...
[69]
A burrowing ecosystem engineer positively affects its microbial prey ...
We conclude that even in a simple two‐species system a predator can positively impact prey via ecosystem engineering. ... trophic and nontrophic interactions ...
[70]
[PDF] Exposure to a novel predator induces visual predator recognition by ...
Jul 10, 2020 · Our results show that predator-naïve prey species' ability to recognise novel predators is inducible through experience. Keywords Antipredator ...
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[PDF] Dispersal, facilitation, and burrow architecture in banner-tailed ...
May 1, 2010 · Increases in burrow entrances after large rainfall events would speed evaporation within mounds and reduce spoilage of critical seed caches.
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[PDF] Using Burrow Builders for Pocket Gopher Control
For plains pocket gophers, a population of six to eight animals per acre is considered high density. Pocket gophers damage more than $10 million of major field ...
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One burrow at a time: saving the gopher tortoise | US Forest Service
Nov 22, 2023 · Gopher tortoises have a tremendous impact on the environment with the burrows they create, providing protection from elements, fire and ...
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Understanding the keystone gopher tortoise | Texas A&M NRI
The gopher tortoise is an important keystone species; its burrows provide habitat for over 360 species across its range.
[75]
Glossary - KU Ichnology
Skolithos ichnofacies: trace fossil association used to denote loose shifting media, typically in the lower shoreface. Burrows are frequently deep and require ...
[76]
Trace fossils and substrates of the terminal Proterozoic–Cambrian ...
Three principal sources of evidence exist: body fossils, trace fossils (trails, tracks, and burrows of animal activity recorded in the sedimentary record), and ...
[77]
9.6: Trace Fossils - Geosciences LibreTexts
Jul 25, 2025 · Trace fossils are classified into ichnogenera and ichnospecies based on the physical traits of the burrow. We do it this way because ...
[78]
The rise and early evolution of animals: where do we stand from a ...
The trace-fossil record provides a wealth of information to track the rise and early evolution of animals. It comprises the activity of both hard- and ...
[79]
Exploring computed tomography in ichnological analysis of cores ...
Jan 13, 2020 · Orientation of trace fossils within sediment has been widely used to interpret palaeoenvironmental conditions, as it can be affected by ...
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https://www.sjvgeology.com/articles/chamberlain_trace_fossils.pdf