Fact-checked by Grok 2 weeks ago

Tree hollow

A tree hollow is a cavity that develops in the trunk or branches of a living or dead tree, primarily through the gradual decay of heartwood initiated by injury, branch loss, or natural aging, with wood-decay fungi playing a central role in the decomposition process.^[1]^[2] This formation is a slow phenomenon, often requiring centuries in species like eucalypts, as fungi and microorganisms break down internal wood while the outer layers remain structurally sound.^[3]^[4] Tree hollows serve as critical microhabitats, offering shelter, breeding sites, and protection from predators and extreme weather for diverse wildlife including birds, mammals, bats, and reptiles.^[5]^[6] Their scarcity in younger forests underscores their status as keystone features in mature ecosystems, where loss from land clearing exacerbates declines in hollow-dependent species.^[7]^[8]

Formation and Characteristics

Natural Formation Mechanisms

Tree hollows arise primarily from the progressive decay of heartwood, the non-living central core of the trunk or branches, which trees do not actively defend against microbial invasion due to the low structural and functional cost of such loss.^[9] This decay is typically initiated by wounds that breach the protective sapwood and bark, allowing entry for wood-decaying fungi and bacteria.^[9] Abiotic factors, such as lightning strikes, wind-induced branch breakage, fire scorch, and frost cracking, commonly create these initial entry points by damaging outer tissues and exposing inner wood to environmental moisture and pathogens.^[10] Biotic agents accelerate and shape hollow development once decay commences. Fungi, including basidiomycetes specialized in lignocellulose breakdown, colonize and enzymatically degrade the heartwood, often in symbiosis with insects like termites or beetles that further fragment the wood.^[11] ^[10] Woodpeckers and other cavity-excavating birds exploit pre-softened wood, pecking to enlarge cavities; certain species, such as the red-cockaded woodpecker (Picoides borealis), actively disseminate decay fungi on their beaks to facilitate excavation in living trees.^[12] This mutualism enhances cavity formation rates, as fungi weaken wood fibers while birds remove debris, though primary hollows in old-growth forests often originate from unaided fungal rot over decades.^[12] ^[13] Hollow maturation is gradual, requiring trees of advanced age—typically over 100–200 years depending on species and site conditions—for sufficient heartwood accumulation and decay progression.^[13] In fire-prone ecosystems, recurrent low-intensity burns can promote hollow formation by charring bark and stimulating compartmentalization responses that isolate decay to inner zones, preserving tree longevity while fostering cavity development.^[10] Variability in formation stems from tree genetics, soil nutrients, and climate, with faster decay in humid environments versus slower progression in arid ones.^[9]

Influencing Factors and Variability

Tree species significantly influence hollow formation due to variations in wood durability, heartwood chemistry, and branching patterns; species with denser, more decay-resistant heartwood, such as certain oaks, develop hollows more slowly compared to softer-wooded eucalypts prone to fungal ingress.^[14] Age and diameter at breast height (DBH) are key intrinsic factors, with hollow prevalence rising sharply beyond DBH of 40-50 cm and tree ages exceeding 100-150 years, as larger branches shed and expose heartwood to decay.^[15] Fast-growing individuals form hollows earlier than slow-growers, since larger early branches create entry points for pathogens upon abscission.^[16] Extrinsic biological agents, including wood-decay fungi and bacteria, drive decay once wounds occur, with fungal inoculation via branch breaks or injuries initiating heartwood rot that expands into cavities over decades.^[1] Physical disturbances like lightning strikes, wind damage, fire scarring, or termite activity provide initial breaches, accelerating hollow development in otherwise resistant trees; for example, cyclones and fires in Australian savannas boost hollow abundance by compromising bark integrity.^[10] Climate modulates decay rates, with warmer, humid conditions in tropical or wet forests promoting faster fungal activity and higher hollow incidence than in arid or cold-temperate zones where desiccation slows decomposition.^[17] Variability manifests spatially across forest types and regions; wet eucalypt forests exhibit greater hollow occurrence than dry counterparts due to sustained moisture favoring decay, while urban proximity may reduce it through altered microclimates and reduced disturbance regimes.^[17] Temporal dynamics show hollow formation lagging tree maturity by 50-200 years depending on species and site, with ongoing variability from stochastic events like storms introducing patchiness in hollow distribution within stands.^[18] Site-specific soil moisture and topography further contribute, as hummock-hollow microtopography alters local hydrology and thus decay susceptibility.^[19]

Classification of Hollows

Non-Excavated Hollows

Non-excavated hollows, also referred to as decay-formed or passive cavities, develop through the gradual decomposition of heartwood in living trees, primarily driven by fungal activity without intervention from excavating animals.^[20] This process typically initiates when mechanical damage, such as branch breakage or trunk wounds, exposes inner wood to wood-decay fungi, which colonize and break down the non-functional heartwood while the outer sapwood layers continue to provide structural support and transport.^[20] Fungal heart-rot fungi are the primary agents, releasing enzymes that degrade lignin and cellulose, leading to cavity enlargement over time.^[20] Formation of these hollows occurs predominantly in mature or veteran trees, often requiring 100 to 200 years or more to produce cavities suitable for vertebrate use, depending on species and environmental conditions.^[18] For instance, in temperate Eucalyptus forests of Australia, hollow development via decay and physical damage alone can span centuries in the absence of excavators, with abundance increasing in larger, older trees on productive soils.^[18] In Afromontane forests, such as those in Rwanda's Volcanoes National Park, fungal decay accounts for the majority of cavities (83%), concentrated in species like Hagenia abyssinica, facilitated by high precipitation that promotes microbial activity.^[21] Physical factors like storms, lightning, or fire can accelerate initiation by creating entry points for fungi, but the core mechanism remains biological decay rather than active removal of wood.^[18] Unlike excavated hollows, non-excavated ones often form as basal or side cavities, with side cavities emerging from branch stubs after over a century of decay.^[20] These cavities contribute to tree longevity by compartmentalizing decay, preventing spread to vital sapwood, though extensive hollowing can eventually compromise stability in advanced stages.^[20]

Excavated Hollows

Excavated hollows, also known as primary cavities, form through the chiseling action of certain vertebrates that actively drill into tree trunks or branches using specialized bills. These structures differ from decay-induced hollows by originating directly from mechanical excavation rather than fungal or bacterial decomposition alone. Primary excavators preferentially target dead, dying, or partially decayed wood, where softer heartwood facilitates penetration, often beginning in snags or trees infected by pathogens that weaken structural integrity.^[22]^[23] Woodpeckers of the family Picidae dominate as primary excavators worldwide, excavating cavities for nesting, roosting, or foraging on insects within the wood. Species such as the pileated woodpecker (Dryocopus pileatus) in North America create large cavities up to 45 cm deep in snags, while the three-toed woodpecker (Picoides tridactylus) in boreal forests excavates smaller holes in conifers with lifespans exceeding several years post-abandonment. Other excavators include certain nuthatches and chickadees, though less extensively; for instance, white-breasted nuthatches (Sitta carolinensis) occasionally drill into soft wood. These birds may produce multiple cavities annually, selecting sites based on food availability and decay stage, with aspen species hosting 96% of cavities in some ecosystems due to rapid sapwood decay.^[24]^[25]^[23] Characteristics of excavated hollows include oval or rectangular entrances tailored to the excavator's body size, with internal chambers smoothed by repeated use and debris removal. Entrance diameters range from 3-15 cm depending on species, and cavities often extend downward to prevent flooding. In regions like Australia, primary excavators are absent, relying instead on decay processes, highlighting regional variability in hollow formation. These cavities persist after initial use, providing substrate for secondary colonizers and contributing to habitat complexity, though their availability declines in intensively managed forests lacking mature snags.^[26]^[27]^[28]

Artificial Hollows

Artificial hollows are human-engineered cavities intended to substitute for or supplement scarce natural tree hollows, primarily to support cavity-dependent wildlife in ecosystems altered by logging, urbanization, or fires. These structures mimic the shelter, nesting, and roosting functions of natural hollows but are constructed using materials like wood, plastic, or composites, or by modifying living trees through mechanical means. Deployment occurs in restoration projects, with designs tailored to species requirements such as entrance size, depth, and internal volume to reduce predation and optimize microclimate.^[29]^[30] Nest boxes represent the most widespread form of artificial hollows, installed on trunks or branches at heights of 3-15 meters to emulate arboreal cavities. In Australia, where approximately 300 vertebrate species—about 15% of the total—rely on hollows for breeding or refuge, nest boxes have been deployed extensively for birds like owls and parrots, mammals such as squirrel gliders, and bats. For example, Wildlife Queensland has installed carved hollows and boxes for endangered mahogany gliders, achieving occupancy rates by target species within months. However, nest boxes often exhibit higher exposure to weather and predators compared to natural hollows, with studies reporting drier but less thermally stable interiors that can lead to chick mortality in extreme conditions. Maintenance is critical, as untreated boxes degrade within 5-10 years, and improper design attracts invasive species like feral honeybees or starlings.^[31]^[32]^[33] Direct excavation techniques create cavities by drilling or chainsaw-cutting into live trees, accelerating habitat provision without full reliance on decay processes that take decades. A 2021 Australian innovation using specialized drills produced hollows in eucalypts within hours, targeting post-2019-2020 bushfire recovery to generate up to one million sites for displaced fauna like possums and micro-bats. Peer-reviewed trials of mechanically excavated hollows in eucalypt forests demonstrated uptake by multiple species, including woodpeckers and invertebrates, within one year, though success depends on tree health and fungal inoculation to promote internal decay. Inoculation with wood-decay fungi, such as Fomes fomentarius, further hastens cavity maturation by simulating natural rot, with field experiments showing enhanced saproxylic insect colonization. These methods prioritize live trees over snags to minimize structural risk, but they require arborist expertise to avoid compromising tree stability.^[34]^[30]^[1] Emerging approaches include 3D-printed hollows and modular inserts that replicate spout or chimney shapes preferred by certain birds, offering customizable decay resistance. Lifecycle assessments of these designs emphasize sustainability, favoring recyclable materials over plywood to extend lifespan beyond 20 years in humid climates. While artificial hollows provide short-term relief—evidenced by occupancy in 20-50% of units within 2-3 years in Australian studies—they do not fully replicate the biodiversity of mature hollows, which support complex food webs via detritus accumulation. Long-term efficacy hinges on integration with habitat protection, as isolated installations fail without surrounding forage and connectivity.^[35]^[36]^[37]

Ecological Importance

Role as Wildlife Habitat

Tree hollows serve as critical microhabitats for diverse wildlife, providing sheltered sites for breeding, roosting, denning, hibernation, and protection from predators and extreme weather conditions.^[7] ^[38] These cavities enable species lacking the ability to excavate their own shelters to persist, functioning as a keystone resource that supports high levels of biodiversity in forest ecosystems.^[39] ^[40] Among birds, tree hollows are primary nesting and roosting sites for cavity-dependent species, including woodpeckers, owls, nuthatches, and parrots, which rear young in these secure environments.^[41] In Australia, at least 81 bird species in New South Wales depend on hollows for such purposes.^[42] Mammals, such as squirrels, possums, gliders, martens, porcupines, and bats, utilize hollows for dens, hibernation, and refuge, with 46 species in New South Wales relying on them.^[41] ^[42] Reptiles and amphibians also inhabit these spaces; for instance, 34 reptile species and 8 frog species in New South Wales use hollows for shelter and egg-laying.^[42] Invertebrates further contribute to the habitat's complexity, with hollows hosting nutrient-rich communities that sustain food webs for vertebrate occupants.^[36] The availability of hollows influences wildlife population dynamics, as their scarcity limits breeding success and species diversity, underscoring their role in maintaining ecological balance.^[18]

Contributions to Biodiversity and Ecosystem Function

Tree hollows function as keystone habitats that disproportionately support biodiversity in forest ecosystems by providing shelter, nesting, and roosting sites for a wide array of species, including cavity-nesting birds, mammals, bats, and reptiles.^[43] These structures host specialized saproxylic communities, such as beetles and invertebrates, which rely on the decaying wood within hollows for development and reproduction, thereby increasing local species richness.^[13] Studies indicate that hollow-bearing trees sustain rich assemblages of often highly specialized organisms, contributing to overall habitat heterogeneity and facilitating coexistence among taxa that cannot utilize ground-level or foliar resources.^[43] Beyond habitat provision, tree hollows enhance ecosystem functioning through nutrient cycling and food web dynamics. The accumulation of organic matter and detritus in hollows supports decomposition processes driven by fungi and invertebrates, recycling nutrients back into the soil and broader forest system.^[44] Complex food webs within these nutrient-rich microhabitats promote stability, as evidenced by interactions among predators, prey, and decomposers that maintain invertebrate diversity and prevent dominance by generalist species.^[36] This internal ecosystem processing aids in carbon sequestration and soil fertility, with hollows acting as long-term repositories for biomass turnover.^[7] Hollows also influence surrounding vegetation and microclimates, indirectly bolstering biodiversity. By harboring epiphytes and moisture-retaining detritus, they create localized conditions that support additional plant and microbial life, altering light and humidity profiles around host trees.^[45] In managed or fragmented landscapes, the scarcity of natural hollows underscores their role in ecosystem resilience, where retention of such features correlates with higher vertebrate occupancy and reduced extinction risks for dependent species.^[18] Empirical data from surveys of thousands of trees confirm that cavity abundance drives arthropod diversity, linking structural availability to functional outcomes like pollination and pest regulation.^[44]

Threats and Population Dynamics

Anthropogenic Pressures

Human activities, particularly commercial logging and deforestation, have significantly reduced the abundance of hollow-bearing trees by targeting mature and senescent individuals that develop cavities over centuries. In ecologically unsustainable forestry practices, operations directly remove and damage trees with hollows, exacerbating shortages for dependent wildlife such as birds and mammals. For instance, in Australian native forests, logging contributes to a continuing net loss of these trees, with associated road-building and extraction further fragmenting habitats and preventing hollow formation in regenerating stands.^[46]^[47]^[48] Land conversion for agriculture and urbanization compounds this decline by clearing vast areas of old-growth forests where hollow prevalence is highest. In the Neotropics, deforestation for farming has destroyed nesting cavities essential for cavity-nesting birds, potentially leading to local extirpations of species reliant on large-diameter trees. Urban forest fragments retain fewer hollow-bearing trees due to historical clearing and ongoing development, limiting ecosystem integrity despite the presence of remnant veterans. In New South Wales, Australia, more than half of native forests and woodlands have been lost to such pressures, with persistent logging accelerating extinction risks for hollow-dependent fauna.^[49]^[50]^[51] Firewood harvesting and pruning for fuel or aesthetics further deplete hollow resources, especially in rural and peri-urban woodlands. Selective removal of dead or decaying branches eliminates potential cavity sites, while unregulated collection targets snags and veterans critical for long-term hollow supply. In regions like New Guinea, combined logging and hunting disrupt cavity availability, destroying breeding refuges for arboreal species.^[3]^[52] In managed urban environments, hollow-bearing trees face removal due to perceived structural hazards, despite evidence that decay compartmentalization often stabilizes them. Safety assessments frequently prioritize human infrastructure over retention, leading to proactive felling of cavity hosts that pose minimal actual risk. This practice, informed by conservative risk models, diminishes urban biodiversity hotspots where alternative habitats are scarce.^[53]^[54]

Natural Disturbances and Long-Term Decline

Natural disturbances, including wildfires, windstorms, insect outbreaks, and pathogenic infections, exert a dual influence on tree hollow formation and persistence by damaging bark and cambium to initiate decay while also felling mature trees that already bear cavities.^[55] In temperate forests, such events generate tree-related microhabitats like rot-holes and cavities, which peak in abundance 100–150 years post-disturbance during low-severity or late-seral stages, as surviving trees accumulate structural legacies conducive to fungal ingress and heartwood decomposition.^[55] However, high-severity disturbances disproportionately eliminate large-diameter individuals, reducing the substrate for future hollow development.^[56] Wildfires exemplify this dynamic, promoting hollows through basal scarring and limb breakage that expose inner wood to decomposers, yet excessive frequency or intensity yields net losses. In southeastern Australian forests, hollow density exhibits a unimodal response to fire frequency, peaking at intermediate intervals of 7–30 years where medium-sized cavities increase by up to 1.82 per site from one to two fires, but declining beyond three events due to cumulative mortality.^[57] Similarly, large hollows correlate positively with moderate fire severity (gains of 1.75 per site from moderate to high), but very high severity erodes availability by felling hollow-bearing trees (HBTs), with no significant short-term difference in HBT abundance post-single burns but reductions under repeated high-severity regimes.^[57]^[56] Insect infestations and wind events further modulate hollow stocks; bark beetles and wood-borers facilitate rot by colonizing wounded tissues, yet outbreaks synchronized with drought can synchronize tree die-off, curtailing HBT recruitment.^[58] Windstorms contribute snags and fractured crowns that evolve into cavities over decades, but in disturbance-prone stands, they compound fire effects by increasing fuel loads and vulnerability to subsequent burns.^[59] Long-term declines in hollow availability arise when disturbance intervals shorten relative to hollow ontogeny, which demands 100–200+ years for viable cavities in non-excavated trees, engendering lag effects where depleted old-growth cohorts fail to replenish supply.^[4] Climatic shifts amplify this by elevating drought stress and fire severity, negatively associating with basal scars (declines of up to 8.47 trees per site at higher temperatures) and constraining growth rates needed for durable wood accrual.^[57] In fire-adapted systems, projections indicate sustained HBT erosion if regimes trend toward extremes, limiting habitat for obligate cavity users and underscoring disturbances' role in balancing formation against attrition.^[56]^[57]

Conservation and Management

Protective Strategies

Protective strategies for tree hollows emphasize the retention of mature, defective, and senescent trees during forestry operations and land management to sustain long-term habitat availability for wildlife. In managed forests, practices such as variable retention harvesting preserve aggregates of old-growth trees, ensuring a continuous supply of cavity-bearing structures across decay classes and mitigating declines from clear-cutting.^[60] Forest managers prioritize selecting cull trees—those exhibiting early decay indicators like fungal conks, dead branch stubs, or soft wood—for protection, as these are poised to form hollows within decades.^[61]^[62] Regulatory frameworks in various regions mandate the identification and safeguarding of hollow-bearing trees prior to logging or development, often requiring assessments to balance safety risks against ecological value. For instance, in some jurisdictions, trees reaching approximately 67% of their species' maximum diameter are flagged for protection to preempt hollow loss from premature harvest.^[63] Limiting post-disturbance salvage logging of fire-killed or storm-damaged snags further prevents acute shortages, as these provide immediate cavity resources while live trees mature.^[64] In fragmented landscapes, including urban areas, community education and policy enforcement discourage the removal of hazardous yet habitable trees, favoring structural reinforcement over felling where feasible. Integrated wildlife management plans distribute retained snags and cavities evenly across habitats, targeting 4–10 per hectare in woodlands to support cavity-dependent species without compromising timber yields.^[65] Monitoring via environmental DNA sampling from hollows aids in verifying usage and refining retention priorities, enhancing strategy efficacy amid ongoing habitat pressures.^[66]

Mitigation and Restoration Approaches

Retention of hollow-bearing trees during timber harvesting operations serves as a primary mitigation strategy to counteract losses from logging. Forestry guidelines in regions like New South Wales, Australia, mandate retaining at least 10 hollow-bearing trees and 10 recruitment trees (those likely to develop hollows) per 2 hectares in native forests to sustain habitat availability.^[67] Similar retention practices in alternatives to clearcutting emphasize preserving large, old trees to maintain structural legacy elements essential for cavity-dependent species.^[68] These measures address the slow natural formation of hollows, which can require over 100 years in many eucalypt species, by prioritizing trees already containing cavities or showing decay.^[18] Active restoration often employs artificial hollow creation to provide immediate habitat where natural cavities are scarce, particularly in regenerating or second-growth forests. Chainsaw-carved hollows installed in medium-sized live trees have demonstrated increased visitation rates by hollow-dependent fauna, such as birds and mammals, within short periods post-installation, facilitating faster habitat recovery in revegetation projects.^[69] Techniques involve excavating cavities mimicking natural dimensions and entrances, with studies reporting rapid colonization by multiple species, though long-term persistence depends on tree health and environmental conditions.^[70] Complementary approaches include selective snag creation—killing healthy trees to form standing dead wood—while avoiding interference with existing hollows, targeting trees over 10 inches in diameter for durability.^[71] In broader ecosystem restoration, legacy tree selection prioritizes individuals with pre-existing cavities to accelerate the development of old-growth characteristics in managed woodlands.^[72] Such strategies integrate with snag management in upland and riparian areas, promoting fungal decay and woodpecker excavation to enhance future hollow formation without relying solely on artificial interventions.^[73] Empirical monitoring post-restoration, including thermal profiling of artificial versus natural hollows during wildfires, underscores the need for designs that match native microclimates to ensure viability under disturbances.^[74] While artificial structures offer temporary mitigation, their efficacy as substitutes diminishes over decades, necessitating coupled long-term silvicultural planning to foster self-sustaining hollow abundance.^[75]

Global Patterns and Regional Variations

Temperate and Boreal Forests

In temperate forests of Europe and North America, tree hollows primarily form through fungal decay of heartwood, often accelerated by mechanical damage from storms, lightning, or branch failure, requiring 50 to 100 years or more for significant cavities to develop in mature trees such as oaks and beeches.^[4] ^[76] Woodpecker excavation contributes secondary hollows, but decay-driven processes dominate, leading to larger, more persistent cavities compared to faster-forming but shallower bird-excavated ones.^[39] These hollows are critical microhabitats, supporting cavity-nesting birds like owls and woodpeckers, as well as mammals such as bats and squirrels, though abundance is reduced in managed forests where harvesting removes old-growth trees before hollow formation completes.^[13] In Central European managed stands, hollows constitute rare habitats, limiting biodiversity dependent on them.^[77] Boreal forests, dominated by conifers like pines and spruces alongside deciduous aspen and birch, exhibit lower proportions of decay-formed hollows, with most cavities resulting from primary excavation by woodpeckers and subsequent reuse by other species.^[39] Broadleaf trees, particularly trembling aspen, serve as key substrates for cavity formation due to their susceptibility to heart rot fungi, providing essential roosting and nesting sites for bats and cavity-nesting birds amid the biome's harsh winters.^[78] Densities can reach approximately 30 hollows per hectare in primeval boreal stands, as observed in Mongolian taiga, though occupancy rates for wildlife remain moderate at 42-46% in natural cavities versus higher in artificial nest boxes.^[79] ^[80] Fire disturbances and climate influence hollow persistence, with decaying retention trees enhancing nesting opportunities for birds across both temperate and boreal zones, underscoring the role of deadwood in sustaining cavity-dependent communities.^[81] ^[57] Regional variations highlight that temperate forests often feature more diverse hollow types due to longer-lived hardwoods, fostering greater structural complexity, whereas boreal systems rely more on dynamic cavity turnover driven by avian excavators and periodic stand-replacing fires, resulting in fewer but functionally vital large cavities in aspen-dominated patches.^[82] In both biomes, intensive forestry reduces hollow-bearing tree abundance, with meta-analyses indicating that retained decaying trees significantly boost cavity-nester populations, emphasizing the need for legacy tree preservation to mimic natural dynamics.^[81] Unlike tropical regions with abundant rapid-decay hollows, temperate and boreal hollows demand extended timelines for development, making them vulnerable to short-rotation management that prioritizes even-aged stands over uneven-aged, old-growth conditions essential for their formation.^[4]

Tropical and Savanna Ecosystems

In tropical and savanna ecosystems, tree hollows form primarily through interactions between biological decay agents like termites and fungi, physical damage from cyclones or lightning, and environmental factors such as fire regimes. Termite activity significantly accelerates hollow development by hollowing out tree cores, with surveys in northern Australian tropical savannas indicating that most eucalypt trees develop internal hollows due to this process.^[83] In mesic savanna woodlands, hollow abundance reaches approximately 88 per hectare, influenced by higher rainfall and deeper soils that support larger, older trees.^[84] Fire, prevalent in savannas, can inhibit hollow formation by killing large trees and reducing recruitment of hollow-bearing individuals, though moderate fire intervals may promote decay in surviving trees.^[85] Tropical rainforests exhibit hollow formation via branch breakage, heartwood decay from fungi and insects, and buttress root damage, creating refuges that support a succession of colonizers from invertebrates to vertebrates.^[86] In these dense forests, large emergent trees provide deep cavities offering thermal refuge from humidity and predators, essential for arboreal species. Savanna hollows, often in scattered keystone species like baobabs or eucalypts, contrast with the clustered availability in rainforests but sustain disproportionately high biodiversity relative to tree density, as isolated hollow-bearing trees act as biodiversity hotspots.^[87] West African savanna trees develop external splits and internal hollows through synergistic fire scarring and termite excavation, with fire creating entry points for termites to degrade heartwood.^[88] Hollow-dependent fauna in these ecosystems include a wide array of birds (e.g., parrots and owlets), mammals (e.g., bats, possums, and small carnivores), and reptiles, with up to 77% of surveyed bird species utilizing hollows in some tropical studies.^[7] In Australian tropical savannas, overlapping use by declining arboreal mammals like possums highlights competition for limited large cavities, exacerbated by termite-enhanced but fire-limited supply.^[89] Neotropical psittacids, such as the blue-fronted amazon, rely heavily on large tree cavities for nesting, with 93% of parrot species globally dependent on them, underscoring vulnerability in logged or fragmented tropics.^[90] These structures contribute to ecosystem function by facilitating multi-species interactions, including predation and commensalism within cavities, thereby maintaining trophic diversity in resource-scarce savanna matrices.^[91] Compared to temperate forests, tropical and savanna hollow abundance per hectare can exceed that in eucalypt-dominated temperate woodlands, driven by rapid tropical decay rates despite shorter tree lifespans in fire-prone areas.^[92]

Urban and Fragmented Landscapes

In urban landscapes, the availability of tree hollows is often limited by the predominance of younger, fast-growing street trees and intensive management practices that prioritize safety and aesthetics over retention of mature or defective specimens. Large old trees, which disproportionately provide hollows essential for cavity-dependent fauna, constitute a declining proportion of urban canopies, with models projecting an 87% reduction in hollow-bearing trees over 300 years under conventional replacement regimes that favor rapid-maturing species lacking decay-prone heartwood.^[93] Empirical surveys along urban gradients reveal variable densities, such as a mean of 37.5 hollow-bearing trees per hectare, but standing dead trees—key contributors to hollow abundance—are frequently removed due to perceived hazards, exacerbating shortages for species like owls, bats, and invertebrates.^[50] Fragmented landscapes, characterized by isolated patches of remnant vegetation amid agricultural or developed matrices, compound hollow scarcity through disrupted succession and reduced gene flow for hollow-forming processes like fungal decay and woodpecker excavation. In such habitats, the loss of contiguous old-growth stands hinders the recruitment of replacement hollow trees, which require centuries of uninterrupted growth; studies indicate that fragmentation elevates edge effects, accelerating windthrow and decay but also increasing removal rates for safety, thereby limiting habitat for obligate users.^[94] Wildlife reliant on hollows, including near-threatened pythons and woodpeckers, face heightened extinction risks in these settings, as isolation restricts dispersal and foraging, with cavity-nesting birds showing sensitivity to patch size and connectivity deficits.^[95]^[96] Despite these pressures, urban and fragmented hollows sustain disproportionate biodiversity value, hosting taxa absent from non-decayed trees and supporting ecosystem services like pest regulation via insectivorous residents. Risk assessments highlight that hollow prevalence correlates with escalating structural instability categories (A to C), underscoring the need for targeted retention where feasible, such as in parks or cemeteries, to bolster urban wildlife resilience without compromising public safety.^[97] Management data from municipal surveys affirm that conserving select hollow trees enhances perceived ecological benefits, countering biases in traditional arboriculture that undervalue decay as a hazard rather than a habitat driver.^[98]

References

[1]
Making hollow trees: Inoculating living trees with wood-decay fungi ...
Artificial boxes and carved hollows typically provide habitat for cavity fauna in the mid-late stages of a cavity's development, with a hollow accessible to ...
[2]
[PDF] Wood decay in living and dead trees: A pictorial overview
Other wood-decay fungi that are less pathogenic may eventually ... The formation of a hollow tree is the result of the protective response of living.
[3]
Hollow-bearing trees explained - Wilderness Society
Tree hollows are cavities formed in the trunk or branches of a living or dead tree. · Hollows are mostly found in old eucalypt trees, and are rare in many other ...
[4]
Hollows in living trees develop slowly but considerably influence the ...
Aug 24, 2015 · The decomposition rate of tree hollows determines the speed of hollow formation but, except for the measurement of respiration rate of heart ...
[5]
[PDF] Tree Hollows and Wildlife - DBCA Library
Tree hollows provide shelter and breeding sites for many animals, including possums, bats, birds, and reptiles. Animals choose hollows based on size and shape.
[6]
Deep tree hollows: important refuges from extreme temperatures
Tree hollows provide important habitat for fauna, offering protection from predators and extreme temperatures. They also provide a safe environment for rearing ...
[7]
Disentangling the ecological value of tree hollows to wildlife along ...
Nov 1, 2024 · Tree hollows are functional microhabitats meeting the requirements of animals for breeding, roosting, concealment, resting, hibernation, and/or ...
[8]
Into the hollow: Exploring the value and role of naturally occurring…
Oct 31, 2023 · Because hollows take so long to form, the effects of land clearing on native biodiversity are amplified. While planting new trees is a step in ...<|separator|>
[9]
Why are so many trees hollow? - PMC - PubMed Central
Trees may become hollow because it's not economical to chemically defend the central wood, as the cost of this is low compared to the small structural loss.
[10]
Predictive models of the occurrence of hollows in commercial trees ...
Hollows develop mainly due to the action of fungi and insects (Gibbons and Lindenmayer, 2002), with the help of abiotic and stochastic factors, such as fire, ...
[11]
Why are trees hollow? Termites, microbes and tree internal stem ...
Dec 20, 2024 · However, deadwood biomass pools are highly dynamic and are broken down by biotic (e.g. invertebrates or microbes) and abiotic (e.g. fire or UV) ...
[12]
Experimental evidence of a symbiosis between red-cockaded ...
Mar 30, 2016 · We report the first experimental evidence of a symbiosis between fungi and a cavity excavator, the red-cockaded woodpecker (RCW, Picoides borealis).
[13]
Influence of tree hollow characteristics and forest structure on ...
Tree age and diameter are important factors that facilitate the development of tree hollows (Ranius et al., 2009). In contrast to other dead wood structures ...
[14]
[PDF] When a tree falls: Controls on wood decay predict standing dead ...
May 9, 2018 · Furthermore, wood durability might also predict resistance to internal decay in living trees, identifying species that are likely to be hollow ...
[15]
Hollows in living trees develop slowly but considerably influence the ...
Aug 9, 2025 · Factors that favor fungal invasion and tree use by woodpeckers include tree species, growth history, site characteristics such as proximity ...<|control11|><|separator|>
[16]
(PDF) Development of tree hollows in pedunculate oak (Quercus ...
Hollows formed at earlier ages in fast-growing trees than in slow-growing trees, which may be because hollows are formed when big branches shed, and branches ...
[17]
Does hollow occurrence vary with forest type? A case study in wet ...
This research investigates the variation in hollow occurrence across different forest types, focusing on wet and dry Eucalyptus obliqua forests.
[18]
How an unprecedented wildfire shaped tree hollow occurrence and ...
May 3, 2024 · The indirect nature of hollow formation is dependent on the aging and senescing of trees and the absence of cavity excavating fauna means ...
[19]
[PDF] Drivers of individual tree growth and mortality in an uneven-aged ...
At a smaller scale, hummock and hollow microtopography results in more localized variability in soil moisture. 2.2. Field procedures. In 1989 when the NASA ...
[20]
Tree Cavities - an overview | ScienceDirect Topics
Fungal decay is a key process in tree cavity formation, which often starts from branch loss or infection through trunk injury (Shigo, 1984; Bull et al., 1997; ...
[21]
Density and characteristics of tree cavities inside and outside ...
Oct 27, 2022 · Tree cavities, formed by animal excavation or processes of fungal decay and mechanical damage, may provide nesting, roosting, or resting ...
[22]
[PDF] Managing Cavity Trees - for Wildlife - USDA Forest Service
After a standing dead tree has decayed to the point where most of the branches have fallen, it is called a snag if it is at least 20 feet tall, a stub if ...
[23]
Woodpeckers and other excavators maintain the diversity of cavity ...
Nov 5, 2021 · Woodpeckers nest in several tree species, but principally in aspen (96% of cavities), and selected unhealthy or recently dead trees (decay ...
[24]
Three-toed Woodpecker cavities in trees: A keystone structural ...
Apr 1, 2018 · Our results show that cavities made by Three-toed Woodpeckers have rather long lifespan but also that their active use by other cavity-breeding species is ...
[25]
The multilayered influence of woodpeckers as ecosystem engineers
Jul 22, 2021 · As a whole, woodpeckers within the genus Melanerpes excavate cavities within a mix of live and partially decayed trees, though preferentially in ...
[26]
Woodpecker cavity establishment in managed forests: relative rather ...
Feb 14, 2020 · Woodpeckers are considered environmental architects, being able to excavate their own cavities, often located in old, large trees and snags ( ...Methods · Figure 1 · Figure 3
[27]
Woodpeckers, decay, and the future of cavity‐nesting vertebrate ...
May 31, 2011 · We studied survivorship of 2805 nest cavities and found similar persistence of cavities created by woodpeckers and those created by decay in Canada.<|separator|>
[28]
Use of tree hollows by Carnaby's Cockatoo and the fate of large ...
There are no primary excavators in Australia. While some species such as Rainbow Lorikeet Trichoglossus haematodus, Galah Cacatua roseicapilla and Sacred ...
[29]
Artificial Tree Hollows for Animal Habitat
It is estimated that 15% of Australian vertebrate species use natural tree hollows for nesting, raising young and housing.
[30]
Artificial tree hollow creation for cavity-using wildlife – Trialling an ...
Dec 1, 2017 · A novel technique of artificial hollow provision is described. Uptake of mechanically created hollows was fast and by several species.
[31]
[PDF] NEST BOXES AND ARTIFICIAL HOLLOW GUIDELINES
Australian fauna rely on hollows in a variety of ways. Some species, such as nesting birds, prefer one single hollow for their breeding cycle, whilst some ...
[32]
Wildlife Queensland creates artificial tree hollows for endangered ...
Wildlife Queensland has adopted a multifaceted approach by installing nest boxes to create artificial habitats and, more recently, by carving hollows.<|separator|>
[33]
Spout hollow nest boxes provide a drier and less stable ...
Apr 1, 2021 · Spout hollow nest boxes provide a drier and less stable microclimate than natural hollows ... Artificial tree hollows (e.g., nest-boxes) are ...
[34]
Drill invention fast-tracks creation of tree hollows for wildlife ...
Sep 12, 2021 · An innovative drilling technique that fast-tracks the creation of hollows in tree trunks aims to create a million homes for wildlife displaced by the Black ...
[35]
Which design is better? A lifecycle approach to the sustainable ...
Jul 28, 2025 · This article develops a lifecycle-based design approach to sustainably provide artificial tree hollows for habitat restoration.
[36]
It takes a community to maintain a tree hollow: Food web complexity ...
Jul 19, 2022 · Veteran trees are keystone structures and play vital roles in ecosystems. Tree hollows, which are filled with a mix of leaves and wood that ...
[37]
Artificial Hollows - BRRVLN
Oct 25, 2023 · What is an Artificial Hollow? Artificial hollows take many forms. These include traditional timber nest boxes, 3D printed boxes that mimic ...
[38]
Deep tree hollows: important refuges from extreme temperatures
Nov 1, 2016 · Tree hollows provide important habitat for fauna, offering protection from predators and extreme temperatures. They also provide a safe environment for rearing ...
[39]
Tree cavity patterns and implications for forest management in ...
They provide sites for breeding and sheltering for many taxa and, thus, are important to forest biodiversity (Gibbons and Lindenmayer, 2002).
[40]
Interacting Factors Driving a Major Loss of Large Trees with Cavities ...
Large trees with cavities provide critical ecological functions in forests worldwide, including vital nesting and denning resources for many species.
[41]
Tree Death Brings New Life - National Wildlife Federation
Oct 9, 2021 · Porcupines, squirrels, martens and other mammals also shelter in hollow trees. In winter, brown creepers, small nuthatchlike birds, and some bat ...
[42]
Hollows as Homes - The Australian Museum Blog
Nov 4, 2016 · In New South Wales, terrestrial vertebrate species that are reliant on tree hollows for shelter and nesting include at least 46 mammals, 81 ...
[43]
Is Active Management the Key to the Conservation of Saproxylic ...
Trees with hollows are key features sustaining biodiversity in wooded landscapes. They host rich assemblages of often highly specialised organisms.
[44]
Tree hollow parameters and their effects on saproxylic arthropod ...
Apr 7, 2025 · Tree hollows are keystone structures for biodiversity in forest ecosystems, promoting diverse saproxylic communities.<|separator|>
[45]
Tree hollows can affect epiphyte species composition
May 17, 2017 · Tree hollows often harbor animals and microorganisms, thereby storing nutritive resources derived from their biological activities.
[46]
Loss of hollow-bearing trees in native forests and woodlands due to ...
Oct 3, 2021 · Practices which lead to the loss of hollow-bearing trees are: Logging and associated operations which can remove and damage trees with hollows ...
[47]
Continuing net loss of native hollow-bearing trees and coarse ...
Oct 10, 2021 · Smith AP and Lindenmayer D (1988) Tree hollow requirements of Leadbeater's possum and other possums and gliders in timber production ash ...
[48]
Interacting Factors Driving a Major Loss of Large Trees with Cavities ...
Oct 5, 2012 · ... forest retained adjacent to logging cutblocks [52]. Figure 2 ... tree hollow dynamics simulation model. Forest Ecol Manage 123: 179 ...
[49]
Deforestation Limits Nesting Habitat for Cavity-Nesting Birds
Aug 23, 2023 · “When humans cut down forests for farming, it destroys their nesting habitat,” Ke said. “There are some species that might just disappear if we ...
[50]
Distribution and abundance of hollow-bearing trees in urban forest ...
The retention of hollow-bearing trees is therefore vital to the maintenance of ecosystem integrity in urban environments.
[51]
More than half of NSW's forests and woodlands are gone as ...
Aug 5, 2024 · It's a huge job, but it can be done. Here's an outline of what's needed. Learn more · Glider poking head from tree hollow.
[52]
New Guinea animals losing vital tree cavities to logging, hunting ...
Dec 17, 2013 · Logging and hunting wildlife living in tree cavities will destroy potential breeding sites and will affect species having a safe habitat away ...<|separator|>
[53]
Potential hazard characteristics of trees with hollows, cavities and ...
Dec 10, 2022 · This article is a study of risk assessment of trees with hollows, cavities and fruiting bodies for the improvement of the management and protection of urban ...
[54]
Risk assessment of hollow-bearing trees in urban forests - PMC
Dec 14, 2023 · Given the noticeable trend of increasing numbers of hollow trees from categories A to C, the presence of hollows should be considered during ...Missing: variability | Show results with:variability
[55]
Natural Disturbances are Essential Determinants of Tree-Related ...
Mar 27, 2023 · Natural disturbances play a key role in shaping the structural heterogeneity of forest ecosystems and habitat provisioning for biodiversity.
[56]
Effects of fire severity and fire frequency on the abundance of ...
Fire can create new hollow-bearing trees by excavating hollows or causing branches to break thus exposing existing heartwood decay (Gibbons and Lindenmayer, ...Missing: non- | Show results with:non-<|separator|>
[57]
Wildfire and Climate Impacts Tree Hollow Density in a Temperate ...
Jul 4, 2023 · Frequent severe wildfires may also indirectly facilitate hollow formation by moderating tree death (i.e., many hollows occur in large dead trees ...
[58]
Natural disturbance - Trees for Life
Below we will look more closely at larger scale, 'catastrophic' disturbance, particularly the effects wind, fire, water, insects and disease.
[59]
How deadfall from windstorms can affect wildfire behavior
Apr 16, 2025 · Windstorms can damage structures, infrastructure and natural ecosystems. They may lead to defoliation, branch loss and the destruction of trees and shrubs that ...Missing: storms cavities wildlife
[60]
Persistence of tree cavities used by cavity‐nesting vertebrates ...
Feb 4, 2013 · With variable retention harvesting, a range of tree age and decay classes can be retained to ensure a steady supply of suitable cavity trees as ...
[61]
Snags, Cavity Trees and Downed Logs | Oklahoma State University
Cavity trees are live trees with holes or other structures big enough to shelter animals. This includes trees with only limb cavities. Limb cavities are more ...
[62]
[PDF] Incorporating Wildlife Needs into Forest Management Plans
Some damaged young trees also can be reserved to provide future cavity trees. Trees with fungal conks, dead branches, old scars, and soft or decaying wood ...
[63]
Save our native mature trees – pillars of biodiversity
May 12, 2017 · They provide habitat for wildlife, nurseries for seeds to grow and facilitate vital soil processes. Mature trees with natural hollows are ...
[64]
Optimal management of a flammable multi-stand forest for timber ...
Cavity tree decline can be mitigated by protecting old-growth forest, limiting salvage logging of fire-killed cavity trees (Lindenmayer et al., 2003) and ...
[65]
[PDF] Woodland Wildlife Management - Purdue Extension
Evenly distributed snags and cavity trees are needed in each habitat type (Figures 1 and 2). Forest interior select species with long life spans (favor white ...
[66]
Wildlife Queensland launches initiative to protect threatened tree ...
Nov 19, 2024 · Wildlife Queensland's strategy to protect hollow-bearing trees · Collect eDNA samples to identify species that inhabit tree hollows, from tiny ...
[67]
[PDF] Identifying hollow-bearing and recruitment trees
The PNF Code for northern and southern NSW requires that you must retain 10 hollow bearing trees and 10 recruitment trees per 2 hectares. Where the total number.
[68]
Retention of large, old trees in alternatives to clearcutting with a ...
Nov 1, 2020 · Ground-based surveys are the most commonly used approach to survey hollow-bearing trees and estimate hollow availability (Koch, 2008, Stojanovic ...
[69]
[PDF] Installing chainsaw-carved hollows in medium-sized live trees ...
hollow-dependent wildlife. • Chainsaw hollows may facilitate improved outcomes of revegetation and restoration programs by reducing the amount of time it takes ...Missing: mitigation | Show results with:mitigation<|separator|>
[70]
Artificial tree hollow creation for cavity-using wildlife
Dec 1, 2017 · Five species used the hollows over a 15-month cavity monitoring period; feathertail glider (Acrobates pygmaeus) (in 75% of the hollows), brown ...
[71]
Wildlife Conservation Land Program
When creating snags in a forested stand, select trees 10 inches or larger in diameter to promote longevity. Avoid killing living trees with hollows or dens ...
[72]
[PDF] Restoring Old-Growth Characteristics - Harvard Forest
In the selection of legacy trees, preference should be given to trees in your woods that already contain important habitat features, such as cavities and dens.
[73]
[PDF] INTERACTIONS BETWEEN FUNGI & CAVITY NESTERS - dnr.wa.gov
An important component of maintaining or restoring complex forest habitat in upland and riparian areas is the management of snags (standing dead trees).
[74]
Thermal Profiles of Chainsaw Hollows and Natural Hollows during ...
Loss of large hollow-bearing trees is a major threat to the persistence of wildlife in agricultural landscapes. To combat this problem, local governments, ...
[75]
Artificial habitat structures for animal conservation: design and ...
Feb 15, 2022 · Artificial habitat structures can act as temporary stopgaps while natural habitat structures recover; for instance, artificial tree hollows can ...
[76]
[PDF] Hollows in living trees develop slowly but considerably influence the ...
This is the author manuscript accepted for publication and has undergone full peer review but has not been through the copyediting, typesetting, ...
[77]
Influence of tree hollow characteristics and forest structure ... - PubMed
Dec 8, 2021 · Tree hollows are among the rarest habitats in today's Central European managed forests but are considered key structures for high biodiversity ...Missing: temperate | Show results with:temperate
[78]
Relationships between roosting habitat of bats and decay of aspen ...
Bats are one group of species that rely on tree cavities in these forests, particularly for communal maternity roosting.
[79]
The Abundance of Tree Holes and Their Utilization by Hole-Nesting ...
Aug 6, 2025 · The average density of tree holes in the study area approached 30 holes/ha, while that of hole-nesting birds was 2.4 nests/ha only. The riparian ...
[80]
Full article: Occupancy rates of excavated cavities and nest boxes in ...
We found a high occupancy rate in nest boxes (81%), but lower rates in natural cavities excavated in forest interior, forest edges and retention trees (42-46%).
[81]
Decaying trees improve nesting opportunities for cavity‐nesting ...
Decaying trees improve nesting opportunities for cavity‐nesting birds in temperate and boreal forests: A meta‐analysis and implications for retention forestry ...
[82]
Full article: Tree-related microhabitat assemblages in boreal forests ...
Unlike temperate forests, tree diameter was not a significant driver in boreal forests. The variability of TreM assemblages within the boreal biome underscores ...
[83]
[PDF] Tree hollow abundance in the tropical savannas of northern Australia
Termite activity was strongly associated with a higher abundance of hollows. Most trees in northern. Australian savannas have hollow cores or pipes due to ...
[84]
Cyclones, fire, and termites: The drivers of tree hollow abundance in ...
Jul 1, 2018 · Tree hollow abundance in mesic tropical savanna woodlands is high (88 hollows ha−1). · Important predictors of abundance are rainfall, soil depth ...
[85]
The influence of fire and termites on tree hollow development in an ...
Feb 1, 2025 · In temperate southern Australia, a reduction in tree hollows has been associated with reduced abundance of hollow-dependent fauna (e.g. Smith ...
[86]
https://www.britannica.com/topic/rainforest-tree-hollows-1673047
[87]
(PDF) The importance of scattered trees for biodiversity conservation
Areas with scattered trees support greater levels of biodiversity than open areas, as well as communities that are more similar to those inhabiting habitat ...
[88]
Origin of trunk damage in West African savanna trees
Mar 10, 2011 · Two main types of hollow tree are frequently found in savannas: trees with external splits (externally damaged trees), and trees with no or ...
[89]
Overlapping den tree selection by three declining arboreal mammal ...
Aug 25, 2020 · In Australia's tropical savannas, development of tree cavities is enhanced by high termite activity, and, conversely, reduced by frequent fires.
[90]
The availability of Blue-fronted Amazon nesting cavities is at risk in ...
Eighteen percent of the world's avifauna uses tree cavities for nesting, including 93% of the 374 psittacid species (van der Hoek et al. 2017). Approximately ...
[91]
The Multi‐Functional Use of Large Tree Cavities by Arboreal ...
Nov 7, 2024 · For vertebrate animals, cavities can provide shelters from predators and stable microclimate conditions, offering breeding, resting, feeding, or ...
[92]
[PDF] Hollow formation in the Eucalyptus miniata – E. tetrodonta open ...
Despite their less substantial forest structure, the total density of hollow trees, and hollows, is greater in tropical eucalypt forests than is typical of ...
[93]
The Future of Large Old Trees in Urban Landscapes - PMC
Jun 18, 2014 · ... hollows but rather relative hollow occurrence per tree. ... hollow and tree-dependent biodiversity are to be maintained in urban landscapes.
[94]
[PDF] Hollow-Bearing Trees as a Habitat Resource along an Urbanisation ...
Apart from tree age and species, the formation of hollows is also influenced ... Hollow formation is influenced by a variety of environmental factors related to ...<|separator|>
[95]
Tree cavity availability in urban cemeteries and city parks
Jan 22, 2019 · Tree cavities provide shelter, nesting sites and food storage for many species of birds, mammals and insects. While tree cavities are ...Missing: peer | Show results with:peer
[96]
Does forest loss and fragmentation reduce woodpecker-associated ...
Woodpeckers inhabiting large trees are particularly susceptible to forest loss and fragmentation. They play a critical role in the ecosystem, as they not only ...Missing: wildlife | Show results with:wildlife<|separator|>
[97]
Risk assessment of hollow-bearing trees in urban forests - Nature
Dec 14, 2023 · Does the risk class (A, B, C, and D) depend on the presence of hollows or decay? Is there a correlation between the presence of hollow cavities ...
[98]
Old trees are perceived as a valuable element of the municipal ...
Jan 12, 2022 · Hollows are found in old trees, whether living or not. A tree hollow, usually in the form of a semi-enclosed cavity, forms naturally in the ...