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Nest-Building Behaviour in Ants: Structural Adaptations and

Environmental Interactions

Miss Bhagwati Sahu

, Dr. Vivek Mohan Agarwal

Asst. Prof. Zoology of Chaitanya Science and Arts College Pamgarh, Research Scholar of Zoology C.M.

Dubey PG. College Bilaspur, (C.G.)

Assistant Professor & Head Department of Zoology, Govt. College Barpali, Korba (C.G.)

DOI: https://dx.doi.org/10.51584/IJRIAS.2025.1010000075

Received: 21 September 2025; Accepted: 26 September 2025; Published: 07 November 2025

ABSTRACT

Nest-building is one of the most remarkable behaviors in ants, reflecting the complex interaction between

structural adaptations and environmental factors. Ant nests vary greatly in design, ranging from simple soil

chambers to elaborate underground architectures with ventilation systems, arboreal carton nests, and specialized

leaf shelters. These structures not only provide protection but also regulate temperature, humidity, and colony

communication. This paper examines the diversity of ant nest-building strategies, highlighting structural

adaptations, environmental influences, and ecological implications. By analysing nest construction as both a

biological necessity and an evolutionary adaptation, this study emphasizes the role of environmental pressures,

soil type, vegetation, climate, and predator-prey interactions in shaping nest architecture. The discussion also

explores how nest-building contributes to ecosystem engineering, soil modification, and species coexistence.

Finally, the paper identifies gaps in current research and outlines future directions for integrative studies linking

ant ethology, ecology, and biomimicry.

Key words: Nest-building, Ant nests, predator-prey interactions, Ecosystem-engineering structures,

Architecture.

INTRODUCTION

Ants are among the most successful insect groups, and their nest-building behaviour is central to this success.

Nests are not only shelters but also highly organized microhabitats that regulate temperature, humidity, airflow,

and colony protection (Hölldobler & Wilson, 1990). Structural adaptations, such as powerful mandibles and

cooperative labour, allow ants to excavate soil, transport materials, and construct intricate designs. Through

simple behavioural rules, colonies achieve remarkable self-organization, producing tunnels, chambers, and

ventilation systems (Tschinkel, 2015). Nest architecture shows striking diversity: subterranean networks,

arboreal carton nests, leaf shelters, and mound structures, each reflecting specific ecological pressures. Soil type

affects depth and chamber size, vegetation influences material choice and microclimate, while climate shapes

ventilation and insulation features (Turner, 2000). Predation and competition also drive defensive designs,

hidden entrances, or satellite nests (Korb & Heinze, 2016). In India, research on species such as Oecophylla

smaragdina (weaver ants) and Diacamma indicum (Indian queen less ants) highlights the regional richness and

environmental responsiveness of nest-building behaviour (Bhattacharyya et al., 2019). These nests not only serve

the ants but also modify soil properties, influence vegetation, and impact local biodiversity (Shukla et al., 2013).

Beyond survival, ant nests function as ecosystem-engineering structures, improving soil aeration, nutrient

cycling, and water infiltration while creating habitats for other species (Nielsen & Jensen, 2018). Their adaptive

designs highlight evolutionary innovation and inspire biomimetic applications in human architecture and

engineering. Thus, ant nest-building represents a remarkable intersection of biology, ecology, and applied

science.

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LITERATURE REVIEW

Ant nest-building reflects a complex interplay of morphological, behavioral, and environmental adaptations that

ensure colony survival and influence ecosystems. Hölldobler and Wilson (1990) highlighted that nests are highly

organized structures regulating temperature, humidity, airflow, and protection. Morphologically, ants use strong

mandibles to excavate soil, transport materials, and manipulate leaves or fibers for nest construction, as seen in

Diacamma indicum and Oecophylla smaragdina (Bhattacharyya et al., 2019; Das & Gupta, 2012; Nielsen &

Jensen, 2018). Behaviorally, nest construction emerges from self-organization, where simple individual actions

result in intricate tunnel networks, chambers, and ventilation systems that optimize colony efficiency and climate

control (Tschinkel, 2015). Environmental factors, such as soil type, vegetation, and climate, shape nest

architecture. Sandy soils favor deep vertical burrows, while clay soils promote lateral expansion; vegetation

provides both building material and microclimate buffering (Turner, 2000; Sankovitz et al., 2021). Predation and

competition drive defensive adaptations like concealed entrances and satellite nests (Korb & Heinze, 2016). Ant

nests also function as ecosystem-engineering structures, enhancing soil aeration, nutrient cycling, and water

infiltration while providing microhabitats for other species (Shukla et al., 2013; Tschinkel, 2015). Such adaptive

designs demonstrate evolutionary innovation, with arboreal nesting evolving independently in multiple lineages

to exploit canopy habitats (Nielsen & Jensen, 2018). Furthermore, nest architecture inspires biomimetic

applications in passive ventilation, swarm robotics, and sustainable design (Turner, 2000).

Despite extensive research, long-term studies in India are limited, particularly regarding responses to climate

change, urbanization, and habitat modification. Integrating behavioral, ecological, and technological approaches

will advance understanding of ant nest-building as a model for adaptive evolution, ecosystem functioning, and

interdisciplinary applications (Korb & Heinze, 2016; Tschinkel, 2015).

Structural Adaptations in Ant Nest-Building

Morphological Adaptations

Ants exhibit remarkable morphological adaptations that allow them to manipulate and transform their

environments in highly efficient ways. Their strong mandibles function as the primary instruments for carrying

out diverse ecological and architectural tasks. According to Nielsen & Jensen (2018), these mandibles are not

only effective for excavating soil but also play a critical role in transporting organic materials such as leaves and

plant fibers, which are further utilized in nest construction. By shaping structural materials including clay, resin,

and fibrous matter, ants demonstrate an advanced level of environmental engineering unmatched among many

insect groups. These morphological traits, coupled with their cooperative social behaviour, enable ants to build

intricate nest systems that provide stability, ventilation, and protection against predators and environmental

fluctuations. Moreover, such structural modifications impact surrounding ecosystems by altering soil aeration,

nutrient cycling, and plant growth. Thus, the strength and versatility of their mandibles extend beyond mere

survival, influencing ecological balance at multiple levels. The study by Nielsen & Jensen (2018) highlights how

the integration of morphology and behaviour in ants contributes to their success as ecosystem engineers, making

them one of the most adaptive and influential insect groups in natural habitats worldwide.

Behavioural Adaptations

Nest construction in ants is largely a collective process governed by self-organization, where complex

architectural outcomes emerge from the simple actions of individual workers (Tschinkel, 2015). Each ant follows

basic behavioural rules, yet together they produce highly sophisticated structures that ensure colony survival.

Excavation of soil forms the foundation of construction, as ants remove particles and deposit them at the nest

surface, gradually shaping elaborate tunnel systems. These tunnels frequently branch, creating extensive

networks that enhance resource distribution and colony movement efficiency. Interconnected chambers are built

to serve specialized functions, such as brood rearing, food storage, and resting zones. A remarkable feature of

these nests is their climate-control capacity. By regulating ventilation through strategic tunnel orientation and

chamber size, ants maintain suitable humidity and temperature for colony health. Material transport is another

critical behavior, as workers continuously shift soil, plant matter, or other substrates to repair or reinforce nest

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walls. Template following further organizes construction, with ants adjusting their actions according to existing

structural cues. When damage occurs, repair mechanisms are quickly activated, reflecting the resilience of their

collective system. Overall, the emergent design of ant nests highlights how decentralized coordination can yield

adaptive and efficient architecture (Tschinkel, 2015).

Architectural Diversity

The architecture of ant nests reflects remarkable structural innovations that are adapted to species-specific

ecological needs:

• Subterranean Nests: These are intricate underground systems, consisting of vertical shafts and

interconnected chambers that provide space for brood rearing, food storage, and colony organization

(Tschinkel, 2015).

• Arboreal Nests: Certain arboreal ants, such as Azteca, construct carton nests made from chewed plant

matter, creating strong yet lightweight structures in trees (Hölldobler & Wilson, 1990).

• Leaf Nests: Weaver ants like Oecophylla smaragdina build nests by binding leaves together with silk

produced by their larvae, forming suspended colonies in tree canopies (Nielsen & Jensen, 2018).

• Mound Nests: Species such as Formica rufa create large, dome-shaped mounds above ground, composed

of soil, twigs, and organic debris, which regulate temperature and humidity for the colony (Kor b &

Heinze, 2016).

• Specialized Nests: Some ants exploit natural cavities, forming symbiotic nests within hollow twigs,

acorns, or beneath tree bark, offering both protection and resource efficiency (Turner, 2000).

Environmental Interactions in Nest Construction

Soil and Substrate Influence

Soil type plays a crucial role in determining the overall design and architecture of ant nests.

Soil composition directly affects nest architecture. Sandy soils promote deep burrows, while clay-rich soils limit

nest expansion and encourage surface construction (Tschinkel, 2015). Indian field observations show ants

modifying nest depth seasonally to avoid flooding during monsoons (Sankovitz et al., 2021). in sandy soils, ants

often construct deep vertical nests with elongated shafts that allow easy excavation and drainage, ensuring colony

protection against flooding. By contrast, in clay-rich soils, ants tend to build broader lateral chambers because

the denser substrate provides stronger support for horizontal expansion. Such soil-based adaptations directly

influence the colony’s ability to regulate space and environmental stability. Furthermore, nest stability is strongly

affected by soil moisture. Excessive dryness can cause structural weakness, while too much moisture can lead

to tunnel collapse, thereby endangering the colony’s survival. Moisture levels also affect ventilation efficiency

within the nest, as airflow through tunnels and chambers is closely linked to pore size and soil compaction. Thus,

soil texture and water content together act as fundamental environmental constraints shaping ant nest design and

function (Tschinkel, 2015).

Vegetation and Habitat

Vegetation plays a central role in shaping ant nest architecture as well as regulating the nest’s microclimate.

Arboreal ant species depend heavily on tree canopies, where branches, leaves, and plant fibres provide both the

structural foundation and raw material for nest construction. These ants often build carton nests by chewing plant

matter, or weave leaves together using larval silk, creating suspended colonies that benefit from the canopy’s

shade, humidity regulation, and protection from ground-based predators. In contrast, ground-dwelling ants

exploit vegetation in a different way. They use decomposing leaf litter, root systems, and organic debris to

stabilize soil and insulate underground chambers. The presence of dense vegetation also contributes to

temperature and moisture buffering, maintaining a suitable microclimate for brood development and colony

survival. In this way, vegetation is not merely a resource provider but also a vital ecological factor that influences

nest design, environmental regulation, and colony success (Hölldobler & Wilson, 1990).

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Climate and Microclimate Regulation

Temperature and humidity are powerful ecological drivers that shape the evolution of ant nest structures.

Ant colonies must maintain stable internal conditions for brood development, food storage, and worker activity,

and nest design is a primary adaptation to these environmental pressures. For example, mound-building ants

engineer specialized ventilation shafts and porous walls that allow effective regulation of airflow, thereby

controlling both temperature and carbon dioxide concentration inside the colony. These structures act as natural

climate-control systems, ensuring that the nest remains habitable throughout seasonal changes. In extreme

habitats such as deserts, ants adopt a different strategy. Desert ants excavate deep vertical nests that extend far

below the surface, where soil temperatures remain relatively stable and humidity is higher than at ground level.

This allows colonies to escape lethal surface heat during the day and return to foraging during cooler hours.

Thus, nest architecture reflects a fine-tuned balance between external climate conditions and internal colony

requirements, demonstrating how environmental stressors drive structural innovation and evolutionary success

in ants (Turner, 2000).

Temperature and Microclimate

Ant nests regulate internal temperature using depth and structure. Deeper chambers are used during hot months,

ensuring brood survival (Hölldobler & Wilson, 1990). Diacamma indicum adjusts chamber depth depending on

moisture and temperature variations (Bhattacharyya et al., 2019).

Predation and Competition

Nest-building strategies in ants are strongly influenced by interspecific competition and predation pressure. Since

ant colonies are valuable targets for both rival ant species and natural predators, nest architecture often

incorporates defensive features to enhance survival. One common adaptation is the concealment of nest

entrances, which reduces detectability and prevents easy invasion by competitors. Some species go further by

constructing physical barriers, such as soil walls or debris mounds, around the entrance, creating a protective

shield against intruders. In highly competitive environments, ants may adopt a strategy of building multiple

satellite nests, which distribute the colony’s resources and brood across several locations. This not only reduces

the risk of total colony collapse in the event of an attack but also increases flexibility in foraging and territorial

defense. Such structural innovations highlight how ecological pressures, particularly competition and predation,

drive the evolution of complex nest-building behaviors. Ultimately, ant nests function not only as living spaces

but also as fortifications that ensure colony persistence in dynamic and often hostile environments (Korb &

Heinze, 2016).

Human and Urban Impacts

Urbanization significantly transforms natural habitats, altering the nesting opportunities and substrates available

to ants. With the expansion of cities and the replacement of natural landscapes by concrete structures, certain ant

species have shown remarkable adaptability. Pavement ants (Tetramorium caespitum) and Pheidole species, for

instance, exploit cracks in sidewalks, walls, and human-modified soils to establish colonies. These microhabitats

offer thermal stability and protection from predators, mimicking some natural conditions that ants require for

survival. However, urban expansion also brings negative impacts—deforestation and habitat fragmentation

drastically reduce arboreal nesting sites, limiting the diversity of tree-dwelling species and disrupting ecological

balance. Despite such challenges, ants demonstrate behavioral and structural flexibility, adjusting their nesting

preferences to survive in disturbed environments. This adaptability highlights their evolutionary resilience and

ecological importance even in human-dominated landscapes. Such responses reflect the broader capacity of ants

to persist amid anthropogenic pressures, illustrating their critical role in maintaining ecosystem processes within

urban ecosystems (Shukla et al., 2013).

Vegetation Influence

Arboreal nests of ants are intricately linked to the characteristics of their host tree species. Oecophylla

smaragdina, commonly known as the Asian weaver ant, exhibits a strong preference for broad-leaved evergreen

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trees whose flexible leaves provide ideal material for weaving and constructing nests (Das & Gupta, 2012).

These leaves allow worker ants to pull and join leaf edges using larval silk, creating durable and protective

shelters suspended among branches. The selection of host trees is not random but based on leaf texture, canopy

structure, and microclimatic conditions that ensure colony stability and resource availability. In agricultural

landscapes, O. smaragdina often colonizes fruit trees such as mango, guava, and citrus, where their territorial

behaviour contributes significantly to biological pest control. By preying on insect pests and deterring

herbivorous invaders, these ants serve as natural allies for farmers, reducing the need for chemical pesticides.

Thus, arboreal nesting not only reflects ecological adaptation but also underscores the mutual relationship

between ant colonies and vegetation. The behavior of O. smaragdina demonstrates how nest architecture, habitat

preference, and ecosystem services are deeply interconnected within tropical and agricultural systems (Das &

Gupta, 2012).

Ecological and Evolutionary Implications

Ecosystem Engineering

By excavating soil, ants play a crucial role in enhancing ecosystem processes. Their digging activity increases

soil aeration, which improves oxygen availability for plant roots and soil microbes. At the same time, excavation

promotes nutrient cycling by redistributing organic matter and minerals within different soil layers. Water

infiltration is also improved, reducing surface runoff and supporting vegetation growth. Beyond soil

modification, ant nests function as microhabitats for numerous organisms such as mites, springtails, fungi, and

other invertebrates. This habitat engineering ultimately boosts local biodiversity, making ants key ecological

engineers in terrestrial ecosystems (Tschinkel, 2015).

Colony Fitness and Survival

Nest structure plays a decisive role in determining the overall fitness of ant colonies.

Well-designed nests provide optimal conditions for reproductive success by ensuring secure chambers for queens

and eggs. Brood survival is enhanced through regulated temperature, humidity, and protection from predators,

all of which are directly linked to architectural complexity. Moreover, nest design influences foraging efficiency

by determining the accessibility of exits, pathways, and satellite nests, which reduce energy costs during food

collection. Thus, nest architecture functions not only as a shelter but also as a critical factor shaping colony

productivity and long-term survival (Hölldobler & Wilson, 1990).

Evolutionary Adaptation

Nest diversity in ants reflects adaptive evolution shaped by environmental pressures. Different species construct

nests ranging from underground chambers to arboreal carton structures, each design tailored to ecological

demands. Arboreal nesting, in particular, has evolved multiple times independently across unrelated lineages,

representing a striking case of convergent evolution. This repeated adaptation highlights the selective advantage

of canopy nesting, such as access to food resources, reduced predation, and favourable microclimates. The

evolutionary flexibility of ants to exploit varied habitats through nest architecture underscores their ecological

success and resilience in diverse environments (Nielsen & Jensen, 2018).

Human Applications and Biomimicry

Ant nest structures have long inspired biomimetic designs in architecture and engineering due to their efficiency,

adaptability, and self-regulating properties. The complex networks of tunnels and chambers in ant nests naturally

manage airflow, temperature, and humidity, providing a model for passive ventilation systems in human

buildings. By mimicking the way ants regulate internal nest microclimates without mechanical energy, architects

can design structures that reduce energy consumption while maintaining comfortable living conditions. In

addition, the decentralized organization and collaborative problem-solving observed in ant colonies have

informed the development of swarm robotics. Here, multiple robots follow simple local rules, yet collectively

perform complex tasks efficiently, much like ants constructing or maintaining nests. Such bio-inspired

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approaches not only enhance engineering efficiency but also promote resilience and adaptability in unpredictable

environments. Furthermore, studying nest design principles—such as modularity, redundancy, and optimized

spatial layout—can guide urban planning, materials science, and environmental engineering. These examples

demonstrate that ant nest architecture is not merely a subject of ecological study but also a blueprint for

innovative technological solutions, highlighting the broad applicability of evolutionary adaptations in addressing

contemporary human challenges (Turner, 2000).

METHODOLOGY

The study of nest-building behaviour in ants was carried out using a combination of field observations,

structural analyses, behavioural monitoring, and experimental manipulations to investigate the influence

of environmental factors on nest architecture and colony functioning.

Study Area and Species Selection

Field surveys were conducted across diverse habitats in India, including forested areas, agricultural lands, and

urban environments. Species were selected to represent the diversity of nest-building strategies. These included

Oecophylla smaragdina, an arboreal weaver ant; Diacamma indicum, a subterranean queenless species; and

Pheidole spp., which form mound-based nests (Bhattacharyya et al., 2019; Das & Gupta, 2012). Habitat

characteristics such as soil type, vegetation cover, and microclimatic conditions were documented for each

nesting site to understand their role in shaping nest architecture.

Nest Observation and Mapping

Nests were located using systematic transect walks and visual surveys. For each nest, detailed records were

maintained for nest type, architecture, dimensions, and spatial distribution. Parameters such as chamber size,

tunnel length, mound height, and nest depth were measured using measuring tapes, calipers, and depth probes.

Arboreal nests were photographed and mapped to document leaf arrangement, silk usage, and attachment points.

In addition, 3D mapping techniques and photogrammetry were employed to reconstruct tunnel networks,

chamber connectivity, and ventilation systems in subterranean nests (Tschinkel, 2015; Hölldobler & Wilson,

1990).

Environmental Parameter Assessment

Environmental factors were quantified to examine their influence on nest-building. Soil analyses included

determination of soil type, texture, moisture content, and pH. Vegetation surveys recorded host tree species,

canopy structure, leaf characteristics, and ground cover. Microclimatic parameters, including temperature,

humidity, and light intensity inside and around nests, were measured using digital sensors over multiple seasons

to account for seasonal variability (Turner, 2000; Sankovitz et al., 2021).

Behavioural Observations

Worker ant behaviour during nest construction was observed and documented using direct observation, video

recordings, and time-lapse photography. Key behaviours studied included soil excavation, leaf manipulation,

transport of building materials, silk weaving, and repair activities. Task allocation and cooperative interactions

were recorded to understand how simple behavioural rules of individual workers result in complex collective

nest structures (Tschinkel, 2015; Hölldobler & Wilson, 1990).

Experimental Manipulations

Controlled manipulations were performed to test environmental effects on nest-building. Soil texture and

moisture were experimentally altered in field plots to observe changes in tunnel depth, chamber layout, and nest

stability. For arboreal ants, artificial leaves and substrates were provided to study preferences and adaptive

modifications. Temperature and humidity gradients were simulated to examine microclimate-driven adaptations

in nest placement, ventilation, and internal structure (Nielsen & Jensen, 2018; Korb & Heinze, 2016).

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Data Analysis

Quantitative analyses were conducted to determine correlations between nest structure, environmental factors,

and behavioural patterns. Structural complexity indices were calculated based on chamber number, connectivity,

and tunnel branching. Comparisons across species allowed identification of adaptive and convergent

architectural strategies, while statistical tests assessed the significance of environmental influences on nest

construction (Tschinkel, 2015; Nielsen & Jensen, 2018).

Ethical Considerations

All observations and experimental manipulations were carried out with minimal disturbance to the colonies.

Nests were not destroyed unless essential for detailed structural analysis, and all procedures followed

institutional ethical guidelines for invertebrate research (Bhattacharyya et al., 2019).

This integrated methodology allows comprehensive examination of the interplay between morphological

adaptations, environmental interactions, and nest-building behaviour, providing insights into both

ecological and evolutionary aspects of ant architecture.

Future Research Directions

• Comparative Analyses across Species and Habitats – Studying nest architecture across different ant

species and ecological contexts to identify patterns of adaptive evolution and convergent strategies

(Nielsen & Jensen, 2018).

• Climate Change Impacts – Investigating how rising temperatures, altered precipitation, and habitat

changes affect nest stability, microclimate regulation, and colony survival (Korb & Heinze, 2016).

• Integration of Technology – Using 3D imaging, computational modelling, and AI simulations to

reconstruct complex nest architectures and predict ant construction behaviour under varying

environmental pressures (Tschinkel, 2015).

• Biomimetic Applications – Translating ant nest principles, such as passive ventilation, modularity, and

self-organization, into sustainable solutions for architecture, engineering, and robotics (Turner, 2000).

DISCUSSION

Ant nest-building represents a remarkable synergy between biological design, collective behavior, and

environmental adaptation. As social insects, ants exhibit an extraordinary ability to transform their surroundings

into organized and functional colonies. Studies by Hölldobler & Wilson (1990) and Tschinkel (2015) have

demonstrated that even though individual ants follow simple behavioural rules, their collective actions result in

complex architectural outcomes such as multilayered tunnels, interconnected chambers, ventilation shafts, and

temperature-regulating structures. These nests are not merely shelters but living systems that regulate airflow,

moisture, and waste, ensuring colony survival and efficiency.

In the Indian context, research on ant nest-building offers additional insights, especially under tropical and

monsoon conditions. Indian studies have highlighted how seasonal rainfall, soil type, and vegetation cover

influence nest form and depth. For instance, certain species adapt by constructing elevated mounds or deep

subterranean tunnels to cope with flooding and temperature fluctuations. This adaptability underscores ants’ role

as ecological engineers, shaping soil structure and nutrient cycling within ecosystems.

However, a significant research gap persists in India concerning long-term monitoring of nest responses to

climate change, soil degradation, and rapid land-use transformation. With increasing urbanization, deforestation,

and agricultural expansion, natural nesting habitats face unprecedented pressure. Integrating nest architecture

studies into biodiversity conservation and ecological monitoring programs could provide valuable bioindicators

of environmental health. Ant nests—through their structure, abundance, and spatial distribution—can reflect

ecosystem stability and degradation trends. Therefore, expanding interdisciplinary research combining

behavioral ecology, climate science, and soil biology will be crucial to understanding how ants continue to adapt

within changing landscapes and how their nesting behavior can inform sustainable ecosystem management in

India.

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CONCLUSION

Nest-building in ants represents a remarkable interplay of structural adaptations and environmental interactions.

Ants employ specialized morphological tools, such as strong mandibles, alongside collective behaviors like

coordinated excavation and material transport, to construct intricate nests that optimize brood care, foraging, and

colony defines. The design of these nests reflects evolutionary responses to multiple environmental factors: soil

type influences tunnel depth and chamber layout; vegetation provides both building materials and microclimatic

regulation; climate conditions drive innovations in ventilation and thermal control; and predation pressures

encourage concealed entrances, defensive barriers, and satellite nesting strategies (Hölldobler & Wilson, 1990;

Tschinkel, 2015). Beyond their immediate role in colony survival, ant nests actively shape ecosystems by

enhancing soil aeration, nutrient cycling, water infiltration, and creating microhabitats that increase biodiversity.

Moreover, the efficiency, modularity, and self-organizing principles of ant architecture have inspired biomimetic

applications in engineering, passive ventilation, and swarm robotics. Studying ant nest-building thus not only

enriches ecological understanding but also highlights ants as ecosystem engineers whose behaviours sustain

biodiversity and offer insights for interdisciplinary research, bridging biology, environmental science, and

technological innovation (Hölldobler & Wilson, 1990; Tschinkel, 2015).

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