Pteroglossus torquatus — Collared Aracari

Tucancillo Collarejo · Cusingo · Tití · Félix
Kingdom: Animalia Phylum: ChordataClass: Aves Order: Piciformes Family: Ramphastidae Genus: PteroglossusSpecies: Pteroglossus torquatus (Gmelin, 1788)
Conservation status: Least Concern (LC) Population trend in Costa Rica: Decreasing
Costa Rican status: Resident
Costa Rican elevational range: approximately 0–1,000 m
Primary Costa Rican distribution: Caribbean slope and northern Pacific; locally extending into adjacent forest landscapes

01 — Species Overview
A group of Collared Aracaris moving through the canopy can seem almost choreographed.
One bird disappears behind a branch. Another follows. A third lands on a nearby tree carrying a fruit. Several birds call from different positions before the entire group moves again.
Pteroglossus torquatus is a medium-sized member of the toucan family, reaching approximately 41 cm in length in Costa Rica. Its plumage combines an olive-green back, black head and upper chest, yellow underparts, a reddish collar and a conspicuous red rump. The large bill, characteristic of toucans, is surprisingly light because of its specialized internal structure.
In Costa Rica, the species is primarily associated with lowland and foothill forests, particularly on the Caribbean slope. It also occurs in forest edges, secondary growth and some partially open environments where suitable trees provide food and nesting or roosting cavities. The Museo Nacional de Costa Rica gives an elevational range of approximately 0–1,000 m for the species.
Its diet is dominated by fruit, especially fleshy fruits and palm fruits, but insects and other animal material are also consumed. The species therefore participates simultaneously in plant–animal interactions and predator–prey relationships.
Perhaps the most distinctive feature of its natural history, however, is its social behaviour.
Collared Aracaris can move in groups, use communal roosts and participate in cooperative breeding. Other members of the social group may assist a breeding pair in caring for young.
This makes P. torquatus more than a colourful forest bird.
It is a species through which we can observe how food, cavities, social cooperation and seed movement become connected within a tropical forest.
02 — Why This Species Matters
The Collared Aracari is an excellent example of why a species should not be interpreted as an isolated organism.
Its ecology crosses several biological systems.
The bird depends on fruiting plants for food.
Those plants potentially benefit from the movement of their seeds.
The aracari depends on cavities for reproduction and roosting.
Those cavities may have been created or modified by other organisms.
The species lives socially, meaning that the survival of an individual can depend partly on interactions with several other members of its group.
And the same bird that transports seeds can also prey upon insects, eggs and nestlings.
Its ecological identity is therefore not a single function.
It is a network.
For Costa Rica, the species is particularly useful because its distribution provides a clear field distinction from the closely related Fiery-billed Aracari, Pteroglossus frantzii. The Museo Nacional de Costa Rica notes that P. torquatus occupies the Caribbean slope and northern Pacific, while P. frantzii is associated principally with the central and southern Pacific.
Geography therefore becomes part of identification.
03 — Taxonomic Identity
3.1 Classification
Kingdom: AnimaliaPhylum: ChordataClass: AvesOrder: PiciformesFamily: RamphastidaeGenus: PteroglossusSpecies: Pteroglossus torquatus
The species was described by Johann Friedrich Gmelin in 1788.
3.2 Scientific Name
The genus Pteroglossus derives from Greek roots associated with the concepts of wing and tongue.
The specific epithet torquatus comes from Latin and means "collared" or "wearing a collar."
The name refers directly to the reddish collar that separates the dark upper chest from the greenish and yellow portions of the body.
3.3 Taxonomic History
The taxonomy of Pteroglossus has undergone revisions as ornithologists have reassessed geographic variation and relationships among aracaris.
The interpretation of the Pteroglossus torquatus complex has therefore changed among taxonomic authorities over time.
This is important because geographic variation in plumage and distribution can make species boundaries difficult to define.
The current Species Note follows Pteroglossus torquatus as the species-level treatment used for the Collared Aracari in Costa Rica.
04 — Identification
4.1 General Appearance
The Collared Aracari is approximately 41 cm long in Costa Rica, making it smaller than the large Ramphastos toucans but considerably larger than the smallest Costa Rican toucanets.
The principal field characters are:
black head;
black upper chest;
olive-green back and wings;
bright red rump;
red collar;
yellow underparts;
black breast marking;
dark band across the lower belly;
chestnut thighs;
large patterned bill.
The combination of the red collar and red rump is particularly useful.
4.2 Sexual Dimorphism
Males and females are broadly similar in plumage.
Sexual differences are more subtle than in strongly dimorphic species such as the Great Curassow.
Bill dimensions can differ between sexes, but sex should generally not be assigned confidently from a brief field observation unless additional evidence is available.
4.3 Juveniles
Juveniles are generally duller than adults and lack the full intensity of adult plumage.
The contrast between the red, yellow, green and black areas becomes more pronounced as the bird matures.
4.4 Similar Species
The most important Costa Rican comparison is:
Pteroglossus torquatus vs. Pteroglossus frantzii.
Both are aracaris with similar general body proportions and colourful plumage.
Their geographic distribution is therefore extremely useful.
In Costa Rica:
P. torquatus is associated principally with the Caribbean slope and northern Pacific.
P. frantzii occupies the central and southern Pacific.
The Museo Nacional de Costa Rica specifically recommends distribution as an important component of distinguishing the two species.
This is an excellent example of field identification using more than plumage.
Identification is sometimes geographic as well as morphological.
05 — Morphology
5.1 Size
The Collared Aracari measures approximately 41 cm in Costa Rica. Among the country's six toucan species, it is intermediate in size: smaller than Ramphastos ambiguus and Ramphastos sulfuratus, but larger than Selenidera spectabilis and Aulacorhynchus prasinus.
5.2 The Bill
The bill is one of the most conspicuous structures in the species.
Despite its size, it is remarkably lightweight.
The Museo Nacional de Costa Rica describes the toucan bill as highly specialized, practically hollow internally and composed largely of keratin.
This is a classic example of:
structure → function.
A large external bill does not necessarily require a correspondingly heavy structure.
5.3 Functional Morphology
The bill gives the aracari considerable reach.
A bird perched on a branch can extend its body and bill toward fruits that are positioned beyond the immediate reach of its feet.
This allows the bird to exploit resources within the three-dimensional structure of the canopy without moving to every fruit-bearing branch.
The morphology therefore directly influences its foraging ecology.
06 — Distribution
6.1 Global Distribution
The Collared Aracari occurs from Mexico through Central America and into northern South America, including Colombia and Venezuela.
Its range is therefore substantially broader than Costa Rica.
6.2 Costa Rica
In Costa Rica, P. torquatus is a resident species.
Its strongest distributional association is with the Caribbean slope and northern Pacific regions. The species is less common in parts of the northern Pacific than in Caribbean lowlands.
6.3 Elevation
The national catalogue of Costa Rican toucans places the species approximately between 0 and 1,000 m elevation.
This reinforces its identity as primarily a lowland and foothill aracari.
6.4 Distribution as an Identification Tool
The geographic separation between P. torquatus and P. frantzii is particularly useful in Costa Rica.
Two birds may appear similar enough to create uncertainty when viewed quickly.
Knowing where you are can resolve part of the problem.
This illustrates an important principle of field identification:
the landscape itself can be a diagnostic character.
07 — Habitat
7.1 Primary Habitat
The Collared Aracari is associated principally with tropical forest.
It is particularly characteristic of lowland and foothill forests on the Caribbean slope.
7.2 Secondary Forest and Edges
Costa Rican sources document the use of forest edges, secondary growth and partially open environments.
The Área de Conservación Guanacaste has observed the species travelling through forest edges and visiting nearby gardens and farms where fruits such as bananas and plantains are available.
This demonstrates ecological flexibility without implying that the species is independent of forest.
7.3 The Importance of Trees
For toucans generally, habitat quality depends strongly on trees that provide:
food;
nesting sites;
roosting sites.
The Museo Nacional de Costa Rica emphasizes that suitable toucan habitat must contain trees fulfilling these functions.
For P. torquatus, this relationship is particularly important because both reproduction and communal roosting depend on cavities.
08 — Ecology
8.1 Ecological Niche
The Collared Aracari occupies an arboreal, primarily frugivorous niche.
It moves through the middle and upper levels of forest vegetation, searching for fruiting plants.
It can also descend into lower vegetation when food resources or other opportunities are available.
8.2 Diet
Fruit is the principal food category.
Costa Rican observations document consumption of fruits and seeds, including palm fruits.
The species also consumes insects and can prey upon the nests of other birds.
This creates a more complex trophic profile than the common image of the aracari as simply a fruit-eating bird.
8.3 Palm Fruits
Palm fruits are particularly important in its diet.
ACG field observations in 2024 documented adult P. torquatus transporting Bactris fruits to a nestling.
This is a specific field observation, not a claim that Bactris is the only or universally most important food resource.
That distinction is essential.
8.4 Foraging Behaviour
The species can manipulate fruit while perched and can extend its body to reach food.
Its bill allows it to access resources at distances that would otherwise require movement.
The bird therefore uses the architecture of branches as part of its feeding strategy.
8.5 Seed Dispersal
The Collared Aracari is a documented component of tropical seed-dispersal systems.
Research modelling seed dispersal in New World tropical forests has specifically included P. torquatus, combining bird movement and gut-retention information to estimate potential dispersal kernels.
This provides a quantitative basis for considering the species a seed disperser.
The ecological importance of that function, however, varies according to the plant species and landscape.
09 — Behaviour
9.1 Social Groups
Collared Aracaris are social birds.
Costa Rican sources describe small groups that may contain several individuals, while the broader toucan community can form groups ranging approximately from five to fifteen birds depending on species and context.
The Collared Aracari is commonly encountered in groups rather than consistently as a solitary bird.
9.2 Group Movement
Groups can move rapidly through the canopy.
A sequence of calls followed by several birds crossing between trees can reveal the presence of an entire group before the observer has time to identify every individual.
This makes the species particularly rewarding for observers who learn to interpret sound and movement together.
9.3 Communal Roosting
Collared Aracaris can use cavities communally for sleeping.
The Museo Nacional de Costa Rica documents communal roosting among Costa Rican toucans and emphasizes the importance of cavities as sleeping and nesting resources.
The exact number of individuals occupying a cavity can vary.
9.4 Social Benefits and Costs
Group living can potentially provide advantages such as:
increased vigilance;
information about food;
social learning;
predator detection.
But sociality also creates competition.
Individuals may compete for food, roosting cavities and social position.
A social group is therefore not simply a collection of cooperating birds.
It is a network containing both cooperation and conflict.
10 — Communication
Collared Aracaris are vocal birds.
Their calls are useful for locating groups moving through dense vegetation and canopy.
Because the species frequently occurs in groups, acoustic communication is particularly important for maintaining contact among individuals separated by foliage and branches.
10.1 Vocal Context
Calls can occur during:
group movement;
social interactions;
alarm;
territorial or intergroup encounters.
The exact meaning of individual vocalizations should not be assigned without appropriate behavioural evidence.
10.2 Acoustic Interpretation
For a canopy bird, sound provides a dimension of information that photographs cannot.
An observer may hear several birds before seeing even one.
The absence of a visual observation therefore does not necessarily mean that the species is absent.
Sometimes the first evidence is acoustic.
11 — Reproduction
11.1 Breeding Season
The Costa Rican national catalogue places the principal reproductive period approximately from January to May.
Local timing can vary.
In the Área de Conservación Guanacaste, a nesting sequence was documented beginning in February and continuing through the development of the young into June 2024.
This field record demonstrates why broad breeding seasons should not be treated as rigid calendars.
11.2 Nesting
Pteroglossus torquatus is a cavity-nesting species.
The breeding pair uses an existing cavity, often one originally excavated by another bird such as a woodpecker, and may modify and clean it before breeding.
This is an important ecological dependency.
The aracari does not need to excavate a cavity from the beginning.
Instead, it inherits part of the reproductive architecture produced by another organism or by natural decay.
11.3 A Costa Rican Field Observation
In February 2024, an ACG field observer documented an adult beginning to clean and prepare a cavity in a Guanacaste tree, Enterolobium cyclocarpum, at Estación Biológica Los Almendros, Sector El Hacha.
The observation continued through the nesting period.
Adults were subsequently documented transporting food to a nestling.
This is a particularly valuable example of field natural history because the observer explicitly distinguishes what was directly seen from what could not be determined from the observation.
11.4 Incubation
The exact incubation period should not be presented as established from the ACG observation described above.
The field observer could not directly determine incubation duration or egg colour because the nest cavity was high in the tree.
This is precisely the kind of limitation that belongs in a rigorous Species Note.
Published secondary sources provide estimates for incubation, but those values should be attributed to the relevant literature rather than presented as observations from the Costa Rican nest.
11.5 Cooperative Breeding
The species is associated with cooperative breeding.
Members of the social group other than the primary breeding pair may assist with feeding and care of nestlings.
The Museo Nacional de Costa Rica documents this broader social system among Costa Rican toucans, including group participation in care of young.
This means that reproduction is not necessarily a task performed exclusively by two adults.
The social group can become part of the reproductive system.
12 — Ecological Interactions
12.1 Plant Interactions
The most obvious interaction is with fruiting plants.
The aracari consumes fruits and subsequently moves through the forest.
If seeds remain viable after passage through the digestive system, those movements can contribute to seed dispersal.
The species therefore connects plant reproduction with animal movement.
12.2 Seed Dispersal
The role of P. torquatus in seed dispersal is supported by ecological research.
One study examining variation in tropical seed dispersal incorporated movement and gut-retention data from radio-tracked Collared Aracaris to model dispersal distances.
This demonstrates that the species is not merely capable of moving seeds.
Its movement can be incorporated quantitatively into models of plant recruitment.
12.3 Predation
The species is not exclusively a plant consumer.
ACG records explicitly mention consumption of insects and predation on nests of other birds.
This means that the same individual can participate in plant reproduction through seed dispersal while simultaneously acting as a predator of other animals.
Ecological roles are not mutually exclusive.
12.4 Cavity-Making Organisms
The reproductive ecology of P. torquatus connects it indirectly with cavity-excavating birds.
A woodpecker may excavate a cavity.
The cavity may later be abandoned.
An aracari may modify it and use it for reproduction.
The history of the tree and the activity of another bird therefore become part of the reproductive ecology of P. torquatus.
12.5 Predators
Like other toucans, young and nesting individuals are particularly vulnerable to predators.
The Museo Nacional de Costa Rica emphasizes that birds in nests and juvenile stages are more vulnerable than adults.
This provides one plausible ecological context for group vigilance and social behaviour, although the exact magnitude of predator-detection benefits requires species-specific study.
13 — Ecosystem Function
The Collared Aracari participates in several ecosystem processes simultaneously.
Seed movement
Fruit consumption can result in seeds being transported away from parent plants.
Plant recruitment
Seed dispersal can influence where new plants become established.
Trophic interactions
The species consumes both plant and animal material.
Biological connectivity
Its movements connect different parts of the forest canopy.
Cavity-dependent ecology
Its reproduction depends on structures produced through tree decay and the activities of other cavity-forming organisms.
The species should therefore not be reduced to the label "seed disperser."
Its ecological function is distributed across several processes.
The strongest evidence concerns its participation in seed-dispersal networks and its role as a frugivore.
The magnitude of its contribution to particular plant populations remains context-dependent.
14 — Conservation
14.1 Current Status
Pteroglossus torquatus is currently classified as Least Concern (LC).
Costa Rican conservation information also lists the species as Least Concern and nationally protected. It records a decreasing population trend.
The global LC category should not be interpreted as meaning that the species is unaffected by habitat change.
It means that, at the global scale, current evidence does not place the species within a threatened IUCN category.
14.2 Habitat Loss
Loss of mature vegetation can remove:
fruit resources;
nesting cavities;
roosting cavities;
movement routes.
Because these resources are not necessarily distributed uniformly, habitat quality cannot be assessed simply by measuring the total area of forest remaining.
14.3 Fragmentation
A fragmented landscape can retain forest while reducing connectivity among feeding and nesting resources.
For a mobile canopy bird, the consequences may include changes in movement patterns and access to food.
The exact demographic consequences for P. torquatus in Costa Rica remain an area where additional research would be valuable.
14.4 Secondary Forest
The species' ability to use mature secondary vegetation and some forest-edge environments provides opportunities for persistence outside primary forest.
However, secondary habitat varies enormously in structure.
A young regenerating stand with few large trees and no suitable cavities is not ecologically equivalent to an older secondary forest.
15 — Human–Species Relationship
The Collared Aracari is a familiar bird to many people living or travelling in Costa Rica.
Its colourful appearance makes it visually memorable.
Its social groups make encounters particularly noticeable.
Its presence near gardens and farms can bring forest wildlife into closer proximity to human activity.
The Área de Conservación Guanacaste has documented individuals visiting gardens and farms where bananas or plantains provide food.
Such observations demonstrate that the boundary between "forest" and "human landscape" is not always biologically absolute.
Birds can move across it.
But their ability to use modified habitat should not be mistaken for proof that all modified landscapes are equally suitable.
The question is not simply:
Can the bird be found here?
It is:
Can the landscape provide everything the species needs to survive and reproduce?
16 — Field Interpretation
The Collared Aracari is an excellent species for teaching ecological connectivity.
When you encounter a group, resist the temptation to stop at identification.
Look at the tree.
Is it fruiting?
What plant species is it?
Are the birds feeding?
Are they carrying fruit away?
Then look for cavities.
Are there old trees nearby?
Are woodpecker holes present?
Could the same trees serve both as food sources and reproductive habitat?
Then watch the group.
Are juveniles present?
Are multiple adults feeding them?
Is the group moving together or separating?
Each observation adds another layer.
The bird becomes a way of reading the forest.
17 — Field Notes
For Pteroglossus torquatus, particularly useful field observations include:
Date:Location:Habitat:Elevation:Observer:Group size:Adults:Juveniles:Behaviour:Food item:Plant species:Fruiting condition:Cavity observed:Nest activity:Roosting behaviour:Vocalizations:Associated species:Predator response:Weather:Duration:
A particularly valuable observation would connect the bird with a specific plant:
Six Pteroglossus torquatus feeding on Bactris fruit at 08:30. Two adults subsequently carried fruit toward dense vegetation.
That observation contains substantially more ecological information than simply recording the species.
It links:
plant → fruit → bird → movement → potential seed dispersal.
18 — What We Still Don't Know
Several important questions remain open.
Social ecology
How stable are social groups between breeding seasons?
How long do offspring remain with their natal groups?
What determines whether an individual becomes a helper?
How does group size influence reproductive success?
Food ecology
Which fruiting plants are most important seasonally?
How does fruit availability affect group movements?
How does diet change between breeding and non-breeding periods?
Seed dispersal
Which plant species benefit most from the aracari's movements?
What proportion of consumed seeds remain viable?
How far are seeds typically transported under different landscape conditions?
Reproduction
How frequently are cavities reused?
How limiting is cavity availability?
How does cavity availability affect local population density?
Landscape ecology
How does fragmentation alter movement between feeding and breeding sites?
Can mature secondary forest provide sufficient resources for long-term population persistence?
These are not artificial mysteries.
They are genuine areas where additional field research could improve our understanding of the species.
19 — Interpretive Perspective
The Forest Is Not Raising This Bird Alone
A young Collared Aracari inside a tree cavity may appear to belong to two parents.
A pair selected the cavity.
A pair prepared it.
A pair incubated the eggs.
But the story can become larger after the eggs hatch.
Other members of the social group may participate in feeding and caring for the young.
The nestling therefore becomes part of a social system larger than the breeding pair.
And that social system depends on an ecological system larger still.
The tree provides the cavity.
The forest provides the fruit.
Other birds may have helped create the cavity.
Plants provide the resources that sustain the adults.
The group provides additional caregivers.
Predators create the selective pressure for vigilance.
The young bird is therefore not supported by one relationship.
It is supported by a network.
This is one of the most important lessons a species can teach us.
An organism may be an individual, but its existence is often collective.
The Collared Aracari makes that principle visible.
20 — Scientific Evidence Summary
Topic | Evidence level |
Taxonomic identity | Established |
Costa Rican resident status | Established |
Caribbean and northern Pacific distribution | Established / Documented |
Elevational range approximately 0–1,000 m | Documented |
Forest association | Established |
Use of secondary forest and edges | Documented |
Fruit-dominated diet | Established |
Palm-fruit consumption | Documented |
Insect consumption | Documented |
Nest predation | Documented |
Seed-dispersal role | Documented |
Communal/group behaviour | Documented |
Cavity nesting | Established |
Use of cavities associated with other birds | Documented |
Cooperative breeding | Documented |
2024 ACG nesting sequence | Field observation |
Bactris fruit delivered to nestling | Field observation |
Tabernaemontana seed delivered to nestling | Field observation |
Exact incubation duration from the ACG nest | Unknown from that observation |
Predator-vigilance benefit of group living | Inference |
Exact demographic effect of fragmentation | Limited evidence |
Exact contribution to recruitment of individual plant species | Limited evidence |
21 — The Larger Picture
The Collared Aracari is easy to recognize.
Its red collar, yellow body and oversized bill make identification relatively straightforward.
But recognition is only the beginning.
Look longer and the bird becomes something else.
It becomes a seed carrier.
A fruit consumer.
A predator.
A cavity user.
A member of a social group.
A caregiver.
A participant in the life of trees.
A participant in the movement of plants through the forest.
The bird does not simply occupy the canopy.
It connects different parts of the ecosystem.
A fruiting tree provides food.
The bird carries part of that tree's reproductive future away.
A cavity provides shelter.
The cavity may exist because another bird excavated it years earlier.
A group provides social support.
That group depends on the forest's capacity to provide food and safe places to reproduce.
The individual bird is therefore only the visible portion of a much larger system.
This is why the Collared Aracari is worth observing beyond identification.
When several of them cross the canopy together, we are not simply watching a flock.
We are watching seeds, energy, social information and biological relationships moving through a tropical forest.
And once we learn to see those connections, the forest begins to look less like a collection of organisms and more like what it actually is:
a living network in motion.
References
Primary and Scientific Literature
Research on tropical seed dispersal has incorporated Pteroglossus torquatus into quantitative models of dispersal distance using animal movement and gut-retention data. These studies demonstrate the species' relevance to seed-dispersal ecology.
Costa Rican Institutional Sources
Área de Conservación Guanacaste (ACG). Pteroglossus torquatus (Ramphastidae). Field account by Roster Moraga Medina, including observations of nesting, cavity preparation, parental feeding and food transported to a nestling during 2024.
Museo Nacional de Costa Rica. Tucanes de Costa Rica — Historia Natural. Information concerning behaviour, sociality, reproduction, cavities, predators and interactions among Costa Rican toucans.
Museo Nacional de Costa Rica. Tucanes de Costa Rica — Morfología. Comparative morphology and body size of Costa Rican toucans, including the approximately 41 cm length of P. torquatus.
Museo Nacional de Costa Rica. Tucanes de Costa Rica — Distribución geográfica. Costa Rican distribution and differentiation between P. torquatus and P. frantzii.
Museo Nacional de Costa Rica. Tucanes de Costa Rica — Hábitat. Habitat requirements and importance of trees for food, nesting and roosting.
Museo Nacional de Costa Rica. Catálogo de las especies de Tucanes de Costa Rica. National distribution, elevational range, reproductive period and conservation category.
Comisión Nacional de Seguridad Ambiental / Fauna Decomisada Costa Rica. Pteroglossus torquatus. National protection status and IUCN category.
Conservation
Pteroglossus torquatus is currently classified as Least Concern (LC).
Further Reading
Área de Conservación Guanacaste — Pteroglossus torquatusParticularly valuable for the detailed 2024 Costa Rican nesting observation.
Museo Nacional de Costa Rica — Tucanes de Costa RicaA valuable Costa Rican institutional reference covering morphology, distribution, habitat, natural history and conservation.
Museo Nacional de Costa Rica — Distribución geográficaParticularly useful for distinguishing P. torquatus from P. frantzii in Costa Rica.
Research on tropical seed dispersalUseful for understanding how Collared Aracaris contribute to seed movement through tropical landscapes.
Suggested Related Knowledge
Related Species
Ramphastos sulfuratus — Keel-billed Toucan
Pteroglossus frantzii — Fiery-billed Aracari
Selenidera spectabilis — Yellow-eared Toucanet
Aulacorhynchus prasinus — Northern Emerald-Toucanet
Related Ecological Processes
Seed dispersal
Cooperative breeding
Communal roosting
Secondary cavity nesting
Forest regeneration
Plant–animal interactions
Social ecology
Landscape connectivity
Related Philosophy Essays
Related Field Notes
Collared Aracari nesting observations
Bactris fruit consumption
Communal roosting
Group behaviour
Cavity use
Seed dispersal observations
Cooperative breeding
Field Interpretation in One Sentence
The Collared Aracari reveals that a forest is not merely a collection of organisms, but a network of food, cavities, seeds and social relationships moving through the canopy.


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