Umbrellabird

Umbrellabirds (genus Cephalopterus) comprise three species of large, black-plumaged cotingas (family Cotingidae) endemic to the humid rainforests of Central and South America, characterized by a prominent forward-folding crest resembling an umbrella and, in males of most species, an inflatable wattle or throat sac used in courtship displays.^[1]^[2] The long-wattled umbrellabird (C. penduliger) inhabits premontane and cloud forests on the Pacific slopes of the Andes in Colombia and Ecuador, where males gather at leks to produce deep, resonant calls and perform ritualized displays; it feeds mainly on fruits supplemented by insects and small vertebrates.^[3]^[4] The Amazonian umbrellabird (C. ornatus) occupies lowland and montane forests of the Amazon basin, while the bare-necked umbrellabird (C. glabricollis) is restricted to montane forests in Costa Rica and Panama, both species sharing similar frugivorous diets and lek-based mating systems.^[1]^[5] All umbrellabirds face population declines due to deforestation and habitat fragmentation, with the long-wattled classified as vulnerable and the bare-necked as endangered by the IUCN, underscoring their reliance on intact forest canopies for foraging and breeding.^[3]^[6]^[7]

Taxonomy and Systematics

Etymology and Classification

The common name "umbrellabird" refers to the prominent, erectile crest on the male's head, which expands to resemble an open umbrella during displays.^[8] The genus name Cephalopterus derives from New Latin roots cephal- (from Greek kephalē, head) and -pterus (from Greek pterós, winged), alluding to the feather crest extending from the head like wings.^[9] Étienne Geoffroy Saint-Hilaire established the genus in 1809.^[10] Umbrellabirds belong to the genus Cephalopterus in the family Cotingidae, a group of approximately 65 species of primarily frugivorous birds native to Neotropical forests.^[11] The family Cotingidae is placed within the order Passeriformes, specifically the suborder Tyranni (suboscine passerines), distinguished by their syringeal anatomy lacking the complex muscles of oscine songbirds.^[12]^[11] This classification reflects their evolutionary position among New World perching birds adapted to arboreal, fruit-based diets.^[12]

Phylogenetic Position

The genus Cephalopterus, comprising the three species of umbrellabirds (C. glabricollis, C. penduliger, and C. ornatus), is monophyletic based on molecular analyses of nuclear and mitochondrial DNA sequences.^[13] Cotingidae itself forms a monophyletic family within the Tyrannida clade of suboscine passerines (Tyranni), characterized by distinct vocal and morphological traits distinguishing it from oscine passerines.^[14] Within Cotingidae, Cephalopterus nests in the derived "core cotingas" clade, specifically the fruitcrow subgroup (sometimes termed Cephalopterinae), which includes genera such as Perissocephalus, Pyroderus, Querula, and Haematoderus.^[13] This fruitcrow clade is strongly supported as monophyletic (Bayesian posterior probabilities of 0.99–1.00), with Cephalopterus most closely related to Perissocephalus tricolor (the capuchinbird), forming a subclade sister to one containing Haematoderus, Querula, and Pyroderus.^[13] These relationships derive from Bayesian phylogenetic inferences using datasets of over 2,100 base pairs (nuclear introns and cytochrome b) in earlier studies and up to ~7,500 base pairs across six loci (including myoglobin intron, G3PDH, RAG-1/2, cytochrome b, and ND2) in more comprehensive analyses.^[13] The positioning reflects shared frugivorous adaptations and lekking behaviors among these taxa, though intergeneric relationships show moderate support in some nodes (e.g., posterior probability 0.84 for Cephalopterus–Perissocephalus).

Recognized Species

The genus Cephalopterus includes three recognized species of umbrellabirds, all classified within the family Cotingidae and native to Neotropical forests. These species are distinguished by morphological traits such as erectile crests and, in males of two species, prominent throat wattles, adapted for display in lekking behaviors.^[1]^[15]

Species	Distribution	Elevation Range	IUCN Status	Key Threats
C. penduliger	Pacific Andes, Colombia–Ecuador	500–1,800 m	Vulnerable	Deforestation, fragmentation
C. ornatus	Amazon basin, South America	<1,000 m	Least Concern	Localized habitat loss
C. glabricollis	Talamanca mountains, Costa Rica–Panama	1,400–1,900 m (breeding)	Endangered	Logging, agriculture

No subspecies are formally recognized within these species, though genetic studies indicate limited divergence consistent with allopatric distributions.^[20]^[21]^[22]

Physical Characteristics

Morphology and Plumage

Umbrellabirds (genus Cephalopterus) possess a robust, stocky build characteristic of many cotingids, with body lengths ranging from 35 to 50 cm and weights up to 500 g in the largest individuals. They feature broad, rounded wings suited for maneuvering through dense forest canopies, strong legs with acute claws for perching, and a heavy, hooked bill adapted for consuming fruit and seizing prey.^[2]^[23] Adult plumage across species is predominantly glossy black, often exhibiting subtle metallic blue or purple iridescence on the head, upperparts, and sometimes the throat structures, which enhances visibility in low-light understories. The defining morphological trait is an erectile crest of elongated, stiff feathers atop the head, capable of fanning into an umbrella-like hood; these feathers curl outward at the tips in species like the long-wattled umbrellabird (C. penduliger), forming a brush-like fringe.^[15]^[24] In the long-wattled umbrellabird, the crest covers the crown and nape, while the Amazonian umbrellabird (C. ornatus) displays a similar but proportionally shorter structure, with plumage lacking white markings that distinguish it from sympatric woodpeckers during flight. The bare-necked umbrellabird (C. glabricollis) features a more pronounced "Mohican"-style crest extending over the bill, paired with a largely unfeathered, scarlet throat and neck adorned with filamentous tassels rather than a fully feathered wattle. Juveniles generally show duller, less glossy black feathers without full crest development.^[17]^[19]^[25] Throat appendages vary significantly: the long-wattled species has a pendulous wattle up to 35 cm long, covered in short, scaly feathers that inflate to resemble a pine cone during displays, whereas the Amazonian wattle is shorter and triangular with bare patches, and the bare-necked lacks extensive feathering on its inflatable red throat sac. These structures, integral to plumage and morphology, aid in thermoregulation and signaling within humid tropical environments.^[26]^[3]^[27]

Sexual Dimorphism

Umbrellabirds (Cephalopterus spp.) exhibit pronounced sexual dimorphism, with males larger and more ornate than females, reflecting adaptations for lek-based mating displays where males compete for female attention through visual and acoustic signals.^[28] Males possess exaggerated crests forming an umbrella-like hood, glossy black plumage, and species-specific throat structures such as long feather wattles or bare inflatable sacs, which females lack or have in reduced form.^[29] Females display duller, often browner plumage for camouflage, smaller crests, and no prominent display ornaments, prioritizing reproductive investment over costly signaling.^[29] This dimorphism supports female choice based on male quality indicators like crest size and territorial aggression.^[28] In the long-wattled umbrellabird (C. penduliger), males measure 40–42 cm in length with a prominent erectile crest and a controllable feather wattle extending up to 30 cm, absent or minimal in females who reach 35–37 cm.^[30] Male body mass averages 338 g, exceeding that of females, whose plumage is less glossy and crested for reduced visibility during solitary foraging and nesting.^[30] The Amazonian umbrellabird (C. ornatus) shows similar patterns, with males larger (48–55 cm) and featuring a more prominent crest and inflatable throat wattle for booming displays, while females are approximately two-thirds the male size with subdued ornaments.^[31]^[32] For the bare-necked umbrellabird (C. glabricollis), males attain 41 cm in length and 450 g, with a scarlet bare throat sac and puffy crest inflated during courtship, contrasting females at 36 cm and 320 g who lack the sac and have a shorter, less prominent crest atop blackish plumage.^[5]^[33] This size disparity—males 14% longer and 41% heavier—aligns with male-biased dimorphism in lekking cotingids, enhancing display efficacy without corresponding female traits.^[5]

Size and Adaptations

Umbrellabirds (genus Cephalopterus) rank among the largest members of the Cotingidae family, with body lengths ranging from 35 to 55 cm and weights from 320 to 571 g, varying by species and sex. Males are consistently larger than females across all species, reflecting sexual dimorphism that supports their roles in elaborate courtship displays. The Amazonian umbrellabird (C. ornatus) attains the greatest dimensions, with males measuring 48–55 cm in length and weighing 480–571 g, while females average 41–44 cm.^[25] ^[34] In contrast, the bare-necked umbrellabird (C. glabricollis) is smaller, with males at 41 cm and 450 g, and females at 36 cm and 320 g.^[5] ^[35] The long-wattled umbrellabird (C. penduliger) falls between these, with males reaching 41 cm and approximately 338 g, and females around 36 cm.^[30] These sizes equip umbrellabirds for life in the forest canopy, where their robust bodies facilitate perching on branches while foraging for fruit.^[2] Key morphological adaptations center on reproductive displays, driven by sexual selection in lekking systems where males compete visually and acoustically for females. The erectile crest of forward-curving feathers atop the head, prominent in males, is raised during courtship to enlarge apparent body size and signal fitness.^[3] In the long-wattled species, males possess an extensible, inflatable throat wattle up to 30 cm long, which they swing and pulsate to amplify deep booming calls, enhancing communication in dense humid forests.^[3] These traits, absent or reduced in females, impose no evident survival costs in their frugivorous niche but confer mating advantages, as evidenced by aggregation at leks for synchronized displays.^[36] The bare-necked species lacks a pronounced wattle but retains the crest for similar signaling purposes.^[37] Additionally, their heavy, hooked bills are adapted for cracking large fruits, supporting a diet of canopy-available resources.^[38]

Distribution and Habitat

Geographic Range

The genus Cephalopterus comprises three species of umbrellabirds, each with restricted and largely non-overlapping distributions confined to humid forests of Central and northwestern South America.^[20]^[21]^[22] The long-wattled umbrellabird (C. penduliger) inhabits the Pacific slope and adjacent lowlands of southwestern Colombia, from Chocó Department to Nariño Department, and western Ecuador, from Esmeraldas Province south to El Oro Province.^[6]^[20] Its range follows the western Andean foothills, typically at elevations of 500–2,200 meters, though it occasionally descends to sea level.^[15] The Amazonian umbrellabird (C. ornatus) occupies the largest area, encompassing much of the Amazon Basin and adjacent Andean foothills across southern Venezuela (including upper and middle Orinoco drainage), Guyana, northern Bolivia, Amazonian Brazil, Colombia (south to Guainía), Ecuador, and Peru.^[17]^[21] This species ranges from lowlands up to 1,500 meters elevation, with local populations near tepui bases in southern Venezuela.^[17] The bare-necked umbrellabird (C. glabricollis) is limited to the humid highlands of Costa Rica and extreme western Panama, primarily in the Talamancan montane forests at elevations of 1,400–2,500 meters.^[22]^[39] Rare vagrant records extend to southern Nicaragua, but the core population remains within protected areas like Monteverde and La Amistad reserves.^[39]

Habitat Preferences

The long-wattled umbrellabird (Cephalopterus penduliger) inhabits humid premontane and cloud forests on the Pacific slopes of the Andes, primarily at elevations from 140 to 1,800 meters, where it utilizes the mid- to upper canopy of tall trees for foraging and movement.^[26]^[3] These forests feature high rainfall and dense vegetation, supporting the large fruit crops essential to its diet, with the species avoiding fragmented or secondary growth areas that lack sufficient canopy connectivity.^[15] The Amazonian umbrellabird (C. ornatus) occupies two distinct habitat regimes: seasonally flooded várzea forests along riverbanks and on islands in the Amazon basin lowlands, and subtropical moist montane forests in the eastern Andean foothills of Ecuador and northern Peru, up to about 1,500 meters.^[17]^[16] It prefers areas with emergent trees and proximity to watercourses, which facilitate seed dispersal and provide abundant fruiting trees during non-breeding periods, though it largely shuns dry or heavily disturbed woodlands.^[16] The bare-necked umbrellabird (C. glabricollis) is restricted to subtropical moist lowland and lower montane forests in the Caribbean versant of Costa Rica and western Panama, typically between 100 and 1,000 meters, favoring primary old-growth stands with intact subcanopy and upper understory layers over secondary regrowth.^[18]^[5] These habitats include high-rainfall regions with large-diameter trees suitable for leks and nesting, and the species shows seasonal elevational shifts, descending to lowlands outside breeding periods from March to June when montane fruit availability peaks.^[40] Across species, umbrellabirds exhibit a marked preference for undisturbed, humid tropical forests with continuous canopy cover, as canopy gaps from logging or agriculture disrupt their arboreal lifestyle and reduce fruit abundance, leading to localized population declines where such degradation exceeds 30% of available habitat.^[4]^[41]

Seasonal Movements

The bare-necked umbrellabird (Cephalopterus glabricollis) performs annual altitudinal migration tied to fruit productivity, spending most of the year in lowlands at 100–500 m elevation and ascending to 800–1,500 m or higher during the breeding season from March to June.^[42]^[43] These movements occur in groups that include breeding males, reflecting a strategy to track seasonal fruit abundance across elevational gradients in Costa Rica and western Panama.^[43] In contrast, the long-wattled umbrellabird (C. penduliger) is largely resident within humid premontane and cloud forests on Pacific slopes, with only occasional, non-seasonal wanderings from highland habitats (above 1,000 m) into adjacent lowlands below 500 m.^[15] Some populations show sedentary behavior year-round, though local altitudinal shifts may occur in response to food resources without a fixed seasonal pattern.^[15] The Amazonian umbrellabird (C. ornatus) shows no pronounced seasonal movements, remaining sedentary in riverine and floodplain forests of the Amazon basin and Orinoco region, as well as Andean foothills up to 1,500 m, where it exploits stable fruiting cycles without elevational migration.^[17] Across the genus Cephalopterus, such altitudinal patterns underscore frugivory's influence on distribution, though habitat fragmentation increasingly constrains these shifts.^[42]

Ecology and Behavior

Diet and Foraging

Umbrellabirds of the genus Cephalopterus are predominantly frugivorous, with their diet consisting mainly of fruits and berries from plant families including Marcgraviaceae, Cecropiaceae, Arecaceae, Lauraceae, Annonaceae, and Urticaceae.^[44]^[5] They regurgitate large seeds and pits, facilitating seed dispersal across forest habitats, particularly for palms like Oenocarpus in the case of the long-wattled umbrellabird (C. penduliger).^[4]^[45] While fruits dominate adult diets, animal matter supplements foraging, including arthropods such as insects, and small vertebrates like lizards, frogs, and snakes, especially when provisioning nestlings.^[15]^[46] For the bare-necked umbrellabird (C. glabricollis), observations include opportunistic foraging at army ant swarms to capture flushed insects, though such behavior appears infrequent.^[47]^[48] The Amazonian umbrellabird (C. ornatus) targets fruits and arthropods in the upper forest canopy.^[17] Foraging occurs primarily in the forest canopy and subcanopy, where individuals hop between branches or perch to pluck items, often singly, in pairs, or small groups; they exhibit secretive habits and are more frequently detected by vocalizations than visual sightings during feeding.^[49]^[17] Flight between foraging sites is lumbering and direct, reflecting their large size and adaptation to dense humid forests.^[17] Seasonal shifts may emphasize protein-rich animal prey during breeding to support nestling growth, reverting to fruit-heavy intake otherwise.^[4]^[46] Umbrellabirds (Cephalopterus spp.) exhibit a classic lek mating system, characterized by males aggregating at communal display sites to perform vocal and visual courtship displays for female mate selection, with no provision of resources or paternal care.^[50] This polygynous or promiscuous structure relies on female choice based on male display quality, typical of many cotingids.^[51] In the long-wattled umbrellabird (C. penduliger), males form leks of 5–15 individuals on ridge tops in Chocóan montane forests, defending small territories of approximately 25 m² within arenas spanning about 1 ha; peak activity occurs from August to February, with displays concentrated in early mornings and late afternoons.^[50] Males produce deep booming calls audible up to 500 m and extend their prominent wattles during displays, while competing via displacements; floater males circulate between leks without fixed territories.^[3] ^[50] Females visit leks infrequently for copulation before departing to nest solitarily.^[50] The bare-necked umbrellabird (C. glabricollis) shows analogous lek behavior, with males in small groups performing throaty calls and cat-like screeches during rival chases, escalating to frenetic wing-spreading and displacements upon female presence.^[52] ^[53] Similarly, the Amazonian umbrellabird (C. ornatus) engages in lekking, often in exploded formations where males display in dispersed canopy sites to attract observing females.^[54] Socially, umbrellabirds are predominantly solitary outside breeding periods, foraging independently in forest canopies, though males may form coordinated off-lek groups during high lek activity to synchronize foraging and return to display sites, potentially enhancing mating success.^[28] Post-mating, females handle all nesting and chick-rearing alone, building platform nests and provisioning with fruit.^[3] This minimal social structure aligns with their frugivorous ecology and altitudinal movements to leks.^[50]

Vocalizations and Displays

Males of the genus Cephalopterus engage in lekking behavior, congregating at traditional display sites in the forest canopy to perform courtship rituals that attract females, who visit briefly to select mates without forming pair bonds.^[15]^[3] Displays typically occur year-round but intensify during breeding seasons, involving physical posturing and vocalizations amplified by throat structures or wattles.^[55] In the long-wattled umbrellabird (C. penduliger), males raise their crests, inflate and swing the elongated throat wattle—which can double in length during displays—and emit a resonant, foghorn-like "boooh" or mooing call, often accompanied by wing-snapping noises.^[15]^[3] These vocalizations, described as emphatic "aaugh" bursts or low-frequency moos, carry far through humid forests and are produced from perches 20–30 m high at leks, where 4–10 males may gather.^[55] The wattle functions partly as a resonance chamber, enhancing the deep, booming quality akin to a distant foghorn.^[3] The Amazonian umbrellabird (C. ornatus) features similar lek displays along rivers or forest edges, with males extending wattles and delivering deep, melodious booms or low-pitched moos that project ventriloquially over distances.^[56]^[17] Both sexes produce churring or growling notes, but male advertising calls dominate leks.^[17] For the bare-necked umbrellabird (C. glabricollis), displays occur in exploded leks spanning 2–4 km² at 750–1,300 m elevation, peaking from mid-March to mid-May. Males execute dawn "double-hoots"—low-frequency hoots carrying hundreds of meters—while bowing and throwing back their heads from canopy perches, followed by daytime "hoot-growl" or "inflate-hum" calls every 15–20 minutes during perch-to-perch flights within 50 × 100 m territories.^[49] Intensity escalates with female presence, prompting frenetic movements among 4–5 males per group.^[49]

Reproduction

Umbrellabirds (genus Cephalopterus) employ a lek mating system, in which males congregate at traditional display sites to perform vocal and visual courtship displays, including crest erection and booming calls, while females visit leks to select mates without subsequent male involvement in parental care.^[50] This system is documented across species, with leks often located in canopy gaps or forest edges during breeding periods aligned with local rainy seasons.^[51] Breeding phenology varies by species and region; for the long-wattled umbrellabird (C. penduliger) in southwestern Ecuador, nest initiation occurs from late December to January, with eggs hatching in February.^[57] Limited records for the Amazonian umbrellabird (C. ornatus) indicate laying in July in Mato Grosso, Brazil, while the bare-necked umbrellabird (C. glabricollis) breeds from March to June in Costa Rica and Panama.^[17]^[58] Nesting is performed solely by females, who construct bulky platform nests 5–25 meters above ground in the forest canopy or subcanopy, using interwoven twigs, lined with moss, leaves, and fibers.^[46] Clutch size consists of a single white or cream-colored egg, measuring approximately 5 cm in length.^[4] Incubation, lasting 27–28 days, is conducted exclusively by the female, with no male attendance observed at the nest site.^[59] Females cover the egg for 65–88% of daylight hours, averaging 79%, during bouts of 10–40 minutes, leaving periodically to forage.^[46] Nestlings, also cared for only by the female, remain in the nest for about 30–33 days before fledging; provisioning includes large arthropods, small vertebrates such as lizards, frogs, and snakes, and possibly fruit pulp.^[15]^[57] Post-fledging dependence extends for several weeks, with females continuing to feed juveniles. Data on reproductive success remain sparse due to the species' elusive nature and remote habitats, but predation and nest abandonment pose significant risks.^[60]

Conservation and Threats

Population Status

The long-wattled umbrellabird (Cephalopterus penduliger) is classified as Vulnerable by the IUCN, with an estimated global population of 7,500–15,000 mature individuals and a decreasing trend inferred from ongoing habitat loss and fragmentation.^[6] Earlier estimates placed the total at 10,000–20,000 individuals, but recent assessments emphasize the mature breeding segment as more critical for viability.^[15] In Colombia, the population is particularly low, comprising fewer than 2,500 individuals.^[15] The bare-necked umbrellabird (Cephalopterus glabricollis) holds Endangered status under the IUCN, with a global population estimated at fewer than 2,500 mature individuals and a continuing decline driven by deforestation in its restricted range across Costa Rica and Panama.^[18] Breeding populations in Costa Rican Important Bird Areas were assessed at 740–1,430 mature individuals in 2007, while broader estimates for Costa Rica suggest 1,900–7,100 birds, though these figures likely overestimate the fragmented global total due to survey limitations in Panama.^[61] In contrast, the Amazonian umbrellabird (Cephalopterus ornatus) is listed as Least Concern by the IUCN, reflecting a more widespread distribution in the Amazon basin, though its population size remains unquantified and is described as uncommon with patchy occurrence.^[16] No specific numerical estimates are available, but its larger range buffers it against the severe declines seen in congeners.^[16]

Primary Threats

The primary threats to umbrellabirds in the genus Cephalopterus stem from anthropogenic activities, with habitat destruction being the dominant factor across species. Deforestation for agricultural expansion, including cattle ranching, palm oil and banana plantations, and logging, has fragmented and reduced the humid and montane forests essential to their survival, leading to population declines. In western Ecuador, for example, the vast majority of original lowland forests have been lost, severely impacting the long-wattled umbrellabird (C. penduliger), which relies on contiguous canopy habitats for foraging and leks.^[6]^[62] Hunting and trapping exacerbate these pressures, particularly for the long-wattled and bare-necked umbrellabirds (C. glabricollis), where birds are targeted for bushmeat or the illegal pet trade due to their conspicuous size and displays. These activities are facilitated by increased human access via roads and settlements in formerly remote areas.^[6]^[18]^[63] Emerging risks include climate change, which may alter forest microclimates and fruit availability critical to their diet, though quantitative impacts remain understudied relative to direct land-use changes. The Amazonian umbrellabird (C. ornatus) faces lower immediate threats but shares vulnerability to logging in its range.^[64]^[17]

Conservation Efforts and Outcomes

Efforts to conserve umbrellabirds emphasize habitat protection, ecological research, and community engagement to counter deforestation and fragmentation. For the long-wattled umbrellabird (Cephalopterus penduliger), designated Vulnerable by the IUCN in assessments up to 2023, key initiatives include designation of protected areas such as Colombia's Farallones de Cali National Park and Ecuador's Cotacachi-Cayapas, Mache-Chindul, and Cotacachi-Cayapas Ecological Reserves, which encompass portions of its foothill and montane forest range.^[6] Community-led projects, like the 20-hectare Recinto 23 de Junio reserve in Ecuador established around 2010, involve local farmers in monitoring leks and sustainable land use to preserve breeding sites.^[65] Research on nesting biology and lek dynamics, including a 2003 study documenting solitary female nests in fragmented habitats, has informed targeted anti-hunting and reforestation campaigns in the Chocó region.^[4] The bare-necked umbrellabird (C. glabricollis), also Vulnerable per IUCN evaluations through 2023 with a restricted range in Costa Rica and Panama, benefits from the Umbrellabird Action Network (UCAN), founded to safeguard lowland-to-highland corridors in the Tilarán Mountains since the early 2010s.^[7] A 2005 study on altitudinal migrations revealed seasonal shifts from breeding elevations above 800 meters to foraging lowlands below 800 meters, prompting recommendations for reforestation and connectivity projects to link fragmented forests.^[66] UCAN's site-specific protections, including anti-deforestation patrols, have stabilized some leks, though broader lowland clearance persists.^[67] In contrast, the Amazonian umbrellabird (C. ornatus), rated Least Concern by the IUCN due to its extensive Amazon Basin range exceeding 1 million square kilometers, faces fewer targeted interventions but gains from regional initiatives like Ecuador's Chocó forest safeguards west of Cotacachi-Cayapas Reserve.^[16] Local efforts, such as those by Amarun Pacha in Peru, monitor lek losses to agriculture since the 2010s, advocating sustainable alternatives to economic pressures driving habitat conversion.^[68] Outcomes indicate limited reversal of declines: the long-wattled species' global population, estimated below 15,000 mature individuals in 2021 assessments, continues decreasing at rates exceeding 10% per decade from ongoing fragmentation, despite protected area coverage of less than 20% of suitable habitat.^[3] Bare-necked populations, numbering fewer than 10,000, show localized stability in monitored Costa Rican sites but overall vulnerability to clearance, with no quantified recovery.^[18] Amazonian numbers remain stable, though isolated lek extirpations highlight risks without expanded monitoring.^[16] These efforts underscore the need for scaled-up enforcement and restoration, as current measures mitigate but do not halt primary threats.^[69]