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Nicobar megapode

The Nicobar megapode (Megapodius nicobariensis), also known as the Nicobar scrubfowl, is a medium-sized, ground-dwelling bird in the family Megapodiidae, endemic to the Nicobar Islands in the Andaman and Nicobar Islands territory of India.^[1] It measures 37–43 cm in length, features predominantly dark brown plumage with a paler head, a short tail, yellow bill, red facial skin, and large yellow legs and feet adapted for digging and foraging.^[2] This species is distinguished by its unique reproductive strategy, in which pairs construct and maintain large compost mounds of sand, loam, and organic matter near the coast to incubate eggs using solar and microbial heat, with incubation lasting about 72 days and chicks emerging fully independent.^[1] The Nicobar megapode inhabits subtropical and tropical moist lowland forests, including coastal and inland areas up to 600 m elevation, with highest densities in coastal forests and secondary growth.^[1] It is monogamous and forages on the forest floor for invertebrates, seeds, and fallen fruits, often in dense undergrowth of dry deciduous, evergreen, or mixed vegetation such as Areca, Casuarina, and Pandanus.^[2] There are two subspecies: the nominate M. n. nicobariensis in the central Nicobar Islands (e.g., Camorta, Trinkat, Katchall) and M. n. abbotti in the southern islands (e.g., Great Nicobar, Little Nicobar), with an extent of occurrence of approximately 8,700 km² across 13 islands.^[1] Classified as Vulnerable on the IUCN Red List, the species has a global population of 300–600 mature individuals (as of 2023), considered declining following a historical reduction of 66–75% due to the 2004 Indian Ocean tsunami, which destroyed coastal nesting mounds and habitats.^[1]^[3] Ongoing threats include habitat degradation from plantations, the proposed Great Nicobar development project, invasive species, and hunting by local communities; only about 37% of potential coastal breeding habitat remains available.^[4]^[5] Conservation measures encompass protection under Schedule I of India's Wildlife Protection Act (1972), occurrence in national parks like Campbell Bay and Galathea, and targeted monitoring and nest-mound restoration efforts by organizations such as the Wildlife Institute of India.^[1]

Taxonomy

Etymology and discovery

The scientific name Megapodius nicobariensis reflects its characteristic morphology and geographic origin. The genus name Megapodius derives from the Ancient Greek words megas (μέγας), meaning "large," and pous (πούς), meaning "foot," alluding to the bird's notably large and powerful feet adapted for scratching and mound-building.^[6] The specific epithet nicobariensis denotes its endemic occurrence in the Nicobar Islands, following standard binomial nomenclature for species tied to particular localities.^[7] The Nicobar megapode was first scientifically described in 1846 by Edward Blyth, the curator of the museum of the Asiatic Society of Bengal in Calcutta. Blyth's description appeared in the Journal of the Asiatic Society of Bengal (volume 15, part 1, no. 169, p. 52), based on specimens collected from the Nicobar Islands in the Bay of Bengal. These specimens were obtained through early exploratory efforts in the region, including contributions from missionaries like Reverend Jean Pierre Barbe, who documented local fauna amid British colonial expansion into the Andaman and Nicobar territories during the mid-19th century.^[8] Blyth's work was followed by notes he published that year on the Nicobar fauna, highlighting the islands' unique avifauna and distinguishing the megapode from related forms. Early taxonomy of the Nicobar megapode involved confusion with other megapodes in the genus Megapodius, particularly the more widespread dusky megapode (M. freycinet). Some ornithologists initially treated M. nicobariensis as a subspecies of M. freycinet due to overlapping morphological traits and limited specimen availability from the remote Nicobars.^[9] This lumping persisted into the late 20th century, but genetic and morphological studies confirmed its distinct status, leading to its recognition as a full species by the 1970s.^[10]

Systematics and subspecies

The Nicobar megapode (Megapodius nicobariensis) belongs to the family Megapodiidae, a group of mound-building galliform birds characterized by their unique incubation behaviors, and is classified within the genus Megapodius, which comprises the scrubfowl species.^[2] This placement reflects its phylogenetic position as part of a monophyletic clade of Megapodius species, which diverged early within the Megapodiidae family alongside the related genus Eulipoa.^[11] Within this genus, the Nicobar megapode shares close evolutionary ties with other scrubfowls, such as the dusky megapode (M. freycinet), based on analyses of mitochondrial ND2 gene sequences and nuclear rhodopsin intron 1 (RDP1) data that resolve Megapodius as a well-supported group distinct from mound-building megapodes like those in Talegalla and Leipoa.^[11] Two subspecies are currently recognized, reflecting geographic isolation across the Nicobar archipelago. The nominate subspecies M. n. nicobariensis occupies the northern and central Nicobar Islands, including Teressa, Bompoka, Tillanchong, Camorta, Trinkat, Katchall, and Nancowry, while M. n. abbotti (described by Oberholser in 1919) is restricted to the southern islands such as Meroe, Treis, Menchal, Little Nicobar, Great Nicobar, and Kondul.^[1]^[2] These subspecies are separated by approximately 30 km of open sea (Sombrero Channel) between the central and southern island groups, limiting gene flow and contributing to their divergence. M. n. abbotti is diagnosable by its overall darker brown plumage, particularly on the upperparts, underparts, and primaries, compared to the paler nominate form.^[2] Historically, the Nicobar megapode was sometimes treated as conspecific with or a subspecies of the dusky megapode (M. freycinet), reflecting broader lumping of Megapodius taxa in early 20th-century classifications; however, revisions from the 1930s onward, including detailed morphological assessments, elevated it to full species status based on consistent differences in size, vocalizations, and plumage.^[10]^[9] Genetic studies in the 2000s, utilizing mitochondrial DNA sequences, have further confirmed its distinctiveness as a separate lineage within Megapodius, supporting the current taxonomy and highlighting minimal hybridization potential with continental congeners.^[11]

Physical characteristics

Morphology and plumage

The Nicobar megapode (Megapodius nicobariensis) is a medium-sized, stocky bird measuring 37–43 cm in length and weighing 0.6–1.0 kg.^[2]^[12] It possesses large, strong feet equipped with robust claws, well-adapted for scratching and digging in soil and leaf litter.^[13] The bird's overall body structure features strong legs suited for mound-building activities and short, rounded wings indicative of its predominantly terrestrial lifestyle.^[14] The plumage of the adult Nicobar megapode is predominantly dark brown, providing effective camouflage in forested undergrowth. The head is grey with a distinctive rufous crest, bare reddish facial skin, and a yellow bill, contrasting with the otherwise subdued coloration.^[2]^[13] It has a short tail and yellow legs, enhancing its ground-foraging appearance.^[15] Subspecies variations include M. n. abbotti, which is darker in plumage overall, particularly on the upperparts and primaries, compared to the nominate M. n. nicobariensis.^[2] Sexes are similar in size and plumage.^[2]

Sexual dimorphism and juveniles

The Nicobar megapode exhibits no marked sexual dimorphism, with males and females indistinguishable in size and plumage. Both sexes share the same overall dark brown coloration, small grey head, rufous crest, and bare reddish facial skin, making field identification based on morphology challenging.^[2] Juveniles are precocial, hatching fully feathered and capable of immediate locomotion and foraging without parental assistance, though they are noticeably smaller than adults at around 50% of adult body length. Chicks display a quail-like form with brown or buffy body feathers featuring subtle barring patterns for camouflage in leaf litter. They grow rapidly, achieving near-adult size within 2–4 months through independent foraging in dense undergrowth. The species' secretive habits and preference for thick forest cover limit direct observations of juveniles and potential subtle sex differences, with most data derived from opportunistic mound excavations and radio-tracking studies.^[16]^[17]

Distribution and habitat

Geographic range

The Nicobar megapode (Megapodius nicobariensis) is endemic to the Nicobar Islands in the Union Territory of Andaman and Nicobar Islands, India, with no records from the neighboring Andaman Islands or elsewhere.^[1]^[18] The species is non-migratory and confined to this archipelago, where it occupies a total extent of occurrence of approximately 8,700 km².^[1] The species comprises two subspecies with distinct geographic ranges separated by the Sombrero Channel. The nominate subspecies M. n. nicobariensis occurs in the northern and central Nicobar Islands north of the channel, including Tillangchong, Bompoka, Teressa, Camorta, Trinket, Nancowry, and Katchal.^[2]^[19] In contrast, M. n. abbotti is restricted to the southern Nicobar Islands south of the channel, such as Meroe, Treis, Menchal, Little Nicobar, Great Nicobar, and Kondul.^[2]^[19] It is absent from Car Nicobar, Chaura, and Batti Malv.^[4]^[19] Historically, the Nicobar megapode was present on nearly all suitable islands in the archipelago prior to the 2004 Indian Ocean tsunami, with records from a broader set of sites including Megapode Island and Trax Island.^[4]^[2] Post-tsunami assessments confirmed local extirpations on Trax and Megapode Islands, where the latter was fully submerged and the former saw complete loss of suitable habitat.^[4]^[2] Recent surveys have verified ongoing presence across more than 10 islands, with a 2020–2022 study documenting active nesting mounds on 13 islands, including Great Nicobar (64 active mounds), Bompoka (31), and Teressa (35).^[19] These findings indicate persistence in coastal forest habitats within the subspecies' respective ranges, though with varying mound activity levels across sites.^[19]

Habitat requirements

The Nicobar megapode primarily inhabits tropical evergreen forests, secondary growth areas, and coastal scrub vegetation, with the highest densities occurring in littoral and coastal forests. These habitats are characterized by moderately dense canopy cover ranging from 5 to 20 m in height, often dominated by tree species such as Pandanus leram, Macaranga peltata, and Barringtonia asiatica. The bird favors flatter coastal terrains at elevations typically below 300 m, though records extend up to 600 m in some interior forest patches.^[1]^[20] Nesting sites are restricted to sandy coastal areas near beaches, where the bird constructs large incubation mounds using sand, loam, humus, and occasionally coral rubble for structural stability. These mounds are predominantly built within 100–200 m of the shoreline to facilitate access to suitable substrates and maintain optimal decomposition heat for egg incubation.^[1]^[20] For microhabitat requirements, the Nicobar megapode relies on dense understory vegetation in coastal forests for protective cover from predators, allowing it to remain elusive in its ground-dwelling lifestyle. It forages in areas rich with leaf litter and ground debris, scratching the forest floor with its powerful feet to uncover invertebrates, seeds, and fallen fruits, which form the bulk of its diet. This preference for litter-laden undergrowth underscores the bird's adaptation to the nutrient-cycling dynamics of island ecosystems.^[2]^[21] The species exhibits tolerance to periodic natural disturbances in its island environment, such as the impacts from the 2004 tsunami, where populations persisted despite habitat shifts, with nesting mounds observed to relocate closer to shores post-event. This resilience highlights its suitability to dynamic coastal ecosystems, though sustained access to undisturbed litter and understory remains essential for foraging and cover.^[22]

Behavior and ecology

Foraging and diet

The Nicobar megapode maintains an omnivorous diet dominated by invertebrates and plant matter. Invertebrates form a core component, including insects such as beetle larvae, worms, crustaceans, molluscs like snails (Nerita polita and Littorina spp.), and occasionally small reptiles. Plant-based foods consist of seeds (particularly from Macaranga species), fallen fruits, and other debris. Observations confirm that the species ingests small stones or grit to facilitate digestion of hard-shelled items and fibrous vegetation.^[23]^[24] Foraging occurs exclusively on the ground in dense forest leaf litter and soil, where the bird uses its large, powerful feet to scratch and rake through debris in search of food. This technique is efficient for uncovering buried invertebrates and scattered seeds, while also contributing to ecosystem services like soil aeration and seed dispersal. The activity is primarily diurnal, though crepuscular foraging at dawn and dusk has been noted, aligning with the bird's secretive, ground-dwelling lifestyle.^[24]^[25] Limited data suggest year-round availability of insects supports consistent protein intake, with fruits and seeds potentially more abundant during the wet season due to increased forest productivity, though comprehensive seasonal diet analyses remain scarce. Foraging demands are substantial, supporting the high energy requirements for mound construction and maintenance in reproduction.^[21] The Nicobar megapode leads a secretive, predominantly terrestrial lifestyle, spending much of its time concealed in the dense undergrowth of coastal forests where it forages and navigates with caution. It exhibits crepuscular activity patterns, becoming most active at dawn and dusk, during which periods its calls echo through the habitat. At night, individuals roost in low vegetation, such as shrubs or low trees near the forest floor, to avoid predators.^[26]^[19] Socially, the species is organized around solitary individuals or monogamous pairs that maintain loose, overlapping territories averaging 0.81 hectares, often centered on communal nesting mounds rather than rigidly defined areas. These territories are defended year-round by both pair members through displays and confrontations with neighbors or intruders, though unpaired birds exhibit less vocal and aggressive behavior. Flocking is minimal, with birds rarely forming groups larger than pairs, except during brief aggregations of up to eight individuals at disputed boundaries or shared sites.^[27]^[28] Vocalizations play a key role in communication and territorial maintenance, consisting of low-pitched clucks, grunts, and duets that advertise presence and deter rivals, particularly at night or during aggressive encounters. Alarm calls, including distress notes, are emitted when threatened, aiding in evasion or alerting nearby individuals. Interactions outside of these displays are limited, but occasional aggression—such as chases, fights, or physical confrontations—arises at territory edges or over resources like food patches.^[13]^[28]

Reproduction

Mating system

The Nicobar megapode maintains a primarily monogamous mating system, characterized by pair bonds that facilitate access to and defense of incubation mounds, though these bonds are not always lifelong and break-ups occur between or within breeding seasons. Extra-pair copulations are documented, often arising during territorial disputes at shared mounds, adding complexity to the otherwise pair-based structure.^[17] Pair formation typically involves solitary individuals attaching to existing mounds or constructing new ones to join or establish partnerships, with mate selection influenced by the quality and defensibility of the mound and surrounding territory. Courtship between established pairs is minimal or absent, lacking specialized displays; instead, bonds are reinforced through female-initiated vocal duets, joint territorial patrols, and cooperative behaviors such as mound excavation near nest sites. The slight sexual dimorphism, including the female's grey head and rufous crest, likely aids in individual recognition during these vocal and territorial interactions.^[17] Breeding occurs more or less year-round, but activity peaks during the dry season from November to April, with egg-laying concentrated from January to June based on field observations.^[2]^[17]

Nesting and incubation

The Nicobar megapode (Megapodius nicobariensis) constructs large incubation mounds primarily from sand, loam, and decomposing vegetation, with heights typically ranging from 1 to 3 meters and volumes varying from less than 1 m³ to over 10 m³. These mounds are often built and maintained communally by multiple pairs, though individual pairs may also construct and tend their own; unpaired males frequently initiate mound building to attract females. Mound construction occurs year-round in coastal habitats, where the sandy-loam substrate facilitates heat retention, and larger mounds generally provide more stable internal conditions for incubation.^[1]^[28]^[29] Females lay 2–5 white eggs per clutch directly into a chamber within the mound, with eggs deposited at intervals of several days, such as 9 days in observed cases; a single mound may contain up to 20 eggs from multiple females due to communal use. The eggs are incubated externally for 70–80 days, with an average period of about 75 days, relying on heat generated primarily through microbial decomposition of organic matter in the mound, supplemented by solar warming. Incubation success depends on mound size and location, as larger structures maintain more consistent temperatures.^[2]^[30]^[31] Unlike most birds, the Nicobar megapode provides no brooding; instead, both parents regulate incubation by adding or removing mound material to sustain an optimal internal temperature of 33–37°C, with observed averages around 32–34°C near hatching. This thermoregulation is critical, as fluctuations can affect embryonic development, and pairs may contribute to multiple mounds annually to maximize reproductive output. A hierarchy among sharing pairs ensures orderly egg-laying access.^[22]^[28]^[1]

Conservation status

Population estimates

The population of the Nicobar megapode (Megapodius nicobariensis) was estimated at 2,322–4,065 breeding pairs, or approximately 4,500–8,000 mature individuals, prior to the 2004 Indian Ocean tsunami, based on counts of 1,161 active incubation mounds across the Nicobar Islands.^[1]^[22] Following the tsunami, surveys in 2006 recorded 394 active mounds, indicating 394–788 breeding pairs and a 66–75% overall decline from pre-tsunami levels.^[1]^[22] Subsequent assessments in 2009–2011 identified 376–752 breeding pairs, equivalent to 750–1,500 mature individuals, primarily through mound counts suggesting population stability at the reduced level.^[1] Surveys from 2020–2022 across 13 islands documented 162 active mounds, yielding an estimate of 162–324 breeding pairs (assuming 1–2 pairs per mound) and signaling a further 59% decline since 2006.^[3] Earlier work from 2015–2018 similarly reported 149 active mounds, aligning with detections of around 300 individuals.^[32] A 2024–2025 survey by the Wildlife Institute of India across 16 islands estimated 151–302 breeding pairs in the central Nicobar group (a 54% increase from 2006 levels) and 277–554 pairs in the southern group (a 7% decline from 2006), indicating an overall 8.4% increase from post-tsunami estimates but remaining below pre-tsunami numbers.^[33] Population assessments rely on line transect surveys of coastal habitats, typically within 100–600 m of shorelines, to enumerate active incubation mounds, with 1–2 breeding pairs assumed per mound post-tsunami due to reduced nesting success.^[1]^[22]^[3] These methods, covering over 150 km per survey effort, also incorporate point counts and camera traps for direct sightings in Nicobar reserves.^[32] Regarding subspecies, the southern form M. n. abbotti (on Great Nicobar and Little Nicobar) supported higher post-2004 numbers at 297–594 breeding pairs, compared to 97–194 pairs for the northern/central M. n. nicobariensis (Nancowry group).^[1]^[22] Recent data indicate persistent disparities, with Great Nicobar holding 64 active mounds versus 31 on Bompoka in the central group, though both subspecies have experienced severe localized declines exceeding 95% on islands like Little Nicobar and Trinket.^[3] The species is classified as Vulnerable by the IUCN Red List owing to its small and fragmented population.^[1]

Threats and protection

The Nicobar megapode faces multiple anthropogenic and natural threats that have contributed to significant population declines. The 2004 Indian Ocean tsunami devastated coastal habitats and nesting sites across several islands, resulting in an estimated 66-75% reduction in the species' population.^[1] Ongoing habitat loss remains the primary threat, driven by conversion of coastal forests to coconut, banana, and cashew plantations, as well as rice paddies, with approximately 5.5% of forest cover lost between 2000 and 2019.^[1]^[34] Proposed infrastructure developments, including the Great Nicobar Integrated Development Plan announced in 2020, exacerbate this risk by threatening up to 59% of known nesting mounds through land clearance and associated disturbances in key areas like Galathea Bay.^[35] Hunting and egg collection by local communities, often via snaring or shooting, have intensified post-tsunami, posing direct mortality risks.^[1] Invasive species such as feral cats and dogs further endanger the ground-dwelling bird by preying on adults and chicks.^[36] Climate change compounds these pressures, with projected sea-level rise potentially inundating low-lying nesting beaches and increasing vulnerability to extreme weather events.^[37] Legal protections afford the Nicobar megapode the highest level of safeguarding in India, classified as Schedule I under the Wildlife (Protection) Act of 1972, which prohibits hunting, trade, and habitat disturbance.^[38] The species occurs within protected areas, including Great Nicobar National Park, Campbell Bay National Park, and three wildlife sanctuaries, covering portions of its range and restricting development activities.^[1] Monitoring efforts support conservation, with post-tsunami surveys conducted from 2009 to 2011 to assess recovery, followed by updated assessments in 2023 under national programs to track distribution and population trends.^[1]^[38] Active conservation initiatives include community awareness campaigns to reduce hunting and promote habitat stewardship among Nicobarese residents, as outlined in species-specific action plans.^[9] Anti-poaching patrols are enforced within protected areas to deter illegal activities, while proposals for habitat restoration focus on replanting native coastal forests to mitigate losses from agriculture and development.^[1] In response to the Great Nicobar project, long-term monitoring committees were established in 2025 with allocated funding to evaluate impacts and implement mitigation, though their effectiveness remains under scrutiny.^[39] These measures aim to stabilize the vulnerable population, but escalating development pressures as of late 2025 continue to challenge long-term survival.^[35]

References

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Features a dark brown plumage with a slight gloss, giving it a distinctive appearance in the sun. •. Has a red facial skin patch and yellow legs, providing a ...
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### Summary of Nicobar Megapode Behavior from the Memoir
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