Treeswift
Treeswifts (family Hemiprocnidae) are a small group of aerial birds in the order Apodiformes, closely related to true swifts but distinguished by their ability to perch and their ornate crests.[1] The family comprises a single genus, Hemiprocne, with four extant species: the grey-rumped treeswift (H. longipennis), crested treeswift (H. coronata), whiskered treeswift (H. comata), and moustached treeswift (H. mystacea).[1] These birds measure 15–31 cm in length, featuring slender bodies, long forked tails, and elongated wingtips that cross when folded, adaptations suited to their agile flight.[1] Native to tropical regions of the Oriental and parts of the Australian biogeographic realms, treeswifts range from India and southern China through Southeast Asia to New Guinea and the Solomon Islands.[1] They inhabit diverse wooded environments, including open woodlands, forest edges, dense rainforests, evergreen and deciduous forests, and mature mangroves, typically at elevations from sea level to 2,000 m.[1][2] Primarily insectivorous, they forage by hawking prey in the air above the forest canopy, often perching on exposed branches to scan for insects before launching into swift, direct flights.[2][3] Treeswifts exhibit relatively solitary or small-group behavior, with territorial displays involving high-speed chases and distinctive squeal-like calls.[1] They are monogamous breeders, constructing shallow half-saucer nests from bark, feathers, and bryophytes on horizontal branches 4–30 m above ground; clutches consist of a single white or pale grey egg, incubated mainly by the female with biparental care.[1] None of the species are currently classified as endangered or vulnerable by the IUCN, though the grey-rumped treeswift faces localized declines from hunting and pesticide use, highlighting their dependence on intact forest habitats.[1]Taxonomy and evolution
Classification
Treeswifts are classified within the family Hemiprocnidae, a monotypic family comprising a single genus, Hemiprocne.[1] The family Hemiprocnidae was formally established by Harry C. Oberholser in 1906 in the Proceedings of the Biological Society of Washington.[4] The genus Hemiprocne was introduced by Christian Ludwig Nitzsch in 1829.[5] In the broader avian hierarchy, treeswifts belong to the order Apodiformes, which is nested within the neoavian clade Strisores.[6] This placement reflects their close relationship to true swifts (Apodidae) and hummingbirds (Trochilidae), forming a monophyletic group of aerial specialists.[7] Historically, treeswifts were often confused with true swifts due to their similar aerial lifestyles and morphology, leading to their initial inclusion in expanded classifications such as Caprimulgiformes alongside nightjars and allies.[2] This taxonomic separation was clarified with advancements in anatomical and molecular studies, distinguishing Hemiprocnidae as a distinct lineage.[7]Phylogenetic relationships
Treeswifts (family Hemiprocnidae) belong to the order Apodiformes, where they form one of three monophyletic families alongside true swifts (Apodidae) and hummingbirds (Trochilidae). Molecular phylogenies consistently place Hemiprocnidae as the sister group to Apodidae, with this swift-treeswift clade in turn sister to Trochilidae, supported by analyses of mitochondrial DNA, nuclear genes, and combined datasets that confirm the monophyly of Apodiformes within Neoaves.[2][8] Despite these close genetic affinities, treeswifts are morphologically distinct due to perching adaptations, including stronger feet and reversible outer toes, which enable them to rest on branches—a trait absent in the continuously aerial Apodidae.[2] Evidence from molecular clock studies dates the divergence of Hemiprocnidae from Apodidae to approximately 34–42 million years ago, spanning the late Eocene to early Oligocene, based on calibrations using multiple genetic loci and fossil constraints. Earlier splits within Apodiformes, such as between the swift-treeswift clade and hummingbirds, occurred around 70 million years ago in the late Cretaceous. These timelines highlight a gradual evolutionary radiation, with Hemiprocnidae likely originating in Southeast Asia or Australasia following the isolation of these regions.[8] Synapomorphies uniting Apodiformes encompass unique osteological traits, such as specialized jaw and tongue structures adapted for aerial feeding. Phylogenetic mapping of feather taxis across birds reveals complex patterns of diastataxy and eutaxy, with multiple origins and reversals potentially evolving through genetic drift rather than strong selection, as evidenced by parsimony and likelihood analyses.[9] The family includes four recognized species, all assigned to the genus Hemiprocne: the crested treeswift (H. coronata), gray-rumped treeswift (H. longipennis), whiskered treeswift (H. comata), and moustached treeswift (H. mystacea).[1] The fossil record for Hemiprocnidae is notably incomplete, with no confirmed fossils directly attributable to the family, likely due to the small, delicate skeletons of apodiforms that fossilize poorly. Earliest known relatives include stem apodiforms like Eocypselus from early Eocene deposits in Europe and North America (approximately 50–55 million years ago), which exhibit perching foot morphologies akin to modern treeswifts and suggest an Eocene origin for the broader lineage. Debated assignments, such as Scaniacypselus to the Apodidae-Hemiprocnidae split, further underscore gaps in the Paleogene record.[10]Description
Physical characteristics
Treeswifts are small to medium-sized birds, typically measuring 15 to 31 cm in length and weighing between 20 and 80 g, with a lightweight build that facilitates aerial agility.[1][11][12][13] Their slender, streamlined bodies are adapted for life in forested environments, emphasizing efficiency in flight over ground mobility. The wings of treeswifts are long and pointed, with short arm bones and elongated primaries that enable sustained, gliding flight over short bursts of speed.[1] These wings feature crossed tips and outer streamers for enhanced maneuverability among trees. The tail is deeply forked, often extending up to half the body length, aiding in precise aerial control and steering during foraging sallies.[1] Unlike true swifts, treeswifts possess a nonreversible hind toe that remains fixed backward, allowing them to perch securely on branches and twigs for extended periods. This anatomical adaptation distinguishes them morphologically from their swift relatives, which lack effective perching ability. Their plumage is generally soft and fluffy, providing insulation during perching and flight, with males exhibiting an iridescent sheen on the mantle feathers.[14] This softer feather texture contrasts with the more rigid plumage of true swifts, reflecting their semi-arboreal lifestyle.Plumage variations
Treeswifts in the family Hemiprocnidae exhibit notable plumage variations across species, characterized by the presence of crests and facial ornaments that distinguish them from related swifts. Most species feature elongated forehead feathers forming a crest, measuring 2.5–3 cm in height in the grey-rumped treeswift (Hemiprocne longipennis) and crested treeswift (Hemiprocne coronata), which is often raised while perched; the whiskered treeswift (Hemiprocne comata) and moustached treeswift (Hemiprocne mystacea) have subtler crests.[1] Facial ornaments include bold white stripes along the head in the whiskered and moustached treeswifts, enhancing their distinctive appearance.[1] Sexual dimorphism is evident in plumage coloration and patterns, with males typically displaying brighter, iridescent blues and greens on the upperparts and head. For instance, in the crested treeswift, males possess chestnut ear coverts and glossy blue-black flight feathers, while females lack the chestnut tones and show a narrow silver band across the breast.[15] Similarly, the grey-rumped treeswift's males have dull orange-red or chestnut ear coverts, absent in females, who appear paler below with less vibrant gray upperparts.[12] In the whiskered treeswift, males exhibit dark bronze plumage, contrasting with females' blue-green ear coverts and pink-cinnamon underwing coverts, underscoring a pattern where females often have duller browns or less iridescent tones overall.[11] Juvenile treeswifts possess softer, less contrasted plumage adapted for camouflage, featuring cryptic patterns on the head and body. In the crested treeswift, young birds display a finely banded mix of sooty-black, rufous, buff, and white, predominantly dark above with a pale forehead. Grey-rumped treeswift juveniles show scaly underparts and overall muted tones resembling adults but duller.[1] This juvenile plumage is lost during the first post-fledging molt, transitioning to adult-like feathers.[1] Molt in treeswifts follows an annual cycle, with primaries replaced in a proximal-to-distal pattern that supports their aerial lifestyle; breeding plumage emerges more vibrant post-molt, accentuating iridescent hues and ornaments in adults.[16] In the moustached treeswift, for example, flight feather replacement occurs without omission of juvenile primaries, ensuring continuous flight capability during the annual renewal.[16]Distribution and habitat
Geographic range
Treeswifts, belonging to the family Hemiprocnidae, inhabit tropical and subtropical regions across Asia, with their range extending from India and southern China through Southeast Asia to Indonesia, New Guinea, and the Solomon Islands. This distribution encompasses the Oriental biogeographic region and parts of the Australasian region, where the family achieves its greatest diversity.[1][2] The species overlap extensively in the Indo-Malayan realm, a hotspot of avian endemism, though several treeswifts are restricted to specific island groups, such as the Moluccas and Bismarck Archipelago, highlighting patterns of insular speciation.[17][18] Most treeswifts are sedentary, maintaining stable territories year-round, although some populations exhibit minor altitudinal migrations in response to seasonal variations in food availability or weather.[15][11] Fossil records of stem apodiforms, close relatives of modern treeswifts, from Eocene deposits in North America suggest that the lineage originated in the New World before undergoing historical range expansions into Asia, likely facilitated by tectonic and climatic changes.[19]Habitat preferences
Treeswifts, belonging to the family Hemiprocnidae, primarily inhabit open woodlands, forest edges, and areas with scattered trees, where they can perch on exposed branches and hunt insects in flight.[2] These birds favor environments with clear aerial pathways, such as open woodlands and canopy breaks along roads or rivers, in addition to continuous forest cover.[1] They are also commonly found in mature mangrove stands and coastal regions, particularly in subtropical and tropical zones of Asia and the Indo-Pacific.[20] Their altitudinal range extends from sea level to approximately 1,500 meters, with some species recorded up to 2,000 meters in montane forests.[2][1] Treeswifts inhabit a variety of wooded environments from open woodlands and forest edges to dense rainforests, where they forage above the canopy.[2] They show a particular association with flowering trees, which attract abundant insects and support their aerial foraging lifestyle.[2] Treeswifts demonstrate adaptability to human-modified landscapes, including plantations, gardens, and urban edges, enabling them to persist in fragmented or disturbed areas.[18][2] This flexibility contributes to their occurrence in agricultural settings alongside natural woodlands.[18]Behavior and ecology
Flight and locomotion
Treeswifts possess highly maneuverable and agile flight capabilities, enabling them to navigate through forested environments with precision. Unlike the rapid, erratic wingbeats of true swifts in the family Apodidae, treeswifts employ deeper, more deliberate wing strokes that allow for a somewhat slower but still swift pace, often incorporating gliding elements during sustained aerial movement.[2][1] Their long, pointed wings, adapted for efficient aerodynamics, further facilitate this style of locomotion.[21] In addition to powered flight, treeswifts can briefly hover in place, a behavior observed during territorial displays or while scanning from perches. This hovering ability, combined with their gliding proficiency, distinguishes them from their swift relatives and supports their arboreal lifestyle.[22] On the ground or branches, treeswifts adopt an upright perching posture, gripping vertical surfaces with small, weak feet featuring a non-reversible hind toe that aids in clinging to bark or twigs. This foot structure, while not as versatile as in typical perching birds, allows them to rest comfortably on exposed branches or overhead wires for extended periods.[23][24] Treeswifts exhibit crepuscular activity patterns, with heightened aerial and perching behaviors at dawn and dusk. They avoid landing on the ground almost entirely, relying instead on tree perches and only descending briefly during nesting to attend to eggs or young.[25][1]Foraging and diet
Treeswifts are exclusively aerial insectivores, capturing a variety of flying arthropods on the wing, including members of the orders Hymenoptera (such as bees and wasps), Diptera (flies), Hemiptera (bugs), Isoptera (termites), and Coleoptera (beetles).[1] They occasionally glean insects from foliage or the surface of vegetation, but the majority of their diet consists of insects pursued and caught during flight.[2] Prey selection is opportunistic, with treeswifts frequently targeting swarms of alate termites when available, particularly in species like the gray-rumped treeswift.[12] Foraging behavior involves perching on exposed, mid-level bare twigs in emergent trees, from which they launch brief, high-speed swooping flights to intercept prey before returning to the perch.[26] This perch-and-dart strategy contrasts with the continuous flight of true swifts, allowing greater maneuverability in forested environments.[1] Foraging heights vary by habitat and species but are typically elevated; for example, the gray-rumped treeswift forages at a mean height of 35 m above open ground or 47 m above the forest canopy in Malaysian lowlands.[12] Activity peaks during crepuscular periods at dawn and dusk, when low light conditions are aided by their large eyes and mouth bristles that assist in trapping insects.[1][26] Treeswifts often forage solitarily or in pairs but form loose small flocks of 5–20 individuals during peak feeding times, occasionally numbering up to 50 or more in species like the gray-rumped treeswift; larger aggregations of hundreds may occur in the moustached treeswift.[27][13] These flocks may include mixed-species groups with other aerial insectivores, enhancing foraging efficiency over forested or open areas.[26] For hydration, treeswifts occasionally perform fast skimming swoops over open water surfaces to scoop up liquid, a behavior shared with other apodiform birds, though it is not a primary activity.[12]Reproduction
Treeswifts form monogamous pairs that exhibit biparental care throughout the reproductive cycle, including nest construction, incubation, and chick provisioning.[2][1] In tropical regions, breeding occurs year-round for some species but typically peaks during the dry season or warmer months, such as December to October for the crested treeswift (Hemiprocne coronata), with regional variations like January to April in the Indian subcontinent.[15][1] Nests consist of a small, shallow pad or half-saucer, approximately 25–40 mm in diameter, constructed from bark, feathers, bryophytes, and other plant materials bound with saliva and glued to an exposed tree branch 4–30 m above ground; the structure is so fragile that adults straddle it rather than sitting directly on it.[1][26] Each pair lays a single white-to-pale-grey egg, measuring 23–30 mm in length depending on the species.[1] Both parents share incubation duties, with females typically contributing more time, over a period of about 21 days; the total duration from incubation to fledging is at least 50 days for species like the grey-rumped treeswift (Hemiprocne longipennis).[1][12] Chicks hatch altricial, covered in grey down, and are brooded continuously by both parents initially; they fledge after 25–42 days, varying by species—for instance, 28 days for the whiskered treeswift (Hemiprocne comata) and 42 days in one observed case of the crested treeswift—and receive food from adults for up to three weeks post-fledging.[1][15][28] Parents feed nestlings compact "food balls" of captured insects, with both sexes participating actively, though the low clutch size of one egg constrains overall population growth potential in this family.[1][26]Species
List of species
The family Hemiprocnidae consists of four species in the genus Hemiprocne, all assessed as Least Concern by the IUCN Red List.[20][18][29][30]| Common Name | Scientific Name | Distribution Range | IUCN Status |
|---|---|---|---|
| Crested Treeswift | Hemiprocne coronata | India, Sri Lanka, and Bangladesh east to southern China, Myanmar, Thailand, Laos, Cambodia, and Vietnam | Least Concern |
| Grey-rumped Treeswift | Hemiprocne longipennis | Southern Myanmar and Thailand south through peninsular Malaysia, Singapore, Brunei, and Indonesia (Sumatra, Borneo, and smaller islands) | Least Concern |
| Whiskered Treeswift | Hemiprocne comata | Southern Myanmar and Thailand south through peninsular Malaysia, Singapore, Brunei, Indonesia (Sumatra, Borneo, and smaller islands), and the Philippines | Least Concern |
| Moustached Treeswift | Hemiprocne mystacea | Northern Moluccas (Indonesia) east through New Guinea (Indonesia and Papua New Guinea) to the Solomon Islands | Least Concern |