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Psittacula

Psittacula is a of parrots in the family , consisting of long-tailed parakeets characterized by their predominantly , stout bills, and graduated tails, with many featuring distinctive colored heads or neck rings. Native primarily to , , , and various islands, the genus encompasses adapted to a range of habitats including tropical forests, woodlands, mangroves, and even urban areas. Traditionally, Psittacula includes about 14 species, making it one of the largest genera in the family, though recent molecular phylogenetic studies have revealed and proposed splitting it into several monophyletic genera such as Himalayapsitta, Nicopsitta, Belocercus, Palaeornis, and Alexandrinus, with Psittacula stricto limited to four species: the (P. alexandri), the Derbyan parakeet (P. derbiana), the (P. longicauda), and the Nicobar parakeet (P. caniceps). These revisions aim to reflect evolutionary relationships more accurately, incorporating genera like Psittinus and Tanygnathus within the broader , as well as the extinct Mascarene parrots (Mascarinus). Despite these taxonomic changes, the traditional Psittacula remains widely recognized in and conservation contexts. Notable species within the traditional genus include the (P. krameri), a widespread and often known for its adaptability and vocal mimicry abilities, and the (P. eupatria), prized for its size and striking coloration. Many Psittacula parrots are popular in the pet trade due to their intelligence, sociability, and talking potential, but several face threats from habitat loss, , and , with at least three species extinct in modern times, including Newton's parakeet (P. exsul) from Rodrigues Island. Conservation efforts, guided by organizations like , focus on protecting remaining populations in their native ranges, where they play key ecological roles as seed dispersers.

Etymology and Description

Etymology

The genus name Psittacula derives from the modern Latin psittacula, a of the Latin psittacus meaning "," which in turn originates from the ψιττακός (psittakós), denoting a or . This etymological root reflects the group's characteristic long-tailed, parrot-like form within the . The name was introduced by the French naturalist and zoologist in 1800, as part of his efforts to classify birds systematically in Leçons d'anatomie comparée. Cuvier's work marked a foundational moment in early 19th-century , emphasizing anatomical organization to delineate amid the expanding catalog of global avian species discovered through exploration. The type species for Psittacula was subsequently designated as Psittacula alexandri (Linnaeus, 1758), the red-breasted parakeet, by Australian ornithologist Gregory M. Mathews in 1917 within his multivolume The Birds of Australia. This designation clarified the genus's scope during a period when taxonomic revisions were refining Linnaean classifications based on emerging specimen collections and morphological studies. The genus Psittacula forms a core component of the parrot subfamily Psittaculinae within the family Psittaculidae.

Physical Characteristics

Species of the genus Psittacula are medium-sized parrots, typically measuring 33–58 cm in total length, with the long, graduated tail often comprising up to half of the body length. This elongated tail structure contributes to their distinctive silhouette and aids in maneuverability during flight. The genus is characterized by variations in tail and wing morphology that support agile navigation through forested environments. The predominant plumage across Psittacula species is green, providing effective in tropical and subtropical habitats. is prominent, particularly in head coloration, where adult males frequently exhibit colored rings—such as rose-pink or blue—around the neck, while females typically show less pronounced or absent markings. Some species also display "moustache"-like patterns on the and cheeks, enhancing visual distinction between sexes. This dimorphism, influenced by the genus's distribution across and , underscores adaptations to diverse ecological pressures. Psittacula parrots possess stout, curved bills ideally suited for cracking , a primary dietary component, with the upper often featuring a striking coloration in many . varies slightly in size and hue across the , serving not only functions but also roles in displays like mutual feeding. These features, combined with zygodactyl feet, enable precise manipulation of food items in arboreal settings.

Taxonomy and Phylogeny

Classification History

The genus Psittacula was established by the French naturalist in 1800, placing it within the family based on shared morphological traits among parrots, such as beak structure and zygodactyl feet. Cuvier's classification emphasized anatomical similarities, with the later designated as Psittacus alexandri Linnaeus, 1758, in 1917. In the , ornithologists expanded the to encompass a broader range of Asian parakeets, reflecting increased exploration and specimen collection from the Indo-Malayan region. This period saw the incorporation of species previously grouped under names like Palaeornis by in 1825, consolidating Asian taxa such as the into Psittacula through comparative external morphology. A key milestone occurred in the 1830s with the inclusion of species, notably the (Psittacula krameri), originally described as Psittacus krameri by in 1769 and reassigned to Psittacula by in 1837 based on shared ringed neck and tail characteristics. In the 20th century, classifications shifted toward recognizing Psittacula within the subfamily (later elevated to family ), driven by anatomical studies highlighting distinct osteological features like skull shape and limb musculature that differentiated parrots from forms. These refinements, building on works like Tommaso Salvadori's 1891 morphological review, provided a more nuanced understanding of the genus's diversity across Afro-Asian distributions.

Phylogenetic Relationships

Psittacula is classified within the family , a diverse group of parrots that diverged from other psittaciform lineages approximately 28 million years ago during the Oligocene-Miocene transition. Within , Psittacula forms part of a well-supported that includes the genera Tanygnathus (short-tailed fruit doves) and Prioniturus (racket-tailed parrots), with phylogenetic analyses indicating these as close relatives based on shared molecular markers and morphological features. This positioning highlights Psittacula's role in the broader radiation of Asian and Indo-Malayan parrots, distinct from and African lineages. Molecular phylogenetic studies have demonstrated that Psittacula is paraphyletic, with species of the genera Psittinus and Tanygnathus nesting deeply within its radiation, necessitating taxonomic revisions to restore . A comprehensive using mitochondrial and nuclear RAG-1 sequences reconstructed the relationships of Psittacula sensu lato, revealing that basal divergences within this complex occurred around 20–25 million years ago, coinciding with tectonic shifts in that facilitated . These findings underscore the dynamic evolutionary history of the group, driven by vicariance and dispersal across island archipelagos. Key synapomorphies defining the clade include distinctive ringed neck patterns—such as black collars and colored bands—and elongated tails, which represent derived morphological traits adapted for aerial agility in forested habitats. These features, observed across multiple , evolved convergently in some lineages but serve as diagnostic markers for the group's monophyletic core, distinguishing it from shorter-tailed relatives. Early morphological classifications, which emphasized these traits, provided initial insights into groupings but were refined by genetic data to resolve ambiguities. The extinct genus Mascarinus, known from the , is phylogenetically embedded within the Psittacula radiation, suggesting an ancient dispersal event from mainland to oceanic islands followed by isolation and specialization. analyses confirm Mascarinus as a close relative, sharing molecular signatures with extant long-tailed Psittacula species and supporting its inclusion in the broader paraphyletic complex. This connection illustrates how events have shaped the modern diversity of the .

Alternative Taxonomies

Recent molecular phylogenetic studies have revealed that the traditional Psittacula is paraphyletic, prompting proposals for taxonomic revisions to achieve . In a study, et al. analyzed mitochondrial and nuclear DNA sequences from multiple Psittacula and related genera, concluding that the requires splitting into several monophyletic units. They proposed retaining only P. alexandri and P. derbiana in the core Psittacula, while reassigning other to new or resurrected genera, such as Himalayapsitta for P. cyanocephala, P. roseata, P. himalayana, and P. finschii, and Nicopsitta for P. columboides and P. calthrapae. Additional genera suggested include Belocercus for P. longicauda and Alexandrinus for P. krameri and related forms. A phylogenomic further supports these splits, retaining Psittacula sensu stricto for P. alexandri and P. derbiana while confirming deep divergences (e.g., 10 for Psittacula-Tanygnathus ) and embedding extinct Mascarinus within the radiation. Following this research, BirdLife International and the IUCN have adopted several of these splits in their taxonomic frameworks for conservation assessments, particularly post-2020, to better reflect evolutionary distinctiveness and inform threat evaluations. For instance, species like the Malabar parakeet (Nicopsitta columboides) and plum-headed parakeet (Himalayapsitta cyanocephala) are now recognized under their respective new genera in BirdLife's species factsheets. These changes enhance precision in Red List evaluations by highlighting potentially overlooked conservation priorities among the splintered lineages. Alternative proposals have considered mergers rather than splits to resolve the paraphyly identified in the same phylogenetic data, where genera like Tanygnathus and Psittinus nest within the broader Psittacula based on mtDNA analyses. Some researchers incorporating Tanygnathus species (e.g., T. megalorynchos) and Psittinus (e.g., P. abbotti) into an expanded Psittacula to maintain a single genus encompassing all closely related Asian and Indo-Malayan parrots, though this approach has not gained widespread acceptance. Ongoing debates also focus on elevating certain subspecies to full species status within Psittacula. For example, proposals to split the African subspecies (P. k. krameri) from the Asian forms (P. k. borealis and P. k. manillensis) due to genetic divergence and morphological differences observed in phylogenetic reconstructions, potentially recognizing two distinct across the species' range. Such elevations remain contentious, as hybridization potential and limited sampling complicate boundaries.

Distribution and Habitat

Geographic Range

The genus Psittacula exhibits one of the broadest distributions among parrot genera, spanning from across southern to and the islands of the . Native populations are primarily concentrated in tropical and subtropical zones, with the core range influenced by historical biogeographic patterns tied to tectonics in the . The highest species diversity occurs in , particularly the , where over eight are recorded, including widespread forms like the (P. krameri) and endemics such as the Nicobar parakeet (P. caniceps) restricted to the . Additional hotspots include and the Himalayan foothills, supporting like the (P. cyanocephala). In , several occupy island archipelagos, such as the red-breasted parakeet (P. alexandri) in and the , while hosts a few, notably P. krameri across savannas from to . islands feature isolated populations, including the Mauritius parakeet (P. eques). Due to the international pet trade, multiple Psittacula species have established introduced populations far beyond their native ranges. The (P. krameri), in particular, has formed feral colonies in over 35 countries, including widespread presence in (e.g., since the 1960s, with over 12,000 breeding pairs), the (e.g., , UAE), and the Americas (e.g., , ). Other species, like the (P. eupatria), have naturalized in parts of (e.g., , ) and the . These introductions often stem from escaped or released cage birds, leading to self-sustaining populations in urban and suburban environments. Extinct species highlight historical endemism in isolated hotspots, such as the (P. wardi) on Mahé and islands, which disappeared due to human activities by the early . Similarly, Mascarene Island endemics like P. exsul from underscore the genus's vulnerability on oceanic islands.

Habitat Preferences

Psittacula species primarily inhabit tropical and subtropical ecosystems, including moist lowland forests, dry woodlands, and open savannas across their native ranges in and . These parrots favor environments with abundant cover for roosting and nesting, often occurring from up to elevations of 3,000 m, though some species like the Derbyan parakeet (P. derbiana) extend into higher montane coniferous and mixed forests up to 4,000 m. For instance, the slaty-headed parakeet (P. himalayana) thrives in , , and forests between 600 m and 2,500 m in the . Many Psittacula species demonstrate remarkable adaptability to human-modified landscapes, frequently exploiting agricultural fields, orchards, and spaces where natural habitats have been altered. The (P. krameri), for example, has successfully colonized city parks and suburban areas in both native and introduced ranges, benefiting from the availability of exotic fruit trees and reduced predation. Similarly, the (P. eupatria) occupies farmlands, mangroves, and even urban gardens up to 900 m, showcasing the genus's versatility in anthropogenically influenced environments. Microhabitat requirements for Psittacula include access to large cavities for nesting, often in or mature living trees, which provide secure sites for . Proximity to sources, such as or wetlands, is also preferred, particularly for in drier savannas like P. krameri, which inhabits marshes and bogs alongside arid grasslands. The genus tolerates a wide climatic spectrum, from humid tropical rainforests to semi-arid zones, with some exhibiting altitudinal to track seasonal resources; the slaty-headed , the only psittacid known for this behavior, descends to lower elevations in winter.

Behavior and Ecology

Diet and Foraging

Psittacula parrots are primarily granivorous and frugivorous, with their diet consisting mainly of seeds, nuts, fruits, and berries sourced from various trees, including figs (Ficus spp.) and palms. Species such as the rose-ringed parakeet (P. krameri) derive approximately 80% of their intake from seeds, utilizing their robust beaks to husk and crack open hard-shelled items like grains and nuts, while also consuming fleshy fruits and vegetables opportunistically. This plant-based focus supports their role as seed predators in native ecosystems, acting both as seed predators and dispersers depending on seed type and handling. Supplementing their herbivory, Psittacula exhibit opportunistic omnivory by occasionally ingesting small amounts of animal matter, such as and their larvae, as well as from flowers. In human-modified landscapes, they raid agricultural crops like , , and pulses, contributing to their classification as pests in regions including and parts of . Nectar consumption, particularly from blossoms, provides seasonal energy boosts, while intake, though minor in observed populations, aids in protein acquisition during breeding periods. Foraging occurs in gregarious typically numbering 10–100 individuals, enabling coordinated scanning for and predator avoidance; larger aggregations form near abundant resources like fruiting . Activity is diurnal, with heightened at dawn and dusk when visibility aids detection of scattered patches, and birds often travel several kilometers daily in search of optimal sites. Flock dynamics facilitate information sharing about locations via vocalizations, enhancing efficiency in patchy environments. Dietary composition varies seasonally, with flower buds and unripe seeds predominating in dry periods when fruits are scarce, and a shift toward higher and intake during monsoons when fruits abundantly. In arid regions, this adaptability includes targeting nectar-rich blooms during water-stressed months, maintaining nutritional balance across fluctuating availability. Such flexibility underscores their resilience in diverse habitats from savannas to urban fringes.

Reproduction and Life Cycle

Psittacula parrots typically form monogamous pairs that maintain strong pair bonds during the . is seasonal, often occurring from to in regions like , though timing varies by species and latitude to align with food availability. Pairs engage in displays, including mutual and aerial chases, before nest site selection. Nesting occurs primarily in natural tree hollows or cavities in rotten trunks, though some utilize cliffs or human-made structures in altered habitats. The female lays a clutch of 3–6 eggs, with an average of 4 observed in representative like the rose-ringed parakeet (Psittacula krameri). , performed solely by the female, lasts 23–28 days, during which the male provides food to the incubating partner. Hatching produces altricial chicks covered in sparse down, dependent on parental provisioning. Fledging occurs at 6–7 weeks, after which young remain with parents for several months to learn foraging skills. Parental care is biparental, with both sexes feeding regurgitated food to nestlings and fledglings; the male often plays a key role in defending the nest and territory. Juveniles exhibit duller plumage lacking adult coloration, such as the distinctive neck rings in males of some species, aiding camouflage during this vulnerable stage. Sexual maturity is reached at approximately 3 years, enabling pair formation and breeding. In the wild, average lifespan is estimated at 10–15 years, though some individuals live longer (up to 14+ years), influenced by predation, disease, and resource availability. Diet quality during chick rearing can affect growth rates, with protein-rich foods promoting faster development.

Social Structure

Psittacula species, such as the (Ps. krameri), exhibit flock-based societies characterized by nomadic groups that facilitate and resource sharing in variable environments. These birds form cohesive flocks ranging from small groups of 10–20 individuals to larger aggregations of hundreds or even thousands, particularly during non-breeding seasons when they colonially in trees or structures. Hierarchical structures emerge within flocks, often centered on dominant pairs that secure priority access to food sources, reducing intra-group conflicts and enhancing overall stability. Communication among Psittacula individuals relies on a combination of vocalizations and visual displays to maintain group cohesion and coordinate activities. Vocal repertoires include sharp screeches for or flight initiation, rhythmic chatters during interactions, and softer purrs to signal aggregation at roosts. Visual signals, such as fanning to assert dominance or head bobbing during affiliative encounters, complement these calls, allowing to convey intentions without disrupting movement. often occurs in these fluid groups, where shared vigilance improves efficiency in locating scattered resources. Territoriality in Psittacula is primarily focused on defending nesting sites rather than broad areas, with pairs aggressively competing for cavities while maintaining fluid memberships in larger flocks. This selective territorial behavior allows groups to remain adaptable to changing habitats, as individuals can shift between flocks without rigid boundaries. In captivity, Psittacula parrots demonstrate notable tameness, forming strong bonds with human caregivers and exhibiting advanced abilities, where rose-ringed parakeets can imitate human speech phrases with repertoires sometimes exceeding hundreds of units, particularly in males.

Species Diversity

Extant Species

The genus Psittacula includes 13 extant species of parrots, mostly native to and parts of , characterized by their predominantly plumage, long tails, and adaptations to diverse tropical and subtropical environments.
  • Psittacula krameri (): A widespread native to and , now invasive in , , and other regions; notable for pronounced , with adult males displaying a distinctive black and rose-colored absent in females. It has four recognized subspecies, including P. k. borealis in .
  • Psittacula eupatria (): One of the largest in the genus, reaching up to 58 cm in length, with males featuring a bill and variable shoulder patches; native to and .
  • Psittacula alexandri (): Found in , distinguished by the male's maroon breast and blue-grey head; adaptable to modified habitats like gardens and plantations.
  • Psittacula longicauda (long-tailed parakeet): Native to , including the , , , , and nearby islands, recognized by its exceptionally long comprising more than half of its total length of 40-50 cm and predominantly coloration with subtle yellow undertones.
  • Psittacula derbiana (Derby's parakeet): Native to the Himalayan region, featuring all- plumage with a robust build and a distinctive yellowish bill; it inhabits high-altitude forests.
  • Psittacula caniceps (Nicobar parakeet): Endemic to the of , with males showing a pale head and nape contrasting the body; restricted to primary forest habitats.
  • Psittacula eques (): Restricted to , the smallest at about 34 cm, with subtle plumage and a short ; subject to ongoing from near-extinction.
  • Psittacula calthropae (Layard's parakeet): Endemic to Sri Lanka's forests, notable for its short and uniform appearance with minimal sexual differences.
  • Psittacula cyanocephala (plum-headed parakeet): Native to the , where males have a purplish-blue head and hens a one; prefers open woodlands.
  • Psittacula himalayana (slaty-headed parakeet): Distributed in the Himalayas and northern , characterized by the male's slate- head and orange bill base.
  • Psittacula finschii (grey-headed parakeet): A rare in the eastern Himalayas, with both sexes showing a head and body; inhabits remote montane forests.
  • Psittacula roseata (blossom-headed parakeet): Found in , distinguished by the male's pinkish head and variable rose- plumage patches.
  • Psittacula columboides (blue-winged parakeet): Endemic to the Western Ghats of , featuring blue flight feathers and a predominantly body with a short .
Taxonomic debates occasionally propose splits within the genus, but the current consensus recognizes these 13 as valid extant species.

Extinct Species

The genus Psittacula includes several species that became extinct in the 18th and 19th centuries, primarily due to human activities on Indian Ocean islands. These losses highlight the vulnerability of island-endemic parrots to introduced threats and habitat alteration. Psittacula wardi, known as the , was endemic to the granitic islands of Mahé and , where it was reported as abundant in the early . The , characterized by its and long tail, went extinct in the , with the last confirmed individuals shot in 1893. Primary causes included intense hunting for food and persecution by farmers and owners who viewed the birds as pests damaging crops. Psittacula exsul, or Newton's parakeet, inhabited the island of in the Mascarene archipelago and was last reliably observed in the . This , which featured a distinctive ring-necked appearance similar to some extant relatives, suffered from a combination of through for and direct by . Colonial activities accelerated the loss of its native woodland habitat, leading to its rapid decline. The Réunion parakeet, tentatively classified as a subspecies Psittacula eques eques, is known from historical accounts and a single preserved specimen; it formerly occurred on Réunion Island and became extinct around the 1730s. Descriptions from early European visitors indicate it was a green parakeet adapted to forested environments, but no complete skeletal remains exist to confirm its taxonomy fully. Its disappearance is attributed to early colonial deforestation and the introduction of invasive species such as rats, which preyed on eggs and nestlings. Phylogenetic analysis links it closely to the extant echo parakeet (P. eques) on nearby Mauritius, suggesting a shared island radiation. Fossil records of Psittaciformes, the order including Psittacula, from deposits in , such as Sansan in , indicate that parrots had a much wider historical range across before contracting to tropical regions. These remains, dating to approximately 14-12 million years ago, include multiple taxa and suggest ancestral distributions that predate the modern Afro-Asian focus of the genus. Across these extinct Psittacula species, common extinction drivers were colonial-era introductions of predators like rats, widespread for plantations, and opportunistic hunting, which collectively decimated small island populations within decades.

Conservation Status

Major Threats

Psittacula parrots face multiple threats across their native ranges in and , with loss, illegal capture for the pet trade, and conflicts with being the most significant drivers of declines. These pressures are exacerbated by the genus's wide distribution, which facilitates both legal and illegal exploitation. According to assessments by , over 80% of Psittacula are affected by at least one of these threats, contributing to their vulnerable or near-threatened statuses on the . The illegal pet trade poses a severe risk to many Psittacula species, particularly through widespread trapping and export despite regulations. For instance, the (Psittacula krameri) has been heavily targeted, with historical pre-CITES trade volumes estimated in the tens of thousands annually from and , leading to significant population reductions in source regions. Species like the (Psittacula alexandri) and (Himalayapsitta roseata) continue to suffer from cage-bird capture, which accounts for the primary threat to their survival, often resulting in nest and adult mortality rates that outpace reproduction. Habitat destruction, driven by for and , has severely impacted Psittacula populations in , where many species rely on lowland forests. Since the , has lost approximately 15-20% of its , with annual rates averaging 2-3 million hectares, fragmenting habitats essential for nesting and . Lowland species such as Psittacula alexandri have experienced about 16% forest loss over recent generations, intensifying vulnerability to and reduced food availability. Agricultural expansion has led to direct conflicts, as Psittacula parrots raid crops like fruits, grains, and vegetables, resulting in through and deterrents. The (Psittacula krameri) is notorious as a in and , causing yield losses of 10-20% in orchards such as and sunflower, prompting farmers to or birds en masse. Similarly, the (Psittacula eupatria) faces targeted killing in and fields, where flock foraging behaviors amplify damage and fuel retaliatory actions. In introduced ranges outside their native habitats, Psittacula species act as invasives, posing risks through competition and disease transmission to native avifauna. The aggressively competes for nest cavities, displacing species like woodpeckers and in and , with documented cases of fatal interference. Additionally, they can spread pathogens such as (PBFD), which has infected over 60 psittacine species and threatens endemic birds in Pacific islands.

Conservation Measures

Conservation measures for Psittacula species primarily focus on habitat protection, regulation of , and targeted interventions for threatened taxa, given that several species face risks from poaching and habitat degradation. Following 2020 taxonomic revisions, many former Psittacula species are now classified in genera such as Alexandrinus and Himalayapsitta, but conservation assessments by organizations like IUCN and continue to address the broader . Most species in the genus are listed under Appendix II of the Convention on in Endangered Species of Wild Fauna and Flora (), which regulates trade to prevent while allowing sustainable utilization. These listings facilitate monitoring and enforcement against illegal pet trade, a major threat across the genus. Additionally, protected areas in native ranges, such as national parks and biosphere reserves, safeguard key s like tropical forests and woodlands essential for foraging and nesting. For the Vulnerable Echo parakeet (Alexandrinus eques) of , intensive conservation has driven a remarkable recovery from near-extinction, with the increasing from fewer than 20 individuals in the to over 800 by 2021, and downlisted from Endangered to Vulnerable on the . Key measures include a program initiated in 1993, which produced 139 birds for release into the wild until 2004, supplemented by translocations to sites like the Bambous Mountains, where a self-sustaining established by 2018. Habitat management within involves nest box provision, supplementary feeding, and control of such as rats, pigs, and macaques that compete for resources or predate nests. management, particularly for , includes routine treatment of populations on offshore islands like Ile aux Aigrettes. Ongoing efforts recommend further translocations to additional sites, including potential reintroduction to Island, and continued monitoring of breeding success. Near Threatened species like the (Psittacula caniceps) and grey-headed parakeet (Himalayapsitta finschii) benefit from habitat-focused initiatives in the Biosphere Reserve and Himalayan protected areas, respectively, aimed at curbing loss from development and . For the , with an estimated 6,000–15,000 mature individuals, recommended actions include calculating rates of post-2004 and enforcing stricter habitat protection to counter settlement pressures. Trade regulation under Appendix II is emphasized, with calls to assess impacts and strengthen legal against cage-bird capture. Similarly, for the grey-headed parakeet, whose is declining by an estimated 20–29% over three generations due to pet trade in countries like and , measures involve conducting range-wide surveys, monitoring illegal trade routes, and launching awareness campaigns to reduce trapping. Habitat protection targets the 15% loss observed over the same period, prioritizing conservation in fragmented woodlands. Across the genus, collaborative efforts by organizations like the World Parrot Trust and national wildlife authorities emphasize research into population trends and threat mitigation, ensuring measures are adaptive and evidence-based. These strategies have contributed to status improvements for some , such as the , highlighting the potential for recovery when threats are addressed proactively.