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Notopteridae

Notopteridae is a family of ray-finned fishes in the order Osteoglossiformes, commonly known as featherbacks or knifefishes, comprising four genera and eleven species characterized by their elongated, laterally compressed bodies, reduced or absent dorsal fins, and long-based anal fins that are confluent with the caudal fin.^[1] These fishes are primarily freshwater dwellers, occasionally inhabiting brackish waters, and are found in quiet, slow-moving habitats such as floodplains, ponds, backwaters, and rivers across sub-Saharan Africa and Southeast Asia.^[1] Adapted to low-oxygen environments, notopterids are air-breathing species that utilize a physostomous swim bladder connected to the gut for supplemental respiration, enabling survival in hypoxic conditions.^[2] The family includes the African genera Papyrocranus and Xenomystus, with two and one species, respectively, and the Asian genera Chitala and Notopterus, with six and two species, respectively; taxonomic revisions continue to refine species boundaries, particularly within Chitala and the recent recognition of two species in Notopterus.^[1]^[3]^[4] Morphologically, notopterids feature 120–180 scales along the lateral line, 25–45 ventral scutes, and pelvic fins with 3–6 rays when present, with maximum reported lengths reaching 80 cm.^[1] Ecologically, they are mostly nocturnal, carnivorous predators feeding on smaller fishes, crustaceans, and insects, exhibiting sluggish swimming and parental care as guarders during reproduction in the rainy season.^[1]^[5] Valued for food in their native ranges and as ornamental fish in aquaria—smaller species like Xenomystus nigri for homes and larger ones like Chitala ornata for public displays—notopterids face threats from overexploitation, habitat loss, and pollution, with some species assessed as vulnerable or endangered.^[1]^[6]

Taxonomy and phylogeny

Classification

Notopteridae is a family of ray-finned fishes classified within the order Osteoglossiformes, a group of ancient teleosts commonly referred to as bony-tongued fishes due to their toothed basihyal bone.^[1] This placement positions Notopteridae among the basal actinopterygians, sharing the order with families such as Osteoglossidae (arapaimas) and Mormyridae (elephantfishes).^[1] The family name Notopteridae derives from the Greek terms noton (back) and pteron (fin or wing), alluding to the distinctive small dorsal fin present in most members.^[1] It was originally established by the Dutch ichthyologist Pieter Bleeker in 1851, based on morphological features observed in Asian freshwater species.^[7] Subsequent taxonomic revisions, incorporating molecular and morphological data, have refined the family to include 10 valid species across four genera: Chitala (Asian featherbacks), Notopterus (bronze featherbacks), Xenomystus (African knifefish), and Papyrocranus (African knifefishes).^[1]^[8] Diagnostic characters of Notopteridae include a compressed, knife-like body; a small, quill-shaped dorsal fin (absent in Xenomystus); and a long anal fin that is confluent with the reduced caudal fin, featuring over 100 combined rays.^[1] Additionally, the swim bladder possesses paired anterior extensions that contact the inner ear structures, enhancing auditory sensitivity—a trait unique among non-otophysan teleosts in this family.^[9] These features distinguish Notopteridae from other osteoglossiforms and underscore their adaptation to freshwater environments in Africa and Asia.^[1]

Evolutionary relationships

Notopteridae occupies a basal position within the superorder Osteoglossomorpha, a morphologically and molecularly distinct lineage of teleost fishes characterized by primitive traits such as a protrusible tongue with toothed basihyal. Phylogenetic analyses consistently place Notopteridae in the suborder Notopteroidei of the order Osteoglossiformes, where it forms the sister group to Mormyroidea (comprising Mormyridae and Gymnarchidae), with Hiodontidae (order Hiodontiformes) as the successive sister taxon to the entire Osteoglossiformes clade.^[10]^[11] This arrangement positions Notopteroidei as one of the earliest-diverging lineages within Osteoglossomorpha, highlighting its role in understanding the early radiation of basal teleosts. Earlier morphological studies had proposed a direct sister-group relationship between Notopteridae and Hiodontidae based on shared features like a specialized basihyal tooth plate, though modern syntheses refine this to reflect the broader Osteoglossiformes structure.^[12]^[13] Molecular phylogenies, derived from concatenated datasets of mitochondrial genes (such as cytochrome b, 12S rRNA, and 16S rRNA) and nuclear markers (including RAG1 and RAG2), robustly support the monophyly of Notopteridae and its placement within Osteoglossomorpha, with bootstrap values exceeding 95% across multiple studies. These analyses estimate the divergence of Notopteroidei from other osteoglossiform lineages (such as Osteoglossoidei) around 150–200 million years ago during the Late Jurassic to Early Cretaceous, predating the diversification of more derived groups like arowanas and arapaimas. Within Notopteridae, the split between the African (Xenomystinae: Xenomystus and Papyrocranus) and Asian (Notopterinae: Notopterus and Chitala) subfamilies occurred approximately 133 million years ago (95% confidence interval: 111–156 Mya), aligning with molecular clock calibrations using fossil constraints.^[10]^[14]^[15] Key synapomorphies defining Notopteridae include a dorsally elongated swim bladder with extensive vascularization and a duct for air intake, adapted for bimodal respiration in hypoxic freshwater habitats—a trait shared across Osteoglossomorpha but specialized in notopterids for frequent surface gulping. Additionally, the genus Xenomystus within the African subfamily Xenomystinae exhibits electrogenic organs derived from modified axial musculature, producing weak electric organ discharges (up to 1–2 mV) for electrolocation and social signaling, a convergence with but independently evolved from the stronger discharges in closely related Mormyridae.^[16]^[17] These features underscore the family's adaptive radiation in tropical freshwater systems. The biogeographic pattern of Notopteridae, spanning sub-Saharan Africa and Southeast Asia with no intervening populations, implies a Gondwanan origin during the Mesozoic, followed by vicariance driven by continental drift. Molecular evidence indicates that the ancestral notopterid stock inhabited freshwater habitats on the supercontinent prior to its fragmentation, with the African-Asian divergence timed to the rifting of the Indian plate from eastern Gondwana around 133 Mya, allowing limited gene flow until approximately 85–90 Mya via land bridges. This vicariance model is reinforced by the family's obligate freshwater ecology, precluding post-rift marine dispersal, and contrasts with dispersal hypotheses for other osteoglossomorphs.^[15]^[18]^[19]

Physical description

Morphology

Notopteridae fishes possess an elongate, laterally compressed body that assumes a distinctive knife-like shape, facilitating agile navigation through aquatic environments. These fishes can attain lengths of up to 150 cm, with the body adorned by 120-180 scales along the lateral line and 25-45 hardened ventral scutes providing structural support.^[1]^[20]^[21] The fin arrangement is characteristic, featuring a small, quill-like dorsal fin positioned posteriorly, which is absent in the genus Xenomystus; the anal fin spans a long base and fuses seamlessly with the small, rounded caudal fin, resulting in a combined ray count of 100 or more. Pelvic fins, when present, are reduced and bear 3-6 rays, contributing to the overall streamlined profile.^[1]^[20] The head region includes a protrusible mouth adapted for prey capture, small eyes suited to low-light conditions, and the absence of a subopercle bone; a prominent lateral line system runs along the body to detect water movements.^[22]^[1] Coloration varies across genera but typically presents as silvery or bronze tones on the body, often accented by spots, stripes, or bars that enhance camouflage in their habitats; for instance, Chitala chitala is overall silvery in color, with a series of transverse gold or silver bars on the dorsum in subadults and adults (though not always present), while Notopterus notopterus adults exhibit plain brown hues.^[23]^[24]

Adaptations

Notopteridae exhibit several specialized physiological adaptations that enable survival in hypoxic freshwater environments, particularly through modifications to their respiratory and sensory systems. The swim bladder in these fishes is elongated and divided into two lateral chambers by a longitudinal septum, allowing it to serve multiple functions beyond buoyancy regulation. This structure connects directly to the gut via a wide pneumatic duct, facilitating air-breathing as an accessory respiratory organ that supplements gill-based oxygen uptake when dissolved oxygen levels are low. The vascularized walls of the swim bladder enable efficient gas exchange, with oxygen comprising approximately 11% and carbon dioxide around 3% of the gas content under normal conditions, rising to 6.5% CO₂ during stress.^[25]^[26] Additionally, the swim bladder contributes to sound production, likely through vibrations generated by associated musculature, aiding in acoustic communication within turbid habitats. Sensory adaptations in Notopteridae include electroreception in the African genera Xenomystus and Papyrocranus, where ampullary electroreceptors detect weak electric fields for navigation and prey detection in low-visibility waters. These species possess weakly electric organs that generate low-amplitude discharges, enabling electrolocation without the high-voltage output seen in strongly electric fishes. This system is particularly advantageous in the dimly lit, sediment-laden rivers of Africa, where visual cues are limited.^[27]^[28] Locomotion in Notopteridae is adapted for maneuverability in constrained environments, relying primarily on an undulating anal fin that extends along much of the body length to propel the fish forward or backward with equal efficiency. This ribbon-like fin generates a traveling wave that provides thrust, allowing rapid direction changes without body torsion. The pectoral fins are reduced in size relative to the prominent anal fin, serving mainly for stabilization rather than primary propulsion, which enhances streamlining in fast-flowing or vegetated waters.^[29]^[30] The respiratory physiology of Notopteridae supports high tolerance to hypoxia, with the dual-breathing capability via the swim bladder allowing survival in waters with critically low dissolved oxygen, such as seasonally flooded tropical wetlands. This adaptation is crucial for their persistence in environments where oxygen levels can drop below levels sustainable for purely aquatic breathers, enabling extended aerial respiration during droughts or stagnation.^[31]^[32]

Distribution and habitat

Geographic range

The family Notopteridae, commonly known as featherbacks or knifefishes, is distributed across freshwater and occasionally brackish waters in tropical regions of Africa and Southeast Asia, with no native presence in other continents.^[1] This disjunct distribution reflects the family's ancient Gondwanan origins, with the two major clades separated by continental drift and lacking overlap between African and Asian lineages.^[18] In Africa, Notopteridae are represented by the genera Papyrocranus and Xenomystus, which are endemic to West and Central Africa. Papyrocranus afer occurs in the Niger River basin and coastal drainages such as the Oluwa, Tano, Pra, and Lekki Lagoon systems in Nigeria and Ghana, while Papyrocranus congoensis is found in the Congo Basin.^[33]^[34] Xenomystus nigri has a wide distribution across tropical African freshwater systems, including the Nile, Chad, Niger, Ogowe, and Congo basins, as well as coastal rivers from Sierra Leone to Angola.^[35] These distributions are confined to tropical freshwater systems south of the Sahara, with no records extending into East or Southern Africa.^[18] The Asian clade includes the genera Chitala and Notopterus, primarily distributed across the Indian subcontinent, Indochina, and the Malesian region encompassing Indonesia and Malaysia. Notopterus notopterus ranges from the Indus and Ganges basins in India and Pakistan through the Irrawaddy, Salween, Chao Phraya, and Mekong basins in Myanmar, Thailand, Laos, and Vietnam.^[4] Species of Chitala, such as C. chitala in the Ganges and Brahmaputra rivers of India and Bangladesh, C. ornata in the Mekong and Chao Phraya basins of Thailand, Laos, Cambodia, and Vietnam, and C. lopis in Indonesian waters including Sumatra and Borneo (previously considered extinct but rediscovered in Java in 2023), exhibit similar broad but fragmented distributions tied to major river systems.^[18]^[36] Some Chitala species, notably C. ornata and C. chitala, have established introduced populations in Singapore's reservoirs and waterways, likely from aquarium releases, where they now form feral breeding groups.^[37] Biogeographically, the separation of African and Asian Notopteridae clades occurred during the breakup of Gondwana approximately 80–120 million years ago, with the African lineage retaining a basal position and the Asian forms dispersing via the northward drift of the Indian plate, possibly using Madagascar-Seychelles as stepping stones in an "Out-of-India" scenario around 85–90 million years ago.^[18] This tectonic history explains the family's vicariant distribution without intercontinental migration post-divergence.^[8]

Environmental preferences

Notopteridae species primarily inhabit slow-moving or stagnant freshwater environments, including rivers, swamps, floodplains, lakes, canals, ponds, and backwaters, where they avoid fast currents and prefer lowland tropical wetlands.^[1] These fishes are adapted to primary freshwater habitats but show tolerance for brackish conditions, often entering river mouths or coastal areas.^[38] For example, the bronze featherback (Notopterus notopterus) is commonly found in swamp forests, floodplains, and still waters across Southeast Asia. Similarly, African species like the African knifefish (Xenomystus nigri) occupy quiet, vegetated streams, lakes, and coastal river basins in tropical Africa. They thrive in warm water temperatures ranging from 24–28°C, which supports their metabolic needs in tropical regions.^[39] Water pH levels between 6 and 6.5 are typical, with neutral to slightly acidic conditions common in their vegetated habitats that provide cover from predators. Notopteridae exhibit a high tolerance for low dissolved oxygen levels due to their air-breathing capability via a vascularized swim bladder, allowing survival in hypoxic swamps and stagnant pools where oxygen saturation can drop critically low.^[2] Microhabitat preferences vary diurnally: during the day, individuals often rest as bottom-dwellers among roots, submerged vegetation, decaying organic matter, and debris for concealment, while at night they become active surface swimmers in open waters.^[40] This behavior aligns with their nocturnal lifestyle and ecological niche in densely vegetated, low-flow wetlands that offer both shelter and foraging opportunities.^[41]

Ecology and behavior

Diet and feeding

Species in the family Notopteridae exhibit carnivorous and insectivorous diets, preying primarily on aquatic insects, crustaceans, small fishes, and molluscs. Juveniles, including fry and early-stage adolescents, rely heavily on zooplankton such as cladocerans, copepods, and rotifers to support rapid growth, transitioning to larger invertebrates like insect larvae and prawns as they mature. Adults favor more substantial items, including minnows, teleosts, and bivalve molluscs, reflecting an ontogenetic shift in prey size and type that aligns with increasing body size and mouth gape.^[42]^[43]^[44] Foraging behavior in Notopteridae is predominantly nocturnal or crepuscular, with individuals targeting surface-feeding prey in the dimly lit, vegetated shallows of rivers and wetlands. These fishes adopt an opportunistic, sluggish ambush strategy, lying in wait among submerged vegetation or leaf litter to strike at passing prey rather than engaging in prolonged active pursuit, which suits their elongated body form and low-oxygen environments. This mode is particularly effective in turbid, low-visibility waters where visual cues are limited.^[45] Sensory adaptations enhance prey detection during feeding; for instance, passive electroreception via ampullary organs in species like Xenomystus nigri allows localization of bioelectric signals from hidden prey in murky conditions. The family's unique tongue-bite apparatus, involving specialized dentition on the tongue and basihyal, facilitates intraoral prey manipulation and crushing, complementing suction-based capture through rapid hyoid depression. As mid-level predators, Notopteridae occupy an intermediate trophic position in tropical wetland food webs, controlling invertebrate and small fish populations while serving as prey for larger piscivores.^[46]^[47]

Reproduction and parental care

Members of the Notopteridae family are guarders, producing adhesive eggs typically measuring 3-4 mm in diameter that are laid in clusters on submerged vegetation, roots, stones, or other substrates. These eggs are demersal and non-guarded initially but receive protection post-deposition. For instance, in Notopterus notopterus, eggs are spherical with spiraling ridges around the micropyle for adhesion, averaging 3.8-4 mm in size.^[5] In Chitala chitala, eggs range from 3-4.5 mm in diameter and are similarly adhesive.^[48] Spawning in Notopteridae generally occurs during the daytime in warm water temperatures of 25-28°C, with males performing courtship displays such as chasing, circling, and tail fanning to attract females. Females are fractional spawners, releasing eggs in multiple batches over several spawning events; a single female may produce 1,200-20,000 eggs total per reproductive cycle, depending on species and body size. In Notopterus notopterus, females deposit 15-225 eggs per spawning act, totaling up to 1,560 eggs across 20 events in captivity.^[5] Larger species like Chitala chitala exhibit higher fecundity, with absolute fecundity averaging around 13,431 eggs.^[49] Fertilization is external, with males releasing milt over the egg clusters immediately after deposition.^[50] Parental care is exclusively paternal, with males guarding the eggs and newly hatched fry against predators and maintaining the nest site by fanning to oxygenate the clutch; females provide no post-spawning involvement and typically leave the area. In Notopterus notopterus, males aggressively defend the nest, reducing their own feeding during this period, and continue care for several days until hatching.^[5] Similarly, in Chitala ornata and Chitala chitala, males guard adhesive eggs attached to substrates, ensuring high survival rates through territorial behaviors.^[51] This male-only care is a key reproductive strategy in the family, correlating with the production of fewer but larger eggs compared to non-guarding teleosts.^[52] Embryonic development leads to hatching in 5-8 days under typical conditions of 26-27°C, after which larvae absorb their yolk sac within 12-14 days and begin exogenous feeding. In Notopterus notopterus, larvae hatch at approximately 10.5 mm total length after 168-204 hours, with the larval period extending to day 36 before transitioning to juveniles; sexual maturity is reached at 1-2 years, or around 23-27.5 cm in length.^[5] For Chitala chitala, hatching occurs in about 5 days at similar temperatures, with fry receiving continued male protection.^[48] In African species like Papyrocranus afer, development follows a comparable pattern, though specific hatching times are less documented.^[52] Reproduction in Notopteridae is seasonal, peaking during rainy periods that increase water levels and food availability, though some tropical populations may breed year-round in stable environments. For Notopterus notopterus, spawning aligns with the monsoon season (June-August) in natural habitats like reservoirs.^[50] In Papyrocranus afer, gonadal development and spawning peak in July during the rainy season in West African lagoons.^[52] Chitala species in Southeast Asia similarly time breeding to wet seasons from March to July.^[53]

Fossil record

Known fossils

The fossil record of Notopteridae is limited and fragmentary, reflecting the family's elongate and delicate body plan, which rarely preserves as complete skeletons; instead, remains consist predominantly of otoliths, isolated vertebrae, skull fragments, and braincases. The earliest evidence comes from Palaeonotopterus greenwoodi, known from isolated braincases and fragmentary skull elements, including parasphenoid tooth plates, collected from the Cenomanian-aged Tegana Formation (Kem Kem Beds) in southern Morocco, dated to approximately 100–94 million years ago.^[54] These remains, first described from a single braincase and later supplemented by additional fragments, exhibit osteoglossomorph features such as a derived hyomandibular articulation, supporting its placement within Notopteridae despite incomplete preservation. This specimen represents the oldest definitive record of the family. A later Cretaceous otolith attributed to Notopteridarum nolfi was recovered from the Maastrichtian Deccan Intertrappean Beds near Rangapur, India, dated to approximately 70.6–66 million years ago. This specimen was initially described based on its morphological similarity to modern notopterid otoliths, though later revisions confirmed its attribution while noting the overall paucity of pre-Tertiary material.^[55] Tertiary fossils are similarly sparse but more widespread, including scattered otoliths and vertebrae from Eocene to Miocene deposits across Asia and Africa. A notable example is Notopterus primaevus from the Eocene Sangkarewang Formation in Sumatra, Indonesia, represented by an incompletely prepared near-complete body fossil that closely resembles the living genus Notopterus in its vertebral column and fin structure.^[56] Other Tertiary records comprise isolated otoliths from Paleogene and Neogene sediments in regions like the Indian subcontinent and North Africa, underscoring the family's persistence in freshwater environments post-Cretaceous. At least two extinct genera are recognized in the fossil record: Palaeonotopterus and Notopteridarum, highlighting a modest diversity of stem-group taxa alongside the radiation of modern lineages.

Paleobiogeography

The Notopteridae exhibit a fossil record that points to a Gondwanan origin, with early diversification occurring in the Cretaceous period across landmasses corresponding to modern Africa and India. The earliest known notopterid fossil, Palaeonotopterus greenwoodi, dates to the Cenomanian stage (~94 Ma) from deposits in Morocco, suggesting an initial African center of origin for the family within the broader Osteoglossiformes clade. This aligns with molecular estimates placing the divergence of notopterid lineages in the Early Cretaceous, prior to the full fragmentation of Gondwana.^[58] Vicariance events associated with the breakup of Gondwana are implicated in the separation of African and Asian notopterid lineages around 100 million years ago, coinciding with the rifting of the India-Madagascar block from Africa. Indian subcontinent fossils, including notopterid otoliths from Late Cretaceous (~66 Ma) deposits, provide evidence linking early African forms to the subsequent radiation of Asian genera such as Notopterus and Chitala. These otoliths indicate stem-group notopterids persisted on the drifting Indian plate, supporting a vicariant model where tectonic drift facilitated isolation and independent evolution on separated continents.^[59] The fossil record of Notopteridae is notably sparse during the Paleogene, with few documented specimens bridging the Late Cretaceous and Eocene, which implies the family's survival through the Cretaceous-Paleogene (K-Pg) extinction event approximately 66 Ma. This endurance is likely attributable to their air-breathing adaptations, including a supra-branchial organ, which would have enabled tolerance of hypoxic or anoxic conditions prevalent post-extinction in freshwater habitats. The reappearance of definitive notopterid fossils in the Eocene of Sumatra (Notopterus primaevus) underscores this gap, highlighting periods of low preservation or rarity in the interim. Fossil evidence corroborates the independent evolution of African (Papyrocranus, Xenomystus) and Asian lineages following their vicariant split, with no intermediate forms indicating post-separation convergence in morphology or ecology. Alternative hypotheses propose limited dispersal across marine barriers, such as via the Tethys Sea during the Eocene, potentially explaining minor overlaps in distribution patterns, though molecular data favor primarily vicariant processes over long-distance migration.^[58]^[59]

Diversity

African genera

The African genera of Notopteridae comprise Xenomystus and Papyrocranus, both restricted to freshwater systems in equatorial Africa and characterized by elongate, knife-like bodies lacking a dorsal fin, obligate air-breathing capabilities, and electroreceptive sensory systems adapted for detecting weak electric fields in murky waters.^[60] These genera diverged from Asian notopterids in the Early Cretaceous, with their distributions shaped by Africa's ancient river basins.^[61] Unlike electrogenic fishes such as mormyrids, African notopterids possess specialized medullary structures for passive electroreception but lack electric organs for generating discharges.^[62]^[63] Xenomystus is monotypic, represented solely by X. nigri (African knifefish), a nocturnal species reaching a maximum standard length of 30 cm.^[35] It inhabits quiet, vegetated waters in coastal basins from Senegal to Cameroon and inland drainages including the Nile, Chad, Niger, Ogooué, and Congo basins, preferring demersal habitats with pH 6.0–8.0 and temperatures of 22–28°C.^[35] This species feeds primarily at dusk or night on invertebrates such as worms, crustaceans, insects, and snails, emerging from cover to hunt while using electroreception and large eyes for navigation in low-light conditions.^[35] It produces barking sounds via its swim bladder and surfaces periodically to gulp air, aiding survival in low-oxygen environments.^[64] Females deposit 150–200 eggs (2 mm diameter) in vegetation or on surfaces, with males providing parental care by guarding and fanning the eggs.^[40] X. nigri holds minor economic value in the aquarium trade due to its hardy nature and unique appearance, though overcollection poses a localized threat alongside habitat degradation from pollution and deforestation.^[64] Its conservation status is Least Concern globally, reflecting a wide but patchy distribution, though populations in fragmented West African coastal rivers may warrant monitoring.^[64] Papyrocranus includes two species: P. afer (reticulate knifefish) and P. congoensis (Congo knifefish), both scaly and adapted to dense, vegetated, slow-flowing rivers where they use undulating anal fins for forward and backward propulsion. P. afer attains up to 80 cm total length and 1.3 kg, featuring 35–45 ventral spines, a head 5.0–5.1 times standard length, and caudal vertebrae numbering 63–69.^[65] It occurs in West African systems, including the upper Niger, Benue, Niger Delta, and coastal basins from Senegal and Gambia to the Tano and Pra rivers in Ghana, but is absent from the Volta, Chad, Ogooué, and Kwilu-Niari basins.^[65] This demersal species thrives in tropical waters (24–30°C, pH 6.5, dH 10) and supports subsistence fisheries for its flesh. P. congoensis, smaller at a maximum 29.2 cm standard length, is confined to the Central African Congo basin, including the Cuvette Centrale, Ubangi tributaries, lower Congo, and Kouilou-Niari, excluding the upper Congo above Kisangani and Cameroonian sectors.^[66] Both species exhibit anal fin rays of 104–118 and 52–58 caudal vertebrae, respectively, and rely on electroreception for foraging in turbid habitats.^[66] They face threats from habitat loss due to logging, agriculture, and dam construction in equatorial forests, which fragment riverine ecosystems, though their status remains Least Concern owing to relatively stable populations.^[67]^[68] The genus contributes modestly to the aquarium trade, particularly juveniles of P. afer, but wild collection is limited compared to Asian congeners.^[67]

Asian genera

The Asian genera of the family Notopteridae, Chitala and Notopterus, are native to freshwater habitats ranging from the Indian subcontinent through mainland Southeast Asia to the Indonesian islands of Sumatra, Java, and Borneo.^[1] These fishes are characterized by their elongated, knife-like bodies, a reduced dorsal fin positioned posteriorly, and a long anal fin that extends along much of the ventral surface, often confluent with a small caudal fin.^[1] Unlike the African genera, they lack electric organs and rely on air-breathing via a physostome swim bladder connected to the gut, enabling survival in low-oxygen waters.^[69] Spawning generally occurs during the rainy monsoon season, from May to August, with adhesive eggs guarded by males.^[70] The genus Chitala, commonly known as Asian knifefishes or featherbacks, encompasses six species, several of which reach large sizes and hold commercial value in aquaculture, food markets, and the aquarium trade.^[71] These include C. chitala (clown knifefish, up to 122 cm SL, native to rivers in India, Bangladesh, and Myanmar), C. ornata (clown featherback, up to 100 cm SL, found in Southeast Asian basins including the Mekong and Chao Phraya), C. hypselonotus (up to 100 cm TL, restricted to Sumatra and Borneo), C. borneensis (up to 39.3 cm SL, endemic to Borneo), C. blanci (Indochina featherback, up to 120 cm SL, in the Mekong basin), and C. lopis (giant featherback, up to 150 cm SL, historically from Java, Sumatra, Borneo, and Thailand).^[71] C. chitala and C. ornata are particularly prized for their size and taste in local fisheries, while C. ornata is a popular ornamental species due to its striking black-and-white banding; both have been introduced outside their native range, such as C. ornata in Florida, USA, where established populations exist.^[53] Conservation statuses vary: C. chitala is Near Threatened due to overfishing and habitat loss, C. blanci is Near Threatened from dam construction and exploitation in the Mekong, C. lopis was assessed as Extinct in 2019 but was rediscovered in 2023, prompting calls for reassessment (status remains Extinct as of 2025), and the remaining species (C. ornata, C. hypselonotus, C. borneensis) are Least Concern, though regional declines from overharvesting occur.^[70]^[72]^[73]^[74] The genus Notopterus contains a single recognized species, N. notopterus (bronze featherback), though genetic studies indicate it may comprise two allopatric species diverging by approximately 7.5% in mitochondrial DNA, warranting taxonomic revision.^[75]^[4] This species attains a maximum length of 60 cm SL and is widely distributed across South and Southeast Asia, from Pakistan and India to Indonesia, inhabiting rivers, floodplains, and swamps.^[75] It supports important fisheries and aquaculture in countries like Thailand and Indonesia, valued for its fast growth and mild flavor.^[61] N. notopterus is listed as Least Concern globally, with stable populations, though local overfishing poses risks in some areas.

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[17]
From Chromosomes to Genome: Insights into the Evolutionary ...
Their precise phylogenetic position is unresolved, although they probably belong to stem species of Notopteridae. The oldest (and only) fossil assignable to ...
[18]
Emerging patterns of genome organization in Notopteridae species ...
Feb 4, 2019 · Phylogenetic studies support the monophyletic status of Notopteridae, placing them within the suborder Notopteroidei, together with ...
[19]
https://pmc.ncbi.nlm.nih.gov/articles/PMC6361938/
[20]
[PDF] NOTOPTERIDAE) FROM THE PASAK RIVER, CHAO PHRAYA ...
Specimens of a featherback, Chitala lopis (Actinopterygii: Notopteridae), were collected from the Pasak River (Chao Phraya River system), central Thailand, ...
[21]
Chitala chitala summary page
### Morphological Description of Chitala chitala
[22]
Notopterus - Bronze featherback - FishBase
Teleostei (teleosts) > Osteoglossiformes (Bony tongues) > Notopteridae (Featherbacks or knifefishes) > Notopterinae ... Systematic revision of the old ...
[23]
RESPIRATORY FUNCTION OF THE SWIMBLADDER OF ...
The large swimbladder with a wide pneumatic duct is reported to function as an accessory respiratory organ. Circulation of blood is modified to discharge this ...
[24]
Swim-Bladder: Development, Structure and Types | Fishes
Swim-Bladder acts as Adjustable Float; Maintains Proper Centre of Gravity; Swim-Bladder helps in Respiration; Swim-Bladder as Resonator. 1. Introduction to Swim ...
[25]
https://zslpublications.onlinelibrary.wiley.com/doi/10.1111/j.1469-7998.1962.tb01834.x
[26]
[PDF] Electrocommunication in teleost fishes. Behavior and experiments
Gymnarchus has a swim bladder adapted for atmospheric respiration (with a modified circulatory system). The fish gulps air at the surface, often near fal-.
[27]
Swimming in the electric eels and knifefishes
Notopterus notopterus and Notopterus chilata employ the body and anal fin as a single propulsive unit and generate a body wave. Rapid bouts of burst swimming ...
[28]
Swimming in the electric eels and knifefishes - ResearchGate
Aug 6, 2025 · The kinematics of steady forward swimming in six species of Gymnotidae and three species of Notopteridae are described.
[29]
Aquatic Oxygen Uptake of the Freshwater Dual Breathing ...
Aug 7, 2025 · Oxygen uptake (VO 2) through the gills in relation to body weight has been studied in Notopterus notopterus at 24.0±1.0ºC under continuous ...
[30]
Evaluating establishment success of non-native fishes introduced to ...
Aug 26, 2021 · Hypoxia tolerance score: 1 – intolerant of low dissolved oxygen, 2 – moderately tolerant, 3 – are air-breathers. Closed circles represent number ...<|control11|><|separator|>
[31]
Papyrocranus afer - Facts, Diet, Habitat & Pictures on Animalia.bio
This fish is native to tropical West Africa. It occurs in the Niger basin and other coastal river basins including the Tano River and Pra River basins in Ghana, ...
[32]
[PDF] Reproductive parameters of Reticulate Knife fish, Papyrocranus afer ...
Apr 15, 2019 · Two genera, Papyrocranus and. Xenomystus are endemic to West and Central Africa while the remaining two (Chitala and Notopterus) are endemic ...
[33]
Genetic evidence for the recognition of two allopatric species of ...
Jul 1, 2020 · The fish genus Notopterus Lacepède, 1800 (Notopteridae) currently includes only one species, the Asian bronze featherback Notopterus ...
[34]
[PDF] The non-native freshwater fishes of Singapore
Apr 16, 2020 · Of the 123 species, 42 species (~34%) from 15 families have established breeding populations in Singapore, with the dominant family being the ...
[35]
https://www.fishbase.se/summary/Xenomystus-nigri
[36]
Bronze featherback • Notopterus notopterus • Fish sheet - Fishipedia
Notopterus notopterus inhabits lakes, flooded plains, canals, ponds, oxbow lakes, as well as estuarine areas with brackish water. The species also colonizes ...
[37]
Xenomystus nigri (African Knifefish) - Seriously Fish
Habitat. Said to be found most often in stiller, overgrown waters. Its sizeable natural range, various biological adaptations and ease of captive maintenance ...
[38]
African knifefish • Xenomystus nigri • Fish sheet - Fishipedia
This animal evolves in areas characterized by a strong presence of vegetation (aquatic and marsh plants, decaying organic matter, roots...). Main ...
[39]
[PDF] FOOD AND FEEDING HABITS OF THREATENED NOTOPTERUS ...
As shown in Fig. 1-3, the preferred diets were crustaceans, insects, molluscs, minnows and fishes. Although there were some differences in the diet ...Missing: sources | Show results with:sources
[40]
[PDF] Zooplankton as dietary component of selected freshwater fish - Bioflux
Feb 25, 2018 · Irrespective of the fish species, in all instances, cladocerans, copepods and rotifers were found to be common dietary elements in the fish gut ...
[41]
food and feeding habits of a feather back, notopterus notopterus ...
The principal foods were graded as: bivalve molluscs, animal remains ... References (0). Reproduction and development of the asian bronze featherback ...
[42]
Chitala ornata, Clown featherback : fisheries, aquarium - FishBase
A predator on surface-feeding fishes, crustaceans and insects, with a crepuscular or nocturnal activity pattern. Moves into inundated forest during the high ...
[43]
Comparable Ages for the Independent Origins of Electrogenesis in ...
May 14, 2012 · ... within the Osteoglossomorpha is the family Notopteridae (BP = 100%). ... Phylogenetic position of the enigmatic Lepidogalaxias ...<|separator|>
[44]
tongue-bite apparatus» in the knifefish family Notopteridae ...
Osteoglossomorph fishes are unique in possessing a specialized feeding mechanism, the tongue-bite apparatus (TBA) involving the hyoid apparatus.
[45]
[PDF] CHITOL: Chitala chitalaI Hamilton, 1822
Breeds between June and July by building nest and eggs are 3-4.5 mm in diameter (Rahman, 1989). Eggs are 3-4.5 mm in diameter and young receive parental care ( ...
[46]
(PDF) Growth coefficient and fecundity of Chitala ... - ResearchGate
Aug 6, 2025 · Mean absolute fecundity was calculated as 13431±2144 in sampled fish. To the best of authors' knowledge, length-weight relationship and ...
[47]
(PDF) Reproduction and development of the asian bronze ...
Aug 6, 2025 · Reproduction and development of the asian bronze featherback Notopterus notopterus (Pallas, 1769) (Osteoglossiformes, Notopteridae) in captivity.
[48]
(PDF) Reproductive Biology of Invasive Knifefish (Chitala ornata) in ...
Jul 4, 2019 · They spawned fewer and larger eggs than native fish species in the lake and provides parental care. The adult sex ratio was male skewed ...
[49]
[PDF] ERSS - Clown Knifefish (Chitala ornata)
The piscivorous and invertivorous Chitala ornata is native to Southeast Asia. The species has become established in the United States in the state of Florida, ...
[50]
https://www.researchgate.net/publication/323682803_Reproduction_and_development_of_the_asian_bronze_featherback_Notopterus_notopterus_Pallas_1769_Osteoglossiformes_Notopteridae_in_captivity
[51]
https://www.researchgate.net/publication/334224509_Reproductive_Biology_of_Invasive_Knifefish_Chitala_ornata_in_Laguna_de_Bay_Philippines_and_its_Implication_for_Control_and_Management
[52]
https://www.globalscienceresearchjournals.org/articles/reproductive-parameters-of-reticulate-knife-fish-papyrocranus-afer-gunther-1868-from-lekki-lagoon-lagos-nigeria.pdf
[53]
African, but Not Asian, Notopterid Fishes Are Electroreceptive
The unexpected finding of this structure in the medulla of the African members of the family Notopteridae strongly suggests that they are electroreceptive.Missing: electrogenic | Show results with:electrogenic
[54]
Genetic and morphological evidence of a single species of bronze ...
The origin of the Notopteridae is tracing back to the Early Cretaceous when the African (Papyrocranus and Xenomystus) and Asian (Notopterus, Chitala) clades ...Missing: etymology | Show results with:etymology
[55]
African, but not Asian, notopterid fishes are electroreceptive
The unexpected finding of this structure in the medulla of the African members of the family Notopterdae strongly suggests that they are electroreceptive.Missing: electrogenic | Show results with:electrogenic
[56]
Comparable Ages for the Independent Origins of Electrogenesis in ...
Whereas the African Notopteridae and Siluriformes are only passively electroreceptive, the Gymnotiformes and the Mormyroidea secondarily and independently ...Missing: conservation threats
[57]
Xenomystus nigri summary page
### Summary of Xenomystus nigri (African Knifefish)
[58]
https://doi.org/10.1016/j.ympev.2009.01.020
[59]
Papyrocranus afer, Reticulate knifefish : fisheries, aquarium - FishBase
With 35-45 ventral spines; head 5.0-5.1 times SL; eye diameter 4.6-5.3 times head length; length of nasal tentacle twice the eye diameter.Missing: electric organs
[60]
Papyrocranus congoensis summary page
- **Description**: Papyrocranus congoensis is a freshwater fish in the Notopteridae family (featherbacks or knifefishes), subfamily Xenomystinae. Obligate air-breathing. Anal soft rays: 104-118; caudal vertebrae: 52-58.
[61]
https://www.sciencedirect.com/science/article/pii/S2351989423004213
[62]
https://www.sciencedirect.com/science/article/pii/0304394082902257
[63]
(PDF) Growth and survival of bronze featherback (Notopterus ...
Oct 9, 2023 · The aim of this study was to determine the survival and growth of N. notopterus reared in bucket and fed with different diets.<|control11|><|separator|>
[64]
Chitala chitala, Clown knifefish - FishBase
IUCN Red List Status (Ref. 130435: Version 2025-1). Near Threatened (NT) ; Date assessed: 28 May 2010. CITES. Not Evaluated. CMS (Ref. 116361). Not Evaluated ...
[65]
Fish Identification
### Summary of Chitala Species (Notopteridae)
[66]
https://www.fishbase.se/summary/Papyrocranus-congoensis
[67]
Chitala lopis, Giant featherback : fisheries - FishBase
Considered extinct by IUCN in 2019. Asia: Borneo, Sumatra and Java in Indonesia, Malaya, and Thailand (Ref. 7050). Found in the Mekong basin (Ref. 12693).
[68]
Rediscovery of the giant featherback Chitala lopis (Notopteridae) in ...
However, the taxonomy of Chitala species is unclear, with 3 nominal species (C. lopis, C. hypselonotus and C. borneensis) historically synonymized under a.<|control11|><|separator|>
[69]
Fish Identification
### Summary of Notopterus Species from Notopteridae