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Anabantoidei

Anabantoidei, commonly known as labyrinthfishes, is a suborder of ray-finned fishes within the order Anabantiformes, distinguished by the possession of a unique labyrinth organ—a modified gill arch that functions as an accessory air-breathing structure, enabling survival in oxygen-poor waters.^[1]^[2]^[3] This suborder comprises three families: Anabantidae (climbing gouramies, 33 species), Helostomatidae (kissing gouramis, 1 species), and Osphronemidae (gouramies and related fishes, approximately 135 species), totaling around 169 species across 19 genera.^[1]^[2]^[3]^[4]^[5] These primary freshwater fishes exhibit a disjunct distribution across Africa and South and Southeast Asia, with biogeographic patterns potentially linked to the Gondwanan breakup or later dispersal events.^[6]^[2] Species sizes vary dramatically, from the diminutive Parosphromenus ornaticauda at about 19 mm to the giant gourami Osphronemus goramy reaching up to 70 cm.^[2] Notable adaptations include diverse reproductive strategies, such as bubble-nest building, mouthbrooding, and substrate guarding, with parental care evolving independently at least three times within the group.^[6]^[2] Many species, particularly in the aquarium trade, are popular for their vibrant colors and behaviors, though habitat loss poses conservation challenges for several taxa.^[2]

Taxonomy and classification

Definition and characteristics

Anabantoidei is a suborder of ray-finned fishes within the order Anabantiformes, comprising 174 species distributed across three families and 18 genera.^[7] These primary freshwater fishes are characterized by their monophyletic lineage, distinguished from other anabantiforms by specific morphological and physiological adaptations that enable survival in challenging aquatic environments.^[6] The broader order Anabantiformes encompasses eight families, 26 genera, and around 287 species, with Anabantoidei forming the core group alongside the suborder Channoidei.^[7] Members of Anabantoidei exhibit a general perch-like body plan, with forms ranging from elongated to disc-shaped, and sizes varying significantly from about 2 cm in small species such as certain Bettas to up to 70 cm in the giant gourami Osphronemus goramy.^[5] This diversity in body morphology supports their adaptation to diverse freshwater habitats, where a key feature is their air-breathing capability via a specialized labyrinth organ, allowing supplemental respiration in low-oxygen waters and differentiating them from strictly gill-dependent fishes.^[8] The temporal range of Anabantoidei extends from the Eocene epoch to the present, reflecting their evolutionary persistence as primary freshwater inhabitants with a disjunct distribution across sub-Saharan Africa and southern to southeastern Asia.^[9] This biogeographic pattern underscores their ancient origins and adaptation to isolated continental freshwater systems.^[6]

Families and genera

The suborder Anabantoidei comprises three primary families: Anabantidae, Helostomatidae, and Osphronemidae.^[3]^[4]^[5] Recent phylogenetic studies based on molecular data have consolidated this classification, merging the former family Belontiidae into Osphronemidae and recognizing only these three families within the suborder, superseding earlier schemes that divided Anabantoidei into five families (including separate Belontiidae and Badidae).^[2]^[6] As of August 2025, the suborder includes 174 species, reflecting ongoing discoveries particularly in Osphronemidae.^[7] The family Anabantidae, known as climbing perches or climbing gouramies, includes 33 species across 4 genera: Anabas, Ctenopoma, Microctenopoma, and Sandelia.^[3] These genera are distributed in freshwater habitats of tropical Africa and Asia, with Anabas comprising 3 species noted for their climbing abilities.^[3] The family exhibits moderate diversity, with many species showing adaptations for overland movement. Helostomatidae is a monotypic family containing a single species, Helostoma temminckii, the kissing gourami, in the genus Helostoma.^[4] Native to Southeast Asia, this species is distinguished by its unique feeding behavior and is the sole representative of its family. The largest family, Osphronemidae (gouramies, bettas, and allies), encompasses 140 species in 14 genera, including Betta, Trichopodus, Osphronemus, Macropodus, Trichopsis, Belontia, Parosphromenus, Pseudosphromenus, Trichogaster, Ctenops, Sphaerichthys, Parasphaerichthys, Luciocephalus, and Malpulutta.^[5]^[7] Key genera include Betta (fighting fishes, over 75 species), Trichopodus (gouramies, 5 species), and Osphronemus (giant gouramies).^[5] This family dominates the suborder's diversity, particularly in Southeast Asia, where high endemism is evident among its genera. Overall, Anabantoidei includes 174 species across these families, with the majority concentrated in Southeast Asian freshwaters.^[7]

Family	Genera Count	Species Count	Key Genera Examples	Distribution Focus
Anabantidae	4	33	Anabas (3 spp.), Ctenopoma	Tropical Africa, Asia
Helostomatidae	1	1	Helostoma	Southeast Asia
Osphronemidae	14	140	Betta (>75 spp.), Trichopodus (5 spp.), Osphronemus	Primarily Southeast Asia

Anatomy and physiology

Labyrinth organ

The labyrinth organ, a defining feature of the Anabantoidei suborder, is a highly vascularized, sponge-like structure derived from the modified epibranchial bone of the first gill arch. It is housed within paired suprabranchial chambers located above the gills in the opercular cavity, forming a series of intricate, plate-like folds or chambers lined with thin respiratory epithelium that greatly expands the surface area for gas exchange. This labyrinthine arrangement consists of bony or cartilaginous plates supported by connective tissue, permeated by a dense network of blood vessels that facilitate oxygen diffusion from inhaled air.^[10]^[11] Development of the labyrinth organ occurs post-hatching and is not functional at birth, with juveniles initially relying on gill-based aquatic respiration. In species such as Betta splendens and Trichopodus trichopterus, the organ begins to form within the first few weeks, transitioning to supplemental air-breathing capability around 32–42 days post-fertilization, influenced by environmental oxygen levels; intermittent hypoxia can accelerate this onset by up to 4 days in B. splendens. The structure matures over several weeks to months, sometimes even after sexual maturity, as capillary density and epithelial complexity increase to support efficient aerial respiration.^[12] The primary function of the labyrinth organ is to enable supplemental aerial respiration, allowing Anabantoidei to gulp air at the surface and extract oxygen through the vascularized folds, thereby sustaining survival in hypoxic aquatic environments or during brief periods of emersion—up to several hours if the substrate remains moist. Unlike true lungs, it operates via a double-pump mechanism involving buccal and opercular movements to draw in and distribute air, providing an efficient but accessory means of O₂ uptake that complements reduced gill function. This adaptation is crucial for inhabiting oxygen-poor waters, though the organ's efficiency varies with its developmental stage and environmental demands.^[11]^[10] Variations in the labyrinth organ occur across Anabantoidei species, with size and complexity often larger and more elaborate in those inhabiting low-oxygen, stagnant waters to enhance O₂ extraction, while reduced or simpler forms appear in species from oxygen-rich, fast-flowing habitats, such as certain members of the Osphronemidae family. Epithelial thickness and fold density also differ phylogenetically and by location within the organ. The organ is absent in non-Anabantoidei anabantiform fishes, underscoring its role as a suborder-specific innovation.^[11]^[13]

Body structure and other adaptations

Members of the suborder Anabantoidei display a diverse array of body morphologies, ranging from compact, orbicular forms to highly elongate, torpedo-like shapes adapted to various aquatic niches. For instance, species in the genus Betta exhibit moderately deep bodies with a Vertebrate Shape Index (VSI) of approximately 40–80, while the pikehead gourami Luciocephalus pulcher represents an extreme, with a VSI of 153–163 achieved through rapid elongation of the head and precaudal region via increased vertebral count and postorbital bone lengthening.^[14] Their scales are typically strongly ctenoid, providing a rough texture that aids in movement through vegetated or structured habitats, as observed in genera like Microctenopoma and Ctenopoma. Mouth position varies among species, from terminal in surface-oriented forms like Trichogaster to inferior in bottom-dwelling ones like Anabas, reflecting dietary specializations from planktonic feeding to predation on larger prey.^[15] Sensory adaptations in Anabantoidei include a well-developed lateral line system, consisting of neuromasts along the body that detect hydrodynamic vibrations and water movements, enabling precise navigation in low-visibility environments such as dense vegetation or murky waters.^[16] In climbing species like the climbing perch Anabas testudineus, robust pectoral fins facilitate terrestrial locomotion and ascent of obstacles, allowing the fish to traverse land between water bodies during dry periods, though pelvic fins play minimal role in this process.^[17] Physiologically, Anabantoidei demonstrate notable tolerance to environmental stressors, including low pH levels and temperature fluctuations common in their tropical freshwater habitats. Species such as Betta splendens thrive in pH ranges from 5.0 to 8.0 and temperatures of 24–30°C, with juveniles particularly resilient to rapid shifts that would stress non-adapted teleosts.^[18] Additionally, some species engage in cutaneous respiration through their highly vascularized skin during emersed states or hypoxic conditions in juveniles, supplementing aquatic gas exchange and aiding survival on moist substrates.^[19] This skin-based oxygen uptake supports early development prior to full labyrinth organ functionality. Sexual dimorphism is prevalent across Anabantoidei, with males typically exhibiting brighter coloration, larger body sizes, and elongated unpaired fins for courtship displays. In Betta splendens, males possess extended dorsal, anal, and caudal fins up to several times the length of females', along with vivid, uniform body hues, contrasting with the shorter fins and subdued patterns of females.^[20] Similarly, in Trichogaster species, males develop longer dorsal fins reaching the caudal peduncle and more intense blue-red pigmentation compared to females.^[15] This dimorphism extends to genera like Ctenopoma, where males are larger overall and display more iridescent green tones on the body and fins.^[21]

Distribution and habitat

Geographic range

The Anabantoidei display a characteristic disjunct distribution pattern, with native populations confined to freshwater systems in sub-Saharan Africa and across South, Southeast, and parts of East Asia, but absent from regions such as Madagascar, the Americas, and Australia.^[6] In Asia, the suborder's primary range spans from South Asia—including India and Sri Lanka—through Southeast Asia to the Philippines and Indonesian archipelago, encompassing countries like Pakistan, Myanmar, Thailand, Malaysia, and Indonesia.^[22] Some species extend into East Asia, with records in southern China, Japan, and Korea, exemplified by the round-tailed paradise fish (Macropodus ocellatus).^[23] This Asian distribution reflects a core area of diversification, where the suborder achieves its highest species richness. Africa hosts a more restricted subset of the suborder, primarily the subfamily Ctenopominae within Anabantidae, distributed in sub-Saharan river basins such as the Congo, with genera like Ctenopoma (18 species) and Microctenopoma (12 species) representing key components.^[24]^[25]^[26] Endemism hotspots occur within the Indonesian islands, particularly Borneo and Sumatra, where diversity is elevated; Sumatra alone records 32 anabantoid species, including multiple endemics in the genus Betta (with at least 21 species documented), underscoring the archipelago's role as a center of speciation.^[27]^[28] Human-mediated introductions have expanded the suborder beyond its native range, notably the climbing perch (Anabas testudineus), which has established invasive populations in Papua New Guinea, the Torres Strait Islands of Australia, and parts of the Americas, including Colombia and the United States, primarily through the aquarium trade and accidental releases.^[29]^[30]^[31] Similarly, the Siamese fighting fish (Betta splendens) has become invasive in subtropical waters of South America and the United States.^[32] Regarding historical biogeography, a Gondwanan vicariance origin was initially hypothesized to explain the African-Asian disjunction, but molecular phylogenetic analyses, incorporating mitochondrial and nuclear DNA sequences, support an Asian cradle of diversification followed by dispersal to Africa during the Late Mesozoic to Early Tertiary period, with lineage divergence estimated at 40–60 million years ago—too recent for Gondwanan breakup.^[6]^[24]

Habitat preferences

Anabantoidei species predominantly inhabit stagnant or slow-flowing shallow freshwater environments, such as swamps, rice paddies, and lowland rivers, where dissolved oxygen levels are often critically low, typically below 2 mg/L.^[2]^[33] These waters maintain warm temperatures ranging from 22°C to 30°C and exhibit a broad pH tolerance from acidic (around 4) to neutral (up to 8), allowing adaptation to varied chemical conditions in tropical ecosystems.^[2]^[34] The labyrinth organ enables survival in these hypoxic conditions by facilitating aerial respiration.^[2] Within these habitats, Anabantoidei favor microhabitats characterized by dense aquatic vegetation, including floating plants like water hyacinth and submerged roots, which provide essential cover from predators and resting sites near the water surface.^[35]^[36] Certain species, such as climbing perches (Anabas testudineus), utilize marginal vegetation and land bridges formed by overhanging plants or moist substrates to traverse dry land during migrations between water bodies.^[37] In acidic peat swamp forests, genera like Betta thrive in blackwater environments with soft, humic-stained waters and tangled root systems, where pH can drop to 3.5–4.0, supporting specialized acid-tolerant physiologies.^[38]^[39] Many Anabantoidei demonstrate remarkable adaptations to environmental extremes, including drought resistance through burrowing into mud aestivation, as seen in Anabas species that encase themselves in secreted mucus cocoons to endure prolonged dry periods.^[37] Populations in floodplain regions also exhibit flood tolerance, synchronizing life cycles with seasonal inundations and receding waters to exploit ephemeral habitats.^[2] However, these specialized habitats face significant threats from deforestation and drainage for agriculture and mining, which fragment swamp forests, alter hydrology, and reduce available stagnant, vegetated refugia, exacerbating vulnerability for acid-tolerant and hypoxia-adapted species.^[40]^[41]

Ecology and behavior

Feeding habits

Members of the Anabantoidei suborder exhibit predominantly carnivorous and insectivorous diets, consuming small aquatic invertebrates such as zooplankton, insect larvae, and occasionally small fish.^[42] Species like the African bush fish (Ctenopoma acutirostre) display adaptations for carnivory, including a shorter intestinal length relative to standard length (IL/SL ratio of 0.74) and robust jaw structures suited for capturing live prey.^[43] Juveniles are typically planktivorous, relying on zooplankton, while adults shift toward piscivory and broader invertivory as they grow.^[42] Some taxa, such as those in the genus Trichogaster, incorporate herbivorous elements, feeding on algae and diatoms alongside animal matter, reflecting opportunistic omnivory in mature individuals.^[44] Foraging strategies vary across the suborder, often leveraging the labyrinth organ's accessory respiration to access prey in low-oxygen environments. Surface-oriented species, including dwarf gourami (Trichogaster lalius) and pearl gourami (Trichopodus spp.), employ water-spitting jets to dislodge aerial insects from overhanging vegetation, enabling short-range hunting with precision strikes.^[45] In contrast, bottom-dwelling members of the Anabantidae, such as the climbing perch (Anabas testudineus), forage along substrates using food-stocking behaviors where prey items are temporarily held in the mouth before swallowing, facilitating efficient capture of benthic invertebrates and detritus.^[46] Aerial prey capture through leaps is observed in several taxa, allowing access to insects above the water surface beyond spitting range. Within freshwater food webs, Anabantoidei occupy mid-level trophic positions as predators of primary consumers, controlling invertebrate populations while serving as prey for larger piscivores. This role is exemplified by their diet's emphasis on zooplankton and insects in juveniles, transitioning to small fish in adults, which influences community dynamics in tropical Asian and African habitats.^[42] Dietary variations highlight ecological specialization; the giant gourami (Osphronemus goramy) is largely herbivorous, grazing on aquatic plants and algae, supported by a digestive tract adapted for plant material processing.^[47] Conversely, Betta species function as ambush predators, lying in wait among vegetation to strike at passing invertebrates or small fish with rapid lunges.^[48] These adaptations underscore the suborder's diversity in exploiting niche resources.^[49]

Reproduction and parental care

Anabantoidei exhibit remarkable diversity in reproductive strategies, with over eight distinct modes ranging from free-spawning without care to elaborate forms involving complex parental investment, a variation that exceeds that seen in many other fish suborders. This diversity is phylogenetically structured, with parental care recognized as a plesiomorphic trait present in 16 of the 19 genera, having evolved independently at least three times and correlating strongly with clade-specific adaptations.^[50] Such strategies enhance offspring survival in often hypoxic tropical freshwater environments, where the labyrinth organ aids early fry in accessing atmospheric oxygen during low-oxygen brooding periods. Common reproductive modes include bubble-nest building, observed in genera such as Betta and Trichopodus, where males construct floating foam nests from air bubbles and oral secretions to house adhesive eggs. In these species, the male fertilizes eggs released by the female beneath the nest, collects them into the foam, and provides care by guarding and fanning the clutch to ensure oxygenation, typically for 2-3 days until hatching, after which fry remain under protection for up to a week. Mouthbrooding represents another advanced mode, practiced by certain Osphronemidae like Sphaerichthys osphromenoides and some Betta species (e.g., Betta macrostoma), where males or sometimes both parents incubate eggs and early fry in their mouths for periods extending 10-14 days, releasing them once independent. Substrate spawning with care occurs in taxa such as Sandelia, involving males guarding adhesive eggs deposited on leaves or vegetation, fanning them vigorously for oxygenation over 1-4 weeks until fry achieve independence.^[50] Less derived modes, like free-spawning without care in Anabas and Ctenopoma, involve pelagic or demersal egg release into open water, though even here, infanticide risks underscore the adaptive value of care in related clades. Parental care in Anabantoidei is predominantly male-mediated, a pattern that aligns with the suborder's phylogenetic history, where males fan eggs to prevent fungal growth and hypoxia while defending against predators.^[50] This investment lasts from hatching to fry dispersal, often 1-4 weeks depending on species and environmental conditions, allowing larvae to develop the functional labyrinth organ for air-breathing. In nest-building forms, males may provision fry with mucus or regurgitated food, while mouthbrooders ensure constant humidity and protection during incubation.^[50] Biparental care appears in select cases, such as Osphronemus, but male dominance prevails across most genera, reflecting evolutionary pressures for efficient resource allocation in variable habitats. Breeding is typically triggered by environmental cues like monsoonal rains and elevated temperatures (above 25°C), which simulate flooding and increase oxygen availability in shallow waters, prompting gonadal maturation. Courtship displays vary but commonly feature male fin flaring, opercular extensions, and chasing to attract females and establish territory, as seen in Betta and Trichopodus, where these behaviors escalate into spawning embraces.^[8] This high variation in modes and care, basal nest-building in some clades like Anabantidae, underscores the suborder's adaptive radiation tied to Southeast Asian and African freshwater ecosystems.^[50]

Evolutionary history and phylogeny

Fossil record

The fossil record of Anabantoidei is notably sparse, primarily due to the challenges of taphonomy in freshwater environments, which limits preservation of complete skeletons.^[6] The earliest known fossils date to the probable Eocene deposits of the Sangkarewang Formation in the Ombilin Basin of Sumatra, Indonesia, approximately 50 million years ago. These include an articulated skeleton described as Ombilinichthys yamini gen. et sp. nov., a fossil closely related to the giant gourami Osphronemus within Osphronemidae, initially reported as Osphronemus goramy from Tertiary sediments but now attributed to Eocene layers based on associated fish fauna.^[51] These specimens represent the oldest definitive anabantoids, featuring typical perciform traits but lacking detailed preservation of the labyrinth organ. A significant advancement in the record comes from the late Oligocene (Chattian stage, approximately 26–23.5 million years ago) of the Nima and Lunpola basins in central Tibet, where the fossil climbing perch Eoanabas thibetana gen. et sp. nov. was discovered. This basal anabantid exhibits key diagnostic features, including a posterior notch on the opercle bounded by spines, a V-shaped strut on the inner opercular surface, and a well-developed labyrinth organ with cribrate bony lamellae in the suprabranchial chamber, confirming air-breathing adaptations akin to modern Anabas.^[9] As the earliest known member of Anabantidae, Eoanabas extends the family's geological range back by over 20 million years and supports an Asian origin for the suborder, with subsequent dispersal westward via docked India toward Africa.^[9] Later fossils include isolated opercular bones attributed to Anabas from Pleistocene deposits in Java, Indonesia, providing evidence of the persistence of climbing perches in Southeast Asia into the Quaternary. The evolutionary timeline of Anabantoidei, inferred from molecular clocks calibrated against these fossils, suggests origination around the Late Cretaceous (~72 million years ago), with major family divergences occurring in the Paleogene (e.g., Anabantidae crown ~46 million years ago) and diversification accelerating post-Eocene in Asian freshwater systems.^[52]^[10] Gaps persist, particularly in complete skeletal material and African occurrences, where no pre-Quaternary fossils are known, underscoring the suborder's predominantly Asian fossil footprint.^[6]

Phylogenetic relationships

The phylogenetic relationships within Anabantoidei were first comprehensively resolved using molecular data in a 2006 study analyzing mitochondrial DNA (mtDNA) sequences from 57 species across 19 genera, which recovered the suborder as monophyletic with African lineages in Anabantidae positioned as basal to the derived Asian clades comprising Helostomidae and Osphronemidae.^[6] Subsequent analyses incorporating nuclear DNA markers in the 2010s and genomic data from ultraconserved elements (UCEs) in the 2020s have confirmed this topology, with high support for key intrafamilial clades such as the monophyly of Osphronemidae and the sister-group relationship of Betta to Trichopodus within it.^[52] These updates, drawing on multi-locus datasets, have refined genus-level boundaries without altering the overall family-level structure, for instance by synonymizing Colisa under Trichogaster based on shared molecular and morphological synapomorphies.^[52] Diversification within Anabantoidei occurred through a rapid radiation primarily in Southeast Asia during the Paleogene, with the crown-group age estimated at approximately 72 million years ago (95% HPD: 63–80 Ma), aligning with paleontological evidence of early labyrinth fish fossils from the Eocene.^[6]^[52] The disjunct African-Asian distribution is attributed to post-radiation dispersal events rather than vicariance, with Southeast Asian origins followed by colonization of African freshwaters by Anabantidae lineages around the Oligocene (~30 Ma).^[52] Evolutionary transitions in brood care modes, such as mouthbrooding, show phylogenetic correlation and independent origins at least twice—once in the basal Anabantidae and again within derived Osphronemidae branches—highlighting adaptive shifts tied to specific lineages.^[6] Cladistic analyses consistently support the monophyly of Anabantoidei within the order Anabantiformes, where it forms the sister group to Channoidei (snakeheads), a relationship robustly upheld by both mitochondrial-nuclear concatenations and phylogenomic approaches. No major revisions to this core framework have emerged since 2020, though ongoing genomic studies continue to provide finer resolution at lower taxonomic levels, reinforcing the stability of the subordinal phylogeny.^[52]

Human interactions

As food fish

Several species within the Anabantoidei suborder are commercially exploited for human consumption, particularly in Southeast Asia, where they contribute to local fisheries and aquaculture. The giant gourami (Osphronemus goramy), which can reach up to 70 cm in length and 1-2 kg in weight at harvest, is one of the most significant, farmed extensively in earthen ponds across Indonesia and Thailand.^[53] This species yields a high-protein flesh valued for its mild flavor, with annual production in Indonesia alone exceeding 150,000 tonnes in the early 2020s, primarily supporting domestic markets.^[54] The climbing perch (Anabas testudineus) is another key food fish, harvested from wild stocks and cultured in ponds and cages in countries like Indonesia, Thailand, and Vietnam, often reaching market sizes of 200-500 g.^[30] It is commonly dried or salted for preservation and trade, providing an affordable protein source in rural areas.^[55] The kissing gourami (Helostoma temminckii), growing to about 30 cm, is farmed on a smaller scale in southern Indochina and Indonesia for local consumption, with its firm white meat prepared fresh. Aquaculture of Anabantoidei species has expanded rapidly in Southeast Asia, driven by demand for freshwater fish protein, with Indonesia and Thailand leading production. Combined output from major species like giant gourami and climbing perch reached approximately 250,000-280,000 tonnes annually as of 2021, utilizing low-input pond systems that integrate with rice paddies for sustainable yields.^[56]^[57] These fish offer nutritional benefits, including high protein content (around 15-20% per 100 g edible portion), making them a staple in diets where animal protein access is limited.^[53] However, challenges persist, including overfishing of wild populations to supply seed for farms and disease outbreaks in intensive systems, which have reduced natural stocks in some riverine habitats.^[58] In traditional Asian cuisine, Anabantoidei feature prominently in dishes such as steamed giant gourami with herbs in Indonesian pepes ikan or curried climbing perch soups in Thai and Vietnamese meals, emphasizing their versatility in home and market cooking.^[53] Exports to Europe and the United States remain minimal, limited by preferences for marine species and logistical costs, with most production consumed locally. Sustainability concerns arise from habitat conversion for aquaculture ponds, which has led to declines in wild Anabantoidei populations in floodplain areas of Indonesia and Thailand, prompting calls for better stock management.^[59]

As aquarium fish

Several species within Anabantoidei are highly popular in the ornamental fish trade due to their vibrant colors, interesting behaviors, and relative ease of care in home aquariums. The Siamese fighting fish (Betta splendens) stands out as one of the most sought-after, selectively bred over generations for striking colors and elaborate fin shapes, although this has intensified male aggression, making single-male housing essential to prevent fights.^[60] Other favorites include the dwarf gourami (Trichopodus lalius), prized for its compact size and iridescent blue-red hues, and the paradise fish (Macropodus opercularis), valued for its hardy nature and active swimming.^[61]^[62] The history of Anabantoidei in aquariums dates to the 19th century, with the paradise fish becoming the first tropical freshwater species introduced to Europe in 1869 by French naturalist Pierre Carbonnier, sparking interest in exotic fishkeeping beyond goldfish.^[63] The Siamese fighting fish followed in the late 1800s, initially imported via scientific expeditions, and has been domesticated and bred in Thailand since at least the early 19th century for competitive fighting, a tradition that transitioned to ornamental breeding.^[64]^[65] Global trade in these species is substantial, with Thailand exporting approximately 20-24 million Siamese fighting fish annually as of 2018, valued at around $4 million USD as of 2018, contributing to the broader Asian ornamental fish market exceeding $1.4 billion in 2023.^[66]^[67] In 2025, betta exports generated approximately $11 million USD, reflecting growth in the sector.^[68] Dwarf gouramis and paradise fish add to this volume through exports from Southeast Asia, supporting a hobbyist demand that sustains millions of specimens traded yearly worldwide.^[69] Care for Anabantoidei typically requires a minimum tank size of 20-30 gallons (75-113 liters) for community setups to accommodate their active nature and territoriality, with water temperatures maintained at 24-28°C (75-82°F), pH 6.5-7.5, and low flow to mimic their natural slow-moving habitats.^[70]^[71] Aquariums should be densely planted with floating surface cover like Indian fern or lettuce to provide security and oxygen access via their labyrinth organ, while separating aggressive males—especially in Betta and Macropodus—prevents injury.^[72]^[73] Breeding these fish in captivity is straightforward for enthusiasts, often involving males constructing bubble nests at the water surface using saliva and plant material to house eggs, which females then fertilize; success rates improve in slightly warmer conditions (26-28°C) and with separate rearing tanks for fry to avoid predation.^[60]^[62] Challenges in keeping Anabantoidei include disease susceptibility, such as the iridovirus (Megalocytivirus) affecting dwarf gouramis, which causes lethargy, lesions, and near-100% mortality in infected stock, often introduced via contaminated imports.^[74] Additionally, intensive selective breeding for aesthetic traits in Siamese fighting fish has led to deformities like excessively elongated fins that hinder mobility and increase infection risks, shortening lifespans compared to wild forms.^[18]^[75]

Conservation status

The conservation status of Anabantoidei species varies, but many face significant risks due to their restricted ranges in Southeast Asian wetlands. According to the IUCN Red List, at least two species are classified as critically endangered, five as endangered, and eight as vulnerable within the genus Betta alone, reflecting broader threats across the suborder.^[76] For instance, Betta miniopinna is critically endangered owing to its extremely limited distribution in peat swamp forests of Singapore and Indonesia, where ongoing habitat loss has confined it to fragmented remnants.^[76] Similarly, Betta persephone holds endangered status due to its small population size and severe habitat degradation in Malaysian peat swamps.^[76] Wild populations of Betta splendens are assessed as vulnerable, primarily from pollution and habitat alteration in their native Thai rice paddies and ditches. Major threats to Anabantoidei include habitat destruction through deforestation and agricultural expansion, particularly in Indonesia and Malaysia, where peat swamp conversion for palm oil plantations has decimated specialized blackwater ecosystems.^[38] Overcollection for the international aquarium trade exacerbates declines in endemic species, while introductions of non-native species compete with natives in altered wetlands.^[37] Climate change further compounds these pressures by drying out peat swamps through altered rainfall patterns and increased temperatures, reducing suitable acidic, low-oxygen habitats essential for labyrinth organ function.^[38] Conservation efforts focus on habitat protection and species recovery programs. Protected areas in Borneo and Sumatra, such as peat swamp reserves, safeguard key populations of endemic taxa like Betta and Parosphromenus species.^[76] Captive breeding initiatives have gained momentum, notably on Bangka Island, Indonesia, where community-led programs in 2025 aim to bolster critically endangered Betta burdigala through ex situ propagation and habitat restoration to prevent imminent extinction.^[38] These actions involve local partnerships to monitor wild sites and reduce illegal collection. Research priorities emphasize the over 15 data-deficient Anabantoidei species, many of which are narrow endemics on islands like Sumatra and Borneo, where urgent assessments are needed to inform targeted protections amid rapid environmental change.^[77]

References

[1]
https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=172582
[2]
(PDF) Labyrinth fishes (Anabantoidei) - ResearchGate
Jan 14, 2018 · Labyrinth fishes (Anabantoidei) are grouped into three families (Osphronemidae, Helostomatidae, and Anabantidae) within the teleost Order ...Abstract And Figures · References (21) · Recommended Publications
[3]
Molecular Phylogenetics and Evolutionary Diversification of ...
Labyrinth fishes (Perciformes: Anabantoidei) are primary freshwater fishes with a disjunct African-Asian distribution that exhibit a wide variety of ...
[4]
Order ANABANTIFORMES: Families ANABANTIDAE ...
Aug 28, 2025 · 8 families · 26 genera · 287 species. Family ANABANTIDAE. Climbing Gouramies. 4 genera · 33 species. Anabas. Cloquet. 1816 from the Greek anabás.
[5]
FAMILY Details for Osphronemidae - Gouramies - FishBase
Nov 29, 2012 · Family Osphronemidae - Gouramies. Order, : Anabantiformes · Osphronemidae. Class, : Teleostei. No. in FishBase, : Genera : 14 | Species ...
[6]
Balancing the competing requirements of air-breathing and display ...
Air-breathing fish of the Anabantoidei group meet their metabolic requirements for oxygen through both aerial and aquatic gas exchange.
[7]
Fossil climbing perch and associated plant megafossils indicate a ...
Apr 13, 2017 · It comprises a fossil climbing perch (Anabantidae) and a diverse subtropical fossil flora from the Chattian (late Oligocene) of central Tibet.
[8]
FAMILY Details for Anabantidae - Climbing gouramies
- **Number of Genera and Species in Anabantidae:**
[9]
FAMILY Details for Helostomatidae - Kissing gouramies
### Summary of Helostomatidae - Kissing Gouramies
[10]
[PDF] Labyrinth fishes (Anabantoidei) - TimeTree.org
Labyrinth fishes form a natural (monophyletic) group of teleost fishes, the Suborder Anabantoidei. They are arranged into three families: Osphronemidae (~120.Missing: definition characteristics
[11]
https://doi.org/10.1111/jfb.13357
[12]
https://doi.org/10.1111/jfb.12322
[13]
https://www.journals.uchicago.edu/doi/full/10.1086/658996
[14]
[PDF] Body shape transformation along a shared axis of anatomical ...
In labyrinth fishes (Anabantoidei), rapid evolution of a highly derived torpedo-shaped body in the common ancestor of the pikehead (Luciocephalus aura and L.
[15]
[PDF] Trichogaster trichopterus, - the NOAA Institutional Repository
All gouramis belong to the suborder Anabantoidei and are commonly known as the labyrinth fishes. These fish are characterized primarily by an accessory ...
[16]
Sensory ecology of the fish lateral-line system: Morphological and ...
This review starts out by briefly describing naturally occurring hydrodynamic stimuli and the morphology and neurobiology of the fish lateral-line system.Missing: Anabantoidei adhesive
[17]
Terrestrial locomotion in the climbing perch, Anabas testudineus ...
Aug 7, 2025 · No evidence of involvement of the pelvic fins in locomotion was obtained; the pectorals seemed only to help in tilting the head from side to ...
[18]
https://pmc.ncbi.nlm.nih.gov/articles/PMC9334006/
[19]
Anabantidae - an overview | ScienceDirect Topics
Many of the species in the families Callicthyidae and Clariidae, and all of the species in the five families comprising the anabantoid suborder (Anabantidae, ...
[20]
Air‐breathing behaviour of the African anabantoid fish Ctenopoma ...
Aug 7, 2025 · Frequency of aerial respiration in Ctenopoma muriei increased as dissolved oxygen (DO) decreased; breathing behaviour was spatially synchronous.
[21]
Something Fishy about Siamese Fighting Fish (Betta splendens) Sex
Siamese fighting fish show sexual dimorphism; males have long ventral and tail fins with vivid and uniform body coloration, whereas females have shorter fins ...
[22]
A New Species of Nest Building Ctenopoma (Teleostei ... - jstor
extensive sexual dimorphism in size (males larg- er); coloration (males distinctively patterned and brightly colored); and fin development (un- paired and ...
[23]
https://www.inaturalist.org/taxa/326133-Macropodus-ocellatus
[24]
Round-tailed Paradisefish (Macropodus ocellatus) - iNaturalist
The round-tailed paradise fish (Macropodus ocellatus) is a species of gourami native to eastern Asia, where it is found in China, Japan, and Korea.
[25]
(PDF) Molecular Phylogenetics and Evolutionary Diversification of ...
Aug 6, 2025 · Our results suggest that the evolution of breeding behavior in anabantoids is highly correlated with phylogeny, and that brood care evolved ...<|control11|><|separator|>
[26]
(PDF) The labyrinth fishes (Teleostei : Anabantoidei, Channoidei) of ...
Aug 7, 2025 · Betta cracens, new species, differs from other members of the B. pugnax group in having the most slender body (body depth 21.2-24.2% SL), more ...Missing: examples | Show results with:examples
[27]
Diversity, distribution, and conservation status of Betta fish (Teleostei
According to the IUCN data in 2020, a total of 21 Betta species were recorded from inland waters of Sumatra, Indonesia, where 28.6% are critically endangered, ...
[28]
Climbing perch - Business Queensland
Dec 14, 2018 · Climbing perch (Anabas testudineus) is an invasive animal in Queensland. Climbing perch is a restricted animal under Queensland legislation.
[29]
Anabas testudineus, Climbing perch : fisheries, aquaculture, aquarium
Obligate air-breathing (Ref. 126274); Found mostly in canals, lakes, ponds, swamps and estuaries (Ref. 41236, 57235). Adults occur in medium to large rivers ...
[30]
[PDF] First record of the non-native three spot gourami, Trichogaster ...
Five specimens of the non-native three spot gourami (Osphronemidae: Trichogaster trichopterus) were collected from a small pond near the northern coast of ...<|separator|>
[31]
Siamese fighting fish: Current Biology - Cell Press
May 8, 2023 · Siamese fighting fish are becoming a problematic invasive species in South America, Australia and the United States. Their ability to survive in ...
[32]
[PDF] Cardio-respiratory ontogeny and the transition to bimodal respiration ...
While the gills may start differentiating during this time, cutaneous respiration dominates, as the skin is thin and well vascularized (Hughes et al., 1986).
[33]
Betta brownorum • Fish sheet - Fishipedia
Oct 11, 2023 · B. brownorum inhabits very shallow forest puddles and ditches within peat swamps: black, very soft and acidic waters (22–26 °C), no to slow ...
[34]
[PDF] Anabas testudineus, Climbing Perch - IUCN Red List
Aug 10, 2019 · Range Description: The species is widely distributed in Pakistan, India, Bangladesh, Nepal, Bhutan (most likely), Sri Lanka,. Myanmar ...
[35]
Labyrinth Fishes: Anabantoidei | Encyclopedia.com
Habitat: Siamese fighting fish live in standing water with dense plant life, especially rice paddies and canals. They may dig into the mud when the water level ...Missing: deforestation | Show results with:deforestation
[36]
[PDF] Climbing Perch (Anabas testudineus) - U.S. Fish and Wildlife Service
Often found in areas with dense vegetation. [Rainboth 1996]. Can tolerate extremely unfavorable water conditions and is associated mainly with turbid ...Missing: microhabitats | Show results with:microhabitats
[37]
Save This Species: The Bettas of Bangka Island - The Revelator
May 7, 2025 · Betta burdigala is found in only one watershed on the small island of Bangka, Indonesia. The fish are acidic, blackwater swamp specialists and can only thrive ...
[38]
Betta livida
Habitat is highly acidic tea-coloured water of between pH 3.5 to 3.7. Found in a peat swamp forest, mostly in well shaded shallow waters (less than a meter ...
[39]
[PDF] DNA Barcoding Indonesian freshwater fishes: challenges and ...
Including Peninsular Malaysia and the islands of Sumatra, Java and Borneo (Fig. 1), this hotspot exhibits one of the highest species richness and endemism for ...<|separator|>
[40]
[PDF] Distribution Range Update of Red Wine Betta, Betta burdigala in ...
Jul 3, 2025 · Both legal and illegal tin extraction have led to widespread deforestation, soil erosion, and hydrological disruption across the island (Adi et ...
[41]
Comparative Morphology of the Digestive Tract of African Bush Fish ...
Aug 13, 2023 · This study describes, for the first time, the structure of the digestive tracts of two Anabantoidei fishes: African bush fish (Ctenopoma ...
[42]
https://pmc.ncbi.nlm.nih.gov/articles/PMC10451994/
[43]
study on food and feeding biology of trichogaster fasciata bloch ...
Jun 25, 2018 · It mainly feeds on algae, diatoms, zooplankton, insects and decayed organic matter (Khongngain et al. 2017 ). The body weight (BW) of each ...
[44]
exploring the function and constraints of water shooting in dwarf ...
Several species of gouramis from the genera Trichogaster and Trichopodus are able to shoot water above the surface, a behaviour referred to as 'jetting' (Hall, ...
[45]
The influence of hunger on food‐stocking behaviour of climbing ...
Aug 7, 2025 · The present study demonstrated that the climbing perch Anabas testudineus collected and stocked food pellets in its mouth, a behaviour that ...
[46]
Taking the Leap: Archerfish Snag More Prey with Amazing Jumps
Apr 19, 2017 · However, a fish that jumps is frequently successful at nabbing its meal, capturing the prey 70 percent of the time, according to the new study.
[47]
Decoupled Jaws Promote Trophic Diversity in Cichlid Fishes
Functional decoupling of oral and pharyngeal jaws is widely considered to have expanded the ecological repertoire of cichlid fishes.
[48]
Giant gourami - Facts, Diet, Habitat & Pictures on Animalia.bio
Gourami tend towards herbivory, preferring algae-based foods, but will eat meaty foods. An algae-based flake food, along with freeze-dried bloodworms, tubifex, ...
[49]
BETTA FISH HUNTING for Food | Mini Ambush Predators - YouTube
Mar 10, 2019 · When you Feed your Betta Fish they don't just eat their Food, they Hunt for it, this family of Bettas crawl over this blanket of Lace Fern ...Missing: diet foraging
[50]
[PDF] Are Kissing Gourami Specialized for Substrate-Feeding? Prey ...
foraging habit. However, because of anatomical modifications of the gill-rakers, Helostoma has been considered a midwater filter-feeding species. We offered ...
[51]
https://www.tandfonline.com/doi/abs/10.1080/02724634.2014.906444
[52]
https://doi.org/10.1111/jbi.14781
[53]
Giant gourami - Cultured Aquatic Species
So far, with an estimated production in excess of 145 000 tonnes in 2017, Indonesia is the most important producer of giant gourami and its production ...
[54]
Gonadal development and sexual maturity onset of giant gourami ...
Sep 5, 2025 · ... Indonesia with annual production reaching 152,669 tons in 2020 [23]. This species is extensively dispersed across the primary Indonesian ...
[55]
Climbing perch (Anabas testudineus) | MegaFishingThailand
Very common fish in Thailand and can be found in almost every body of water, lake drainage ditch or paddy field. Popular as a food fish they are easy to catch ...
[56]
[PDF] Genetic Variation of Giant Gourami (Osphronemus goramy ...
Apr 11, 2025 · Indonesia is one of the leading producers of giant gourami in Southeast Asia, with an annual production of 183,354 tonnes recorded in 2019 (FAO ...
[57]
Similarity and Genetic Variation of Climbing Perch, Anabas ...
Climbing perch, A. testudineus, is an economically important freshwater fish and emerged as a prominent species in aquaculture in Indonesia [2] and in South ...Missing: giant gourami
[58]
(PDF) The effect of anthrophogenic changes on the sustainability of ...
Aug 6, 2025 · Anthrophogenic activities contributed significantly to the degradation of floodplain fisheries resources, especially in relation to a decrease ...<|control11|><|separator|>
[59]
An Introduction to Labyrinth Fish- Gouramis and Bettas - Fishkeeping
May 2, 2010 · This gourami reaches an adult size of 12 inches and so is unsuitable for most tanks. They are difficult to sex so if you are looking to breed ...Missing: examples Osphronemus
[60]
Quick guide to anabantoids - Practical Fishkeeping
Fighting fish (Betta spp.) This is the largest and best known group of anabantoids with 66 species and counting, although the name 'fighting fish' is almost ...Missing: definition characteristics taxonomy
[61]
Paradise Fish (Macropodus chinensis, Labrus opercularis)
M. opercularis is a classic aquarium fish, having been introduced to the hobby by a French soldier named Gerault in 1869. Of the initial 100 specimens shipped, ...Missing: history | Show results with:history
[62]
The Chinese Paradisefish: never give up! - Aqualog.de
Jan 12, 2015 · In 1869 the true Paradisefish, Macropodus opercularis, was the first exotic aquarium fish after the Goldfish to reach Europe and triggered a ...
[63]
History of Betta Fish: The Origins of Siamese Fighting Fish
May 27, 2025 · The first stop on this journey was Europe. By the 1890s, Siamese fighting fish had been introduced to European aquarists. In fact, records ...
[64]
[PDF] Siamese-fighting-fish.pdf - SEAFDEC Institutional Repository (SIR)
From 2014 to 2018, the average volume per year and average value per year of exported betta fish were 23.92 million and US$ 4.29 million, respectively.Missing: annually | Show results with:annually
[65]
Siamese fighting fish need more space in pet shops: new behaviour ...
Feb 18, 2025 · Thailand alone exported around 20 million Siamese fighting fish in 2018, with major markets including the USA, China, Singapore, France and ...Missing: annually | Show results with:annually
[66]
Asia Pacific Ornamental Fish Market | Industry Report, 2030
The Asia Pacific ornamental fish market size was estimated at USD 1433.8 million in 2023 and is expected to grow at a CAGR of 9.7% from 2024 to 2030.Missing: Anabantoidei gourami betta $100
[67]
Recent Trends in Breeding and Trade of Ornamental Gourami in India
Mar 13, 2019 · In contrast, the dwarf gourami Trichogaster lalius (Hamilton, 1822) is a popular gourami species in the ornamental fish industry due to its ...
[68]
Gouramis Care Guide - Aqueon
Gouramis are a group of fish in the families Osphronemidae, Helostomatidae and Anabantidae. They have a labyrinth organ that acts sort of like a lung.
[69]
Dwarf Gourami (Colisa lalia) - Care, Feeding, Breeding, and Tank ...
Sep 28, 2024 · Dwarf Gouramis need a 60L tank with gentle flow, pH 6.5-8, 23-28°C, and a varied diet. They are peaceful, but males can be territorial.
[70]
https://www.aqueon.com/resources/care-guides/gouramis
[71]
Paradise Fish 101: Care, Tank Size, & Tank Mate - Aquarium Source
These fish thrive in room-temperature water (around 65-78°F), so if your home stays within that range, you can skip the heater. They're actually a bit tougher ...
[72]
Iridovirus and Gouramis - General Chat - Aquarium Co-Op Forum
Jan 13, 2025 · Iridovirus affects only dwarf gouramis (Trichogaster lalius). It's estimated that about half of all dwarf gouramis in the pet trade are infected with this ...Is this Dwarf Gourami Disease? - Aquarium Co-Op ForumIll Dwarf Gouramis, Iridovirus??? - Diseases - Aquarium Co-Op ForumMore results from forum.aquariumcoop.com
[73]
Care and Use of Siamese Fighting Fish (Betta splendens) for ...
Native to Southeast Asia, betta have been selectively bred for their fighting ability for hundreds of years, which has resulted in the species' characteristic ...
[74]
Viral Aquarium Fish Diseases | Tropical Fish Hobbyist Magazine
... may be noticed. Dwarf Gourami Iridovirus (DGIR). About 10 years ago, aquarium dealers and hobbyists noticed a greatly increased mortality rate in dwarf gouramis ...
[75]
The IUCN Red List of Threatened Species
**Summary of Betta Species by IUCN Category:**
[76]
The IUCN Red List of Threatened Species
**Summary of IUCN Red List Data for Anabantoidei (Labyrinth Fishes):**