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Gilled lungfish
Gilled lungfish
from Wikipedia

Gilled lungfish
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Dipnoi
Order: Ceratodontiformes
Family: Protopteridae
Genus: Protopterus
Species:
P. amphibius
Binomial name
Protopterus amphibius
(Peters 1844)[2]
Synonyms[3][4]
  • Rhinocryptis amphibia Peters 1844

The gilled lungfish (Protopterus amphibius, also known as the East African lungfish, is a species of African lungfish. It is native to Kenya, Somalia and Mozambique.[5][6] Records from Tanzania require confirmation[6] and may be the result of introductions.[5]

Taxonomy

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Protopterus amphibius was named in 1844 by German naturalist Wilhelm Peters[7] and is synonymous with Rhinocryptis amphibia.[8][9] The gilled lungfish is part of the Protopteridae family and in the order of Lepidosireniformes.[7] Protopterus amphibius is most closely related to the other three species of African lungfish in the Protopteridae family.[10] Protopteridae are sister groups with tetrapod vertebrates.[11]

Description

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Protopterus amphibius has an elongate body which reaches a length of 44cm (17 in), making it the smallest extant lungfish.[6][12] It has fused dorsal-caudal-anal fins and small cycloid scales.[12] This lungfish is uniform blue or slate grey in colour. It has small inconspicuous black spots and a pale grey belly.[6]

Near the ventral snout, the gilled lungfish has paired olfactory organs containing lamellae and a nasal sac allowing water takeup in shallow water or wet land.[13] Their heavy tooth plates give them a unique jaw morphology, often used for identification.[11] While it shares many anatomical features with other African lungfish, its size makes it easy identifiable.[12]

Ventilation

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Like all African lungfish, the gilled lungfish has two lungs, six gill arches, and five gill clefts. It is an obligate airbreather and can survive dry spells through aestivation, a burrowing process in which it creates a protective mucous cocoon.[6][12] Aestivation involves reduction of the lungfish's metabolic rate and requires usage of stored fuel, and is therefore a type of dormancy induced by the dry season.[11] Both its gills and its lungs are used for ventilation, with frequency of each dependent on ambient gas tensions. Gill and lung usage is stimulated by hypoxia and hypercapnia individually, and depressed by hyperoxia and hypocapnia individually. Combinations of hypoxia and hypercapnia depress gill usage and increase lung usage.[14] Generally, African lungfish become less dependent on gill usage as they mature.[5]

Life history

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Apart from increased dependency on external gills in juveniles, nothing is known about the breeding biology of Protopterus amphibius or the biology of the young. The three other members of Protopterus spawn in nests whose structure is influenced by environmental factors. These three species exhibit parental care for their young.[15] African lungfish have a lifespan of about 18 years in captivity; there is no information about the lifespan of wild African lungfish.[16] Little is known about the feeding habits of Protopterus amphibius specifically, but the feeding habits of other Protopterus suggest the gilled lungfish may be an omnivorous carnivore feeding on invertebrates and plant material. They likely use olfaction to locate prey and likely slowly stalk their prey until it is close enough for suction. Protopterus species ingest food through a process of chewing, spitting out, and sucking back in their prey.[15]

Habitat

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The gilled lungfish resides in inland freshwater ecosystems in tropical zones. It lives largely within the riverbeds of the Zambezi River system of East Africa and the region's wetlands. It is a primarily demersal fish.[17] Its abundance is relatively unknown.[5]

Conservation

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Fisheries for Protopterus species are located in Uganda and Kenya.[18] The gilled lungfish is a species of Least Concern on the IUCN Red List because of its wide distribution and lack of major threats, although their data deficiency should be noted.[5] The largest threats to the gilled lungfish are the damming of the Zambezi, which will reduce the size of the delta they inhabit, pollution in areas they inhabit, and habitat reduction due to agricultural encroachment on wetlands.[5]

References

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Revisions and contributorsEdit on WikipediaRead on Wikipedia
from Grokipedia
The gilled lungfish (Protopterus amphibius) is a species of African distinguished as the smallest extant member of its group, attaining a maximum total length of 44.3 cm. Native to freshwater environments in eastern , it features an elongated, cylindrical body with a circular cross-section and embedded scales, enabling it to navigate swampy habitats. As an obligate air-breathing , it utilizes both functional gills and paired lungs for respiration, allowing survival in low-oxygen waters. Belonging to the family Protopteridae within the subclass Dipnoi (lungfishes), P. amphibius is classified under the order Ceratodontiformes and represents one of four African lungfish species in the genus . Its range extends across tropical regions from southward to and the River delta, primarily in swamps, floodplains, and coastal river systems between approximately 10°N and 19°S latitude. These demersal habitats experience seasonal flooding and drying, to which the species is well-adapted through estivation, forming a protective cocoon in the mud to endure months without water. Biologically, the gilled lungfish exhibits notable respiratory and survival traits as a sarcopterygian , bridging aquatic and terrestrial adaptations in vertebrate . It feeds on small , , and plant matter in vegetated shallows, with long, thread-like pectoral and pelvic fins facilitating movement. Reproduction details remain limited, but like other protopterids, it likely spawns during wet seasons in temporary pools. The species supports minor local fisheries but faces no major threats, earning a Least Concern status on the as of 2009.

Taxonomy and phylogeny

Classification

The gilled lungfish, Protopterus amphibius (Peters, 1844), belongs to the family Protopteridae of African lungfishes. Its taxonomic hierarchy is as follows: Kingdom: Animalia, Phylum: Chordata, Class: Dipnoi, Order: Lepidosireniformes, Family: Protopteridae, Genus: Protopterus, Species: Protopterus amphibius. Common names for the species include East African lungfish and gilled lungfish. The genus Protopterus includes four extant species, all native to Africa: P. aethiopicus, P. amphibius, P. annectens, and P. dolloi; these are distinguished primarily by variations in body elongation, paired fin lengths, and scale patterns. Protopterus amphibius was first described by Wilhelm Peters in 1844 from specimens collected in Mozambique.

Evolutionary history

Lungfishes, or dipnoans, belong to the sarcopterygian clade of lobe-finned fishes and represent the closest living relatives to tetrapods, with their lineage originating during the period approximately 400 million years ago. This early radiation occurred amid fluctuating environmental conditions, including periods of low oxygen levels in aquatic habitats, which likely drove the evolution of air-breathing lungs as an adaptation for survival in hypoxic waters. Fossil evidence from deposits, such as the Dipterus, illustrates primitive lungfish morphology with paired lungs that facilitated bimodal respiration, linking ancient forms to modern lungfishes and underscoring their role as a group. The African lungfish family Protopteridae, including the genus , diverged from the lineage (Lepidosirenidae) around 120 million years ago (95% CI: 94–165 Ma), coinciding with the fragmentation of the supercontinent during the . This vicariance event separated ancestral populations, with Protopteridae adapting to freshwater systems across amid cycles of and wetland expansion, as evidenced by fossils dating back at least 100 million years from deposits in northern . Genetic studies further confirm the basal phylogenetic position of within the African , distinguishing it from more derived Australian (Neoceratodus) and lineages that diverged earlier, around 140–200 million years ago. A distinctive feature of the gilled lungfish amphibius is its retention of into adulthood, unlike other species and lungfishes that typically resorb these structures post-larval stage. This trait may represent a primitive condition preserved from ancestral sarcopterygians or a neotenic , allowing enhanced oxygen uptake in low-oxygen environments while complementing lung-based aerial respiration. Such retention highlights P. amphibius as a key model for understanding the evolutionary persistence of juvenile features in response to ecological pressures.

Physical characteristics

Morphology

The gilled lungfish, Protopterus amphibius, exhibits an elongated, eel-like body form adapted to its swampy habitat. The body is covered in small, embedded scales, and it lacks a separate , with the dorsal, caudal, and anal fins instead fused into a continuous marginal fin fold along the posterior two-thirds of the body. The pectoral and pelvic fins are modified into slender, fleshy, tentacle-like appendages that extend well beyond the body, facilitating navigation through dense vegetation and muddy substrates. This species is the smallest among extant , reaching a maximum total length of 44.3 cm (17.4 in). Adults typically weigh less than 0.5 kg, based on length-weight relationships for the . The overall proportions emphasize a long tail that comprises approximately 60% of the total body length, contributing to its streamlined profile and aiding in burrowing into soft sediments during periods of environmental stress. Coloration is predominantly uniform blue to slate on the dorsal surface, accented by small, inconspicuous spots, while the ventral side is pale . The head is darker below, often with subtle spots or vermiculations. Juveniles display slightly more pronounced spotting compared to adults, though the pattern remains subdued overall. The head features a terminal equipped with specialized plates rather than individual teeth, enabling the crushing of hard-shelled prey such as mollusks and crustaceans. The eyes are small and positioned laterally, reflecting adaptations to the low-light conditions of turbid swamp environments where in dim waters is advantageous.

Respiratory and sensory adaptations

The gilled lungfish Protopterus amphibius is an obligate air breather possessing both functional gills and paired lungs for bimodal respiration. The gills are reduced compared to non-air-breathing fishes but remain operational throughout life, with adults uniquely retaining external gills among Protopterus species for supplemental aquatic gas exchange. Smaller individuals rely more heavily on gill-based aquatic respiration (up to 70% of oxygen uptake), while larger adults depend predominantly on pulmonary ventilation via the paired, elongated lungs that extend dorsally along the body cavity. These lungs, derived from the swim bladder, are vascularized sacs lined with capillaries that facilitate efficient aerial oxygen uptake, allowing the fish to surface periodically in hypoxic swamp waters. Sensory adaptations suit the species' murky, vegetated habitats. The olfactory organs are well-developed, featuring paired chambers at the ventral snout with multiple lamellae containing neurons, supporting cells, and mucous cells that enhance chemosensory detection of prey and environmental cues. The system, with mechanoreceptors along the head and body, detects water movements, vibrations, and pressure changes, aiding navigation and foraging in low-visibility conditions. Vision is secondary, with small lateral eyes providing limited acuity suited to dim, turbid waters.

Distribution and habitat

Geographic range

The gilled lungfish ( amphibius) is endemic to , with confirmed native populations restricted to coastal river systems and associated floodplains in , , and . Specific localities include the in southern , the Tana River basin in northeastern , and the River delta along with adjacent coastal swamps in . These sites represent isolated but stable habitats tied to drainage basins, spanning roughly from 0° to 18°S latitude. The species was originally described in 1844 based on syntypes collected from , , marking the southern extent of its known range. Since then, its distribution has remained relatively stable, though fragmented across discontinuous riverine floodplains that limit broad connectivity. Historical records align closely with contemporary surveys, indicating no major range contractions or expansions over the past century. Presence in Kenya's coastal drainages, including the lower Tana River, is confirmed. Uncertain records suggest possible occurrence in , including the floodplains of the Rufiji and Ruaha Rivers as well as the Lake Rukwa basin, but these have not been verified through positive identifications in recent surveys as of 2023. Dispersal is constrained to freshwater environments, with no evidence of marine tolerance, though seasonal flooding may facilitate limited between nearby populations within the same basin.

Environmental preferences

The gilled lungfish (Protopterus amphibius) inhabits freshwater environments characterized by swamps, floodplains, and slow-moving rivers, where it maintains a demersal lifestyle in shallow, vegetated waters. These habitats provide essential cover from predators. Preferred substrates consist of muddy bottoms, which facilitate burrowing for rest and concealment during dry periods. The species tolerates the warm conditions of tropical freshwater systems and low dissolved oxygen concentrations, relying on its air-breathing adaptations to persist in hypoxic settings. Seasonally, the gilled lungfish favors wet periods with inundation, which expands shallow, vegetated zones ideal for activity. It commonly occupies microhabitats like stagnant pools and narrow channels amid dense cover, promoting survival through enhanced resource access.

Ecology and behavior

Feeding habits

The gilled lungfish ( amphibius), like other members of its , exhibits an omnivorous diet with a preference for animal matter, consisting mainly of small such as mollusks, , crustaceans, and worms, along with small and occasionally frogs. Juveniles tend to focus more on , while adults may engage in under certain conditions, based on patterns in closely related species. Although largely focused on animal prey, it incorporates plant parts, , and even or mud in stomach contents, likely incidental to benthic . This dietary flexibility supports to variable environments. Specific diet composition for P. amphibius remains poorly documented, with most data extrapolated from congeners. As benthic dwellers, gilled lungfish employ a suction-feeding strategy to capture non-evasive prey through coordinated hyoid and movements. They are nocturnal foragers, relying on chemosensory and mechanosensory cues in low-visibility conditions to locate food, often probing the substrate with their paired fins to detect and manipulate prey. This slow, deliberate locomotion and feeding approach aligns with ambush-like predation in muddy or vegetated habitats, minimizing energy expenditure in oxygen-poor waters. The feeding apparatus features specialized tooth plates that function as rasping surfaces for crushing hard-shelled prey like mollusks and exoskeletons, enabling efficient processing of durable food items. Their relatively small mouth gape limits them to prey items typically under 10 cm, emphasizing a focus on smaller, accessible targets rather than larger or more mobile ones. In webs, gilled lungfish occupy a mid-level trophic position (3.4 ± 0.6, based on relatives), serving as predators of benthic and small vertebrates while acting as prey for piscivorous birds such as shoebills and , as well as larger and potentially crocodilians. This role contributes to nutrient cycling in seasonal aquatic systems. Feeding activity varies seasonally, with reduced or ceased intake during estivation in the , when the fish enter and rely on stored energy reserves. In contrast, wet seasons see heightened to replenish fat stores and support growth, reflecting the abundance of prey in flooded habitats.

Reproduction and parental care

The gilled lungfish ( amphibius) reaches at a total length of approximately 25-30 cm, typically after 2-3 years of age, based on patterns observed in closely related African lungfish species such as P. annectens and P. aethiopicus where maturity occurs around 29-30 cm. The breeding season is likely tied to the rainy period from November to March in its East African range, aligning with floodplain flooding that facilitates nesting, though this remains unconfirmed through direct observations for P. amphibius itself. Mating behavior in P. amphibius is presumed to mirror that of its congeners, with males constructing nests in shallow mud pits or among flooded vegetation to attract females. These nests, often simple depressions lined with aquatic plants, may serve multiple females, suggesting possible polygamous spawning events where a single male fertilizes eggs from several partners over the season. During spawning, females deposit adhesive eggs directly into the male-prepared nests, where occurs as the male releases over the . sizes are estimated at 500-1,000 eggs per spawning event, drawing from data in related species like P. annectens, where absolute ranges from 447 to 1,200 eggs; eggs are typically white, spherical, and measure 3.5-4 mm in diameter. Post-spawning, males provide exclusive by guarding the eggs and resulting larvae within the nest for 4-8 weeks, fanning the clutch to oxygenate it and defending against predators. Larvae hatch with prominent and initially rely on aquatic respiration, but they transition to air-breathing as they grow beyond 23-25 mm in length, developing functional lungs while gill dependence diminishes over early juvenile stages. Despite these inferences, the breeding biology of P. amphibius remains poorly studied, with no confirmed field observations of rituals, precise , or nest site specific to this , limiting detailed understanding compared to its better-documented relatives.

Life history strategies

Estivation process

The estivation process in gilled (Protopterus spp.) is triggered by the recession of water in their seasonal habitats during the dry period, prompting the to into the moist mud at the bottom of drying riverbeds or swamps. As water levels drop, the lungfish secretes copious amounts of from its skin glands, which hardens upon exposure to air to form a protective cocoon that seals in body moisture and minimizes evaporative water loss. This onset phase typically spans 6–8 days, during which the curls into a spherical position within the burrow, leaving a narrow tube-like passage from the cocoon's surface to its mouth for access to atmospheric air. Physiologically, estivation involves a profound suppression of , with oxygen consumption reduced by approximately 70% compared to active aquatic states, allowing survival on minimal energy reserves derived from and . To manage nitrogenous waste without access to , the shifts from ammonotelism to ureotelism, accumulating high levels of in its plasma—up to 10-fold increases—which acts as an osmoregulatory agent to maintain cellular hydration and prevent . Respiratory adaptations include the functional shutdown of the gills, which become encased in dried and exhibit collapsed secondary lamellae, while the lungs handle infrequent air breaths through the cocoon's tube, with ventilatory frequency initially rising before stabilizing at low levels. Cardiovascular changes accompany this, including a halving of to 11–16 beats per minute after 60 days. The cocoon itself is a multilayered structure of hardened, impermeable containing living epithelial cells that provide a semi-porous barrier against , with the outer layer featuring fine filaments potentially aiding in sensory detection of environmental . This , up to several centimeters thick, includes a small anterior opening connected to the burrow's tube, enabling pulmonary gas exchange while deterring predators and pathogens, though bacterial and fungal colonization on the exterior remains a concern. Estivation typically endures 6–12 months in natural conditions, aligning with the dry season's length, though laboratory studies demonstrate survival up to 2 years under controlled humidity. Emergence occurs with the onset of rains, when increased softens the cocoon, allowing the to break free. Recovery involves rapid rehydration, with surging up to 22-fold within hours of immersion to clear accumulated waste, alongside swift resumption of function and feeding to restore body mass lost at rates exceeding 10% over the period. However, risks are substantial; incomplete cocoon formation or prolonged exposure to extreme heat can result in high mortality from , , or upon arousal, with survival rates varying based on burrow depth and environmental conditions.

Growth and longevity

The gilled lungfish (Protopterus amphibius) undergoes distinct life stages characterized by rapid early development adapted to its ephemeral aquatic habitats. Larvae hatch from eggs measuring 3.5–4 mm in diameter after approximately one week, with prominent that facilitate oxygen uptake in oxygen-poor waters and are reabsorbed during into juveniles, which develop bimodal respiration using internal gills and lungs. Juveniles exhibit rapid growth during the first year under favorable conditions, influenced by seasonal flooding and food availability. Growth rates for P. amphibius are notably influenced by environmental factors. In aquaculture settings with ad libitum feeding, specific growth rates in mass reach 1.52% per day, reflecting accelerated somatic development before slowing after the juvenile phase as energy allocation shifts toward maintenance and reproduction. Seasonal flooding enhances growth by increasing prey availability and oxygen levels, while dry periods induce estivation, temporarily halting linear increment but preserving energy through depressed metabolism. However, specific details on growth for P. amphibius remain limited compared to other Protopterus species. Adult gilled lungfish typically attain lengths of 30–44 cm, with maximal recorded size at 44.3 cm total length; is minimal, though females may achieve slightly larger sizes. for African lungfish is estimated at 10–15 years in the wild and exceeding 20 years in captivity, though specific data for P. amphibius are limited. Mortality is highest among juveniles due to predation by birds, , and in shallow floodplains. Adults demonstrate resilience through estivation, enduring months in mud cocoons, but remain vulnerable to disruption from or that prevents formation.

Conservation status

The gilled lungfish ( amphibius) is assessed as Least Concern on the , based on a 2009 evaluation that highlights its wide distribution and apparent resilience, though the species is in several regions due to insufficient monitoring data. Populations remain stable in core habitats like the River basin, where surveys indicate high local abundances. In contrast, population data from are sparse and unreliable, primarily because ongoing political instability has severely limited assessments and field research. While no confirmed global population decline exists, possible local reductions have been reported in areas with intensive human activity, though these require further validation. Recent records suggesting range expansion into Tanzanian waters, such as the floodplain, need verification to distinguish between genuine distributional shifts and artifacts of improved sampling efforts. The primary threats to P. amphibius stem from habitat loss driven by wetland conversion to ; for instance, significant portions of Kenyan wetlands have been transformed for and vegetable farming, fragmenting seasonal swamps critical for the species. Damming along the , including major projects like Kariba and , has altered natural flood regimes, reducing the extent and timing of inundation and diminishing breeding and foraging grounds. Anthropogenic pollution, including and sediments from agricultural and industrial activities in rivers like the Tana, exacerbates degradation in coastal basins. Overfishing represents a minor risk, confined to localized subsistence harvest without evidence of large-scale commercial pressure. Climate change adds uncertainty by modifying wet-dry cycles, which could disrupt estivation triggers and increase vulnerability in flood-dependent habitats.

Conservation efforts

The gilled lungfish ( amphibius) benefits from current conservation measures embedded in broader initiatives across its range in eastern . The receives incidental via the designation of the Zambezi Delta as a Ramsar Wetland of International Importance, where management plans emphasize the conservation of ecosystems supporting diverse assemblages, including , against drainage and . Research gaps persist for the gilled lungfish, particularly in conducting comprehensive population surveys in northern range areas such as and , where data on local abundances remain sparse despite the species' occurrence in seasonal swamps. Additional needs include targeted breeding studies to evaluate reproductive viability under changing hydrological conditions, as current knowledge of spawning behaviors is limited. Genetic analyses are also recommended to delineate potential variations across fragmented populations, aiding in tailored protection strategies. Management actions for the gilled lungfish involve community-based fisheries regulations that limit harvest during vulnerable dry periods, implemented in parts of its range to promote sustainable use and reduce incidental capture. Habitat restoration projects, such as those under IUCN's programs in eastern , focus on countering damming impacts by restoring connectivity, which supports migration and estivation sites. Internationally, the gilled lungfish is not directly listed under but benefits indirectly from trade bans on species through regional agreements, helping curb unregulated export. Monitoring occurs via African initiatives, including assessments and WWF's efforts to track ancient lineages like Protopteridae. The species' Least Concern status reflects stable populations but underscores the value of these collaborative frameworks. Future recommendations emphasize enhanced data collection beyond 2025, leveraging tools like environmental DNA (eDNA) for non-invasive surveys in remote floodplains. Integration into climate adaptation plans for floodplains is advised, prioritizing resilient wetland management to mitigate drought and flood variability affecting the species' habitats.

References

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