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Mudskipper
Mudskipper
Mudskipper
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Mudskipper
Atlantic mudskipper (Periophthalmus barbarus) in The Gambia
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Order: Gobiiformes
Family: Oxudercidae
Subfamily: Oxudercinae
Gunther 1861[1]
Genera

Apocryptes
Apocryptodon
Boleophthalmus
Oxuderces
Parapocryptes
Periophthalmodon
Periophthalmus
Pseudapocryptes
Scartelaos
Zappa

Synonyms
  • Periophthalminae
  • Periophthalmidae

Mudskippers are any of the 25 extant species of amphibious fish from the subfamily Oxudercinae of the goby family Oxudercidae.[2] They are known for their unusual body shapes, preferences for semiaquatic habitats, limited terrestrial locomotion and jumping, and the ability to survive prolonged periods of time both in and out of water.

Mudskippers can grow up to 30 cm (12 in) long, and most are a brownish green colour that ranges anywhere from dark to light. During mating seasons, the males will also develop brightly coloured spots in order to attract females, which can be red, green or blue. Unlike other fish, the mudskipper's eyes protrude from the top of its flat head. Another notable feature is that their side pectoral fins are located more forward and underneath their elongated bodies. These fins are jointed and function similarly to limbs, which allow the mudskipper to crawl from place to place. Although they have the typical body form of any other gobiid fish, pectoral fins allow the mudskipper to actively "skip" across muddy surfaces (hence the common name) and even climb low-hanging tree branches and scrubs. Mudskippers can leap distances of up to 61 centimetres (24 in) by laterally flexing and pushing with their tails.[citation needed]

Taxonomy

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Oxudercinae is sometimes classified within the family Gobiidae (gobies).[3] Recent molecular studies do not support this classification, as oxudercine gobies appear to be paraphyletic relative to amblyopine gobies (Gobiidae: Amblyopinae), thus being included in a distinct "Periophthalmus lineage", together with amblyopines.[4] Mudskippers can be defined as oxudercine gobies that are "fully terrestrial for some portion of the daily cycle" (character 24 in Murdy, 1989[3]). This would define the species of the genera Boleophthalmus, Periophthalmodon, Periophthalmus, and Scartelaos as "mudskippers". However, field observations of Zappa confluentus suggest that this monotypic genus should be included in the definition.[5]

Behavior

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Mudskippers typically live in burrows in intertidal habitats, and exhibit unique adaptations to this environment that are not found in most intertidal fishes, which typically survive the retreat of the tide by hiding under wet seaweed or in tide pools.[6] These burrows are most often characterised by their smooth and vaulted ceilings. The way the males dig these burrows has been found to be directly linked to their ability to survive submerged in almost anoxic water. It has also been found to play a crucial role in the development of the eggs within the burrow. Mudskippers are quite active when out of water, feeding and interacting with one another, for example, to defend their territories and court potential partners. Once the male has completed digging his burrow he will resurface and will begin attempting to attract a female through assorted yet typical displays. These displays consist of body undulations, different postures and energetic movements. Once the female has made her choice she will then proceed to follow the male into the burrow where she will lay hundreds of eggs and allow them to be fertilized. After fertilization occurs, the period of cohabitation between the male and female is rather short. Eventually, the female will leave and it is the male that ends up guarding the egg-filled burrow from predators.[citation needed]

Mudskippers are amphibious. When leaving the water and moving into a more dry environment on land, they are still able to breathe using water that is trapped inside their large gill chambers. They are also able to absorb oxygen from the lining of their mouth and throat, allowing them to stay out of water for long periods of time. It has been discovered that they spend up to three quarters of their life on land. They are found in tropical, subtropical, and temperate regions, including the Indo-Pacific and the Atlantic coast of Africa.[citation needed]

Adaptations

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Periophthalmus gracilis (from Malaysia to northern Australia)
Periophthalmus barbarus (from western Africa)
Barred mudskipper (Periophthalmus argentilineatus)

Compared with fully aquatic gobies, these specialised fish present a range of anatomical and ethological adaptations that allow them to move effectively on land as well as in the water.[7][8]

Terrestrial movement

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As their name implies, these fish use their fins to move around in a series of skips.

Although mudskippers' fins do not have a joint homologous to the elbow, the joint between the radials and the fin rays serves a functionally analogous role.

— Integrative and Comparative Biology[9]

The mudskipper pectoral fin differs from most actinopterygian fishes in that the radials of the mudskipper pectoral fin are elongated and protrude from the body wall. This unusual morphology creates a pectoral fin with two fin segments (the radials and the rays) and two movable hinge joints: a 'shoulder' joint where the cleithrum meets the radials and an 'intra-fin' joint where the radials meet the rays.[7] In addition, ... the abductor superficialis muscle of the pectoral fin is divided into two sections (rather than being a single muscle, as is common with the rest of the Oxudercinae gobies) with one section inserting on the dorsal rays and the other section inserting on the ventral rays.

— The Journal of Experimental Biology[10]

Breathing

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Mudskipper at Point Calimere Sanctuary, Tamil Nadu, India
Mudskipper at Coringa Mangrove Sanctuary, Andhra Pradesh, India
Periophthalmodon septemradiatus territorial defense call and jumping ability[11]

Mudskippers have the ability to breathe through their skin and the lining of their mouth (the mucosa) and throat (the pharynx); this is only possible when the mudskippers are wet, limiting them to humid habitats and requiring that they keep themselves moist. The ability to breathe through their skin is associated with increased capillary density in their skin.[12] This mode of breathing, similar to that employed by amphibians, is known as cutaneous respiration.[6] Another important adaptation that aids breathing while out of water is their enlarged gill chambers, where they retain a bubble of air. These chambers close tightly when the fish is above water, due to a ventromedial valve of the gill slit, keeping the gills moist, and allowing them to function while exposed to air. Gill filaments are stiff and do not coalesce when out of water.[6]

Diet

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The different species have adapted to various diets on the mudflats. Boleophthalmus boddarti is detritivorous, while others will eat small crabs, insects, snails and even other mudskippers.[13]

Burrowing

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Digging deep burrows in soft sediments allows the fish to thermoregulate,[14] avoid marine predators during the high tide when the fish and burrow are submerged,[15] and lay their eggs.[16] When the burrow is submerged, several mudskipper species maintain an air pocket inside it, which allows them to breathe in conditions of very low oxygen concentration.[17][18][19]

Ammonia regulation

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To reduce toxic ammonia production, mudskippers can suppress amino acid breakdown when exposed to air.[20] Another method they use involves the partial breakdown of amino acids leading to the production of alanine, which also reduces ammonia production.[20]

Mudskippers can reduce the membrane permeability of their skin and acidify the water in their burrows to reduce levels of ammonia from the environment.[21][22]

Blinking

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Mudskippers evolved the ability to blink independently from terrestrial tetrapods. Their eyes are located high on their head compared to other gobies, and they blink by lowering their eyes as a membrane called the dermal cup rises to cover them. Although other fully aquatic goby species do not have the ability to blink, mudskippers did not evolve different muscles or glands to blink with; their blinking is accomplished with the same muscles in a different configuration. Rather than having specialized glands to produce eye lubricant, the fluid film on their cornea is likely composed of mucus secreted by their skin and water from their environment, possibly stored in the infraorbital space behind the dermal cup membrane. Mudskippers likely evolved blinking in response to conditions of terrestrial life, such as to keep their eyes wet out of water (they blink more frequently in high evaporation conditions and only when colliding with things in water) and to clean and protect the eye from debris, which may adhere to the eye or approach at a faster, more dangerous speed when in air versus when in water. Their eyes are not elevated when they are still juveniles, which are fully aquatic.[23]

Species

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The genus Periophthalmus is by far the most diverse and widespread genus of mudskipper. Eighteen species have been described.[24][25][26] Periophthalmus argentilineatus is one of the most widespread and well-known species. It can be found in mangrove ecosystems and mudflats of East Africa and Madagascar east through the Sundarbans of Bengal, Southeast Asia to Northern Australia, southeast China, Taiwan, and the Ryukyus, to Samoa and Tonga Islands.[3] It grows to a length of about 9.5 cm [3] and is a carnivorous opportunist feeder. It feeds on small prey such as small crabs and other arthropods.[27] However, a recent molecular study suggests that P. argentilineatus is in fact a complex of species, with at least three separate lineages, one in East Africa, and two in the Indo-Malayan region.[28] Another species, Periophthalmus barbarus, is the only oxudercine goby that inhabits the coastal areas of western Africa.[3]

References

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Revisions and contributorsEdit on WikipediaRead on Wikipedia

Mudskipper

View on Grokipedia
from Grokipedia
Mudskippers are a diverse group of approximately 42 species of small, amphibious gobies in the Oxudercinae ( Oxudercidae), renowned for their ability to spend extended periods out of water on intertidal mudflats and habitats. Native to tropical and subtropical coastal regions of the Indo-West Pacific and eastern Atlantic, these fish, typically measuring 5–30 cm in length, exhibit a unique lifestyle bridging aquatic and terrestrial environments, foraging for , defending burrows, and even performing displays on land. Key to their amphibious existence are specialized morphological and physiological adaptations, including robust, muscular pectoral fins that enable quadrupedal walking, "skipping," and , mimicking early locomotion. They respire through both gills and when emersed, supplemented by vascularized buccal cavities for air , and possess a thick, mucus-covered to minimize loss in humid intertidal zones. Ecologically, mudskippers play vital roles in ecosystems as predators of small and as prey for birds and , while their burrowing behaviors aerate and influence cycling. Recent genomic studies highlight mudskippers as models for understanding transitions to land, revealing genetic adaptations in hypoxia tolerance, , and muscle function shared with tetrapods, independent of . Despite their resilience, many species face threats from habitat loss due to coastal development and , underscoring their value as bioindicators for in dynamic estuarine systems.

Introduction

Overview

Mudskippers are small, comprising the subfamily Oxudercinae within the family and order Gobiiformes. This group includes approximately 32 distributed across 10 genera, all characterized by their ability to thrive in both aquatic and terrestrial environments. These typically measure between 2 and 30 cm in length, with elongated bodies, protruding eyes mounted on top of their flattened heads for enhanced vision on land, robust pectoral fins that function like limbs for propulsion across mudflats, and a covered in small, scales that retain moisture during emersion. They primarily inhabit the muddy intertidal zones of estuaries, mangroves, and coastal flats in the region—from eastern to Polynesia and —and the eastern Atlantic coast of , where expose them to alternating wet and dry conditions. Mudskippers have long fascinated biologists as living models for the evolutionary transition from water to land, mirroring aspects of the ancient fish-to-tetrapod shift by demonstrating adaptations for air breathing, terrestrial movement, and environmental stress tolerance. Recent genomic studies, including a 2023 analysis of chromosome-level assemblies from multiple mudskipper , have identified key genetic changes in hypoxia-inducible factors and related pathways that enhance oxygen uptake and survival in low-oxygen mud environments. The historical discovery of mudskippers traces back to 18th-century naturalist observations in , particularly the Indo-Malayan archipelago, where their distinctive skipping gait on mudflats inspired the common name and early scientific descriptions in the genus .

Habitat and Distribution

Mudskippers inhabit intertidal zones characterized by soft, muddy substrates that facilitate burrowing and foraging during low tides. Preferred environments include mangrove forests, estuaries, salt marshes, and tidal flats, where they can exploit the dynamic interface between aquatic and terrestrial realms. These s provide ample from decaying and , essential for their diet, while the soft mud allows for the construction of burrows that serve as refuges from predators and . The global distribution of mudskippers is centered in the tropical and subtropical regions of the Indo-West Pacific, spanning from through the to , , and . This range closely aligns with the extent of ecosystems, though natural populations are absent from the western Atlantic despite suitable mangroves there. A smaller subset of occurs in the eastern Atlantic, primarily along the West African coast from to , including offshore islands such as . Mudskippers exhibit broad microhabitat tolerances suited to their fluctuating intertidal lifestyle, including variations from 0 to 40 parts per thousand (ppt), which enables survival in both freshwater-influenced estuaries and hypersaline conditions during tidal exposure. Water and substrate temperatures typically range from 20°C to 35°C, with chronic thermal limits extending to 11–14°C on the lower end and up to 37°C on the upper end, allowing activity across daily and seasonal cycles. These adaptations support periodic emersion on mudflats during low , where they remain active for hours. Habitat degradation, particularly mangrove deforestation, poses significant threats to mudskipper populations by reducing available burrowing sites and food resources. Recent assessments indicate that approximately half of global forests are at risk of collapse due to deforestation and coastal development, with loss rates in key regions exceeding 1% annually in recent decades; this directly impacts mudskipper viability by fragmenting intertidal habitats. In areas like , restoration efforts highlight mudskippers as indicators of recovering ecosystems, underscoring the need for conservation to mitigate these pressures.

Taxonomy and Evolution

Classification

Mudskippers are classified within the kingdom Animalia, phylum Chordata, class , order Gobiiformes, family , and Oxudercinae. This hierarchical placement positions them among the ray-finned fishes, specifically within the diverse group of gobies and their relatives, emphasizing their aquatic origins despite amphibious behaviors. The family Oxudercidae encompasses approximately 42 species across 10 genera, with the Oxudercinae containing the mudskippers and key representatives including (commonly known as true mudskippers, comprising the most species-rich group), Boleophthalmus, Scartelaos, Periophthalmodon, and Zappa. While the polyphyletic nature of mudskippers—when defined loosely by amphibious traits—has been debated in recent taxonomic revisions, phylogenetic analyses consistently support the of Oxudercinae as a cohesive . Recent molecular studies as of 2025 confirm the separation of Oxudercidae from , with new species descriptions such as Rhinogobius phuocbinhensis in 2024 reinforcing ongoing taxonomic refinements. Historically, mudskippers were included under the broad family , but molecular phylogenetic studies in the 2000s prompted their reclassification into the distinct family to reflect evolutionary divergences within Gobioidei. This separation was formalized based on analyses of nuclear and , highlighting unique synapomorphies in oxudercines. More recent genomic sequencing efforts, including chromosome-level assemblies from 2023, have reinforced the monophyly of the subfamily and provided insights into the genetic basis of shared amphibious adaptations, resolving prior uncertainties in taxonomic boundaries.

Phylogenetic Relationships

Mudskippers, belonging to the subfamily Oxudercinae within the family (order Gobiiformes), occupy a phylogenetic position as derived gobioids closely related to other amphibious and semi-terrestrial gobies. Molecular phylogenies based on multi-locus datasets, including nuclear and mitochondrial genes from 29 oxudercine , place Oxudercinae as a monophyletic group nested within Gobioidei, with sister taxa including the semi-amphibious and various aquatic gobies like those in the Glossogobius. These analyses reveal convergence in amphibious traits, such as pectoral modifications for , across multiple genera, rather than a single evolutionary origin. The evolutionary origins of mudskippers trace back to the Eocene epoch, approximately 50 million years ago, coinciding with the global expansion of ecosystems during a period of warmer climates and rising sea levels that created extensive intertidal habitats. Fossil records of early gobioids from deposits, dating to 60–35 million years ago, indicate the emergence of the group in coastal, muddy environments similar to modern mudflats, though direct mudskipper fossils remain scarce due to the soft-sediment preservation challenges in intertidal zones. This timeline aligns with the diversification of , driven by adaptations to fluctuating salinity and oxygen levels in emerging mangrove fringes. Genomic studies highlight key genetic adaptations in mudskippers, with genes involved in hypoxia response, such as (hypoxia-inducible factor 1-alpha), showing signatures of positive selection that enhance tolerance to low-oxygen conditions on exposed mudflats. Similarly, muscle-related genes like myh7 (myosin heavy chain 7), associated with slow-twitch development for sustained terrestrial activity, exhibit adaptive evolution when compared to aquatic gobies and early tetrapods. These findings, derived from whole-genome comparisons, underscore parallel genetic mechanisms in mudskippers and tetrapods for water-to-land transitions, including enhanced oxygen sensing and contractile efficiency. Debates persist regarding the of terrestriality within mudskippers, with analyses and broader phylogenomic data indicating multiple independent transitions to amphibious lifestyles— at least twice within Oxudercinae—rather than a single monophyletic event. For instance, highly terrestrial genera like and Boleophthalmus represent separate evolutionary shifts from less amphibious ancestors, supported by incongruences between mitochondrial and nuclear markers that suggest reticulate evolution or incomplete lineage sorting. This polyphyletic pattern for terrestriality emphasizes driven by similar selective pressures in intertidal niches.

Physical Characteristics

Morphology

Mudskippers exhibit an elongated, cylindrical body form that is adapted for their semi-terrestrial lifestyle, with lengths typically ranging from 5 to 20 cm across species, though the (Periophthalmodon schlosseri) can reach up to 27 cm. The body is to elongated, featuring a robust structure supported by strong axial musculature. A key feature is the fusion of the pelvic fins into a ventral disc, which provides adhesion to substrates. The pectoral fins are muscular and elongated, with 18 to 22 rays in many species, contributing to the overall body support. The skin of mudskippers is thin and is covered by small, embedded scales that are often obscured by a mucous layer. Mucous glands are abundant throughout the , secreting a protective that aids in maintaining hydration. The epidermal structure consists of multiple layers, including an outermost keratinized layer and a germinativum, with vascularization enhancing cutaneous functions. Skeletal adaptations include a reinforced cranium to withstand terrestrial impacts, with the featuring a compact, sturdy that supports elevated structures. The eyes are positioned on short, protrusible stalks atop the head, allowing for independent movement and retraction into sockets. Pectoral skeletons comprise a cleithrum, proximal radials, and segmented fin rays, with variations in ray count correlating to locomotor demands across species. Internally, mudskippers possess a vascularized buccal cavity as a prominent feature, lined with richly supplied that forms part of the bucco-opercular chamber. This cavity is expansive relative to body size, integrated with the pharyngeal and opercular regions to support basic anatomical partitioning.

Sensory Adaptations

Mudskippers exhibit remarkable visual adaptations that facilitate effective surveillance in both aquatic and terrestrial environments. Their eyes are prominently bulging and perched atop the head, allowing for independent movement and a nearly 360-degree , which is crucial for spotting aerial and terrestrial predators while on mudflats. This positioning contrasts with typical submerged , enhancing above-water vigilance. The ocular structure includes a flattened lens paired with a steeply curved , optimizing focus for aerial vision and providing high comparable to that of some terrestrial vertebrates. To counteract in air, mudskippers employ a specialized mechanism involving a or dermal cup that periodically wets and protects the , ensuring sustained functionality during extended emersions. Corneal epithelial cells further display microstructural adaptations, such as densely packed microvilli, which aid in maintaining optical clarity under varying levels. Olfactory and gustatory systems in mudskippers are highly developed to detect chemical signals and food sources within muddy substrates. The olfactory organs are enlarged and feature a novel structure in species like the Atlantic mudskipper (Periophthalmus barbarus), characterized by expanded nasal cavities lined with densely packed lamellae that increase surface area for chemoreception in both water and air. Genomic analyses reveal significant expansions in chemosensory gene families, including olfactory receptors (ORs) and vomeronasal receptors, which underpin this enhanced sensitivity and likely evolved to support foraging and in intertidal zones. Complementing olfaction, are distributed across the head, lips, and , enabling direct sampling of potential prey in without reliance on oral . These extraoral taste receptors, embedded in the skin, allow mudskippers to assess through tactile contact, a critical for their detritus- and invertebrate-based diet in opaque environments. The system in mudskippers is notably modified for dual-media functionality, consisting primarily of superficial neuromasts embedded in the skin rather than enclosed organs typical of fully aquatic . This reduction eliminates the need for water-filled canals, which would desiccate on land, while preserving sensitivity to hydrodynamic vibrations and air currents through mechanoreceptors. Distributed along the head and body flanks, these neuromasts detect subtle movements of nearby objects or conspecifics, aiding in predator avoidance and spatial orientation during terrestrial excursions. Such adaptations maintain the system's utility across habitats, though with diminished range compared to -based systems in submerged species. Hearing in mudskippers is adapted via structures that confer sensitivity to low-frequency sounds, essential for intraspecific communication and . The saccule and utricle, key organs, exhibit morphologies that support detection of frequencies below 1 kHz, aligning with the low-frequency pulses produced during agonistic displays. Audiograms from the Atlantic mudskipper (Periophthalmus barbarus) indicate hearing thresholds similar to those of benthic aquatic fish like the banded , with best sensitivity around 200-500 Hz, but extended capabilities for airborne sound propagation near the water's edge. These features enable mudskippers to perceive substrate-borne vibrations and vocalizations effectively, bridging auditory processing between aquatic and aerial realms without specialized modifications.

Behavior

Locomotion

Mudskippers exhibit a distinctive form of known as "crutching," in which they use their robust pectoral fins, functioning like arms, to synchronously lift and vault the anterior portion of their body forward in a halting, tripod-like . This method allows them to navigate muddy or soft substrates effectively, with the trunk and remaining relatively stiff during slower movements. At higher speeds, mudskippers transition to a skipping , where the pectoral fins propel the body while the provides additional by flexing laterally, enhancing performance on deformable surfaces. These gaits enable traversal of intertidal zones. In water, mudskippers employ a less specialized form of , primarily involving undulating body movements powered by the axial musculature and caudal fin, supplemented by pectoral fin paddling at slower speeds. This swimming style is generally less efficient than their terrestrial crutching due to their elongated, goby-like , which is optimized for bottom-dwelling rather than sustained open-water , resulting in shorter stride lengths and reliance on higher-frequency strokes. For escaping predators, mudskippers perform coordinated jumps using simultaneous thrusts from the pectoral fins and lateral flexion of the body and tail, propelling them distances of up to 60 cm—several times their body length. The caudal fin plays a key role in generating forward momentum during these leaps. Terrestrial locomotion imposes a higher metabolic cost on mudskippers compared to aquatic movement, with elevated oxygen demands that limit sustained activity. can extend exercise duration by reducing anaerobic reliance, highlighting the role of atmospheric oxygen in modulating expenditure during land-based travel.

Social Interactions

Mudskippers display pronounced territorial behaviors, particularly among males, who vigorously defend their burrows and surrounding areas from intruders. These displays often involve visual signals such as erecting and waving the , rapid tail beating against the substrate, and acrobatic leaps to assert dominance. In some species, territorial maintenance includes the construction of mud balls around entrances, which may serve to reinforce boundaries or deter rivals. Such behaviors are especially evident in the genus , where aggression levels are notably high, correlating positively with territory size—larger territories are associated with more intense defensive actions. Courtship in mudskippers incorporates a suite of visual and tactile signals to attract potential mates, including push-up movements using the pectoral fins, chasing displays across the , and body postures that highlight coloration changes. These interactions are often accompanied by substrate vibrations generated through tail beats or body undulations, providing an auditory component that propagates through the wet ground. Tactile elements, such as gentle nips or fin touches, further facilitate close-range communication during these encounters. In non-breeding periods, mudskippers typically maintain solitary lifestyles or form small, loose aggregations on mudflats, with interactions limited to occasional territorial skirmishes rather than sustained group cohesion. Aggression varies by species; for instance, individuals exhibit heightened confrontational responses compared to less territorial genera like Boleophthalmus. Communication modalities adapt to the environment: visual cues dominate on exposed land surfaces for clear signaling during daylight, while chemical pheromones play a key role in aqueous settings for longer-range detection. Acoustic signals, often low-frequency pulses and tones transmitted via substrate vibrations, complement these during aggressive or bouts, as observed in Periophthalmodon septemradiatus. A 2025 neoichnological study in PALAIOS analyzed trace fossils from Periophthalmus and Periophthalmodon trackways, revealing patterns indicative of social interactions, such as overlapping paths and defensive maneuvers preserved in mud and sand substrates.

Reproduction

Mudskippers typically exhibit polygynous mating systems, in which territorial males construct and defend burrows to attract multiple females through elaborate displays, such as fin flaring, push-ups, tail stands, and acrobatic leaps, often enhanced by nuptial coloration during the breeding season. These displays occur primarily on exposed mudflats during , with males leading receptive females into the burrow for spawning. Spawning takes place within the male's , where the female deposits eggs on the or walls of an air-filled subterranean chamber, and the male externally fertilizes them before the female departs. Clutch sizes vary widely among species and with female body size, ranging from a few hundred to over 30,000 eggs per spawn; for instance, in species, mean can reach approximately 6,600 eggs, while in Apocryptes bato it averages around 17,590 eggs. Breeding seasons are often synchronized with periods or warmer months, such as February to May in tropical populations. Following spawning, males provide extensive by guarding the burrow and maintaining oxygenation in the egg chamber, which involves periodic visits to swallow atmospheric air and release it over the clutch to aerate the developing embryos. Embryonic incubation in this aerial environment typically lasts 7 to 14 days, protecting eggs from aquatic predators and while allowing . To initiate , the male floods the chamber with water during high tide, submerging the eggs and prompting rapid larval emergence, often within seconds to minutes, after which he may assist in releasing the larvae to the . Hatched larvae enter a pelagic phase, drifting in coastal waters for 30 to 50 days while feeding on , before undergoing into juveniles at total lengths of 10 to 15 mm, at which point they transition to the . Juveniles grow rapidly in mangroves or mudflats, reaching at sizes of about 10 to 11 cm and ages of 6 to 12 months, depending on and conditions, with a lifespan potentially up to 5 years.

Physiological Adaptations

Respiration

Mudskippers employ a buccal-opercular mechanism to facilitate air during emersion, wherein they gulp atmospheric oxygen into their expanded buccopharyngeal and opercular cavities, which serve as primary sites for . This process involves rhythmic movements of the mouth and opercular flaps to draw in and circulate air, allowing oxygen to diffuse across the highly vascularized lining of the bucco-opercular cavity. When out of water, mudskippers derive a substantial portion of their total oxygen uptake from air, with some species fully satisfying resting metabolic demands aerially, depending on species and environmental conditions, with the remainder supplemented by residual aquatic respiration if gills remain moist. The gills of mudskippers are reduced in size and filament density compared to fully aquatic gobies, limiting their efficiency in air but enabling them to retain moisture and contribute modestly to oxygen uptake through diffusion when emersed. Cutaneous respiration via the highly vascularized skin provides an additional pathway for oxygen diffusion, particularly across the thin epithelium of the head and body, while the richly perfused buccopharyngeal lining functions analogously to a primitive lung by enhancing aerial gas exchange. These adaptations collectively support sustained terrestrial activity in hypoxic intertidal zones. Mudskippers can tolerate emersion for several hours to days provided the substrate remains moist, preventing of their respiratory surfaces and maintaining functionality. In response to hypoxia, they exhibit upregulated gene expression in metabolic pathways for anaerobic ATP production and defenses under air exposure. During tidal transitions, mudskippers switch from predominantly -based aquatic respiration to air at low tide, accompanied by an immediate increase in upon the first aerial gulp to optimize oxygen delivery and circulatory support for terrestrial exertion. This physiological shift enables efficient exploitation of exposed mudflats. In burrow-dwelling species, air-breathing behaviors also briefly aid in aerating eggs by replenishing oxygen in nest chambers.

Osmoregulation

Mudskippers demonstrate exceptional osmoregulatory capabilities, enabling them to inhabit intertidal zones with salinities fluctuating between brackish and near-marine conditions. As teleosts, they act as both hypo- and hyper-osmoregulators, maintaining at 260–300 mOsm across external salinities of 4–27‰, which prevents excessive loss or gain despite osmotic gradients. In hypo-osmotic environments such as freshwater or dilute , mudskippers actively uptake essential ions like Na⁺ and Cl⁻ to sustain internal balance. This process occurs primarily through specialized ionocytes in the gills, with supplementary uptake via the skin, which features epidermal ion-transporting cells that enhance ionoregulatory efficiency during immersion. During emersion, mudskippers shift to terrestrial strategies to minimize water loss and toxicity. They reduce urine production, which decreases excretion rates by 29–76%, conserving body water while limiting nitrogenous waste accumulation. is excreted mainly as NH₄⁺ across the gills and , but in confined environments, aerial volatilization of NH₃ facilitates , supported by active proton excretion that acidifies burrow water and reduces NH₃ permeability through the . Burrows are integral to , providing a humid microhabitat that retards cutaneous loss and during extended emersion periods. The retained in burrow walls and pools supports hydration, allowing mudskippers to endure tidal cycles without severe osmotic stress.

Feeding Mechanisms

Mudskippers are primarily carnivorous and insectivorous, with diets consisting mainly of small crustaceans such as and , insects like dipterans, worms, and occasionally small fish or organic ; some species opportunistically consume or plant matter. This opportunistic scavenging allows them to exploit the nutrient-rich , where prey availability fluctuates with tides and seasons, leading to shifts such as increased insect intake during wet periods and more crustaceans in dry conditions. Their foraging techniques are adapted for both mudflats and shallow , featuring protruding that enable scooping prey directly from the substrate. Lacking a true , mudskippers rely on a novel "hydrodynamic " mechanism: they retain in the buccal cavity upon emerging from , which is then protruded and retracted to envelop terrestrial prey via , followed by gulping to transport it intra-orally. This -based feeding is supplemented by rapid protrusion and hyoid depression, allowing effective capture on land where traditional aquatic is less viable. On land, much of mudskippers' foraging occurs terrestrially, employing "head-first" lunges toward visually detected prey from distances up to 2 m, facilitated by their elevated, independently rotating eyes for wide-field aerial vision. Prey capture success is higher on land for small, mobile items due to these , contrasting with slower aquatic approaches limited to closer ranges. Digestive adaptations support their amphibious lifestyle, including a short gut relative to body length that enables rapid processing of protein-rich prey, consistent with carnivorous habits. Feeding is size-selective, constrained by gape limitations to small prey, optimizing intake in patchy environments. The intestinal mucosa's tolerance to varying levels from intertidal prey aids efficient absorption without specialized structures.

Ecology and Conservation

Ecological Role

Mudskippers occupy a mid-level trophic position in intertidal ecosystems, functioning primarily as carnivorous or omnivorous predators that consume small invertebrates such as crustaceans, , polychaete worms, and copepods, thereby exerting top-down control on these populations and influencing community structure. Stable isotope analyses indicate their role as secondary to tertiary consumers within food chains. In this role, they also serve as important prey for avian predators like and , as well as terrestrial mammals and larger crabs, integrating energy flow across trophic levels. As ecosystem engineers, mudskippers significantly influence sediment dynamics through extensive burrowing in anoxic mudflats and mangrove zones, where their activities aerate compacted substrates and promote the oxygenation of deeper layers, fostering microbial decomposition and nutrient remineralization. This bioturbation enhances and cycling by facilitating the exchange of porewater with overlying tides, reducing sulfide toxicity and supporting in these nutrient-limited environments. Neochronological studies of their trace fossils, such as sinuous trails and networks, serve as modern analogs for interpreting ancient incursions into terrestrial habitats, highlighting their role in evolutionary and paleoecological reconstructions. High population densities of mudskippers, reaching up to 12 individuals per square meter in optimal conditions, contribute to substrate stabilization by redistributing sediments and preventing excessive during tidal cycles, thereby maintaining integrity for associated benthic communities. Their and burrowing behaviors bridge aquatic and terrestrial food webs, transferring and energy from detrital sources to higher predators, which sustains across the intertidal gradient in systems. Mudskippers form loose associations with burrowing and benthic within their networks, where shared spaces may offer mutual benefits such as enhanced against visual predators and supplemental from algal biofilms lining walls. These interactions, observed in diverse assemblages, underscore mudskippers' contributions to multi-species bioturbation that amplifies overall resilience.

Predators and Threats

Mudskippers face predation from a variety of terrestrial and aquatic species in their and habitats. On land, they are commonly preyed upon by birds such as (e.g., purple herons, Ardea purpurea) and , as well as snakes like the dog-faced water snake (Cerberus schneiderii), which actively hunt them during . In water, larger predatory fish target them during aquatic phases, while crabs, including larger species that share the , occasionally consume smaller individuals. To evade these predators, mudskippers employ rapid escape behaviors, including high-speed jumps using their powerful pectoral fins and tail, which allow them to propel themselves away or even past threats on mudflats. They also retreat into self-dug burrows for , particularly during high or tidal changes, enhancing their in exposed environments. Anthropogenic threats significantly endanger mudskipper populations through and . Coastal urbanization and aquaculture expansion have led to substantial loss, with approximately 20-22% of global mangrove forests disappearing since the (as of 2020 data), though recent rates have slowed to about 0.16% annually due to conservation efforts. from industrial effluents and clogs their gills, impairs respiratory function, and causes structural damage to gill tissues, while accumulate and disrupt overall . Climate change exacerbates these pressures by altering mudskipper habitats and physiological limits. Rising sea levels, which accelerated in due to and ice melt, inundate tidal zones and shift distributions, potentially displacing mudskipper populations from optimal foraging areas. Warmer temperatures stress their emersion tolerance, with exposure to 32°C increasing oxygen uptake rates and reducing movement efficiency, signaling potential metabolic overload under projected future warming. Overcollection for the aquarium trade and as fishing bait further depletes local populations, particularly in where species like Periophthalmus are harvested intensively for export and local use, leading to reduced abundances in accessible mudflats.

Conservation Status

The majority of mudskipper species are classified as Least Concern (LC) on the , reflecting their widespread distribution and relatively stable populations across Indo-Pacific mangrove and intertidal habitats, with assessments primarily conducted between 2017 and 2024. For instance, the common mudskipper (Periophthalmus kalolo) is rated LC due to its broad range and lack of immediate threats. However, a few species face higher risks; the great blue spotted mudskipper (Boleophthalmus pectinirostris) is listed as Vulnerable () under criterion A2bcd, primarily owing to ongoing habitat loss from coastal development and , with its population inferred to have declined by at least 30% over the past three generations. The version 2025-2 (as of October 2025) incorporates updates following 2023 assessments and confirms these statuses without significant revisions for mudskippers. Mudskippers are indirectly protected through mangrove reserves that encompass their critical habitats, such as the Wonorejo in , where multiple species thrive amid conservation efforts to preserve intertidal zones. In , extensive mangrove areas along Queensland's coast, including protected regions under the , provide safeguards for species like the barred mudskipper (Periophthalmus argentilineatus), supporting their ecological needs. No mudskipper species is currently listed under , as they remain for international trade, though monitoring of ornamental and medicinal trade is recommended for potentially exploited populations. Ongoing research employs genomic tools to assess and , enabling better tracking of and connectivity among mudskipper populations in fragmented . Mangrove restoration projects, including propagule planting in degraded areas, aim to rehabilitate mudskipper , with varying success depending on site and species selection, though targeted efforts have shown improved survival and recovery, contributing to net gains in some regions as of 2025. Challenges persist, including data gaps for Atlantic species like Periophthalmus barbarus, where limited assessments hinder comprehensive risk evaluation despite its LC status. Enhanced establishment of intertidal protected zones is essential to address habitat threats and ensure long-term viability.

Species Diversity

Major Genera

Mudskippers belong to the subfamily Oxudercinae within the family , encompassing 10 genera and approximately 42 species distributed across tropical and subtropical coastal habitats. The genera are distinguished primarily by differences in pectoral fin morphology, which supports varying degrees of , and burrow architectures adapted to intertidal environments. A 2023 phylogenetic analysis using mitochondrial COI gene sequences confirmed three major genetic clades among common mudskipper populations, aligning with genus-level groupings based on these morphological traits. The genus , comprising 22 species, represents the most diverse and widespread group, often referred to as "true" mudskippers due to their advanced terrestriality. These species exhibit elongated pectoral radials that enable skipping, crawling, and climbing on land, with eyes positioned dorsally for above-water vigilance. They inhabit mudflats and mangroves across the and eastern Atlantic, where individuals actively forage on exposed substrates during low tide. Boleophthalmus, with 5 species, consists of burrow-dwelling mudskippers predominantly found in the region. These construct complex burrow systems, often J- or Y-shaped, for refuge, spawning, and feeding, and employ a mud-flap technique where they scoop sediment into their mouths to filter out and diatoms. Their pectoral fins are adapted for short terrestrial excursions around burrow entrances, emphasizing a semi-aquatic lifestyle tied to burrow maintenance. The genus Scartelaos includes 4 that show a tendency toward herbivory compared to other mudskippers, incorporating significant amounts of matter, diatoms, and into their diet alongside small . These build simpler vertical burrows and are distributed in estuaries, using their robust jaws for scraping algal films from substrates. Lesser genera include Apocryptes (1 species, restricted to rivers and estuaries), Parapocryptes (2 species, mangrove dwellers with cryptic burrowing habits), Periophthalmodon (1 species, including larger forms up to 27 cm in length), Apocryptodon (3 species), Oxuderces (2 species), Pseudapocryptes (1 species), and Zappa (1 species). These smaller genera exhibit varying burrow types, from simple shafts to networked systems, reflecting niche adaptations in coastal ecosystems.
GenusNumber of SpeciesKey DistributionNotable Trait
Periophthalmus22Indo-Pacific, AtlanticHigh terrestriality via pectoral fins
Boleophthalmus5Mud-flap feeding, complex burrows
Scartelaos4Herbivorous lean, algal scraping
Periophthalmodon1Larger body sizes up to 27 cm
Apocryptodon3Estuarine habitats
Others (6 genera)7Varied burrowing and cryptic forms

Notable Species

The barred mudskipper (Periophthalmus argentilineatus) is a prominent species commonly inhabiting mangrove forests and intertidal mudflats across the Indo-West Pacific region, ranging from the east coast of Africa through India to Pacific islands such as Fiji and Papua New Guinea. This amphibious goby, reaching a maximum length of 19 cm, exhibits remarkable locomotor abilities, including acrobatic jumps propelled by its pectoral fins to capture aerial prey or evade threats during terrestrial excursions. These jumps, analyzed kinematically, demonstrate enhanced propulsion in aerial prone postures compared to non-aerial movements, highlighting adaptations for efficient movement on deformable mud substrates. Boleophthalmus boddarti, an African mudskipper species endemic to coastal mudflats in regions like the , is notable for constructing tall mud chimneys around its s, which serve to regulate burrow microclimates and deter intruders during . These structures, often exceeding 20 cm in height, facilitate feeding by creating protected areas amid soft sediments. The species faces significant conservation concerns due to degradation from oil spills in West African mangroves, which contaminate sediments and disrupt burrow-building behaviors essential for survival. The Japanese form of Scartelaos histophorus, known as the bearded or walking mudskipper, inhabits intertidal sand and mud flats from to and , including estuarine swamps and tidal zones. This species features pronounced eye stalks that elevate independently, providing a 180° for detecting predators and prey on land without head movement. Studies on its visual adaptations reveal corneal modifications, such as increased epithelial thickness and changes, that enhance aerial acuity and reduce in air, critical for its semi-terrestrial lifestyle in Japanese coastal habitats. The Atlantic mudskipper (Periophthalmus barbarus), a smaller reaching about 10 cm in length, occurs along tropical Atlantic coasts of and exhibits less pronounced terrestrial behaviors compared to its Indo-Pacific congeners, spending more time in shallow water than on exposed mudflats. Recent 2022 studies highlight morphological differences, including reduced fin musculature for emersion and lower vascularization for , underscoring its intermediate amphibious adaptations relative to more land-oriented like Periophthalmus novemradiatus.

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