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Pleuronectidae

The Pleuronectidae, commonly known as righteye flounders, are a family of flatfishes in the order Pleuronectiformes characterized by having both eyes positioned on the right side of the head in adults, with a strongly compressed, oval to elongate body adapted for a benthic lifestyle.^[1]^[2] These fishes lack fin spines, possess a dorsal fin that extends onto the head, and exhibit no swimbladder in adults, enabling them to undergo rapid color changes to camouflage against the seabed.^[1]^[3] Pleuronectidae comprise approximately 24 genera and 59 species (as of 2018), including notable members such as plaice (Pleuronectes platessa), halibuts, and dabs, distributed across the Arctic, Atlantic, Indian, and Pacific Oceans, primarily in marine environments but occasionally entering brackish or freshwater habitats.^[1]^[2]^[4] They inhabit soft-bottom substrates like mud, sand, or silt at depths ranging from shallow coastal waters (as low as 6 m) to 1,000 m, where they prey on small benthic invertebrates and fishes using their small mouths and elongate gill rakers.^[1]^[3] Taxonomically, the family falls within the suborder Pleuronectoidei of the class Teleostei, with a fossil record dating back to the lower Tertiary Eocene; recent phylogenetic studies recognize five subfamilies within the family.^[2]^[1]^[4] Most species are pelagic spawners with eggs lacking oil globules, and nearly all hold commercial importance in fisheries due to their abundance and edibility.^[1]

Taxonomy

History of Classification

The family Pleuronectidae was first established by Constantine Samuel Rafinesque in 1815, who described it as a distinct group within the flatfishes characterized by both eyes positioned on the right side of the head, a small mouth, and elongate dorsal and anal fins with numerous rays.^[5] This initial classification placed Pleuronectidae as a family under the order Pleuronectiformes, encompassing marine species with a compressed body adapted for benthic life across Arctic, Atlantic, and Pacific waters.^[5] During the late 19th and early 20th centuries, ichthyologists refined the internal structure of Pleuronectidae through morphological analyses. Theodore Gill, in his 1893 work "Families and Subfamilies of Fishes," contributed significantly by proposing detailed subfamily divisions based on fin ray counts, scale patterns, and vertebral morphology, distinguishing groups such as Pleuronectinae from emerging taxa like Hippoglossinae.^[6] These revisions built on earlier efforts by figures like Cuvier and Valenciennes, incorporating comparative anatomy to address ambiguities in species placement and emphasizing the family's diversity in northern temperate seas. Subsequent works, such as those by Norman in the 1930s, further adjusted subfamily boundaries using osteological traits, solidifying Pleuronectidae as a core component of right-eyed flatfishes.^[7] Molecular phylogenetics in the 2010s prompted major shifts in the higher-level placement of Pleuronectiformes, including Pleuronectidae. Betancur-R et al. (2017) analyzed extensive genomic data from nearly 2,000 fish species, revealing that flatfishes form a monophyletic clade within the series Carangaria; this classification recognizes Pleuronectiformes as an order nested in Carangaria, supported by shared molecular synapomorphies like specific ribosomal RNA sequences.^[8] Later studies, such as Harrington et al. (2016), reinforced this using ultraconserved elements, confirming a single origin of cranial asymmetry and integrating Pleuronectidae into a broader carangarian framework that challenges traditional morphology-based orders. Fossil discoveries have enriched the evolutionary history of Pleuronectidae, with extinct genera providing insights into its Cenozoic diversification. For instance, Chibapsetta, an extinct right-eyed flounder known from skeletal remains, represents early members of the family and highlights adaptations predating modern species diversity. Miocene records, including related taxa like those from Japanese formations, document the family's expansion into subtropical habitats during this epoch, with fossils showing transitional morphologies between primitive and derived pleuronectids.^[9]

Current Taxonomy

Pleuronectidae is classified in the kingdom Animalia, phylum Chordata, class Actinopterygii, order Pleuronectiformes, suborder Pleuronectoidei, and family Pleuronectidae. This placement reflects molecular phylogenetic revisions that integrate flatfishes into the series Carangaria, emphasizing their evolutionary ties to percomorph fishes based on genomic data.^[8] The family encompasses 59 species distributed across 24 genera, representing a diverse yet cohesive group within the suborder Pleuronectoidei. Phylogenetic analyses utilizing multilocus DNA sequencing, including four nuclear and three mitochondrial genes, have robustly supported the monophyly of Pleuronectidae with high bootstrap values and Bayesian posterior probabilities. These studies demonstrate close evolutionary relationships to other flatfish lineages in the suborder, such as the sand flounders (Paralichthyidae), forming a well-supported clade within Pleuronectoidei that diverged from more basal flatfish groups like Psettodoidei.^[10]^[8] A key morphological distinction of Pleuronectidae from lefteye flounders (Bothidae) lies in the direction of ocular migration during larval metamorphosis: in Pleuronectidae, the left eye migrates dorsally to join the right eye, resulting in both eyes positioned on the right (ocular) side when the fish lies on its left (blind) side as an adult. This asymmetry contrasts with Bothidae, where the right eye migrates to the left side, and is corroborated by developmental studies highlighting the genetic and cellular mechanisms driving this trait. Such differences underscore the adaptive radiation of flatfishes for benthic lifestyles while maintaining phylogenetic proximity within Pleuronectoidei.^[11]^[10]

Subfamilies and Genera

The family Pleuronectidae is classified into five subfamilies based on morphological and molecular phylogenetic analyses: Paralichthodinae, Poecilopsettinae, Pleuronectinae, Rhombosoleinae, and Samarinae.^[12] This division reflects monophyletic groupings supported by multilocus phylogeny, encompassing diverse forms adapted to various marine environments.^[12] Note that some taxonomic catalogs retain traditional subfamilies (e.g., Atheresthinae, Hippoglossinae), but the 2018 revision is adopted in sources like WoRMS. The Paralichthodinae includes large Pacific flounders, such as genera like Paralichthodes, adapted to deeper waters. Poecilopsettinae comprises small, deep-sea righteye flounders in genera like Poecilopsetta, often with ornate patterns. Pleuronectinae, the core group, includes plaice-like forms in genera such as Pleuronectes (e.g., P. platessa, European plaice) and Limanda, dominant in northern temperate regions. Rhombosoleinae features southern hemisphere species like Peltorhamphus (New Zealand flounders), with robust bodies. Samarinae contains Indo-Pacific genera like Samariscus and Yamatakea, characterized by smaller sizes and tropical distributions. Pleuronectidae's diversity shows regional endemism, with the northwestern Pacific as a hotspot.^[13]

Morphology and Anatomy

Body Plan

Pleuronectidae, known as righteye flounders, exhibit a highly asymmetric body plan adapted for a benthic lifestyle, with both eyes positioned on the right (ocular) side of the head and the left (blind) side oriented downward when the fish rests on the substrate.^[1]^[14] This dextral asymmetry arises during larval development through the migration of the left eye across the dorsal surface to join the right eye, resulting in a flattened, laterally compressed form that allows the fish to lie flush against the seafloor.^[15] The body shape is typically oval to diamond-like, with strong lateral compression that enhances camouflage and maneuverability near the bottom; larger species may appear more elongate or robust.^[16]^[3] The fin structure supports this compressed morphology, featuring a long dorsal fin that originates anterior to or on the head and extends continuously to the caudal fin, often with no spines throughout.^[1]^[3] The anal fin is similarly continuous and spineless, running along the ventral margin from near the anus to the tail, and its length correlates with that of the dorsal fin, varying from short in rounded-bodied forms to elongate in more oblong species.^[16] Pelvic fins are present but often reduced or short, particularly on the blind side; pectoral fins are asymmetrical, with the eyed-side fin typically larger and more developed than the blind-side counterpart.^[3] The caudal fin is generally rounded or truncate, not confluent with the dorsal and anal fins. Scales on the ocular side are typically ctenoid, bearing comb-like spines that provide texture and aid in sediment interaction for burial and camouflage, while those on the blind side are smooth cycloid or weakly ctenoid, facilitating contact with the substrate.^[17] This bilateral difference in scale morphology is pronounced in many species, such as Platichthys flesus, where eyed-side scales are tuberculate with multiple spines, contrasting with the smoother blind-side scales.^[17] Size within the family varies widely, from small species, such as the dab (Limanda limanda), reaching a maximum of about 25 cm total length, to the massive Atlantic halibut (Hippoglossus hippoglossus) attaining up to 4.7 m.^[3]^[18]^[19] This range reflects diverse ecological niches, from shallow-water dwellers to deep-sea giants.^[14]

Sensory Adaptations

Pleuronectids exhibit remarkable visual adaptations suited to their benthic habitat, where they lie camouflaged on the seafloor. During metamorphosis, the left eye migrates to the right (eyed) side of the head, resulting in both eyes positioned dorsally to provide binocular vision for detecting prey overhead while the original left eye position becomes reduced or vestigial on the blind side. This arrangement enhances depth perception and stereoscopic acuity in low-light conditions, with opsin genes like rh1 upregulated for scotopic vision and lens proteins such as cryaa and crygs4 supporting optical clarity in both eyes post-migration. Genetic analyses indicate positive selection on visual perception genes, including dmbx1a and opn3, facilitating this asymmetry as an evolutionary response to ambush predation.^[20]^[21] Camouflage in Pleuronectidae relies on dynamic coloration mediated by chromatophores, enabling rapid adjustments to match sedimentary substrates for predator avoidance and prey ambush. Melanophores, iridophores, and lipophores expand or contract under neural control to produce mottled brown, gray, or ochre patterns that disrupt the body outline against sand, gravel, or mud. In juvenile plaice (Pleuronectes platessa), for instance, two independent pattern components—fine spots and coarser blotches—are flexibly combined within minutes of environmental change, with spots predominating on high-contrast gravel and blotches on pebbled surfaces to optimize crypsis. This chromatophore-driven adaptability is crucial in variable benthic environments, where effective matching reduces visibility to predators.^[22] The lateral line system in Pleuronectidae displays bilateral asymmetry reflective of their flattened body plan, with cranial canals present on both eyed and blind sides to detect hydrodynamic disturbances. While some canals, such as the supraorbital and infraorbital, show reductions on the blind side in up to 43% of genera, the system retains functionality for sensing water vibrations from approaching predators or nearby prey movements. This mechanosensory capability compensates for the blind side's limited vision, allowing orientation and escape responses in turbid or dark conditions where visual cues are unavailable. Evolutionary patterns of canal reduction and branching highlight independent adaptations across genera, enhancing survival in sediment-disturbed habitats.^[23] Olfactory and gustatory senses in Pleuronectidae are well-developed to locate and assess buried or hidden food in sediments, particularly on the blind side. Nostrils facilitate olfaction, detecting chemical cues from prey like crustaceans, which prompts localization and enhances visual strikes in diurnal species such as plaice and dab. Taste buds distributed across the blind-side head, lips, and fin rays enable direct sampling of substrates, distinguishing palatable items through chemoreception upon contact. These adaptations are integral to foraging efficiency, as demonstrated by increased biting responses to shrimp-derived odors in experimental settings.^[24]

Distribution and Habitat

Global Distribution

Pleuronectidae, the family of righteye flounders, exhibit a broad global distribution primarily across the Northern Hemisphere's major ocean basins, including the Arctic, Atlantic, Indian, and Pacific Oceans.^[1] They are notably absent from the southernmost Antarctic waters, with no recorded species in the high-latitude Southern Ocean regions dominated by other flatfish families.^[25] This distribution reflects their adaptation to temperate and colder marine environments, spanning from coastal zones to continental slopes. The latitudinal range of Pleuronectidae extends from polar regions to subtropical and tropical waters. In the Arctic, species such as the Greenland halibut (Reinhardtius hippoglossoides) are found circumpolar in the North Atlantic and Arctic Ocean, inhabiting cold, deep waters.^[26] This wide latitudinal coverage underscores the family's ecological versatility across thermal gradients. Most Pleuronectidae species are benthic dwellers on coastal and continental shelf habitats, typically at depths from shallow inshore areas to around 500 meters. However, certain deep-water forms, particularly in the subfamily Hippoglossinae (such as halibuts), extend to 2,000 meters or more, exemplified by the Greenland halibut occurring at 400–2,000 meters.^[26] Endemism is particularly pronounced in the temperate North Pacific, where the majority of the family's species diversity is concentrated, with many genera restricted to regions off North America and Asia.^[27] For instance, Japanese waters host a high number of endemic genera, contributing significantly to the regional richness of approximately 17 genera.^[7]

Habitat Preferences

Pleuronectidae species exhibit a strictly demersal lifestyle, residing on or near the seafloor where they rely on camouflage and ambush predation. They predominantly occupy soft-bottom substrates, including mud, sand, and gravel, which provide suitable conditions for burial and foraging on benthic prey. This preference for fine to coarse sediments is evident across the family, with species like the common dab (Limanda limanda) favoring sandy bottoms and the winter flounder (Pseudopleuronectes americanus) utilizing muddy to sandy areas.^[1]^[28]^[29] Depth preferences vary widely within the family, from shallow coastal zones to deep-sea environments. Many species, such as the European plaice (Pleuronectes platessa), inhabit continental shelves at depths of 0–100 m, typically 10–50 m, while larger forms like the Atlantic halibut (Hippoglossus hippoglossus) extend to abyssal depths exceeding 2,000 m. This range allows adaptation to diverse benthic habitats, from nearshore sands to deep mud slopes.^[30]^[31] Salinity tolerance is primarily marine, with most species thriving in full seawater conditions of 30–35 ppt, though some exhibit euryhalinity. For instance, the European flounder (Platichthys flesus) routinely enters brackish estuaries and even freshwater, tolerating salinities as low as 0 ppt. Temperature ranges are generally cold-temperate, from 0–20°C, supporting species like plaice in 2–15°C waters; however, some members, such as the starry flounder (Platichthys stellatus), occupy warmer margins up to 25°C in subtropical coastal areas.^[1]^[32]^[30]^[33]

Life History

Reproduction

Members of the Pleuronectidae family reproduce via external fertilization, in which females release batches of pelagic eggs into the water column that are subsequently fertilized by males. These eggs are typically spherical, buoyant, and measure 1-2 mm in diameter, with a smooth, transparent chorion and homogeneous yolk.^[34]^[35] Spawning seasonality varies by latitude and species; in temperate regions, it occurs during winter to spring, as seen in European plaice (Pleuronectes platessa), which spawn from December to May at water temperatures around 6°C.^[36] Fecundity is notably high in larger species, with female Atlantic halibut (Hippoglossus hippoglossus) capable of producing up to 2 million eggs per season through repeated batch spawnings over several weeks.^[37] Sexual size dimorphism is common, with females growing larger than males to support higher egg production. Courtship and mating behaviors include promiscuous spawning, where males exhibit heightened swimming activity—often peaking around midnight—to approach females and release milt into the water column near the substrate to fertilize the eggs, sometimes involving multiple males converging on a spawning pair.^[38]^[39]^[40]

Development and Metamorphosis

The eggs of Pleuronectidae species are pelagic and buoyant, typically measuring 1-2 mm in diameter and containing a large yolk reserve to support initial embryonic development.^[41] These eggs are released in offshore waters during spawning and float near the surface, where they are subject to dispersion by currents.^[42] Hatching occurs after 5-20 days, with the exact duration inversely related to water temperature; for example, in the plaice (Pleuronectes platessa), a representative species, eggs incubated at 6-8.5°C hatch in 13-26 days.^[43] Upon hatching, prolarvae emerge at 3-5 mm in length, initially relying on the remaining yolk sac for nutrition before transitioning to exogenous feeding.^[44] The larval stage in Pleuronectidae is characterized by bilaterally symmetrical, upright-swimming forms that resemble typical teleost larvae rather than the leaf-like leptocephali of eels, though they share a planktonic lifestyle.^[45] Larvae grow to 10-20 mm over 30-60 days, feeding primarily on zooplankton such as copepods and rotifers, which they capture using a protrusible mouth and median fins for propulsion.^[46] In species like the plaice, this premetamorphosis phase lasts approximately 67 days on average, during which larvae remain pelagic and are transported by ocean currents toward coastal nurseries.^[46] Pigmentation develops gradually, with early larvae showing minimal markings that intensify as they approach metamorphosis. Metamorphosis in Pleuronectidae marks the transition to the asymmetric adult form and typically spans 2-4 weeks, triggered by thyroid hormones and environmental cues like settlement habitats.^[47] During this process, the left eye migrates dorsally and posteriorly to join the right eye on the upper (ocular) side of the body, while the body flattens through rapid skeletal remodeling and muscle adjustments, positioning the fish to lie on its left side (blind side).^[48] The original left-side eye position becomes the blind side, with that eye becoming non-functional and covered by skin, and the mouth twists to align with the new orientation. Settlement to the benthic environment occurs at 10-30 mm total length; for instance, plaice settle at 13-14 mm once eye migration is complete, shifting from pelagic to demersal habits. This phase is energetically costly, often involving reduced feeding and increased vulnerability.^[47] Mortality during the egg and larval stages in Pleuronectidae is exceptionally high, primarily due to predation by gelatinous zooplankton and fish, as well as starvation and advection away from suitable settlement areas.^[49] Survival from hatching to juvenile settlement is generally less than 1%, with larval predation accounting for the majority of losses during the extended planktonic phase.^[49] In controlled rearing of related flatfish species, survival rates to metamorphosis average 3-6%, but wild conditions yield even lower figures due to environmental stressors.^[50]

Growth and Longevity

Growth in Pleuronectidae begins post-settlement, when juveniles typically measure 12-15 mm, and is characterized by rapid somatic expansion during the first year of life, often reaching 20-30 cm in length for species such as the European plaice (Pleuronectes platessa), before decelerating with advancing age as energy allocation shifts toward maintenance and reproduction.^[51]^[52] This initial fast growth phase supports quick attainment of sizes that reduce vulnerability to predation, with annual increments diminishing thereafter; for instance, plaice exhibit peak growth in the first 6 years, after which rates slow considerably.^[52] Growth patterns reflect indeterminate growth, where individuals continue to increase in size throughout life, albeit at progressively lower rates, as evidenced by otolith annuli that record lifelong accretion without a defined asymptotic limit.^[53] Size at sexual maturity varies widely across the family, influenced by species-specific ecology and sexual dimorphism, with females generally maturing at larger sizes than males. In smaller species like the common dab (Limanda limanda), maturity is typically reached around 25 cm, while in larger forms such as the Atlantic halibut (Hippoglossus hippoglossus), females attain maturity at approximately 100-125 cm.^[54]^[55] Age at maturity aligns with these size thresholds, generally occurring between 2 and 5 years, though environmental factors like temperature can modulate this progression.^[52] Longevity in Pleuronectidae spans a broad range, with smaller species such as plaice and dab typically living 10-20 years, exemplified by plaice reaching about 15 years in the Barents Sea.^[52] Larger species like halibuts exhibit extended lifespans, with Atlantic halibut documented to exceed 50 years, allowing for substantial post-maturational growth and repeated reproductive cycles.^[55] Age determination relies primarily on otolith analysis, where translucent winter rings form annually, providing a reliable record of growth history and confirming the family's indeterminate growth strategy across diverse habitats.^[53]

Ecology and Behavior

Feeding Ecology

Members of the Pleuronectidae family exhibit a carnivorous diet primarily composed of benthic organisms, with composition varying by species size and habitat. Smaller species, such as plaice (Pleuronectes platessa) and dab (Limanda limanda), predominantly consume invertebrates including polychaetes (often 40-70% of diet by weight), amphipods, and bivalve mollusks, reflecting their opportunistic feeding on abundant seafloor resources.^[56]^[57] Larger species, exemplified by the Atlantic halibut (Hippoglossus hippoglossus), shift toward more mobile prey, with fish comprising 40-80% of their intake (e.g., cod, haddock, and herring) alongside cephalopods and large crustaceans.^[56]^[58] Foraging in Pleuronectidae typically involves ambush predation, where individuals rely on cryptic coloration and stillness to blend with the substrate, awaiting prey within striking distance before rapidly expanding their protrusible mouths to capture it.^[59]^[60] This visually oriented strategy is enhanced by their elevated ocular adaptations, allowing detection of prey movement above the sediment.^[60] Ontogenetic shifts in diet are pronounced across the family, with juveniles targeting smaller, sessile or infaunal prey such as polychaetes and tiny crustaceans to support rapid early growth, while adults transition to larger, more mobile items like fish and cephalopods as body size and gape increase.^[56]^[61] For instance, young plaice feed heavily on terebellid polychaetes and shrimps, but larger individuals incorporate more bivalves and occasional fish.^[57] As mid-level carnivores, Pleuronectidae occupy trophic levels of approximately 3.0 to 4.0, channeling energy from detrital-based food webs—via their invertebrate prey—into higher marine trophic strata.^[62] This positioning underscores their role in benthic ecosystem dynamics, linking primary detritivores to piscivorous predators. Members of the Pleuronectidae family exhibit predominantly solitary social behavior as adults, maintaining small home ranges with occasional overlap among conspecifics, as observed in species like the European flounder (Platichthys flesus).^[64] Juveniles often form loose aggregations in nursery habitats to reduce individual predation risk, though true schooling is rare outside of reproductive contexts.^[65] During spawning, some species aggregate in larger groups; for instance, plaice (Pleuronectes platessa) migrate solitarily to spawning grounds but form dense assemblages upon arrival, facilitating broadcast spawning. Similarly, winter flounder (Pseudopleuronectes americanus) show group convergence at spawning sites, with multiple males often interacting around a single female, though strictly paired spawning occurs in only about 22% of events.^[40] Migration patterns in Pleuronectidae are closely tied to reproductive cycles and environmental cues, particularly temperature. Many species undertake inshore-offshore movements for spawning; winter flounder, for example, migrate from offshore feeding grounds to shallow inshore bays and estuaries during winter months.^[66] Plaice exhibit seasonal shifts, moving to deeper offshore waters in winter while returning to coastal spawning areas in spring, with migrations spanning hundreds of kilometers guided by tidal currents and salinity gradients.^[67] Depth changes often correlate with temperature preferences, as individuals seek optimal thermal ranges to avoid extremes, influencing both juveniles settling in warmer shallows and adults relocating during non-reproductive periods.^[64] Anti-predator strategies in Pleuronectidae emphasize crypsis and evasion, with burrowing into sediment being a primary defense mechanism across species, allowing rapid concealment from visual predators.^[64] Rapid color and pattern changes enable matching of substrate textures and shades, enhancing camouflage effectiveness, as demonstrated in juvenile north Pacific flatfishes like English sole (Parophrys vetulus).^[68] Some species, including plaice juveniles, display increased nocturnal activity to exploit reduced predator visibility at night, balancing foraging needs with risk avoidance.^[69] Communication within Pleuronectidae is limited and primarily occurs during courtship, relying on visual fin displays and possibly chemical cues. In winter flounder, males perform pectoral fin fanning to court females and oxygenate egg masses post-spawning, signaling reproductive readiness in close proximity.^[40] Olfactory signals may guide aggregation at spawning sites, as plaice respond to pheromonal cues in locating mates, though broader social interactions lack complex signaling.^[69]

Conservation and Human Impact

Fisheries and Exploitation

Pleuronectidae species are important targets in commercial fisheries, particularly in temperate waters of the North Atlantic and North Pacific Oceans. Major species exploited include the Atlantic halibut (Hippoglossus hippoglossus), European plaice (Pleuronectes platessa), and yellowtail flounder (Limanda ferruginea), among others such as yellowfin sole (Limanda aspera) and Pacific halibut (Hippoglossus stenolepis). Global catches for the family have been substantial, with key species contributing around 258,000 tonnes in 2018 alone, and overall family-level production estimated at approximately 500,000 tonnes annually in the pre-2020 period based on aggregated FAO data.^[70] Fishing for Pleuronectidae primarily employs bottom trawls, which account for the majority of harvests due to the demersal habits of these flatfishes, supplemented by gillnets and longlines for larger species like halibuts. These methods have been in use since the 19th century, when intensive exploitation—particularly of Atlantic halibut—led to early signs of overexploitation in North Atlantic stocks as steam-powered vessels expanded fishing ranges. For instance, 19th-century halibut fisheries in the Gulf of St. Lawrence experienced rapid declines due to unregulated trawling and longlining, setting a precedent for subsequent management challenges.^[71]^[72] Aquaculture of Pleuronectidae remains limited compared to capture fisheries, with Atlantic halibut being the primary species under cultivation, mainly in Norway and Iceland. Production reached about 1,918 tonnes in 2018, valued at roughly US$2.3 million, but faces significant hurdles in larval rearing and metamorphosis, where high mortality rates during the transition to benthic life hinder scalability.^[73] Economically, Pleuronectidae species command high market value as premium whitefish for human consumption, with Atlantic halibut fetching prices up to US$10–15 per kg in export markets. They also appear as bycatch in broader demersal fisheries targeting cod or shrimp, contributing to incidental harvests that can exceed 10–20% of total flatfish landings in some regions.^[70]

Conservation Status

Pleuronectidae species face multiple threats that impact their populations, primarily overfishing, which has led to depleted stocks in several regions, habitat degradation from bottom trawling that disrupts nursery grounds, and climate change effects on larval survival through altered temperature regimes and ocean acidification.^[74]^[75] Overfishing remains the dominant pressure, with historical exploitation reducing biomass in key stocks, while trawling damages benthic habitats essential for juveniles. Climate-induced changes, such as warming waters, can shorten larval durations or increase mortality rates, exacerbating recruitment variability.^[76]^[77] Conservation statuses vary across Pleuronectidae species, reflecting regional differences in exploitation and environmental pressures; for instance, the American plaice (Hippoglossoides platessoides) is classified as Endangered by the IUCN due to ongoing declines in the Northwest Atlantic, while the European plaice (Pleuronectes platessa) is Least Concern globally but with some regional stocks showing depletion.^[78] The Atlantic halibut (Hippoglossus hippoglossus) is assessed as Near Threatened, an improvement from earlier Endangered status following management interventions, though populations remain below historical levels in parts of the Northeast Atlantic. Some stocks, such as North Sea plaice, have shown recovery after implementing quotas, with spawning stock biomass increasing post-2000 due to reduced fishing mortality.^[79] Protective measures include quota systems advised by the International Council for the Exploration of the Sea (ICES) and coordinated by the Food and Agriculture Organization (FAO) of the United Nations, which set total allowable catches for species like plaice and halibut to maintain sustainable yields. Marine protected areas, such as the North Sea Plaice Box—a partially closed zone restricting large beam trawlers to protect juvenile plaice—have contributed to stock rebuilding by reducing discard rates and habitat damage. Bycatch regulations under frameworks like the European Union's Common Fisheries Policy mandate selective gear and observer programs to minimize unintended captures of undersized or non-target flatfishes.^[80]^[81] Looking ahead, warming oceans may drive range shifts poleward for many Pleuronectidae species, potentially benefiting northern stocks but risking local extinctions in southern habitats, with projections indicating 20-30% of flatfish species facing heightened vulnerability from combined climate and fishing pressures. Enhanced adaptive management, including dynamic quotas and expanded protected areas, will be crucial to mitigate these risks and ensure resilience.^[82]^[83]

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Apr 19, 2021 · Our work provides valuable resources and insights for understanding the genetic origins of the unusual body plan of flatfishes.<|separator|>
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Juvenile plaice (Pleuronectes platessa) produce camouflage by ...
Sep 1, 2006 · Plaice (Pleuronectes platessa) is a flatfish well-known for the ability to vary its body pattern, probably for camouflage.
[23]
Lateral line system of flatfishes (Pleuronectiformes): Diversity and ...
Sep 21, 2019 · The IOC is totally absent in the other half of the genera. The blind side SOC is reduced in 40% of genera, the blind side IOC is reduced in 43%, ...
[24]
https://academic.oup.com/icesjms/article/32/3/385/714275
[25]
Pleuronectidae - an overview | ScienceDirect Topics
Order Pleuronectiformes: 11 families, 570 species, including flounders and soles. Order Tetraodontiformes: 9 families, 339 species, including triggerfishes ...
[26]
Greenland Turbot | NOAA Fisheries
A cousin of the Pacific halibut, Greenland turbot are a right-eyed flatfish. As they develop, their left eye migrates across the top of the skull toward the ...
[27]
How the Central American Seaway and an Ancient Northern ...
May 21, 2018 · There are far more Pleuronectidae species in the Pacific Ocean, most of them endemic to the north Pacific Ocean off of North America and ...
[28]
Citharichthys sordidus summary page
### Summary of Citharichthys sordidus Habitat Information
[29]
Pseudopleuronectes americanus, Winter flounder - FishBase
Adults inhabit soft muddy to moderately hard bottoms (Ref. 5951). Feed predominantly in daytime on organisms living in, on or near the bottom.
[30]
Pleuronectes platessa summary page
### Summary of Pleuronectes platessa Habitat Preferences
[31]
Hippoglossus hippoglossus summary page
### Summary of Habitat Information for Hippoglossus hippoglossus
[32]
Platichthys flesus summary page
### Summary of Habitat Information for Platichthys flesus
[33]
Syacium papillosum summary page
### Summary for Syacium papillosum (Dusky flounder)
[34]
(PDF) Reproductive characteristics and egg development in ...
Newly ovulated eggs were spherical and buoyant, with a diameter of 1.075-1.213 mm (avg 1.156±0.025 mm), a colorless transparent chorion, a slightly yellowish ...
[35]
Egg Information Summary - Pleuronectes platessa - FishBase
buoyant (pelagic). Shape of Egg, spherical. Attributes, smooth, not sticky. Color of ... Mean egg diameter in the Barents Sea = 2.17 mm (Ref. 124126). Get ...Missing: Pleuronectidae | Show results with:Pleuronectidae
[36]
Pleuronectes platessa, European plaice - FishBase
Life cycle and mating behavior Maturity | Reproduction | Spawning | Eggs | Fecundity | Larvae. Adult spawn when the temperature is around 6 °C (Ref. 4705) ...
[37]
All Fishing Buy, Flatfishes, Order Pleuronectiformes, Heterosomata ...
Most of the species have pectoral fins, one on eyed side and the other on the blind side. The ventral fins are in front of the pectorals or in line with them; ...<|control11|><|separator|>
[38]
[PDF] Atlantic Halibut, Hippoglossus hippoglossus, Life History and ...
Large females may produce up to 2 million eggs; can spawn numerous batches of eggs within a single reproductive season. Halibut in the GM are thought to ...
[39]
(PDF) An evolutionary explanation of female-biased sexual size ...
Sexual size dimorphism (SSD) is caused by differences in selection pressures and life- . history trade- offs faced by males and females.Missing: Pleuronectidae | Show results with:Pleuronectidae
[40]
Sexual differences in spawning behaviour and catchability of plaice ...
Males were more active than females during the time of spawning, an increase in swimming activity mainly being observed during the hours around midnight.
[41]
(PDF) Behavior of winter flounder, Pseudopleuronectes americanus ...
Spawning by one pair frequently elicited sudden convergence and spawning by secondary males (up to six individuals); consequently, strictly paired spawning was ...
[42]
Pleuronectiform | Flatfish Adaptations & Characteristics - Britannica
Pleuronectiform, (order Pleuronectiformes), any one of about 680 species of bony fishes characterized by oval-shaped, flattened bodies as in the flounder, ...Missing: Carangiformes current
[43]
Potential transport of plaice eggs and larvae between two ...
Their life cycle is characterised by offshore spawning, a period of egg and larval planktonic drift followed by metamorphosis and settlement into shallow, ...
[44]
Effect of temperature on the embryonic development of the plaice ...
In normalyears, eggs and prolarvae of the plaice (Pleuronectes platessa L.) in the southern North Sea develop within the temperature range 6.0–8.5 °C, ...
[45]
Temperature-dependent development rates of plaice (Pleuronectes ...
On average, plaice eggs from the Irish Sea developed more rapidly and hatched up to 2 days earlier compared with North Sea eggs incubated at the same ...
[46]
Life History cycles in flatfish from the Northwestern Pacific, with ...
Nearly 300 species of Pleuronectiformes are found in the western Pacific, of which 120 species are recorded around Japan. In this area the geographical ...
[47]
Protein metabolism, growth and pigmentation patterns during ...
In the present work, development of plaice larvae in premetamorphosis lasted on average 67 days. This corresponds to the stages from hatching to stage 3c (Table ...
[48]
The cost of metamorphosis in flatfishes - ScienceDirect.com
Size at metamorphosis, duration of metamorphosis and synchrony of metamorphosis with settlement vary between and sometimes within species. The order of ...
[49]
Eye migration and cranial development during flatfish metamorphosis
Aug 7, 2025 · Larval development and settlement were largely size dependent, resulting in reduced larval stage duration and earlier settlement at higher ...
[50]
Settlement, abundance, growth and mortality of juvenile flatfish in a ...
Larval mortality and subsequent yearclass strength in the plaice (Pleuronectes platessa L.) J.H.S. Baxter (Ed.), The early life history of fish, Springer, ...
[51]
Recent Developments in Techniques for Rearing Marine Flatfish ...
Jul 29, 2025 · Survival curves show that the highest period of larval mortality occurs between days 5-12 after hatching. Overall survival averages 3-6 ...
[52]
Growth dynamics of European plaice Pleuronectes platessa L. in ...
... Overall, growth conditions in temperate flatfish species appear to be in the order of 0.5-1.0 mm d −1 and observations about stunted growth or starvation ...Missing: maturity | Show results with:maturity
[53]
Long-term variability of the growth rate of Barents Sea plaice
European plaice grow fastest during their first 6 years and are usually <0.5 m, weigh 2-3 kg and have a longevity of approximately 15 years.
[54]
Estimating growth from sex ratio-at-length data in species with ...
Indeterminate growth results from allocating less and less energy to growth ... SSD is very frequent in many fish species including plaice and most flatfish ...
[55]
(PDF) Common dab (Limanda limanda) fisheries biology in the ...
Sexual maturity was found to occur at 184 mm (2.56 years). Thus, the suggested MLS was 185 mm, in order to maintain dab population in a sustainable way, and ...Missing: longevity | Show results with:longevity
[56]
Age determination, bomb-radiocarbon validation and growth of ...
Aug 9, 2025 · Halibut is a long-lived species (38-50 years) with females maturing at larger sizes (103-125 cm) and older ages (7-13 years) than males (55-80 ...
[57]
(PDF) The trophic ecology of flatfishes - ResearchGate
Apr 13, 2021 · Many flatfish primarily consume benthic prey, converting their energy into a form more accessible to higher trophic levels (Link et al., 2015) .
[58]
Feeding ecology and growth of O‐group flatfish (sole, dab and ...
Apr 1, 2005 · O-group plaice diet was dominated by polychaetes (Terebellidae), crustaceans and molluscs at all sizes. O-group sole, dab and plaice did not ...
[59]
Hippoglossus - Atlantic halibut - FishBase
Feed mainly on other fishes (cod, haddock, pogge, sand-eels, herring, capelin), but also takes cephalopods, large crustaceans and other bottom-living animals.
[60]
[PDF] Untitled - ResearchGate
to stimulate the lateral line system of both visual and nonvisual feeders, and can be useful in both prey detection and predator avoidance. While visual systems ...
[61]
Pleuronectiformes (Flatfishes) - Encyclopedia.com
Pleuronectiformes(Flatfishes) Class ActinopterygiiOrder PleuronectiformesNumber of families Approximately 13 Source for information on Pleuronectiformes ...
[62]
(PDF) Feeding of plaice Pleuronectes platessa L. and Sole Solea ...
Sep 25, 2025 · Flatfish, the European plaice, and lemon sole mainly feed upon polychaetes, mollusks, and crustaceans living on or within the sediment ( ...
[63]
Trophic Ecology of Alaska Plaice Pleuronectes quadrituberculatus ...
Mar 10, 2018 · The trophic level is in the range from 3.39 to 4.13; its variability depends on the proportion of bivalves and polychaetes. We described the ...
[64]
The Behaviour of Flatfishes - ResearchGate
We explored whether anti-predator behavior and intrinsic growth are co-evolved traits in 3 co-occurring juvenile flatfish species: English sole Pleuronectes ...Missing: schooling | Show results with:schooling
[65]
Diel movements of juvenile plaice Pleuronectes platessa in relation ...
Aug 5, 2025 · The distribution and movements of juvenile plaice Pleuronectes platessa and their potential predators and competitors were recorded in a ...
[66]
Winter Flounder | NOAA Fisheries
In the winter, adults migrate from offshore areas where they feed to inshore bays and estuaries where they spawn. While inshore, they live on muddy sand ...
[67]
Plaice | Institute of Marine Research
Apr 20, 2020 · Adults migrate each year between spawning areas in central and southern parts of the North Sea and undertake feeding migrations to areas further ...
[68]
Adaptive coloration, behavior and predation vulnerability in three ...
Apr 28, 2008 · ... color or texture may render fish more conspicuous to predators ... The adaptive significance of camouflage as an anti-predator tactic in flatfish ...
[69]
The Behaviour of Flatfishes - Wiley Online Library
Feb 24, 2005 · This chapter contains sections titled: Introduction. Locomotion and related behaviour. Colour change. Reproduction. Feeding. Predation and ...
[70]
[PDF] Flounders, halibuts, soles Capture production by species, fishing ...
plaice(=Long rough dab). Balai(=Plie canadienne). Platija americana. Hippoglossoides platessoides. 1,83(02)014,01. PLA. 21 Canada. 1 903. 1 924. 998. 760. 1 592.
[71]
Small flatfish (flounders, soles) bottom trawling - Fishing Techniques
A single boat bottom otter trawl is one cone-shape trawl towed on the seabed by one boat with its horizontal spread maintained by a pair of otter boards.Missing: methods | Show results with:methods
[72]
The Rise and Fall of the Nineteenth-Century Atlantic Halibut Fishery
Aug 5, 2025 · Overfishing in the 19th century reduced vast beds of oysters in Chesapeake Bay and other estuaries to a few percent of pristine abundances ...
[73]
Sources, Quantities and Cultivation Methods - Seafish
Atlantic halibut (Hippoglossus hippoglossus) is a large, slow growing flatfish species in the family Pleuronectidae, maturing around 10-14 years.
[74]
Vulnerability of flatfish and their fisheries to climate change
Climate change may cut catches by 6-10% by 2050, with overfishing threatening 33% of marine species (51, 55) . Investments of ₹50000 crore ($6 billion) by 2030 ...
[75]
Effects of climate change on coastal ecosystem food webs
Climate change is modifying coastal ecosystem food web structure and function and threatens these essential ecosystem services.
[76]
(PDF) Temperature effect on growth and larval duration of plaice ...
Transport models for planktonic fish eggs and larvae often use temperature to drive growth because temperature data are readily available.
[77]
Effects of warming rate, acclimation temperature and ontogeny ... - NIH
Jul 27, 2017 · At faster rates of warming, larval CTmax significantly increased in herring, whereas no effect was observed in seabass. Higher acclimation ...
[78]
https://www.iucnredlist.org/species/18214783/162705101
[79]
Combined Ex post and ex ante evaluation of the long term ...
Mar 17, 2023 · The North Sea plaice stock is now fished at a level below the management regulation target for this stock (<0.3 per year). The annual rate of F ...
[80]
Latest advice - ICES
You will find the latest official ICES advice on this page. You can also search for our advice by species or ecoregion in the library.
[81]
[PDF] Effects of a partially closed area in the North Sea (''plaice box'') on ...
When more than one ICES rectangle was visited during a trip, effort was split according to the estimated value of the plaice and sole landings in each rectangle ...
[82]
Flatfish Catches to Decline 20% by 2100 Due to Climate Change
Aug 1, 2018 · Climate change will drastically reduce flatfish numbers and alter species distributions by hundreds of kilometers by the end of the century.
[83]
Temperature change effects on marine fish range shifts
May 30, 2023 · The current effects of global warming on marine ecosystems are predicted to increase, with species responding by changing their spatial distributions.