1125:
the ocean is crucial for all marine life as some species are specialized for different regions or use temperature to direct migrations. Thus, species that reside in temperate latitudes tend to have broader preferences and tolerances then species specialized for extreme temperatures. This makes both tropical and polar species more susceptible to changes in climate. Flatfish are negatively affected by these temperature shifts and continue to decline in populations. Globally, their numbers dwindle and have been since the 1970s. The use of technology has allowed researchers to calculate numbers of existing populations as well as estimate the future of the clade. Two of these methods, a vulnerability assessment and simulation model projections, help pinpoint the regions at highest risk of this change and align with previously mentioned latitudinal gradient predictions. Findings suggest that the
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484:
53:
1035:
1210:
536:
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31:
981:: The senses are used to take in both visual and chemical cues (which are important for marine species). Both of these factors have been proved to increase the attractiveness of prey items. In addition to those factors, flatfish have keen motion detection via "wide-set eyes" and a "lateral line system" on their head or body that can be stimulated by motion.
1185:). Climate change is a multivariate problem for scophthalmids as it has led to changes in distribution and net productivity in addition to the aforementioned decreases in population. Yet, these changes in distribution are not exclusive to potentially obvious latitudinal range shifts. The flatfish have also been observed migrating within regions,
758:
propulsive stroke.' This is highly useful to escape predation, initiate gliding, or journey far distances while conserving energy (e.g. traveling between feeding and spawning sites). While the propulsive stroke may be energetically costly, once in the water column the flatfish can exhibit 'swim and glide' behaviors to cruise through the water.
815:
754:, the flatfish use forward and backward movements to propel themselves across the substrate. Oftentimes, these movements are short, with pauses in between strokes (akin to gliding). That being said, the fish can increase their movement speed through "rapid propulsive strokes" that can propel individuals in different directions.
1133:
are at the highest risk of impacts due to climate change (i.e. a tropical region and polar region). As mentioned previously, scophthalmids are almost exclusively located in the
Northeast Atlantic and will suffer heavily from this. Interestingly enough, the rapid warming was exacerbated regionally due
835:
does not explain as to why this is, but it could be speculated that the murkiness caused by burying in muddy substrates can be rather visible to predators, counteracting the primary function of this behavior. Time of day can also be important, as visibility decreases with access to light. Due to this
1124:
is a common term in describing the change of a certain variable with respect to global latitudes. Due to the tilt of the Earth as well as the distance from the sun, this latitudinal gradient is often proportional to the global temperature gradient, both on water and land. The temperature gradient of
811:(ii) Second, the head movements "waft sediment from beneath the body so that it falls back onto the surface of the fish." In this latter have of the movement the fishes do not simply swim into the substrate, they actively displace the substrate so that it falls back on top of them and conceals them.
1087:
tends to take place in the evening, and males and females generally pair due to relative body size. One interesting fact about flatfish is that most species are actually born symmetrical, and their eyes rotate as they mature (can travel 10-25mm on average). As the fish develop, the non-dominant eye
947:
All three feeding types involve the various elements of feeding behavior. Visual predation is the most common in flatfish, but the non-visual nocturnal species rely on chemical cues left in the water (e.g. distress signals, high heart rates, oxygen consumption, cortisol levels, olfactory chemicals,
474:
and excluding one species located in the
Western Atlantic, all other scophthalmids reside in the general Northeastern Atlantic region. However, mapping out a specific organism's changes in habitat over their lifespan would be rather complex. Environmental events can displace fish or incline them to
1205:
species). Because of this, it is harder to migrate, maneuver, and adapt, making their adjustment period relatively slow. All finfish (including scophthalmids), however, are expected to shrink. The decrease in body size relates to the temperature change via oxygen levels. The growing restriction on
1066:
follow the female individual across the sea floor, and then maneuvers either above or below the female depending on the species. If successfully courted, the female (now with the male) rise through the water column. The pair reaches the climax of the courtship ritual as they reach the "apex of the
757:
In addition to the "rapid propulsive strokes," scophthalmids can elevate their heads into an alert position. Using both of these adaptations in tandem, the fishes can change the angle of their projection, and shoot themselves into the water column by lifting their heads and then executing a 'rapid
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committed) is when this method is used on the seafloor where the nets are weighted and dragged. Frowned upon by marine biologists, bottom trawling is harmful to the ecosystem due to the displacement of sediment and organisms as well as the irreversible damage it creates. In addition to negatively
515:
they can exit the water column and sink to the bottom. This is crucial to energy expenditure, especially in migratory events between feeding and spawning grounds that take place every year. During migrations tidal cycles are primary in determining the vertical displacement of flatfish in the
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share a strong inclination to bury themselves and to cease movement, fighting the desire to flee until extremely close to detection. In addition to being inclined to hiding, in the instance they are being caught, flatfish face the disadvantage of their maximum swim speed. Only able to sustain a
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for the family. While decreasing depth can help scophthalmids fight the change in ocean temperature, the deeper they travel the less light can penetrate and both conditions and productivity also decrease. This is a negative constraint on the fish which will eventually reach its limit.
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Highly specialized behaviors are hard to change, making the flatfishes' natural instinct to minimize detection rendered useless. Usually they can be rather cryptic, and have great success avoiding natural predators. Utilizing their low body profile and texture-matching
761:
Besides the lever-system that can propel the fish in the transverse plane, scophthalmids also have unique systems to control their vertical and horizontal movements. Due to their asymmetry, their bodies are on a 90 degree rotation and use their fins differently than most
1239:, and use detection minimization techniques as their primary avoidance behavior. These strategies combine various techniques such as “ burial, highly evolved cryptic capabilities, and low activity,” all of which are useless and therefore detrimental in fighting trawls.
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Little is known in regards to the reproduction of scophthalmids beyond what is widely accepted or generalized in the broader flatfish order. When a female enters a reproductive state (due to internal or external stimuli), courtship is usually initiated by the male. Male
479:
behaviors. While true, movements of the species can be tracked for instances of displacement as well as seasonal cycles of migration–both of which “appear to be triggered by changes in water temperature, light, current velocity, and irregular episodic events.”
839:
Lastly, burying behavior does not just limit predation detection via visual cues. Indeed, burial can reduce "conspicuousness," but flatfish can employ additional tools to solidify their defense. Scophthalmids are believed to be capable of lowering
1169:. Furthermore, some fisheries have overfished certain regions to the point of collapse. In these regions a bounce-back in population is not expected. This change is not just due to temperature change, but additional factors tied to
801:. To initiate this important and unique behavior, flatfish vigorously beat their heads "against the sediment, accompanied by a wave of muscular that with decreasing amplitude down the length of the body." This odd combination of
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spend most of their time below other taxa. Being asymmetrical and spending most of their time on the bottom substrates, scophthalmids have evolved unique locomotive systems to better fit their lifestyle. Turning their bodies into
638:
and have slowly translated their dorsal fin (over the course of evolutionary history) onto the head. Poking eyes out of the substrate and using this dorsal fin akin to a rudder, these three traits all serve the flatfish in its
586:, or ray-finned fish, share traits will all prior marine vertebrates (i.e. a skeleton, paired fins, cycloid or ctenoid scales, covered gills, homocercal tails, protrusible jaws, etc...) The main requirements for the taxa are
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certain speed at relatively lower rates than most roundfish, flatfish tend to respond to the trawls in short bursts and generally remain unresponsive until "approx. <1 meter away." After observing the adaptations
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Location is crucial to these behaviors as some substrates are impenetrable to flatfish. This facet actively selects for specific substrates that the fish can bury in (e.g. sand), and dictate their home ranges. In
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herd in the regard by which they are all migrating in the same direction. This increase in concentration is dangerous as it increases catch per trawl, oftentimes when the flatfish are not even the desired target
1976:
Cheung, W.W.L; Sarmiento, J.L.; Dunne, J.; Frölicher, T.L.; Lam, V.W.Y.; Palomares, M.L.D.; Watson, R.; Pauly, D. (2013). "Shrinking of Fishes
Exacerbates Impacts of Global Ocean Changes on Marine Ecosystems".
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have made to survive in the Ocean, bottom trawling proves to be a major industrial threat to scophthalmids because of its own nature in specifically targeting and taking advantage of flatfish behavior.
1792:
Tittensor, Derek P.; Mora, Camilo; Jetz, Walter; Lotze, Heike K.; Ricard, Daniel; Berghe, Edward Vanden; Worm, Boris (August 2010). "Global patterns and predictors of marine biodiversity across taxa".
552:), so it exhibits any primitive traits of that clade. Being a scientific family, it also has its own set of general traits, as well as a variety of derived traits specific to the different genera of
520:
The location of turbots is cryptic and multivariable, consisting of a mixture of both these cycles, an internal “clock” mechanism, and other factors such as turbulence, hunger, and magnetic fields.
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are native to (i.e. the Baltic, Mediterranean, and Black seas) all fall under this hotspot. It seems that this regional increase of warming can be traced to the “natural variability related to the
527:, making them sink in the absence of currents and upwellings. To preserve energy, this is their natural state. They are rather sedentary and the fish spend most of their time on bottom substrates.
1693:
Stoner, Allan W.; Ottmar, Michele L.; Hurst, Thomas P. (November 2006). "Temperature affects activity and feeding motivation in
Pacific halibut: Implications for bait-dependent fishing".
1231:
affecting the ecosystem, bottom trawling proves to be highly dangerous to scophthalmids as well as other families of flatfish. This method of fishing exploits scophthalmids’ natural
1639:
Ramachandran, V. S.; Tyler, C. W.; Gregory, R. L.; Rogers-Ramachandran, D.; Duensing, S.; Pillsbury, C.; Ramachandran, C. (1996). "Rapid
Adaptive Camouflage in Tropical Flounders".
999:: Stalking is a method different from ambushing. Taxon that use this style of capture lure outside of detection of the prey item, and their primary weapon is stealth and striking.
383:
families of flatfish. These traits can be localized to three main requirements: “i) anterior extension of the two pelvic fins; ii) eyes present on the left-side; iii) presence of
797:
as it is their primary defense in avoiding predators and hunting prey. By burying, the fishes avoid detection from predation risks, and also reduce environmental challenges like
2250:
Burrows, Michael T.; Gibson, Robin N. (July 1995). "The effects of food, predation risk and endogenous rhythmicity on the behaviour of juvenile plaice,Pleuronectes platessaL".
647:
side. This pigmentation serves as a camouflage for the fish and allows them to increase their survivorship via heightened predator avoidance and improved hunting behaviors.
2178:
Handegard, N. O.; Tjøstheim, D. (2005). "When fish meet a trawling vessel: examining the behaviour of gadoids using a free-floating buoy and acoustic split-beam tracking".
2126:
971:: Searching tends to be initiated via hunger, but can also be induced via chemical signals in the water. Generally a slow process, they tend to swim upstream to track
2285:
Lemke, Jena L.; Ryer, Clifford H. (June 2006). "Risk sensitivity in three juvenile (Age-0) flatfish species: Does estuarine dependence promote risk-prone behavior?".
2498:
2012:
Tittensor, D.P.; Mora, C.; Jetz, W.; Lotze, H.K.; Ricard, D.; Berghe, E.V.; Worm, B. (2010). "Global
Patterns and Predictors of Marine Biodiversity Across Taxa".
2537:
1121:
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Female can indicate their "readiness to spawn by rapidly raising and lowering their heads." This in turn inclines males to approach and produce a threat
343:: The features of the family were first recorded and compared with other taxa. The relationship between scophthalmids and other flatfish became apparent.
1197:
Lastly, scophthalmids are simply less mobile than other taxa. Their activity rate is much lower than other fish of similar size and location (alternate
1111:
Visual representation of the Change in
Temperature in the North Atlantic from 1850 to 2000. Each stripe represents the temperature averaged over a year.
716:
Other non-defining traits of scophthalmids that are not specific to the family are their large and prominent lower jaws, sinistral eyes, and the single
2314:"Relative predation vulnerability of three juvenile (Age-0) North Pacific flatfish species: possible influence of nursery-specific predation pressures"
2472:
2511:
2215:
Healey, E. G. (1999). "The skin pattern of young plaice and its rapid modification in response to graded changes in background tint and pattern".
887:. Flatfish take advantage of complex environments while most fish have no choice but to hunt in open water. This gives the fishes relatively high
1945:"Integrating ecophysiology and plankton dynamics into projected maximum fisheries catch potential under climate change in the Northeast Atlantic"
1146:. The highest levels of warming (“LMEs warming at rates 2–4 times the global mean rate” ) was confined to certain areas, especially in the
1107:
349:: This distinction was then further specified when scophthalmids were paired with other left-eyed species of flatfish. This subfamily is called
1206:
oxygen levels proportional to the increase in temperature negatively affects body size, adding yet another side effect to the changing climate.
975:
clues and primarily rely on vision during the hunt (they have extensive vision in all planes due to their protrusive eyes and cranial anatomy).
1762:
de Groot, S. J. (April 1969). "Digestive System and
Sensorial Factors in Relation to the Feeding Behaviour of Flatfish (Pleuronectiformes)".
1610:
1541:
511:, to adult) use the elliptic and cyclic nature of Earth's tides in locomotion. The rising tide carries individuals shoreward and during the
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leave their home ranges. All fishes, including scophthalmids, also face predation risks and prey scarcity, as well as the complexities of
2614:
1005:: Ambush species use their unique adaptations and combine stillness, burying, or cryptic coloration until prey is in striking distance.
831:, it is noted that this preference of substrate drifts towards sandy over coarse, as well as an avoidance of overly muddy substrates.
417:
process in contact with the dorsal margin of the blind side frontal.” Again, these traits are also not perfect as defining a specific
1446:
1275:, they become hard to detect. Furthering their camouflage, they have behavioral modifications that work in tandem to their anatomy.
413:, redefined the subfamily's traits to two more distinct features: “(i) anterior extension of the two pelvic fins; (ii) an elongated
2576:
1165:
In addition to this fact, the increasingly negative population dynamics of all flatfish indicates that the clade has reached its
1862:
Cheung, William W.L.; Oyinlola, Muhammed A. (October 2018). "Vulnerability of flatfish and their fisheries to climate change".
622:(flatfish) share a lot in common with some minor distinctions. Based on the name, it is right to assume that most of genera of
1460:
Froese, R. and D. Pauly. Editors. 2022. FishBase. World Wide Web
Electronic Publication. www.fishbase.org, Version (02/2022).
2516:
503:
The movement of water is crucial to the species as it is a primary form of transportation. All stages of the flatfish (from
2619:
2524:
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are 'flat' and have a generally 'low' body profile. Due to this anatomical difference, flatfish are quite unique and have
163:
52:
1099:." Speculation as to why is not solidified in scientific literature but it is possibly due to competition in paternity.
872:, this force also helps dictate population size and drives growth to be proportional to the various seasons. Being that
2433:
523:
But with all said and done, the flatfish don't go too far. The anatomy of these organisms is intrinsically negatively
643:
lifestyle. Lastly, the generalized coloration of flatfish include a very pale underside with a cryptically pigmented
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factor, it is expected that burial behaviors are less common in scophthalmids at lower depths and during the night.
2609:
1155:
428:
312:
2542:
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will give higher insight onto the specific feeding-type of said species. By the same merit, below are the general
860:
Feeding is crucial to any organisms behavior as it is necessary to sustain life. Unlike other organisms, however,
745:
Scophthalmids live a fairly sedentary lifestyle in comparison to other ray-finned fish, and due to their negative
1246:
sent through the water. This stimulus is received by the fish ("which have good hearing in the range of 300–1000
1186:
557:
331:
epoch. This specific fossil is believed to be from the upper
Tertiary and is the oldest specimen of the family.
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2604:
849:
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1309:
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features. In adult individuals, both eyes lye on the same side of the head on whichever side is dominant (
435:. In addition to these synapomorphies, a third was added, namely the “presence of asymmetrical transverse
1226:
is when nets are dragged through the water to catch as many fish as possible. Bottom trawling (primarily
2563:
1602:
888:
675:
354:
2438:
1905:
Rice, Jake; Cooper, J.Andrew (November 2003). "Management of flatfish fisheries—what factors matter?".
993:: Pursuit is common in faster taxon as the individual uses sheer speed to run down free-swimming prey.
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1986:
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are native to the North Atlantic, as well as the Baltic, Mediterranean, and Black seas. According to
241:
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39:
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2068:"Sound Physiological Knowledge and Principles in Modeling Shrinking of Fishes Under Climate Change"
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987:: Capture consists of three sub-elements that can dictate the style of capture or 'type of hunt.'
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process (often crenulated) in contact with the dorsal margin of the right (blind-side) frontal"
268:
47:
2568:
371:. To make said distinction, various species of similar traits were combined, primarily from the
516:
water column, while on feeding and spawning grounds these behaviors are more influenced by the
466:
environments, but not in freshwater. While certain flatfish can be found in the other regions,
267:) reaches approximately one meter in length. Some species in the family have been historically
2550:
2446:
2343:
2097:
2037:
1817:
1664:
1616:
1606:
1537:
1442:
1361:
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228:
2503:
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Gibson, Robin N.; Stoner, Allan W.; Ryer, Clifford H. (2014). "The behaviour of flatfishes".
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2195:
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1994:
1956:
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1809:
1771:
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1088:'floats' to the dominant side over time, resulting in the intrinsic asymmetry of the order.
1045:
1034:
703:
212:
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295:
family is composed of two main clades, four genera, and eight species. The four genera are
1084:
717:
504:
476:
373:
104:
1250:" ) and the scophthalmids then respond by diving. Because of this, while flatfish do not
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2228:
2083:
2025:
1990:
1918:
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692:
671:
548:
463:
414:
384:
142:
94:
1158:” as well as adjacent terrestrial warming near the coastal seas (caused by industrial/
770:
by changing the angle of their bodies, with particular emphasis on the angle of their
2598:
1305:
1202:
1189:
as well. Averaging to only several meters per decade, this seemingly small change is
798:
595:
517:
185:
179:
2271:
1943:
Cheung, William W. L.; Dunne, John; Sarmiento, Jorge L.; Pauly, Daniel (July 2011).
1891:
1570:
Ryer, Clifford H. (April 2008). "A review of flatfish behavior relative to trawls".
1209:
808:(i) First, the head is driven into the substrate in the first half of the movement.
2109:
2049:
1829:
1676:
1198:
1126:
1096:
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plays a significant role in determining feeding intensity and growth. While making
779:
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591:
535:
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272:
191:
1706:
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1926:
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2395:
2298:
1362:"Interrelationships of scophthalmid fishes (Pleuronectiformes: Scopththalmidae)"
771:
644:
363:: Again, taking a step further the fish are now categorized into a subfamily of
197:
117:
2386:
1775:
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449:
is released by Bruno Chanet, recognizing two main subfamilies of scophthalmids.
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1067:
rise," where gametes are released into the water. Following this dispersal of
1022:
891:
in their feeding behaviors and they enlist a wide range of tactics. Visits to
841:
635:
251:
224:
151:
2347:
2127:"Bottom trawling releases as much carbon as air travel, landmark study finds"
1961:
1944:
1620:
1018:
1014:
972:
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590:(of which make up the fins and are responsible the name of the clade) and a
418:
255:
232:
64:
2263:
2101:
2041:
1821:
30:
1727:
Belkin, Igor M. (April 2009). "Rapid warming of Large Marine Ecosystems".
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1998:
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848:, in turn sending less involuntary cues of their presence and reduce the
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379:
350:
324:
216:
127:
84:
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1813:
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2313:
2151:
Popper, Arthur N. (2003). "Effects of Anthropogenic Sounds on Fishes".
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traits help differentiate scophthalmids from the other flatfish. These
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to best emphasize the differences between the taxon. Being part of the
328:
2092:
2067:
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263:
74:
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688:(ii) "Slightly asymmetric lateral expansions on both pelvic bones"
685:(i) "The anterior extension of the two pelvic fins to the isthmus"
1247:
1106:
1033:
813:
751:
556:
These various features can be distinguished at every level of the
482:
398:
1392:
245:, as well. Some common names found in species of this family are
822:) using cryptic camouflage. Monterey Bay Aquarium, Monterey, CA.
2361:
2464:
1071:, the two return to the seafloor having successfully mated.
271:(predominantly the brill flatfish and the turbot flatfish,
2165:
10.1577/1548-8446(2003)28[24:eoasof]2.0.co;2
1079:
Akin to courtship behaviors, scophthalmids also share a
713:
bent at contact with the dorsal margin of the cranium"
666:) have been crucial in defining the clade. Proposed in
932:"Visual feeders that take principally slow-moving and
353:, and are unique in comparison to other fish in their
1599:
Vertebrates: Comparative Anatomy, Function, Evolution
564:
family, all scophthalmids share the same fundamental
634:
are left-eyed flatfish). Furthermore, flatfish have
2370:
2287:Journal of Experimental Marine Biology and Ecology
2180:Canadian Journal of Fisheries and Aquatic Sciences
393:: After some confusion, it was concluded that the
1242:The first indication of a trawling vessel is the
905:of feeding behavior that exist within the order:
499:at the Scarborough SEA LIFE Sanctuary in England.
261:Of all the scophthalmids, the largest species (
876:is a family of flatfish very similar to other
698:(iv) "The presence of asymmetrical transverse
427:: Both traits proposed in 1993 were confirmed
447:The Interrelationships of Scophthalmid Fishes
411:The Interrelationships of Scophthalmid Fishes
239:, though this name can refer specifically to
8:
1519:
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1507:
1505:
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1083:spawning behavior just like other flatfish.
658:The three primary synapomorphies within the
462:animals and can be found in both Marine and
235:. Fish of this family are known commonly as
1487:
1485:
1483:
1481:
1479:
1477:
1475:
1473:
1471:
1469:
939:"Non-visual nocturnal feeders that consume
606:, and as such share this primary features.
409:) and need to be revisited. Chapleau, from
401:to other subfamilies of flatfish (like the
2358:
1462:FishBase (Family Scophthalmidae - Turbots)
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1454:
29:
20:
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1960:
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1355:
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1347:
1345:
1095:, circling the female "flagging with the
668:Interrelationships of Scophthalmid Fishes
1428:
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323:The first known fossil was dated to the
1294:
1258:as an avoidance behavior, individuals
952:The four elements of feeding behavior
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1938:
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1013:: After capture, all that is left is
7:
1634:
1632:
1630:
1565:
1563:
1561:
1559:
1557:
1555:
1553:
1300:
1298:
720:present in the yolks of their eggs.
662:family (mentioned previously under
2312:Lemke, JL; Ryer, CH (2006-12-20).
2237:10.1111/j.1095-8649.1999.tb00732.x
2066:Pauly, D.; Cheung, W.W.L. (2018).
793:is crucial to the survivorship of
682:anatomical traits are as follows:
14:
1235:. Flatfish find comfort with the
247:turbots, windowpanes, and brills.
1038:Similar stacking positioning in
909:The three general feeding types
546:is a family of ray-finned fish (
421:for any group is rather complex.
51:
1103:Conservation status and threats
2318:Marine Ecology Progress Series
1949:ICES Journal of Marine Science
1764:ICES Journal of Marine Science
782:(akin to a rudder of a boat).
315:yet also subtle distinctions.
254:reveals that this family is a
1:
1707:10.1016/j.fishres.2006.07.005
1584:10.1016/j.fishres.2007.10.005
925:"Visual feeders that consume
691:(iii) "An elongated anterior
1927:10.1016/j.seares.2003.07.001
1884:10.1016/j.seares.2018.06.006
1749:10.1016/j.pocean.2009.04.011
1597:Kardong, Kenneth V. (2015).
1440:Fishes of the World, 4th Ed.
1233:predator avoidance behaviors
805:complete two primary tasks:
766:. Flatfish can change their
2299:10.1016/j.jembe.2005.12.004
670:, the three aforementioned
2636:
2615:Fish of the North Atlantic
1601:(7th ed.). New York:
1534:10.1002/9781118501153.ch12
1438:Nelson, Joseph S. (2006).
1156:North Atlantic Oscillation
833:The Behaviour of Flatfish
829:The Behaviour of Flatfish
594:(a derived trait of this
176:
171:
157:
150:
48:Scientific classification
46:
37:
28:
23:
1776:10.1093/icesjms/32.3.385
1729:Progress in Oceanography
1318:. December 2012 version.
1308:; Pauly, Daniel (eds.).
778:is controlled usnig the
454:Distribution and habitat
2217:Journal of Fish Biology
1907:Journal of Sea Research
1864:Journal of Sea Research
1310:"Family Scophthalmidae"
1213:Trawl Fishing Depiction
844:and well as decreasing
2264:10.1006/anbe.1995.0219
1962:10.1093/icesjms/fsr012
1214:
1183:net primary production
1112:
1053:
948:decomposition, etc.).
941:sedentary benthic prey
934:sedentary benthic prey
823:
742:
598:group). All genera of
540:
531:Anatomy and morphology
500:
2564:Paleobiology Database
2072:Global Change Biology
1979:Nature Climate Change
1603:McGraw Hill Education
1212:
1140:freshwater prevalence
1110:
1037:
817:
736:
674:along with two other
538:
486:
2620:Marine fish families
1999:10.1038/nclimate1691
1528:. pp. 314–345.
1402:Encyclopedia of Life
1187:shifting their depth
1122:Latitudinal Gradient
868:more susceptible to
776:horizontal direction
558:scientific hierarchy
489:Scophthalmus maximus
264:Scophthalmus maximus
242:Scophthalmus maximus
221:North Atlantic Ocean
40:Scophthalmus maximus
2330:2006MEPS..328..267L
2229:1999JFBio..55..937H
2084:2018GCBio..24E..15P
2034:10.1038/nature09329
2026:2010Natur.466.1098T
2020:(7310): 1098–1101.
1991:2013NatCC...3..254C
1919:2003JSR....50..229R
1876:2018JSR...140....1C
1814:10.1038/nature09329
1806:2010Natur.466.1098T
1800:(7310): 1098–1101.
1741:2009PrOce..81..207B
1653:1996Natur.379..815R
1360:Chanet, B. (2003).
1181:, and decreases in
1175:ocean acidification
1134:to factors such as
852:sent to predators.
803:muscle contractions
774:. On the contrary,
397:family is a sister
269:fished commercially
2339:10.3354/meps328267
1695:Fisheries Research
1572:Fisheries Research
1215:
1131:Northeast Atlantic
1113:
1054:
1041:courtship behavior
927:free-swimming prey
846:oxygen consumption
824:
768:vertical direction
743:
541:
539:Turbots (Pictured)
501:
472:Fishes of the Word
256:monophyletic group
252:Cladistic analysis
2610:Pleuronectiformes
2592:
2591:
2551:Open Tree of Life
2364:Taxon identifiers
2186:(10): 2409–2422.
2093:10.1111/gcb.13831
1647:(6568): 815–818.
1612:978-0-07-802302-6
1543:978-1-118-50119-1
1167:maximum potential
1136:human populations
1064:Pleuronectiformes
1046:Green Sea Turtles
878:Pleuronectiformes
664:Taxonomic History
624:pleuronectiformes
619:Pleuronectiformes
611:Pleuronectiformes
570:pleuronectiformes
335:Taxonomic history
229:Mediterranean Sea
205:
204:
146:
128:Pleuronectiformes
2627:
2585:
2584:
2572:
2571:
2559:
2558:
2546:
2545:
2533:
2532:
2530:NBNSYS0000177870
2520:
2519:
2507:
2506:
2494:
2493:
2481:
2480:
2468:
2467:
2455:
2454:
2442:
2441:
2429:
2428:
2419:
2418:
2406:
2405:
2404:
2391:
2390:
2389:
2359:
2352:
2351:
2341:
2309:
2303:
2302:
2282:
2276:
2275:
2252:Animal Behaviour
2247:
2241:
2240:
2212:
2206:
2205:
2203:
2175:
2169:
2168:
2148:
2142:
2141:
2139:
2138:
2123:
2114:
2113:
2095:
2063:
2054:
2053:
2009:
2003:
2002:
1973:
1967:
1966:
1964:
1955:(6): 1008–1018.
1940:
1931:
1930:
1913:(2–3): 229–245.
1902:
1896:
1895:
1859:
1834:
1833:
1789:
1780:
1779:
1759:
1753:
1752:
1735:(1–4): 207–213.
1724:
1711:
1710:
1701:(2–3): 202–209.
1690:
1681:
1680:
1661:10.1038/379815a0
1636:
1625:
1624:
1594:
1588:
1587:
1578:(1–3): 138–146.
1567:
1548:
1547:
1521:
1464:
1458:
1449:
1436:
1405:
1390:
1377:
1376:
1366:
1357:
1320:
1319:
1302:
1244:sound vibrations
1179:oxygen depletion
964:
963:
959:
921:
920:
916:
856:Feeding behavior
850:chemical signals
786:Burying behavior
741:) in locomotion.
739:Pleuronectiforme
704:caudal vertebrae
636:protrusible eyes
367:, and are named
141:
56:
55:
33:
21:
16:Family of fishes
2635:
2634:
2630:
2629:
2628:
2626:
2625:
2624:
2595:
2594:
2593:
2588:
2580:
2575:
2567:
2562:
2554:
2549:
2541:
2536:
2528:
2523:
2515:
2510:
2502:
2497:
2489:
2484:
2476:
2471:
2463:
2458:
2450:
2445:
2437:
2432:
2424:
2422:
2414:
2409:
2400:
2399:
2394:
2385:
2384:
2379:
2366:
2356:
2355:
2311:
2310:
2306:
2284:
2283:
2279:
2249:
2248:
2244:
2214:
2213:
2209:
2192:10.1139/f05-131
2177:
2176:
2172:
2150:
2149:
2145:
2136:
2134:
2125:
2124:
2117:
2065:
2064:
2057:
2011:
2010:
2006:
1975:
1974:
1970:
1942:
1941:
1934:
1904:
1903:
1899:
1861:
1860:
1837:
1791:
1790:
1783:
1761:
1760:
1756:
1726:
1725:
1714:
1692:
1691:
1684:
1638:
1637:
1628:
1613:
1596:
1595:
1591:
1569:
1568:
1551:
1544:
1523:
1522:
1467:
1459:
1452:
1437:
1408:
1391:
1380:
1364:
1359:
1358:
1323:
1304:
1303:
1296:
1291:
1220:
1162:advancements).
1150:. Regions that
1118:
1105:
1077:
1059:
1032:
965:
961:
957:
955:
954:
922:
918:
914:
912:
911:
858:
788:
731:
726:
709:(v) "The first
656:
615:
581:
554:Scophthalmidae.
533:
456:
403:Paralichthyidae
374:Paralichthyidae
337:
327:era during the
321:
289:
283:respectively).
195:
189:
183:
167:
161:
140:
105:Acanthopterygii
50:
24:Scophthalmidae
17:
12:
11:
5:
2633:
2631:
2623:
2622:
2617:
2612:
2607:
2605:Scophthalmidae
2597:
2596:
2590:
2589:
2587:
2586:
2573:
2560:
2547:
2534:
2521:
2508:
2495:
2482:
2469:
2456:
2443:
2430:
2420:
2416:Scophthalmidae
2407:
2402:Scophthalmidae
2392:
2376:
2374:
2372:Scophthalmidae
2368:
2367:
2362:
2354:
2353:
2304:
2293:(2): 172–180.
2277:
2242:
2223:(5): 937–971.
2207:
2170:
2143:
2115:
2055:
2004:
1985:(3): 254–258.
1968:
1932:
1897:
1835:
1781:
1770:(3): 385–394.
1754:
1712:
1682:
1626:
1611:
1589:
1549:
1542:
1465:
1450:
1406:
1395:Scophthalmidae
1378:
1321:
1306:Froese, Rainer
1293:
1292:
1290:
1287:
1282:Scophthalmidae
1277:Scophthalmidae
1219:
1216:
1171:climate change
1152:Scophthalmidae
1148:North Atlantic
1117:
1116:Climate change
1114:
1104:
1101:
1076:
1073:
1058:
1055:
1050:Chelonia mydas
1031:
1028:
1027:
1026:
1008:
1007:
1006:
1000:
994:
982:
976:
953:
950:
945:
944:
937:
930:
910:
907:
874:Scophthalmidae
870:climate change
866:Scophthalmidae
857:
854:
795:Scophthalmidae
787:
784:
764:Actinopterygii
730:
727:
725:
722:
693:supraoccipital
672:synapomorphies
660:Scophthalmidae
655:
652:Scophthalmidae
649:
632:Scophthalmidae
614:
608:
604:Actinopterygii
600:Scophthalmidae
584:Actinopterygii
580:
577:Actinopterygii
574:
566:Actinopterygii
562:Scophthalmidae
549:Actinopterygii
544:Scophthalmidae
532:
529:
497:(Raja clavata)
492:in front of a
468:Scophthalmidae
455:
452:
451:
450:
440:
433:Scophthalmidae
429:synapomorphies
422:
415:supraoccipital
395:Scophthalmidae
388:
385:vomerine teeth
369:Scophthalmidae
358:
344:
336:
333:
320:
317:
313:synapomorphies
311:, all sharing
293:Scophthalmidae
288:
285:
209:Scophthalmidae
203:
202:
174:
173:
169:
168:
162:
155:
154:
148:
147:
138:Scophthalmidae
135:
131:
130:
125:
121:
120:
115:
108:
107:
102:
98:
97:
95:Actinopterygii
92:
88:
87:
82:
78:
77:
72:
68:
67:
62:
58:
57:
44:
43:
35:
34:
26:
25:
15:
13:
10:
9:
6:
4:
3:
2:
2632:
2621:
2618:
2616:
2613:
2611:
2608:
2606:
2603:
2602:
2600:
2583:
2578:
2574:
2570:
2565:
2561:
2557:
2552:
2548:
2544:
2539:
2535:
2531:
2526:
2522:
2518:
2513:
2509:
2505:
2500:
2496:
2492:
2487:
2483:
2479:
2474:
2470:
2466:
2461:
2457:
2453:
2448:
2444:
2440:
2435:
2431:
2427:
2421:
2417:
2412:
2408:
2403:
2397:
2393:
2388:
2382:
2378:
2377:
2375:
2373:
2369:
2365:
2360:
2349:
2345:
2340:
2335:
2331:
2327:
2323:
2319:
2315:
2308:
2305:
2300:
2296:
2292:
2288:
2281:
2278:
2273:
2269:
2265:
2261:
2257:
2253:
2246:
2243:
2238:
2234:
2230:
2226:
2222:
2218:
2211:
2208:
2202:
2197:
2193:
2189:
2185:
2181:
2174:
2171:
2166:
2162:
2159:(10): 24–31.
2158:
2154:
2147:
2144:
2132:
2128:
2122:
2120:
2116:
2111:
2107:
2103:
2099:
2094:
2089:
2085:
2081:
2077:
2073:
2069:
2062:
2060:
2056:
2051:
2047:
2043:
2039:
2035:
2031:
2027:
2023:
2019:
2015:
2008:
2005:
2000:
1996:
1992:
1988:
1984:
1980:
1972:
1969:
1963:
1958:
1954:
1950:
1946:
1939:
1937:
1933:
1928:
1924:
1920:
1916:
1912:
1908:
1901:
1898:
1893:
1889:
1885:
1881:
1877:
1873:
1869:
1865:
1858:
1856:
1854:
1852:
1850:
1848:
1846:
1844:
1842:
1840:
1836:
1831:
1827:
1823:
1819:
1815:
1811:
1807:
1803:
1799:
1795:
1788:
1786:
1782:
1777:
1773:
1769:
1765:
1758:
1755:
1750:
1746:
1742:
1738:
1734:
1730:
1723:
1721:
1719:
1717:
1713:
1708:
1704:
1700:
1696:
1689:
1687:
1683:
1678:
1674:
1670:
1666:
1662:
1658:
1654:
1650:
1646:
1642:
1635:
1633:
1631:
1627:
1622:
1618:
1614:
1608:
1604:
1600:
1593:
1590:
1585:
1581:
1577:
1573:
1566:
1564:
1562:
1560:
1558:
1556:
1554:
1550:
1545:
1539:
1535:
1531:
1527:
1520:
1518:
1516:
1514:
1512:
1510:
1508:
1506:
1504:
1502:
1500:
1498:
1496:
1494:
1492:
1490:
1488:
1486:
1484:
1482:
1480:
1478:
1476:
1474:
1472:
1470:
1466:
1463:
1457:
1455:
1451:
1448:
1447:0-471-25031-7
1444:
1441:
1435:
1433:
1431:
1429:
1427:
1425:
1423:
1421:
1419:
1417:
1415:
1413:
1411:
1407:
1404:
1403:
1398:
1396:
1389:
1387:
1385:
1383:
1379:
1375:(4): 275–286.
1374:
1370:
1363:
1356:
1354:
1352:
1350:
1348:
1346:
1344:
1342:
1340:
1338:
1336:
1334:
1332:
1330:
1328:
1326:
1322:
1317:
1316:
1311:
1307:
1301:
1299:
1295:
1288:
1286:
1283:
1278:
1274:
1268:
1266:
1261:
1257:
1253:
1249:
1245:
1240:
1238:
1234:
1229:
1225:
1224:trawl fishing
1222:Put plainly,
1218:Trawl fishing
1217:
1211:
1207:
1204:
1200:
1195:
1192:
1188:
1184:
1180:
1176:
1172:
1168:
1163:
1161:
1160:anthropogenic
1157:
1153:
1149:
1145:
1141:
1137:
1132:
1128:
1123:
1115:
1109:
1102:
1100:
1098:
1094:
1089:
1086:
1082:
1074:
1072:
1070:
1065:
1056:
1051:
1047:
1043:
1042:
1036:
1029:
1024:
1020:
1016:
1012:
1009:
1004:
1001:
998:
995:
992:
989:
988:
986:
983:
980:
977:
974:
970:
967:
966:
960:
951:
949:
942:
938:
935:
931:
928:
924:
923:
917:
908:
906:
904:
900:
899:feeding-types
896:
895:
894:species pages
890:
886:
883:
879:
875:
871:
867:
863:
855:
853:
851:
847:
843:
837:
834:
830:
821:
816:
812:
809:
806:
804:
800:
799:current shear
796:
792:
785:
783:
781:
777:
773:
769:
765:
759:
755:
753:
748:
740:
735:
728:
723:
721:
719:
714:
712:
707:
705:
701:
696:
694:
689:
686:
683:
681:
677:
673:
669:
665:
661:
653:
650:
648:
646:
642:
637:
633:
629:
625:
621:
620:
612:
609:
607:
605:
601:
597:
593:
589:
585:
578:
575:
573:
571:
567:
563:
559:
555:
551:
550:
545:
537:
530:
528:
526:
521:
519:
514:
510:
506:
498:
495:
494:thornback ray
491:
490:
485:
481:
478:
473:
469:
465:
461:
453:
448:
444:
441:
438:
434:
430:
426:
423:
420:
416:
412:
408:
404:
400:
396:
392:
389:
386:
382:
381:
376:
375:
370:
366:
362:
359:
356:
352:
348:
345:
342:
339:
338:
334:
332:
330:
326:
318:
316:
314:
310:
306:
305:Phrynorhombus
302:
301:Lepidorhombus
298:
294:
286:
284:
282:
281:
276:
275:
270:
266:
265:
259:
257:
253:
249:
248:
244:
243:
238:
234:
230:
226:
222:
219:found in the
218:
214:
210:
201:
200:
199:
194:
193:
188:
187:
186:Phrynorhombus
182:
181:
180:Lepidorhombus
175:
170:
165:
160:
156:
153:
149:
144:
139:
136:
133:
132:
129:
126:
123:
122:
119:
116:
113:
110:
109:
106:
103:
100:
99:
96:
93:
90:
89:
86:
83:
80:
79:
76:
73:
70:
69:
66:
63:
60:
59:
54:
49:
45:
42:
41:
36:
32:
27:
22:
19:
2371:
2321:
2317:
2307:
2290:
2286:
2280:
2258:(1): 41–52.
2255:
2251:
2245:
2220:
2216:
2210:
2201:11250/108691
2183:
2179:
2173:
2156:
2152:
2146:
2135:. Retrieved
2133:. 2021-03-17
2131:the Guardian
2130:
2078:(1): 15–26.
2075:
2071:
2017:
2013:
2007:
1982:
1978:
1971:
1952:
1948:
1910:
1906:
1900:
1867:
1863:
1797:
1793:
1767:
1763:
1757:
1732:
1728:
1698:
1694:
1644:
1640:
1598:
1592:
1575:
1571:
1525:
1400:
1394:
1372:
1368:
1313:
1281:
1276:
1269:
1259:
1254:or directly
1241:
1228:industrially
1221:
1196:
1190:
1166:
1164:
1151:
1147:
1144:land-locking
1143:
1139:
1135:
1127:Indo-Pacific
1119:
1097:pectoral fin
1090:
1080:
1078:
1063:
1060:
1049:
1039:
1030:Reproduction
1010:
1002:
996:
990:
984:
979:Encountering
978:
968:
946:
940:
933:
926:
902:
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820:Peces Planos
819:
818:A Flatfish (
810:
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789:
780:pectoral fin
775:
767:
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711:neural spine
708:
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628:asymmetrical
623:
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596:phylogenetic
592:swim bladder
583:
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458:Turbots are
457:
446:
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410:
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394:
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378:
372:
368:
364:
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309:Scophthalmus
308:
304:
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296:
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192:Scophthalmus
190:
184:
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159:Scophthalmus
158:
137:
111:
101:Superorder:
38:
18:
2486:iNaturalist
2396:Wikispecies
2324:: 267–273.
1273:colorations
1081:three-stage
1075:Life stages
862:temperature
842:heart rates
772:caudal fins
718:oil globule
518:diel cycle.
297:Zeugopterus
198:Zeugopterus
118:Percomorpha
2599:Categories
2137:2022-04-24
1526:Flatfishes
1289:References
1260:indirectly
1023:defecation
889:plasticity
737:Flatfish (
729:Locomotion
280:S. maximus
225:Baltic Sea
164:Rafinesque
152:Type genus
2348:0171-8630
2153:Fisheries
1621:862149184
1237:substrate
1057:Courtship
1019:digestion
1015:ingestion
1011:Ingestion
973:olfactory
969:Searching
700:apophyses
437:apophyses
425:1998-1999
419:monophyly
355:sinistral
319:Discovery
274:S.rhombus
233:Black Sea
143:Chabanaud
71:Kingdom:
65:Eukaryota
2423:BioLib:
2387:Q7924326
2381:Wikidata
2272:53204876
2102:28833977
2042:20668450
1892:90399407
1870:: 1–10.
1822:20668450
1315:FishBase
1203:demersal
1085:Spawning
997:Stalking
903:elements
747:buoyancy
724:Behavior
588:fin-rays
572:traits.
513:ebb tide
509:juvenile
477:spawning
464:Brackish
407:Bothidae
405:and the
380:Bothidae
365:Bothidae
351:Bothidae
325:Cenozoic
287:Taxonomy
217:flatfish
134:Family:
85:Chordata
81:Phylum:
75:Animalia
61:Domain:
2326:Bibcode
2225:Bibcode
2110:3364627
2080:Bibcode
2050:4424240
2022:Bibcode
1987:Bibcode
1915:Bibcode
1872:Bibcode
1830:4424240
1802:Bibcode
1737:Bibcode
1677:4304531
1669:8587602
1649:Bibcode
1399:at the
1265:bycatch
1199:pelagic
1191:drastic
1069:gametes
991:Pursuit
985:Capture
791:Burying
702:on the
641:benthic
525:buoyant
460:benthic
357:nature.
329:Miocene
237:turbots
172:Genera
124:Order:
91:Class:
2582:125580
2569:266155
2556:778687
2517:172713
2504:115125
2346:
2270:
2108:
2100:
2048:
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2014:Nature
1890:
1828:
1820:
1794:Nature
1675:
1667:
1641:Nature
1619:
1609:
1540:
1445:
1369:Cybium
1252:school
1173:(i.e.
1142:, and
1093:signal
1021:, and
1003:Ambush
956:": -->
913:": -->
885:varies
752:levers
654:traits
645:dorsal
613:traits
579:traits
505:larval
307:, and
231:, and
213:family
211:are a
166:, 1810
145:, 1933
2577:WoRMS
2543:52902
2499:IRMNG
2491:61391
2426:16143
2268:S2CID
2106:S2CID
2046:S2CID
1888:S2CID
1826:S2CID
1673:S2CID
1365:(PDF)
676:niche
399:clade
112:Clade
2538:NCBI
2512:ITIS
2478:5085
2473:GBIF
2465:5170
2439:1139
2434:BOLD
2344:ISSN
2098:PMID
2038:PMID
1818:PMID
1665:PMID
1617:OCLC
1607:ISBN
1538:ISBN
1443:ISBN
1256:herd
1201:and
1129:and
1120:The
958:edit
915:edit
901:and
882:diet
680:five
602:are
568:and
443:2003
391:1993
377:and
361:1934
347:1910
341:1900
291:The
277:and
207:The
2525:NBN
2460:EoL
2452:G5B
2447:CoL
2411:ADW
2334:doi
2322:328
2295:doi
2291:333
2260:doi
2233:doi
2196:hdl
2188:doi
2161:doi
2088:doi
2030:doi
2018:466
1995:doi
1957:doi
1923:doi
1880:doi
1868:140
1810:doi
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1772:doi
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