185:
With use of ram, predators are able to change the flow of water around the mouth and focus the flow of water into the mouth. But with too much ram, a bow wave is created in front of the predator which can push the prey away from the predator's body. The mouth aperture and RSI represent the overall tradeoff between having a large gape with lower accuracy but being able to capture larger prey vs. having a smaller gape with increased accuracy but the size of prey is limited. The three main tradeoffs within the fish skull have occurred because of the high kinesis in the skull and the elusiveness of some prey types. However, having kinesis in the skull can enable a predator to evolve new techniques on increasing the performance of prey capture.
160:
capturing elusive prey by swimming towards their prey while using suction to draw prey into the mouth. This diversity in relative use is quantified using the Ram
Suction Index (RSI) that calculates the ratio of use for ram and suction during prey capture. The RSI ratio can be influenced by the morphology of the predator and by the elusiveness of the prey. Ram feeding and suction feeding are on opposite sides of the feeding spectrum, where extreme ram feeding is when a predator swims over an immobile prey item with open jaws to engulf the prey. Extreme suction feeding is demonstrated by sit-and-wait predators that rely on rapid depression of the jaws to capture prey (e.g. frogfish,
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291:
251:
107:, lateral expansion of the suspensorium, and the depression of the lower jaw and hyoid. Suction feeding leads to successful prey capture through rapid movements creating a drop in pressure in the buccal cavity causing the water in front of the mouth to rush into the oral cavity, entrapping the prey in this flow. This mode of feeding has two main phases: expansion and compression. The expansion phase involves the initial opening of the jaws to capture prey. These movements during the expansion phase are similar across all suction feeders with the
317:
151:
qualifications center around the situation that results from a highly kinetic skull. Having a highly mobile skull introduces a tradeoff between the ability to have high speed jaw opening (high kinesis) or higher bite transmission (lower kinesis). While there is a more complex relationship between mechanical advantage and the speed of lower jaw depression, there is consensus that species using high-speed attacks have more cranial kinesis compared to species that exhibit low speed attacks. Species that have a
31:
2215:
377:
86:
2692:
273:
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Ram feeding is a method of feeding underwater in which the predator moves forward with its mouth open, engulfing the prey along with the water surrounding it. During ram feeding, the prey remains fixed in space, and the predator moves its jaws past the prey to capture it. The motion of the head may
155:
diet have also evolved skull morphologies to crush the hard-shelled prey that is a part of their diet. Durophagous species skulls consistently have more fused skulls and shorter jaw lengths. This morphology leads to the skulls being less kinetic than their piscivorous counterparts. Having shorter jaw
122:
linkages that allow greater expansion of the buccal cavity and thereby create a greater negative pressure. Most commonly, this is achieved by increasing the lateral expansion of the skull. In addition, the derived trait of anterior protrusion via the premaxillary bone in the upper jaw is acknowledged
184:
has a larger gape with lower accuracy and lower flow velocity and acceleration. However, with the larger gape the largemouth bass were able to capture larger elusive prey. Using ram feeding in combination with suction feeding can also influence the direction of water into the mouth of the predator.
159:
The third main tradeoff within suction feeding occurs with the incorporation of ram feeding with suction feeding behaviors. Ram feeding involves movement of the predator with its mouth open to engulf the prey. Most species use ram feeding combined with suction feeding to increase the chances of
167:
The mouth aperture represents another tradeoff between the ability to capture large elusive prey with more chances of failure—large gape—or to capture smaller elusive prey with greater success—smaller gape. A predator with a small mouth aperture can generate strong suction force compared to an
490:
150:
The morphologies and behaviors during suction feeding have led to three main proposed tradeoffs that determine the success of prey capture: the rate of jaw opening and closing, the mobility of the bony elements in the skull, and the ratio of ram to suction feeding behavior. The first two
479:
at first locks the head in a ventrally bent position by the alignment of two bars. The release of the locking mechanism jets the head up and moves the mouth toward the prey within 5–10 ms. The trigger mechanism of unlocking is debated, but is probably in lateral
164:). There is wide diversity on how much of each feeding strategy an individual uses, especially when body ram movements are considered. The relative use of ram and suction feeding is species dependent, but it can help determine the accuracy of prey capture.
244:). The fish all open their mouths and opercula wide at the same time (the red gills are visible in the photo below—click to enlarge). The fish swim in a grid where the distance between them is the same as the jump length of the copepods.
1253:
Collar, David C.; Reece, Joshua S.; Alfaro, Michael E.; Wainwright, Peter C.; Mehta, Rita S. (June 2014). "Imperfect
Morphological Convergence: Variable Changes in Cranial Structures Underlie Transitions to Durophagy in Moray Eels".
418:, which can protrude their mouth by several centimeters. This is usually done to extend the striking range of suction feeding, and the retraction of the jaw after protrusion can also help retrieval once the prey has been swallowed.
250:
44:
face a special difficulty as compared to feeding on land, because the density of water is about the same as that of the prey, so the prey tends to be pushed away when the mouth is closed. This problem was first identified by
141:
characteristics also exhibit suction feeding. Although suction may be created upon the mouth opening in such fishes, the criteria for pure suction feeding includes little or no bodily movement towards their prey.
137:, ram feeding is the primary method for prey capture; however, they can modulate between the two methods or use both as with many teleosts. Also, it is commonly thought that fishes with more
290:
198:
in the fluid which pushes the prey away from the jaws, but this can be avoided by allowing water to flow through the jaw. This can be accomplished by means of a swept-back mouth, as in
103:
Suction feeding is a method of ingesting a prey item in fluids by sucking the prey into the predator's mouth. It is a highly coordinated behavior achieved by the dorsal rotation of the
1305:
Durie, C.J.; Turingan, R. (2001). "Relationship between durophagy and feeding biomechanics in gray triggerfish, Balistes capriscus: intraspecific variation in ecological morphology".
1935:
703:
Lauder, George V. (March 1980). "Evolution of the feeding mechanism in primitive actionopterygian fishes: A functional anatomical analysis of
Polypterus, Lepisosteus, and Amia".
156:
lengths, with a more akinetic skull allows for an individual to have a higher bite force, compromising the ability to have a faster jaw opening when the jaw lengths are longer.
1852:
1507:"Sucking while swimming: evaluating the effects of ram speed on suction generation in bluegill sunfish Lepomis macrochirus using digital particle image velocimetry"
355:) around the volume of water to be swallowed. Subsequently, the water flows back through the baleen, keeping back the food particles. The highly elastic and
343:. Lunge feeding could be regarded as a kind of inverted suction feeding, during which a whale takes a huge gulp of water, which is then filtered through the
3396:
2593:
891:
Gidmark, Nicholas J.; Pos, Kelsie; Matheson, Bonne; Ponce, Esai; Westneat, Mark W. (2019), "Functional
Morphology and Biomechanics of Feeding in Fishes",
1133:
Lauder, George V.; Liem, Karel F. (November 1981). "Prey capture by
Luciocephalus pulcher: implications for models of jaw protrusion in teleost fishes".
3138:
568:
In suspension feeding, the water flow is primarily external and the particles themselves move with respect to the ambient water flow, such as in
1587:
Van
Wassenbergh, Sam; Brecko, Jonathan; Aerts, Peter; Stouten, Ilona; Vanheusden, Gwen; Camps, Andy; Van Damme, Raoul; Herrel, Anthony (2010).
168:
individual with a wider gape. This was demonstrated by
Wainwright et al. (2007) by comparing the feeding success of the bluegill sunfish,
1931:
1800:
Muller, M (1996). "A novel classification of planar four-bar linkages and its application to the mechanical analysis of animal systems".
441:
Pivot feeding is a method to transport the mouth towards the prey by an upward turning of the head, which is pivoting on the neck joint.
2965:
1451:
Ferry-Graham, Lara A.; Lauder, George V. (2001). "Aquatic prey capture in ray-finned fishes: A century of progress and new directions".
556:, the water flow is primarily generated by the organism itself, for example by creating a pressure gradient, by active swimming, or by
2995:
2041:
1341:
908:
631:
64:
Most underwater predators combine more than one of these basic principles. For example, a typical generalized predator, such as the
111:
of the skull leading to slight variations. During the compression phase the jaws close and water is compressed out of the gills.
1957:
Gemmell, B. J.; Sheng, J.; Buskey, E. J. (2013). "Morphology of seahorse head hydrodynamically aids in capture of evasive prey".
503:. They use pivot feeding to catch the copepod, which involves rotating their snout at high speed and then sucking in the copepod.
272:
2649:
2586:
1638:"When Giants Turn Up: Sighting Trends, Environmental Influences and Habitat Use of the Manta Ray Manta alfredi at a Coral Reef"
3700:
347:. Biomechanically this is a unique and extreme feeding method, for which the animal at first must accelerate to gain enough
131:
clade. However, a common misconception of these fishes is that suction feeding is the only or primary method employed. In
1844:
2734:
215:
1232:
2634:
1356:
Longo, Sarah J.; McGee, Matthew D.; Oufiero, Christopher E.; Waltzek, Thomas B.; Wainwright, Peter C. (2015-11-23).
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3113:
3083:
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596:
1094:"A functional morphospace for the skull of labrid fishes: patterns of diversity in a complex biomechanical system"
3659:
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2714:
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586:
138:
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1698:
Goldbogen, J. A.; Calambokidis, J.; Shadwick, R. E.; Oleson, E. M.; McDonald, M. A.; Hildebrand, J. A. (2006).
1092:
WAINWRIGHT, PETER C.; BELLWOOD, DAVID R.; WESTNEAT, MARK W.; GRUBICH, JUSTIN R.; HOEY, ANDREW S. (2004-04-22).
854:"Convergence in the Feeding Mechanics of Ecomorphologically Similar Species in the Centrarchidae and Cichlidae"
89:
46:
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3647:
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in high concentrations, the herrings switch to ram feeding. They swim with their mouth wide open and their
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has a smaller gape and was found to have higher accuracy with higher flow velocity and acceleration while
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These are contrasting methods for the removal of food particles from a water flow: for example, by the
380:
808:"Largemouth bass (micropterus salmoides) switch feeding modalities in response to sensory deprivation"
3634:
2930:
2468:
1966:
1809:
1649:
1187:
Martinez, Christopher M.; McGee, Matthew D.; Borstein, Samuel R.; Wainwright, Peter C. (2018-07-10).
1142:
464:
403:
30:
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3366:
3166:
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2842:
2555:
2409:
1749:"Passive versus active engulfment: verdict from trajectory simulations of lunge-feeding fin whales
1636:
Jaine, FRA; Couturier, LIE; Weeks, SJ; Townsend, KA; Bennett, MB; Fiora, K; Richardson, AJ (2012).
399:
3722:
3005:
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2200:
1990:
1729:
1484:
1287:
1224:
1166:
914:
736:
1045:"New insights from serranid fishes on the role of trade-offs in suction-feeding diversification"
1556:"Functional Morphology of the Mouth of the Bowhead Whale and Its Implications For Conservation"
1358:"Body ram, not suction, is the primary axis of suction-feeding diversity in spiny-rayed fishes"
127:
to be engulfed. Protrusible jaws via a mobile premaxilla can only be seen in fishes within the
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Wainwright, P.C.; Carroll, A.M.; Collar, D.C.; Day, S.W.; Higham, T.E.; Holzman, R.A. (2007).
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410:, which possess many forms of coupled linkages in their head. Remarkable examples are the
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133:
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108:
98:
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are specialized on this feeding mechanism. With prey capture times of down to 5 ms (
1970:
1813:
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in the articulations of the mouth. Vertebrate jaw protrusion is known only among modern
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that can extend rapidly, protruding forward to catch prey and bring it to the top jaw.
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58:
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1488:
754:
Holzman, Roi; Day, Steven W.; Mehta, Rita S.; Wainwright, Peter C. (10 June 2008).
581:
1662:
1589:"Hydrodynamic constraints on prey-capture performance in forward-striking snakes"
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900:
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407:
263:
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ram-feeding, swimming against the tidal current with its mouth open and sieving
225:
152:
1875:"The smaller your mouth, the longer your snout: predicting the snout length of
823:
513:
471:
is folded under the head and is aligned with the urohyal which connects to the
3614:
3381:
3315:
3305:
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2512:
2368:
2363:
2353:
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2266:
2154:
2128:
2118:
1324:
Westneat, Mark W. (2005), "Skull
Biomechanics and Suction Feeding in Fishes",
1043:
Oufiero, C. E.; Holzman, R. A.; Young, F. A.; Wainwright, P. C. (2012-08-01).
601:
569:
529:
468:
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17:
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689:
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2763:
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2517:
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2338:
2323:
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1554:
Lambertsen, R. H.; Rasmussen, K. J.; Lancaster, W. C.; Hintz, R. J. (2005).
1188:
716:
481:
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422:
124:
119:
54:
1986:
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952:
935:
831:
789:
771:
732:
680:
663:
1845:"What Do This Dragonfly's Prehensile Mouthparts Have To Do With Its Anus?"
1829:
756:"Jaw protrusion enhances forces exerted on prey by suction feeding fishes"
664:"Patterns of Evolution in the Feeding Mechanism of Actinopterygian Fishes"
641:
3574:
3515:
3461:
3456:
3275:
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2805:
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2010:
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1411:
869:
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442:
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415:
356:
348:
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211:
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50:
114:
Though suction feeding can be seen across fish species, those with more
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2639:
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332:
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128:
34:
1716:
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1555:
1523:
1506:
1464:
1204:
936:"Evolution of Levers and Linkages in the Feeding Mechanisms of Fishes"
3466:
2488:
2478:
2019:
2014:
983:
Bellwood, D.R; Wainwright, P.C; Fulton, C.J; Hoey, A.S (2005-10-12).
557:
541:
344:
325:
68:, combines suction with some amount of protrusion and pivot feeding.
460:) this method is used by the fastest feeders in the animal kingdom.
1267:
57:, have evolved a number of specialized feeding mechanisms, such as
3569:
3498:
2908:
517:
512:
315:
203:
202:
whales, or by allowing water to flow out through the gills, as in
118:
show an increase in suction potential as a result of more complex
29:
1412:"Suction feeding mechanics, performance, and diversity in fishes"
1189:"Feeding ecology underlies the evolution of cichlid jaw mobility"
3471:
3413:
3058:
2611:
2602:
359:
buccal rills are a specialized adaptation to this feeding mode.
2575:
2023:
236:
fully expanded. Every several feet, they close and clean their
61:, ram feeding, suction feeding, protrusion, and pivot feeding.
3510:
3078:
1700:"Kinematics of foraging dives and lunge-feeding in fin whales"
381:
Video of a slingjaw wrasse catching prey by protruding its jaw
65:
1328:, Fish Physiology, vol. 23, Elsevier, pp. 29–75,
1932:"Why Does the Seahorse Have Its Odd Head? Mystery Solved"
985:"Functional versatility supports coral reef biodiversity"
90:
Video of a red bay snook catching prey by suction feeding
806:
Gardiner, Jayne M.; Motta, Philip J. (28 January 2012).
210:. A number of species have evolved narrow snouts, as in
2571:
989:
Proceedings of the Royal
Society B: Biological Sciences
895:, Springer International Publishing, pp. 297–332,
402:
towards the prey, which is achieved via more mobile
37:
capture their prey by sucking them into their mouths
3602:
3535:
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3404:
3395:
3334:
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2939:
2699:
2609:
2382:
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2222:
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2142:
2066:
2057:
369:
78:
463:The secret of the speed of pivot feeding is in a
1693:
1691:
394:Jaw protrusion is the outward movement of the
2587:
2035:
852:Norton, S. F.; Brainerd, E. L. (1993-03-01).
524:concentration (slowed down by a factor of 12)
499:rely on stealth to ambush small prey such as
8:
2013:is a comprehensive database of bony fishes:
657:
655:
653:
651:
3401:
2594:
2580:
2572:
2063:
2042:
2028:
2020:
1873:de Lussanet, M. H. E.; Muller, M. (2007).
3139:Tradeoffs for locomotion in air and water
1906:
1776:
1715:
1671:
1661:
1612:
1571:
1522:
1427:
1373:
1109:
1098:Biological Journal of the Linnean Society
1060:
1016:
951:
779:
679:
421:Another example of protrusion is seen in
612:
486:
246:
1593:Journal of the Royal Society Interface
1038:
1036:
760:Journal of the Royal Society Interface
366:
75:
1843:Gonzalez, Robbie (13 November 2014).
1500:
1498:
1405:
1403:
1401:
1182:
1180:
929:
927:
123:to increase the force exerted on the
7:
2441:
847:
845:
843:
841:
801:
799:
3695:
2966:Electroreception and electrogenesis
1416:Integrative and Comparative Biology
1362:The Journal of Experimental Biology
940:Integrative and Comparative Biology
2431:
2426:
25:
1938:from the original on 26 July 2020
1930:Langley, Liz (26 November 2013).
1747:Potvin, J; Goldbogen, JA (2009).
3694:
3683:
3682:
3665:
3664:
2690:
2436:
2213:
1111:10.1111/j.1095-8312.2004.00313.x
509:Filter versus suspension feeding
489:
375:
289:
271:
249:
84:
2650:Environmental impact of fishing
1934:. National Geographic Society.
1855:from the original on 2021-07-09
1704:Journal of Experimental Biology
1511:Journal of Experimental Biology
1235:from the original on 2020-07-21
1135:Environmental Biology of Fishes
1049:Journal of Experimental Biology
858:Journal of Experimental Biology
934:Westneat, M. W. (2004-11-01).
662:Lauder, George V. (May 1982).
620:Alexander, R. McNeill (1967).
1:
1334:10.1016/s1546-5098(05)23002-9
228:. If they encounter copepods
27:Autonomous feeding of animals
2735:intramembranous ossification
1802:Phil. Trans. R. Soc. Lond. B
1663:10.1371/journal.pone.0046170
1505:Higham, T. E. (2005-07-15).
425:larvae (nymphs), which have
351:to fold its elastic throat (
324:straining water through its
901:10.1007/978-3-030-13739-7_9
623:Functional design in fishes
281:ram-feeding on a school of
172:, and the largemouth bass,
3749:
3114:Fin and flipper locomotion
3084:Sequential hermaphroditism
2971:Jamming avoidance response
2688:
1883:and other pipette feeders"
824:10.1016/j.zool.2011.09.004
597:List of feeding behaviours
387:
96:
49:. As a result, underwater
42:Aquatic feeding mechanisms
3678:
2561:Category:Eating behaviors
2536:
2211:
587:Feeding behaviour of fish
374:
83:
240:for a few milliseconds (
47:Robert McNeill Alexander
3630:Glossary of ichthyology
3192:Diel vertical migration
2541:Antipredator adaptation
1256:The American Naturalist
812:Zoology (Jena, Germany)
717:10.1002/jmor.1051630305
532:of fish, the baleen of
2996:Surface wave detection
2961:Hydrodynamic reception
2635:Diseases and parasites
1899:10.1098/rsif.2006.0201
1822:10.1098/rstb.1996.0065
1769:10.1098/rsif.2008.0492
1605:10.1098/rsif.2009.0385
1001:10.1098/rspb.2005.3276
893:Feeding in Vertebrates
772:10.1098/rsif.2008.0159
626:. London: Hutchinson.
525:
339:by a technique called
329:
38:
3134:Undulatory locomotion
2951:Ampullae of Lorenzini
1959:Nature Communications
1751:Balaenoptera physalus
1453:Journal of Morphology
705:Journal of Morphology
516:
319:
174:Micropterus salmoides
134:Micropterus salmoides
33:
3362:Genetically modified
1887:J. R. Soc. Interface
1757:J. R. Soc. Interface
1560:Journal of Mammalogy
953:10.1093/icb/44.5.378
870:10.1242/jeb.176.1.11
681:10.1093/icb/22.2.275
3167:Aquatic respiration
3054:Life history theory
2556:Carnivorous protist
2410:Intraguild predator
1971:2013NatCo...4.2840G
1881:Centriscus scutatus
1814:1996RSPTB.351..689M
1654:2012PLoSO...746170J
1147:1981EnvBF...6..257L
520:feeding under high
458:Centriscus scutatus
404:mechanical linkages
170:Lepomis macrochirus
3006:Weberian apparatus
2551:Carnivorous fungus
2201:Sexual cannibalism
2186:Animal cannibalism
2051:Feeding behaviours
1979:10.1038/ncomms3840
1429:10.1093/icb/icm032
1375:10.1242/jeb.129015
1155:10.1007/bf00005755
1062:10.1242/jeb.074849
668:American Zoologist
564:Suspension feeding
526:
330:
39:
3710:
3709:
3620:Fish common names
3531:
3530:
3162:Aquatic predation
2986:Capacity for pain
2715:Age determination
2569:
2568:
2546:Carnivorous plant
2422:Aquatic predation
2209:
2208:
2191:Human cannibalism
1808:(1340): 689–720.
1763:(40): 1005–1025.
1717:10.1242/jeb.02135
1573:10.1644/BER-123.1
1524:10.1242/jeb.01682
1517:(14): 2653–2660.
1465:10.1002/jmor.1023
1326:Fish Biomechanics
1307:Florida Scientist
1205:10.1111/evo.13518
1055:(21): 3845–3855.
995:(1582): 101–107.
766:(29): 1445–1457.
465:locking mechanism
386:
385:
95:
94:
16:(Redirected from
3740:
3698:
3697:
3686:
3685:
3668:
3667:
3402:
2694:
2625:Ethnoichthyology
2596:
2589:
2582:
2573:
2415:Pursuit predator
2217:
2196:Self-cannibalism
2064:
2044:
2037:
2030:
2021:
1999:
1998:
1954:
1948:
1947:
1945:
1943:
1927:
1921:
1920:
1910:
1870:
1864:
1863:
1861:
1860:
1840:
1834:
1833:
1797:
1791:
1790:
1780:
1744:
1738:
1737:
1719:
1710:(7): 1231–1244.
1695:
1686:
1685:
1675:
1665:
1633:
1627:
1626:
1616:
1584:
1578:
1577:
1575:
1551:
1545:
1544:
1526:
1502:
1493:
1492:
1448:
1442:
1441:
1431:
1407:
1396:
1395:
1377:
1353:
1347:
1346:
1321:
1315:
1314:
1302:
1296:
1295:
1262:(6): E168–E184.
1250:
1244:
1243:
1241:
1240:
1199:(8): 1645–1655.
1184:
1175:
1174:
1141:(3–4): 257–268.
1130:
1124:
1123:
1113:
1089:
1083:
1082:
1064:
1040:
1031:
1030:
1020:
980:
974:
973:
955:
931:
922:
921:
888:
882:
881:
849:
836:
835:
803:
794:
793:
783:
751:
745:
744:
700:
694:
693:
683:
659:
646:
645:
617:
493:
477:four-bar linkage
379:
378:
367:
293:
275:
253:
88:
87:
76:
21:
3748:
3747:
3743:
3742:
3741:
3739:
3738:
3737:
3713:
3712:
3711:
3706:
3674:
3598:
3527:
3438:
3391:
3330:
3261:
3154:
3148:
3100:
3044:Ichthyoplankton
3010:
2942:
2935:
2931:Digital Library
2926:Teleost leptins
2865:Shark cartilage
2789:pharyngeal slit
2784:pharyngeal arch
2720:Anguilliformity
2705:
2703:
2695:
2686:
2605:
2600:
2570:
2565:
2532:
2523:Surplus killing
2395:Ambush predator
2378:
2302:
2281:
2218:
2205:
2174:
2138:
2053:
2048:
2007:
2002:
1956:
1955:
1951:
1941:
1939:
1929:
1928:
1924:
1893:(14): 561–573.
1877:Syngnathus acus
1872:
1871:
1867:
1858:
1856:
1842:
1841:
1837:
1799:
1798:
1794:
1746:
1745:
1741:
1697:
1696:
1689:
1635:
1634:
1630:
1599:(46): 773–785.
1586:
1585:
1581:
1553:
1552:
1548:
1504:
1503:
1496:
1450:
1449:
1445:
1409:
1408:
1399:
1355:
1354:
1350:
1344:
1323:
1322:
1318:
1304:
1303:
1299:
1252:
1251:
1247:
1238:
1236:
1186:
1185:
1178:
1132:
1131:
1127:
1091:
1090:
1086:
1042:
1041:
1034:
982:
981:
977:
933:
932:
925:
911:
890:
889:
885:
851:
850:
839:
805:
804:
797:
753:
752:
748:
702:
701:
697:
661:
660:
649:
634:
619:
618:
614:
610:
578:
566:
550:
511:
504:
494:
473:shoulder girdle
467:, in which the
439:
412:slingjaw wrasse
392:
376:
370:External videos
365:
314:
307:
303:ram feeding on
301:Indian mackerel
294:
285:
276:
267:
254:
191:
148:
116:cranial kinesis
101:
99:Cranial kinesis
85:
79:External videos
74:
72:Suction feeding
28:
23:
22:
15:
12:
11:
5:
3746:
3744:
3736:
3735:
3730:
3728:Marine biology
3725:
3715:
3714:
3708:
3707:
3705:
3704:
3692:
3679:
3676:
3675:
3673:
3672:
3662:
3657:
3656:
3655:
3650:
3642:
3637:
3632:
3627:
3622:
3617:
3612:
3606:
3604:
3600:
3599:
3597:
3596:
3595:
3594:
3589:
3579:
3578:
3577:
3572:
3567:
3557:
3556:
3555:
3550:
3539:
3537:
3533:
3532:
3529:
3528:
3526:
3525:
3524:
3523:
3518:
3513:
3503:
3502:
3501:
3496:
3491:
3486:
3476:
3475:
3474:
3469:
3464:
3459:
3448:
3446:
3444:Wild fisheries
3440:
3439:
3437:
3436:
3431:
3426:
3421:
3416:
3410:
3408:
3399:
3393:
3392:
3390:
3389:
3384:
3379:
3374:
3369:
3367:Hallucinogenic
3364:
3359:
3354:
3349:
3344:
3338:
3336:
3332:
3331:
3329:
3328:
3323:
3318:
3313:
3308:
3303:
3298:
3293:
3288:
3283:
3278:
3272:
3270:
3263:
3262:
3260:
3259:
3254:
3249:
3244:
3242:Schooling fish
3239:
3234:
3229:
3224:
3219:
3214:
3209:
3204:
3202:Filter feeders
3199:
3194:
3189:
3184:
3179:
3177:Bottom feeders
3174:
3169:
3164:
3158:
3156:
3150:
3149:
3147:
3146:
3141:
3136:
3131:
3126:
3121:
3116:
3110:
3108:
3102:
3101:
3099:
3098:
3097:
3096:
3086:
3081:
3076:
3071:
3066:
3061:
3056:
3051:
3046:
3041:
3036:
3031:
3026:
3020:
3018:
3012:
3011:
3009:
3008:
3003:
2998:
2993:
2988:
2983:
2978:
2973:
2968:
2963:
2958:
2953:
2947:
2945:
2937:
2936:
2934:
2933:
2928:
2923:
2922:
2921:
2916:
2906:
2905:
2904:
2899:
2889:
2884:
2879:
2878:
2877:
2867:
2862:
2857:
2852:
2847:
2846:
2845:
2835:
2830:
2825:
2823:Leydig's organ
2820:
2819:
2818:
2816:pharyngeal jaw
2813:
2803:
2798:
2797:
2796:
2791:
2786:
2781:
2776:
2771:
2769:branchial arch
2761:
2760:
2759:
2749:
2744:
2739:
2738:
2737:
2732:
2722:
2717:
2711:
2709:
2697:
2696:
2689:
2687:
2685:
2684:
2679:
2674:
2669:
2664:
2659:
2658:
2657:
2652:
2647:
2637:
2632:
2627:
2622:
2616:
2614:
2607:
2606:
2601:
2599:
2598:
2591:
2584:
2576:
2567:
2566:
2564:
2563:
2558:
2553:
2548:
2543:
2537:
2534:
2533:
2531:
2530:
2525:
2520:
2515:
2510:
2505:
2500:
2498:Hypercarnivore
2495:
2494:
2493:
2492:
2491:
2481:
2474:Cattle feeding
2471:
2466:
2465:
2464:
2459:
2457:Feeding frenzy
2454:
2449:
2444:
2442:Suction feeder
2439:
2434:
2429:
2419:
2418:
2417:
2412:
2407:
2402:
2397:
2386:
2384:
2380:
2379:
2377:
2376:
2371:
2366:
2361:
2356:
2351:
2346:
2341:
2336:
2331:
2326:
2321:
2316:
2310:
2308:
2304:
2303:
2301:
2300:
2295:
2289:
2287:
2283:
2282:
2280:
2279:
2274:
2269:
2264:
2259:
2254:
2252:Seed predation
2249:
2244:
2239:
2234:
2228:
2226:
2220:
2219:
2212:
2210:
2207:
2206:
2204:
2203:
2198:
2193:
2188:
2182:
2180:
2176:
2175:
2173:
2172:
2167:
2162:
2157:
2152:
2146:
2144:
2140:
2139:
2137:
2136:
2131:
2126:
2121:
2116:
2111:
2106:
2101:
2096:
2091:
2086:
2081:
2076:
2070:
2068:
2061:
2055:
2054:
2049:
2047:
2046:
2039:
2032:
2024:
2018:
2017:
2006:
2005:External links
2003:
2001:
2000:
1949:
1922:
1865:
1835:
1792:
1739:
1687:
1648:(10): e46170.
1628:
1579:
1566:(2): 342–352.
1546:
1494:
1443:
1397:
1368:(1): 119–128.
1348:
1342:
1316:
1297:
1268:10.1086/675810
1245:
1176:
1125:
1084:
1032:
975:
946:(5): 378–389.
923:
909:
883:
837:
795:
746:
711:(3): 283–317.
695:
674:(2): 275–285.
647:
632:
611:
609:
606:
605:
604:
599:
594:
589:
584:
577:
574:
565:
562:
554:filter feeding
549:
548:Filter feeding
546:
510:
507:
506:
505:
495:
488:
438:
435:
390:Jaw protrusion
388:Main article:
384:
383:
372:
371:
364:
363:Jaw protrusion
361:
328:after lunging.
322:humpback whale
313:
310:
309:
308:
295:
288:
286:
277:
270:
268:
266:from the water
259:Mobula alfredi
255:
248:
242:filter feeding
190:
187:
178:L. macrochirus
147:
144:
105:dermatocranium
93:
92:
81:
80:
73:
70:
59:filter feeding
26:
24:
18:Suction feeder
14:
13:
10:
9:
6:
4:
3:
2:
3745:
3734:
3731:
3729:
3726:
3724:
3721:
3720:
3718:
3703:
3702:
3693:
3691:
3690:
3681:
3680:
3677:
3671:
3670:more lists...
3663:
3661:
3658:
3654:
3651:
3649:
3646:
3645:
3643:
3641:
3638:
3636:
3633:
3631:
3628:
3626:
3625:Fish families
3623:
3621:
3618:
3616:
3613:
3611:
3610:Aquarium life
3608:
3607:
3605:
3601:
3593:
3592:fleshy-finned
3590:
3588:
3585:
3584:
3583:
3580:
3576:
3573:
3571:
3568:
3566:
3563:
3562:
3561:
3560:Cartilaginous
3558:
3554:
3551:
3549:
3546:
3545:
3544:
3541:
3540:
3538:
3534:
3522:
3519:
3517:
3514:
3512:
3509:
3508:
3507:
3504:
3500:
3497:
3495:
3492:
3490:
3487:
3485:
3482:
3481:
3480:
3477:
3473:
3470:
3468:
3465:
3463:
3460:
3458:
3455:
3454:
3453:
3450:
3449:
3447:
3445:
3441:
3435:
3432:
3430:
3427:
3425:
3422:
3420:
3417:
3415:
3412:
3411:
3409:
3407:
3403:
3400:
3398:
3394:
3388:
3385:
3383:
3380:
3378:
3375:
3373:
3370:
3368:
3365:
3363:
3360:
3358:
3355:
3353:
3350:
3348:
3345:
3343:
3340:
3339:
3337:
3333:
3327:
3324:
3322:
3319:
3317:
3314:
3312:
3309:
3307:
3304:
3302:
3299:
3297:
3294:
3292:
3289:
3287:
3284:
3282:
3279:
3277:
3274:
3273:
3271:
3269:
3264:
3258:
3255:
3253:
3250:
3248:
3245:
3243:
3240:
3238:
3235:
3233:
3230:
3228:
3225:
3223:
3220:
3218:
3215:
3213:
3210:
3208:
3205:
3203:
3200:
3198:
3197:Electric fish
3195:
3193:
3190:
3188:
3185:
3183:
3180:
3178:
3175:
3173:
3170:
3168:
3165:
3163:
3160:
3159:
3157:
3151:
3145:
3142:
3140:
3137:
3135:
3132:
3130:
3127:
3125:
3122:
3120:
3117:
3115:
3112:
3111:
3109:
3107:
3103:
3095:
3092:
3091:
3090:
3087:
3085:
3082:
3080:
3077:
3075:
3072:
3070:
3067:
3065:
3062:
3060:
3057:
3055:
3052:
3050:
3047:
3045:
3042:
3040:
3037:
3035:
3032:
3030:
3027:
3025:
3022:
3021:
3019:
3017:
3013:
3007:
3004:
3002:
2999:
2997:
2994:
2992:
2989:
2987:
2984:
2982:
2979:
2977:
2974:
2972:
2969:
2967:
2964:
2962:
2959:
2957:
2954:
2952:
2949:
2948:
2946:
2944:
2938:
2932:
2929:
2927:
2924:
2920:
2917:
2915:
2912:
2911:
2910:
2907:
2903:
2900:
2898:
2895:
2894:
2893:
2890:
2888:
2885:
2883:
2880:
2876:
2873:
2872:
2871:
2868:
2866:
2863:
2861:
2858:
2856:
2853:
2851:
2848:
2844:
2841:
2840:
2839:
2836:
2834:
2831:
2829:
2828:Mauthner cell
2826:
2824:
2821:
2817:
2814:
2812:
2809:
2808:
2807:
2804:
2802:
2799:
2795:
2792:
2790:
2787:
2785:
2782:
2780:
2777:
2775:
2772:
2770:
2767:
2766:
2765:
2762:
2758:
2755:
2754:
2753:
2750:
2748:
2747:Chromatophore
2745:
2743:
2740:
2736:
2733:
2731:
2728:
2727:
2726:
2723:
2721:
2718:
2716:
2713:
2712:
2710:
2708:
2702:
2698:
2693:
2683:
2680:
2678:
2675:
2673:
2670:
2668:
2665:
2663:
2660:
2656:
2653:
2651:
2648:
2646:
2643:
2642:
2641:
2638:
2636:
2633:
2631:
2628:
2626:
2623:
2621:
2618:
2617:
2615:
2613:
2608:
2604:
2597:
2592:
2590:
2585:
2583:
2578:
2577:
2574:
2562:
2559:
2557:
2554:
2552:
2549:
2547:
2544:
2542:
2539:
2538:
2535:
2529:
2526:
2524:
2521:
2519:
2516:
2514:
2511:
2509:
2508:Mesocarnivore
2506:
2504:
2503:Hypocarnivore
2501:
2499:
2496:
2490:
2487:
2486:
2485:
2482:
2480:
2477:
2476:
2475:
2472:
2470:
2467:
2463:
2462:Filter feeder
2460:
2458:
2455:
2453:
2452:Bottom feeder
2450:
2448:
2445:
2443:
2440:
2438:
2435:
2433:
2430:
2428:
2425:
2424:
2423:
2420:
2416:
2413:
2411:
2408:
2406:
2403:
2401:
2400:Apex predator
2398:
2396:
2393:
2392:
2391:
2388:
2387:
2385:
2381:
2375:
2372:
2370:
2367:
2365:
2362:
2360:
2357:
2355:
2352:
2350:
2347:
2345:
2342:
2340:
2337:
2335:
2332:
2330:
2327:
2325:
2322:
2320:
2317:
2315:
2312:
2311:
2309:
2305:
2299:
2296:
2294:
2291:
2290:
2288:
2284:
2278:
2275:
2273:
2270:
2268:
2265:
2263:
2260:
2258:
2255:
2253:
2250:
2248:
2245:
2243:
2240:
2238:
2235:
2233:
2230:
2229:
2227:
2225:
2221:
2216:
2202:
2199:
2197:
2194:
2192:
2189:
2187:
2184:
2183:
2181:
2179:cannibalistic
2177:
2171:
2168:
2166:
2165:Breastfeeding
2163:
2161:
2160:Placentophagy
2158:
2156:
2153:
2151:
2148:
2147:
2145:
2141:
2135:
2132:
2130:
2127:
2125:
2122:
2120:
2117:
2115:
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2110:
2107:
2105:
2102:
2100:
2097:
2095:
2092:
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2085:
2082:
2080:
2079:Egg predation
2077:
2075:
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2071:
2069:
2065:
2062:
2060:
2056:
2052:
2045:
2040:
2038:
2033:
2031:
2026:
2025:
2022:
2016:
2015:/fishbase.org
2012:
2009:
2008:
2004:
1996:
1992:
1988:
1984:
1980:
1976:
1972:
1968:
1964:
1960:
1953:
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1937:
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1900:
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1501:
1499:
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1486:
1482:
1478:
1474:
1470:
1466:
1462:
1459:(2): 99–119.
1458:
1454:
1447:
1444:
1439:
1435:
1430:
1425:
1422:(1): 96–106.
1421:
1417:
1413:
1406:
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1398:
1393:
1389:
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1376:
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1343:9780123504470
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1183:
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986:
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954:
949:
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928:
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916:
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910:9783030137380
906:
902:
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894:
887:
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879:
875:
871:
867:
863:
859:
855:
848:
846:
844:
842:
838:
833:
829:
825:
821:
817:
813:
809:
802:
800:
796:
791:
787:
782:
777:
773:
769:
765:
761:
757:
750:
747:
742:
738:
734:
730:
726:
722:
718:
714:
710:
706:
699:
696:
691:
687:
682:
677:
673:
669:
665:
658:
656:
654:
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648:
643:
639:
635:
633:0-09-084770-9
629:
625:
624:
616:
613:
607:
603:
600:
598:
595:
593:
590:
588:
585:
583:
580:
579:
575:
573:
571:
563:
561:
559:
555:
547:
545:
543:
539:
535:
531:
523:
522:phytoplankton
519:
515:
508:
502:
498:
492:
487:
485:
483:
478:
474:
470:
466:
461:
459:
456:
452:
448:
444:
437:Pivot feeding
436:
434:
432:
428:
424:
419:
417:
413:
409:
405:
401:
397:
391:
382:
373:
368:
362:
360:
358:
354:
353:buccal cavity
350:
346:
342:
341:lunge feeding
338:
334:
327:
323:
318:
312:Lunge feeding
311:
306:
305:macroplankton
302:
298:
292:
287:
284:
280:
274:
269:
265:
261:
260:
252:
247:
245:
243:
239:
235:
231:
227:
226:hunt copepods
223:
219:
217:
213:
209:
205:
201:
197:
188:
186:
183:
179:
175:
171:
165:
163:
162:Antennariidae
157:
154:
145:
143:
140:
136:
135:
130:
126:
121:
117:
112:
110:
106:
100:
91:
82:
77:
71:
69:
67:
62:
60:
56:
53:, especially
52:
48:
43:
36:
32:
19:
3733:Biomechanics
3699:
3687:
3587:spiny-finned
3536:Major groups
3257:Intelligence
3237:Scale eaters
3182:Cleaner fish
3064:Mouthbrooder
3016:Reproduction
2991:Schreckstoff
2976:Lateral line
2892:Swim bladder
2882:Spiral valve
2811:hyomandibula
2794:pseudobranch
2677:Hypoxia in -
2528:Trophallaxis
2432:Pivot feeder
2427:Lunge feeder
2421:
2405:Egg predator
2293:Phagocytosis
2143:reproductive
2114:Myrmecophagy
2104:Molluscivore
1962:
1958:
1952:
1940:. Retrieved
1925:
1890:
1886:
1880:
1876:
1868:
1857:. Retrieved
1848:
1838:
1805:
1801:
1795:
1760:
1756:
1750:
1742:
1707:
1703:
1645:
1641:
1631:
1596:
1592:
1582:
1563:
1559:
1549:
1514:
1510:
1456:
1452:
1446:
1419:
1415:
1365:
1361:
1351:
1325:
1319:
1310:
1306:
1300:
1259:
1255:
1248:
1237:. Retrieved
1196:
1192:
1138:
1134:
1128:
1101:
1097:
1087:
1052:
1048:
992:
988:
978:
943:
939:
892:
886:
864:(1): 11–29.
861:
857:
818:(2): 78–83.
815:
811:
763:
759:
749:
708:
704:
698:
671:
667:
622:
615:
582:Cleaner fish
567:
551:
527:
462:
457:
440:
420:
393:
331:
257:
220:
216:water snakes
192:
182:M. salmoides
181:
177:
173:
169:
166:
161:
158:
149:
132:
113:
102:
63:
41:
40:
3701:WikiProject
3660:Prehistoric
3644:Threatened
3335:Other types
3232:Sardine run
3207:Forage fish
3187:Corallivory
3039:Development
3024:Bubble nest
2897:physoclisti
2887:Suckermouth
2860:Root effect
2682:Ichthyology
2349:Planktivore
2334:Detritivore
2329:Coprophagia
2319:Bacterivore
2314:Microbivory
2298:Myzocytosis
2257:Nectarivore
2247:Graminivore
2094:Lepidophagy
2089:Insectivore
2084:Hematophagy
1104:(1): 1–25.
592:Lepidophagy
560:movements.
530:gill rakers
451:sea dragons
408:bony fishes
264:zooplankton
238:gill rakers
189:Ram feeding
153:durophagous
3717:Categories
3316:Groundfish
3311:Freshwater
3306:Euryhaline
3291:Coral reef
3227:Salmon run
3217:Paedophagy
3119:Amphibious
3106:Locomotion
2914:pharyngeal
2902:physostome
2855:Photophore
2801:Glossohyal
2774:gill raker
2757:dorsal fin
2707:physiology
2513:Parasitism
2447:Bait balls
2437:Ram feeder
2369:Plastivore
2364:Lithotroph
2354:Saprophagy
2277:Osteophagy
2267:Palynivore
2224:Herbivores
2155:Paedophagy
2129:Spongivore
2119:Ophiophagy
2059:Carnivores
1859:2021-07-03
1239:2020-09-06
608:References
602:Paedophagy
570:sea lilies
469:hyoid arch
455:shrimpfish
447:sea horses
400:mouthparts
396:premaxilla
97:See also:
3723:Predation
3565:chimaeras
3452:Predatory
3429:Salmonids
3387:Whitefish
3377:Poisonous
3352:Diversity
3286:Coldwater
3222:Predatory
3212:Migratory
3172:Bait ball
3155:behaviour
3074:Pregnancy
3069:Polyandry
2843:papillare
2838:Operculum
2833:Meristics
2779:gill slit
2742:Cleithrum
2672:Fish kill
2662:Fear of -
2655:- as food
2645:Fisheries
2630:Evolution
2620:Diversity
2518:Scavenger
2390:Predation
2359:Xenophagy
2339:Geophagia
2324:Fungivore
2272:Xylophagy
2262:Mellivory
2242:Frugivore
2237:Florivore
2134:Vermivore
2124:Piscivore
2109:Mucophagy
2099:Man-eater
1995:205321320
1533:0022-0949
1473:0362-2525
1384:0022-0949
1276:0003-0147
1213:0014-3820
1193:Evolution
1163:0378-1909
1120:0024-4066
1071:0022-0949
1009:0962-8452
962:1540-7063
919:150135750
878:0022-0949
725:0362-2525
690:0003-1569
536:, or the
497:Seahorses
482:adduction
431:mandibles
427:hydraulic
423:dragonfly
299:of adult
256:Foraging
230:schooling
194:induce a
146:Tradeoffs
139:primitive
129:teleostei
55:bony fish
51:predators
3689:Category
3640:Smallest
3553:lampreys
3516:flatfish
3506:Demersal
3462:mackerel
3457:billfish
3397:Commerce
3326:Tropical
3301:Demersal
3296:Deep-sea
3252:Venomous
3144:RoboTuna
3094:triggers
3089:Spawning
3049:Juvenile
3034:Egg case
2667:FishBase
2469:Browsing
2344:Omnivore
2286:Cellular
2232:Folivore
2011:FishBase
1987:24281430
1965:: 2840.
1936:Archived
1917:17251161
1853:Archived
1787:19158011
1734:17923052
1726:16547295
1682:23056255
1642:PLOS ONE
1623:19828500
1541:16000535
1481:11304743
1438:21672823
1392:26596534
1313:: 20–28.
1292:17433961
1284:24823828
1233:Archived
1229:49311313
1221:29920668
1171:24315046
1079:22855615
1027:16519241
970:21676723
832:22285791
790:18544504
741:26805223
733:30170473
576:See also
501:copepods
445:such as
443:Pipefish
416:sand eel
414:and the
357:muscular
349:momentum
337:plankton
335:feed on
333:Rorquals
283:copepods
234:opercula
222:Herrings
212:gar fish
200:balaenid
196:bow wave
3635:Largest
3548:hagfish
3543:Jawless
3521:pollock
3494:sardine
3489:herring
3484:anchovy
3434:Tilapia
3424:Octopus
3419:Catfish
3406:Farming
3321:Pelagic
3281:Coastal
3268:habitat
3124:Walking
3029:Clasper
2981:Otolith
2943:systems
2941:Sensory
2875:ganoine
2850:Papilla
2701:Anatomy
2640:Fishing
2484:Grazing
2383:Methods
2170:Weaning
2150:Oophagy
2074:Avivore
1967:Bibcode
1908:2373409
1849:Gizmodo
1830:8927640
1810:Bibcode
1778:2827442
1673:3463571
1650:Bibcode
1614:2874232
1489:4996900
1143:Bibcode
1018:1560014
781:2607355
558:ciliary
542:sponges
279:Herring
208:herring
109:kinesis
35:Grouper
3653:sharks
3570:sharks
3499:sprats
3479:Forage
3467:salmon
3347:Coarse
3129:Flying
3001:Vision
2956:Barbel
2870:Scales
2730:dermal
2610:About
2489:Forage
2479:Fodder
2307:Others
1993:
1985:
1942:25 May
1915:
1905:
1828:
1785:
1775:
1732:
1724:
1680:
1670:
1621:
1611:
1539:
1531:
1487:
1479:
1471:
1436:
1390:
1382:
1340:
1290:
1282:
1274:
1227:
1219:
1211:
1169:
1161:
1118:
1077:
1069:
1025:
1015:
1007:
968:
960:
917:
907:
876:
830:
788:
778:
739:
731:
723:
688:
642:456355
640:
630:
534:whales
429:lower
345:baleen
326:baleen
297:School
224:often
204:sharks
3615:Blind
3603:Lists
3382:Rough
3247:Sleep
3153:Other
2919:shark
2909:Teeth
2067:adult
1991:S2CID
1730:S2CID
1485:S2CID
1288:S2CID
1225:S2CID
1167:S2CID
915:S2CID
737:S2CID
538:ostia
518:Krill
120:skull
3648:rays
3582:Bony
3575:rays
3472:tuna
3414:Carp
3372:Oily
3357:Game
3342:Bait
3276:Cave
3059:Milt
2764:Gill
2752:Fins
2725:Bone
2612:fish
2603:Fish
2374:Pica
1983:PMID
1944:2020
1913:PMID
1826:PMID
1783:PMID
1722:PMID
1678:PMID
1619:PMID
1537:PMID
1529:ISSN
1477:PMID
1469:ISSN
1434:PMID
1388:PMID
1380:ISSN
1338:ISBN
1280:PMID
1272:ISSN
1217:PMID
1209:ISSN
1159:ISSN
1116:ISSN
1075:PMID
1067:ISSN
1023:PMID
1005:ISSN
966:PMID
958:ISSN
905:ISBN
874:ISSN
828:PMID
786:PMID
729:PMID
721:ISSN
686:ISSN
638:OCLC
628:ISBN
475:. A
449:and
214:and
206:and
125:prey
3511:cod
3266:By
3079:Roe
2806:Jaw
2704:and
1975:doi
1903:PMC
1895:doi
1818:doi
1806:351
1773:PMC
1765:doi
1712:doi
1708:209
1668:PMC
1658:doi
1609:PMC
1601:doi
1568:doi
1519:doi
1515:208
1461:doi
1457:248
1424:doi
1370:doi
1366:219
1330:doi
1264:doi
1260:183
1201:doi
1151:doi
1106:doi
1057:doi
1053:215
1013:PMC
997:doi
993:273
948:doi
897:doi
866:doi
862:176
820:doi
816:115
776:PMC
768:doi
713:doi
709:163
676:doi
552:In
540:of
398:or
66:cod
3719::
1989:.
1981:.
1973:.
1961:.
1911:.
1901:.
1889:.
1885:.
1879:,
1851:.
1847:.
1824:.
1816:.
1804:.
1781:.
1771:.
1759:.
1755:.
1728:.
1720:.
1706:.
1702:.
1690:^
1676:.
1666:.
1656:.
1644:.
1640:.
1617:.
1607:.
1595:.
1591:.
1564:86
1562:.
1558:.
1535:.
1527:.
1513:.
1509:.
1497:^
1483:.
1475:.
1467:.
1455:.
1432:.
1420:47
1418:.
1414:.
1400:^
1386:.
1378:.
1364:.
1360:.
1336:,
1311:64
1309:.
1286:.
1278:.
1270:.
1258:.
1231:.
1223:.
1215:.
1207:.
1197:72
1195:.
1191:.
1179:^
1165:.
1157:.
1149:.
1137:.
1114:.
1102:82
1100:.
1096:.
1073:.
1065:.
1051:.
1047:.
1035:^
1021:.
1011:.
1003:.
991:.
987:.
964:.
956:.
944:44
942:.
938:.
926:^
913:,
903:,
872:.
860:.
856:.
840:^
826:.
814:.
810:.
798:^
784:.
774:.
762:.
758:.
735:.
727:.
719:.
707:.
684:.
672:22
670:.
666:.
650:^
636:.
572:.
544:.
484:.
320:A
218:.
176:.
2595:e
2588:t
2581:v
2043:e
2036:t
2029:v
1997:.
1977::
1969::
1963:4
1946:.
1919:.
1897::
1891:4
1862:.
1832:.
1820::
1812::
1789:.
1767::
1761:6
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1736:.
1714::
1684:.
1660::
1652::
1646:7
1625:.
1603::
1597:7
1576:.
1570::
1543:.
1521::
1491:.
1463::
1440:.
1426::
1394:.
1372::
1332::
1294:.
1266::
1242:.
1203::
1173:.
1153::
1145::
1139:6
1122:.
1108::
1081:.
1059::
1029:.
999::
972:.
950::
899::
880:.
868::
834:.
822::
792:.
770::
764:5
743:.
715::
692:.
678::
644:.
20:)
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