THE EVOLUTIONARY STEPS OF FISH
Rebecca Kagle
Fish are incredible diverse group, made up of three living classes that hold an important place both in modern ecology and in evolutionary history. Fish are incredibly successful, they arenumerous and have a huge variety of adaptations. They are successful in every water filledhabitat. Furthermore, fish are the first known vertebrates and also the stepping stone to all land-walking vertebrates (tetrapods). Fish are complicated, diverse, and have a long evolutionaryhistory.In order to understand the place of fish as both a transitory group and as a modern one itis important to understand the times in which and from which they evolved. Fish are the firstknown true chordates. The first vertebrate that has been found is the Upper Cambrian fossilAnaspis, which is more than 500 million years old (3). This fossil, while being fragmentary, isthought to be an armored, jawless fish (3). Fish did most of their evolving between five millionand three and a half million years ago. These two periods were known as the Silurian and theDevonian periods. In yhe middle Silurian, the jawless fishes had diversified, but it was not untilthe Devonian that the true variety of fishes really flourished. In fact, the Devonian, is oftenreferred to as the "Age of Fishes". Towards the end of the Devonian the first tetrapods(vertebrates which evolved true legs with which they could walk on land) had evolved from onespecific branch of fish. Fish greatly specialized in their aquatic niche during both the Devonianand the Silurian and part of this evolution led to adaptations to land in the form of amphibians(9).Fish are in no way simply a stepping stone to amphibians, they are a much moresignificant than this. They have evolved to be masters of their domain, the water. They come inmany forms, have the ability to eat a huge variety of foods, and have populated almost every body of water. In fact, fish are the most common vertebrate, with there being approximately24,000 species alive today. This number is mind boggling when put in perspective; the next mostcommon vertebrate are birds with a mere 8,600 species (8). This multitude of species ranges in
size, morphology, agility, and adaptations to environment (10). Fish have been broken down intoa series of classes that separate them based on characteristics. These groups help to classify thewide variety of species that make up fish and help lead to understand of the current andevolutionary niche of fish.The first fish to evolve were the Agnathans (Class agnatha) (7). These jawless fishes arethe first vertebrates. These fish have round mouth parts that could be used for sucking or filter feeding (10). These rasping, sucking mouths are currently found on modern lampreys andhagfishes. These fish were often extremely armored in order to help them protect themselves.One group that evolved before the Silurian were the Ostracoderms which have been described as"small, blunt-headed forms...they fed on debris in the mud, bullet shaped swimmers, andsome...with an astonishing array of spines and crests on their heads." (1). Most of these types of fish are currently extinct with the exceptions of the lampreys and the hagfish.From these bottom feeding, jawless fish came the evolution of jawed fish. Jaws evolvedonly once (rather than evolving multiple times in different species through parallel evolution).Jaws evolved from gill arches which are the bony parts between gill slits. It is thought that a gillarch in an agnathan became fused to its skull (11). The upper part of the gill support became thetop jaw and the bottom part of the gill support became the bottom jaw. Embryology points to thisand the arrangement of nerves in shark heads and most simple fishes shows that jaws are in linewith gill arches. While fish had the first bony jaws, they also have some of the most complicated.While the human head has only one moving part (the jaw), the head of a fish may have morethan twenty-four bones that may move together in feeding (8). The evolution of the jaw isincredibly important because it led to fish to be able to ingest a much wider variety of foods andallowed them to be active hunters as opposed to passive filter feeder (1). This led to a widevariety of adaptations in morphology. Fish became more agile to be better predators, they wereable to reduce their armor because they were less vulnerable, and their muscle density was ableto decrease becuase they no longer led such a sluggish lifestyle (10).The first jawless fishes include the acnathodians and placoderms. Even early in their development the jawless fishes showed great diversity. The acnathodians appeared around four and half million years ago. They were the first jawed fishes to evolve and they tended to be
small, streamlined, and had huge eyes. Placoderms on the other hand appeared in the earlySilurian and dominated the Devonian. These were some strange looking fish! As described byPeter Gore, "the Placoderms had bony armor that covered the head and forepart of the body. Inmany, a movable joint between the head and body armor let the head rock back to open themouth wide. The primitive jaws had jagged bony edges that served as teeth. The tail end usuallylacked protection"(11). At the beginning of the Devonian, Placoderms were small, but they soonincreased in size. The Dunkleosteus, a placoderm, grew to be as large as 35 feet, had welldeveloped jaws, with fang like teeth. The front of the trunk was heavily armored and the hind part was either bare or covered with small scales (11). The Dunkleosteus and all the other large placoderms are extinct, but in the Devonian they dominated both salt and freshwater.While the huge, menacing placoderms have long since vanished from the earth, membersof the Class Chondritchthyes, some of which are equally menacing, have not. This class includesthe sharks, skates, and rays (along with some other fish) and these first evolved between 400 and450 million years ago (1, 4). They probably shared a common ancestor with the placoderms (1).This class is commonly refer to as the cartilaginous fish because they lack true bone, instead theyhave cartilage and calcified cartilage for internal support. This type of skeleton is extremely lightand flexible and helps these species be agile predators. Chondrithyes have evolved two kinds of jaws; sharks have extremely powerful biting and crushing jaws while skates and ray uses their jaws to be bottom feeders searching for mollusks (4). Sharks are often referred to as living fossilssince they have not done much evolving since the Late Paleozoic. This is not to say that thesharks that exist now are they only sharks ever to exist. After the decline of the placoderms, therewas a shark explosion that led to sharks with mouth's filled with coils of serrated teeth, or bony protuberances on their fins. These Paleozoic sharks could be a large as 14m, more than doublethe size of the current Great White Sharks.In terms of currently living fish, Class Chondrichtyes make up only 900 species. The bulk of the fish are from Class Osteichtyes, the true bony fish. There are a reported 19,000 bony fish(this is a very conservative estimate) (5). This is an incredibly diverse class which includeseverything from tuna to eels. These fish evolved more that 410 million years ago (Late Silurian), but did not rise until the middle Devonian as the placoderms and huge sharks began to recede indominance (1). The bony fish, while being varied, all share an extremely important
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