A second major group of chondrichthians, in the subclass Holocephali (hol\ue001o-sef\ue000a-li) (Gr.h o l o s , whole\ue002kephalidos, head), contains about 30 species. A frequently studied example,
large lips. A narrow, tapering tail has resulted in the common name \u201cratfish.\u201d Holocephalans diverged from other chon- drichthians nearly 300 million years ago. Since that time, spe- cializations not found in other elasmobranchs have evolved, in- cluding a gill cover, called ano p e r c u l u m , and teeth modi\ufb01ed into large plates for crushing the shells of molluscs. Holocephalans lack scales.
Frequently, elaborate color patterns on the dorsal surface of these animals provide effective camou\ufb02age (see \ufb01gure 18.9c). The sting ray( D a s y a t i s ) has a tail modi\ufb01ed into a defensive lash; the dorsal \ufb01n persists as a venomous spine. Also included in this group are the electric rays( N a r c i n e andTorpedo) and manta rays( M a n t a )
show posteriorly pointing placoid scales (SEM\ue003500).(b) The teeth of sharks develop as modi\ufb01ed placoid scales. Newer teeth that move from the inside to the outside of the jaw continuously replace older teeth.
Members of the class Osteichthyes (os\ue001te-ik\ue000the-es) (Gr.osteon, bone\ue002ichthyos, \ufb01sh) are characterized by having at least some bone in their skeleton and/or scales, bony operculum covering the gill openings, and lungs or a swim bladder.Any group that has at
least 20,000 species and is a major life-form in most of the earth\u2019s vast aquatic habitats must be judged very successful from an evolutionary perspective.
The \ufb01rst fossils of bony \ufb01shes are from late Silurian deposits (approximately 405 million years old). By the Devonian period (350 million years ago), the two subclasses were in the midst of their adaptive radiations (see table 18.1; see also \ufb01gure 18.3).
\ufb01ns and usually use lungs in gas exchange. One group of sarcoptery- gians are the lung\ufb01shes. Only three genera survive today, and all live in regions where seasonal droughts are common. When freshwater lakes and rivers begin to stagnate and dry, these \ufb01shes use lungs to breathe air (\ufb01gure 18.11). Some(Neoceratodus) inhabit the freshwa- ters of Queensland, Australia. They survive stagnation by breathing air, but they normally use gills and cannot withstand total drying. Others are found in freshwater rivers and lakes in tropical Africa
survive when rivers or lakes are dry by burrowing into the mud. They keep a narrow air pathway open by bubbling air to the surface. After the substrate dries, the only evidence of a lung\ufb01sh burrow is a small opening in the earth. Lung\ufb01shes may remain in aestivation for six months or more. (Aestivation is a dormant state that helps an animal withstand hot, dry periods.) When rain again \ufb01lls the lake or riverbed, lung\ufb01shes emerge from their burrows to feed and reproduce.
A second group of sarcopterygians are the coelacanths. The most recent coelacanth fossils are over 70 million years old. In 1938, however, people \ufb01shing in deep water off the coast of South Africa brought up one \ufb01sh that was identi\ufb01ed as a coelacanth (\ufb01gure 18.12). Since then, numerous other specimens have been caught in deep water around the Comoro Islands off Madagascar. The discovery of this \ufb01sh, Latimeria chalumnae, was a milestone event becauseLatime-
It is large\u2014up to 80 kg\u2014and has heavy scales. Ancient coelacanths lived in freshwater lakes and rivers; thus, the ancestors ofLatimeria must have moved from freshwater habitats to the deep sea.
The subclass Actinopterygii (ak\ue001tin-op\ue000te-rig-e-i) (Gr.aktis, ray\ue002pteryx, \ufb01n) contains \ufb01shes that are sometimes called the ray- \ufb01nned \ufb01shes because their \ufb01ns lack muscular lobes. They usually possess swim bladders, gas-\ufb01lled sacs along the dorsal wall of the body cavity that regulate buoyancy.Zoologists now realize that
there have been many points of divergence in the evolution of the Actinopterygii. One modern classi\ufb01cation system divides the Actinopterygii into two infraclasses.
One group of actinopterygians, the chondrosteans, contains many species that lived during the Permian, Triassic, and Jurassic periods (215 to 120 million years ago), but only 25 species remain today. Ancestral chondrosteans had a bony skeleton, but living members, the sturgeons and paddle\ufb01shes, have cartilaginous skele- tons. Chondrosteans also have a tail with a large upper lobe.
Most sturgeons live in the sea and migrate into rivers to breed (\ufb01gure 18.13a). (Some sturgeons live in freshwater but main- tain the migratory habits of their marine relatives.) They are large (up to 1,000 kg), and bony plates cover the anterior portion of the body. Heavy scales cover the tail. The sturgeon mouth is small, and jaws are weak. Sturgeons feed on invertebrates that they stir up from the sea or riverbed using their snouts. Because sturgeons are valued for their caviar (eggs), they have been severely over\ufb01shed.
Paddle\ufb01shes are large, freshwater chondrosteans. They have a large, paddlelike rostrum that is innervated with sensory organs believed to detect weak electrical \ufb01elds (\ufb01gure 18.13b). They
swim through the water with their large mouths open, \ufb01ltering crustaceans and small \ufb01shes. They are found mainly in lakes and large rivers of the Mississippi River basin and are also known from western North America and China.
The second group of actinopterygians (Neopterygii) \ufb02our- ished in the Jurassic period and succeeded most chondrosteans. Two very primitive genera occur in temperate to warm freshwaters of North America.Lepisosteus, the garpike, has thick scales and long jaws that it uses to catch \ufb01shes.Amia is commonly referred to as the dog\ufb01sh or bow\ufb01n. Most living \ufb01shes are members of this group and are referred to as teleosts or modern bony \ufb01shes. They have a symmetrical caudal \ufb01n and a swim bladder that has lost its connection to the digestive tract.After their divergence from an-
cient marine actinopterygians in the late Triassic period, teleosts experienced a remarkable evolutionary diversi\ufb01cation and adapted to nearly every available aquatic habitat (\ufb01gure 18.14). The number of teleost species exceeds 20,000.
geons (Scaphirhynchus platorynchus). Sturgeons are covered anteriorly by
heavy bony plates and posteriorly by scales.(b) The distinctive rostrum
of a paddle\ufb01sh (Polydon spathula) is densely innervated with sensory
structures that are probably used to detect minute electrical \ufb01elds. Note
the mouth in its open, \ufb01lter-feeding position.
winter \ufb02ounder (Pseudopleuronectes americanus), have both eyes on one
side of the head, and they often rest on their side fully or partially buried
in the substrate.(b) The yellowtail snapper (Ocyurus chrysurus) is a pop-
ular sport \ufb01sh and food item found in offshore tropical water. It reaches a
length of 75 cm and a mass of 2.5 kg.(c) The sarcastic fringehead(Neo-
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