FISHES Jawless fishes CLASS AGNATHA The first vereebrates to evolve were the agnathans, or “jawless fishes.” Their traces are found in rocks of the Late Cambrian period, more than 520 mil- lion years old. These first fishes had no jaws in which to seize and manipulate prey. Nor did they have paired fins, to stabilize their bodies in the water. Cac- ching prey and eating it therefore pre- sented. problems; consequently, these fishes were all small Farely more tha 1 fe/30em long), and restricted to either sucking up microscopic food particles from the mud of the seabed, of feeding on plankton in the surface waters. ‘Agnathans had no bone in their bodies. The internal skeleton was made entirely of gristle or cartilage. Unlike bone, this material decays. So the only way that paleontologists know these ancient fishes existed is because their bodies had an “overcoat” of bone; This consisted of a large bony shield that covered the head, and small bony scales that covered the body. This armor plat- ing was the only protection these small, javwless fishes had against attack by the large, predatory sea scorpions that also lived'in these Paleozoic seas. The bony armor has been preserved in the rocks, and gives the fossil agnathans theit collective name of ost- racoderms, meaning “‘shellskins.” De- -spite their lack of jaws, ostracoderms dominated the seas and freshwaters of the northern hemisphere for about 130 nillion years, from Early Ordovician to Lae Devonian times, To distinct ines evolved (see pp. 18-19) — the pteraspi domorphs (the heterostracans and thelodontids), and cephalaspidomorphs (che osteostracans and anaspids). ‘Only 2 types of agnathan survive today. Neither has the bony armor of their ancestors, ani both. are highly specialized, mainly marine fishes — the wormlike, scavenging hagfishes and the ceel-like, parasitic lampreys ORDER HETEROSTRACL The heterostracans were the frst fishes, and the earliest vertebrates, to evolve. ‘The oldest undisputed remains date from the Early Ordovician, some 500 million years ago (below). ‘These fishes ‘were che most abundant and diverse of all the agnathans. They reached their peak during Late Silurian and Early Devonian times, when a variety of rmatine forms evolved, from mud-eating bottomdwellers to” free-swimming plankton-feeders, Later, they invaded freshwaters. All had the characteristic bony head shield, which could geow throughout the life of the fish. 4 name: Arandaspis tie: Early Ordovician tocatiry: Australia (Northern Teer ica size: possibly 6in/15.em long In 1959, fossils of 4 distinct types of fish were discovered south of Alice Springs, in the heart of the Australian continent. ‘They were entombed in marine sand- stones, laid down in a shallow sea about 500 million years ago. Ie was not until the late 19605 that these remains were recognized as belonging to the earliest- Known vereebrates. “The name of Arandaspis was given t0 the bestpreserved specimens. This was coined to celebrate the aboriginal ‘Aranda tribe of the area, and to reflect the fish’s bony head covering — in Greek aspis means “shield.” ‘Arandastys had a streamlined, deep- bodied shape. With no fins to stabilize it, it would have swum in an erratic, tadpolelike style. Its lower body was covered in oblique rows of small bony scales, each shaped like a cowrie shell, and decorated with pointed tubercles, hich must have given the skin a rough, abrasive texture, like that of a shack, “The front of the body was encased in ahead shield, made from 2 large plates of thin bone —a deep, rounded ventral (underside) plate and a flatter dorsal (topside) plate. There were openings for the eyes, nostrils and the single pair of gill apertures on. either” side. Deep grooves in the shield marked the po- sition of the lateral line canals — the organs in a fish that sense vibrations. ‘Arandaspis’jawless mouth was on the underside of its head, suggesting that it may have fed on or neat the seabed. Like cacher heterostracans, there were prob: ably small, movable plates inside its mouth equipped with ridges of dentine, ‘hase could have formeda flexible pat of “lips,” capable of scooping or suck- ing up particles of food from the mud Name: Preraspis ‘rine: Early Devonian Locatrry: Europe (UK and Belgium) zu in[2dem long Peeraspis is typical ofthe preraspid fami ly of heterostracans, which became very ‘numerous and diverse during the Late Silurian and Early Devonian. Although it lacked paired’ fins, Preraspis was a powerful swimmer, to judge by several hydrodynamic features of is body. Ste bility was provided by bony outgrowths from the back of the head shield — a large spine acted as a kind of dorsal fin, while 2 rigid “wings” oF keels func” tioned as pectoral hydrofoil, The long, flexible tal was also hydro- dynamic, with the lower lobe elongated to provide lift at the front of the body uring swimaning, Additonal lift was provided by the elongated snout, which Sas drawn ou intove bladlike "cor trutn" below which the mouth opened Paleontologists thing that Prvesis and its celatives fed in mid-vater orneat the surface of the sea; among the shoals Of planktonic, shrimplite custacenns ran: Doryaspls ‘tn Early Devonian Toeaurey: Spitsbergen Ser in/isom long ‘This pterapsid (also called Lyktaspis} had 1 much longes snout or rentrum than that of its relatives. There were bony spines set along its length (rather like the “saw’’ of a modern sawfish), and the mouth opened above, rather than below. the rostrum. sae erage ap: pendage must have hid a hydredyramic Rincon sine the shape'of Dayesps supuete wos an etiveswinamer prot ably feeding on plankton ‘An additional function of the ros tedmamong prerapids could have boon to stir up che bottom mud or sand, to. Dumasps had unucualy long, lateral Sass had unuvvaly long, lateral keels growing from the back ofthe head shield. Their leading edges were armed. ‘ih toot spins, These may hae feted as gliding planes, and together ‘vith the rostrum and downcurned tal Would have elevated the front oF the body during swimming. NAME: Drepanaspis ‘rome: Early Devonian Locauiry: Europe (Germany) size: 1 {/30em long, A number of heterostracans, such as Drepanaspis, were well-adapted bottom- dwellers, scavenging in the mud of the seabed for food. The front of the body was broad and flattened, and the eves were set wide on either side of the upturned mouth.oss ORDER THELODONTIDA, The thelodontids were small jawless fishes, related to the heterostracans, although they did not have head shields. Only the tiny bony scales that covered their bodies remain to testify to their ‘existence during Late Silurian and Early Devonian times. NAME: Thelodus ‘Time: Late Silurian yocatiry: Worldwide | s@: Tin/18.em long The mouth of this small thelodontid was on the underside of its flattened hhead, which suggests that it fed on the seabed. But it could obviously swim well also. The lower lobe of the tal was clongated, and fins were developed for stability a dorsal ond an anal fn at the rear, and 2 pectoral flaps at the front. ORDER OSTEOSTRACT The osteostracans (also called the ceph- alaspids, meaning “head shields”) ap- peared in the Late Silurian, about 80 million years after the earliest hetero- stracan fishes. They evolved in the sea, and then colonized freshwaters The osteostracans were flattened bottomadwellers that sucked up food particles from the seabed through a tounded mouth on the underside of the ‘head. The head shield was made of a le plate of undivided bone, which did not grow during the fsh’s adute life (unlike the individual bony head plates ofthe heterostracans) Evidently, osteostracans were also swimmers, since many of them dorsal fin, paired scale-covered flaps where the pectoral fins are norm- ally found, and a strong, upturned tail The anatomy of osteostracans is well known because of a unique develop- ment among these jawless fishes. Bone was laid down inside the body, ina thin layer over the cartilage of the skeleton. From this fossilized bone, the detailed structure of the brain, gills, mouth and even individual nerves and blood ves- sels can be traced. ‘Another innovation was the concen- trated patches of sensory organs devel- ‘oped on both sides, and on top, of the head. These organs were richly supplied with nerves, and must have detected water-borne’ vibrations. Alternatively, they could have been electric organs. NAME: Tremataspis ‘roe: Late Silurian Locatir¥: Europe (Estonia) size: 4in/10cm long This early osteostracan had the typical flattened body and ventral mouth of a bottom-dweller. Its eyes and single nos- tril were on top of the head, near the midline. It fed by sucking up tiny food particles from the seabed, using the gill ‘muscles in the throat asa suction pump. ‘The bony head shield extended half way along the body. Since it was made of ane piece of bone, it is unlikely that it became larger as ‘the animal grew. Paleontologists think that osteostracans hed an unermored larva, and thatthe bony shield developed only when the fish was full-grown, same: Dartmuthia “ine: Late Silurian oeautt¥: Europe (Estonia Sees 4int0em lon ‘The broad head shield is the only pact of Daremuthia hae ts known. It was ‘bomtommfeeder, with sound sek south on the underside ofthe hea ike 5. contemporary —Trematasps (above) These wat a sirall dorsal fin Eaifeny along its back and the pressure: sensitive organs wete well developed on its head and behind the eves: Name: Hemicyclaspis ‘rim: Early Devonian Earope England) ‘em long ‘This osteostracan was 8 more powerful swimmer, and could maneuver itself better inthe water, than either of its bottom-dwelling relatives Tremataspis ‘or Dartmuthia (above). A dorsal fin stabir lized the body, while a paie of scale- covered flaps, like pectoral fins, pro- Vided uplift and kept it on course The comers of the head shield were drawn Out into keel-like cutwaters, And the enlarged upper lobe of the tail pro- duced lift at the rear of the body, 50 keeping the fsh’s head down while it sucked up food from the seabed. FISHES ‘rms: Early Devonian Locatiry: Spitsbergen size: 5in/13em long ‘An least 14 species of Boreaspis are known feom sandstones laid down in the lagoons of Spitsbergen during the Early Devonian. They. differ in. the width of their triangular-shaped head shields, and in che length of the bony spine chat grew our from the cheek area on either side. The snout was elongeted in all species into a bladelike rostrum, Beside its hydrodynamic function, the rostrum was probably used to probe for prey on the muddy lagoon floor. ORDER ANASPIDA ‘The anaspids lacked the heavy head shields of other armored. agnathans. Covered in thin scales, their bodies were slender and flexible, with stabilizing fins. Numerous in the seas of Late Silurian Europe and North America, these active swimmers later invaded rivers and lakes during the Devonian, and survived to the end of that period ‘They are the likely ancestors of the modern lampreys. NAME: Jamoytius: aes Late Silurian Locattry: Europe (Scotland) Size IL ine Tony Named for the English paleontologist J. A. Moy-Thomas, the marine Jamoy. tisha a rartow, ubular shape, wih a Jong fin on its back, a pair of lateral fins sinunesfong eta loa i Uptit was produced by the sro downturned tail. = Jamoytivs had a round, suckerlike mouth, and it was probably a parasite like its living descendant, the marine Inmprey. Th jawess eh ata tlt ‘to other fishes, rasps away their flesh and then sucks their blood, Name: Pharyngolepis ‘Time: Late Silurian Locatrry: Europe (Norway) size: 4in/10em tony This anaspid must have been an awk- ward, ‘inefficient swimmer, since it lacked the basie stabilizing fins of other fishes. A row of crested scales ran along its back, and a pair of bony spines projected from the pectoral area. There ‘was a welldeveloped anal fin, and the tail was downturned. But none of these features would have stabilized this fish's deep body inthe water Pharyngolepis’ feeding method was probably to plow through the bortom sediment, heacefirst, scooping up tiny food particles in its founded mouth 5FISHES Cartilaginous fishes DELTOPTYCHIUS HYBODUS| XENACANTHUS COBELODUS %