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Chapter 4

Craniate Taxa
Not all taxonomies are identical
Taxa consist of species designated by a Latinized binomial name (Linnaeus)
o Enables everyone to understand

Agnatha (Agnathans include 2 contrasting groups: extinct ostracoderms and living hagfishes and lampreys)
Ostracoderms armored craniates oldest known craniates before discovery of Chinese fossils
o Date back to mid-Ordovician or Cambrian
o Dermal armor of broad plates and small tile like scales. (plates largest on the head)
o Lack paired fins
o Stensio anatomy under fossilized head shield of osteostracan (cephalaspidomorph ostracoderm [Cephalaspidomorphi])
Head skeleton flattened denticle-covered bony shield with 4 dorsal apertures
2 were a pair of upward staring eyes
3rd was median/pineal eye
Small anterior opening was a single naris where a nasohypophyseal duct led to olfactory sac etc.
Shield turned under along its lateral edges where it was then replaced under gills by tile like scales
Between anterior edge of shield and scales, a small mouth opened to oro-pharyngeal chamber lined by gills
Curved row of external gill slits extended from corners of mouth to caudal margins of head shield
Head contained an endoskeleton of endochondral bone and cartilage
Little is known of the postcranial skeleton except that the tail was heterocercal like most Paleozoic fishes
o Origin is unknown; possible ties with protochordates and basal craniates; disappeared at the end of Devonian
Living Agnathans hagfish and lamprey
o Hagfish has class status: Myxini in a superclass Agnatha in one cladistic fish taxonomy
o Lampreys tentatively included with ostracoderms in class Cephalaspidomorphi (tentative postulate)
o Hypothesis: hagfish sister group to vertebrates
o Among living taxa, lampreys: sister group to gnathostomes
o Considering fossils, cephalaspidomorphs represent a paraphyletic grouping
o Other postulates: hagfishes and lampreys referred to as cyclostomes
o Living agnathans have prominent notochord = sole axial skeleton
o No paired fins, no skeleton comparable to jaws, no vertebral column comparable to typical vertebrate spine, no bone
o They have 1 (hagfish) or 2 (lampreys) semicircular ducts instead of the 3 found in jawed vertebrates
o Single nostril connected with a single olfactory sac.
o Parasitism adaptations (buccal funnel and rasping tongue)
o Shallow buccal funnel that lacks rasping denticles, surrounded by ring of stubby finger like papillae
o Bottom feeding scavengers who eat invertebrates and viscera of dead or weak fishes
o Eyes, unlike lampreys, are vestigial and covered by opaque skin
o Myxine glutinosa (atlantic hagfish) 6 pairs of pore-like gill pouches that open into a common efferent duct
o Eptatretus stouti has 10-15 pairs of fill pouches that open to the exterior
o Do not enter freshwater and larvae stay within egg mambranes until metamorphosis (adaptation to saltwater)
o Have not changed appreciable since the Carboniferous
o Large buccal funnel lined with horny denticles helps keep it attached to host
o Tongue-like cartilaginous rod with horny teeth rasps flesh of victim leaving only skin and skeleton
o A nasohypophyseal duct leads from median nostril to oldactory sac then terminates blindly in a nasohypophyseal sac
o 7 pairs of gill pouches open separately to exterior via porelike gill slits that conduct water in and out
o Petromyzon marinus anadromous lampreys; live in sea but migrate upstream to lay eggs. In 3 weeks,
ammocoete larvae emerge. Metamorphose in freshwater for several years then migrate to sea = sexual maturity
o Suggesting a relationship between lampreys and gnathostomes (=vertebrata) is the presence of vertebral
elements in trunk, 2 semicircular canals, electroreception and other characters

Gnathostomes: Placoderms
3 groups evolved in Paleozoic : placoderms, chondrichthyans, teleostomes (acanthodians) and osteichthyans
2 abundant groups of unusual jawed fishes: placoderms and acanthodians are present in Paleozoic
Placoderms represent the sister group of chondricthyans + teleostomes and evolved from ostracoderm ancestors
Among teleostomes, acanthodians are the sister group to osteichthyans (lobe finned and ray finned fishes)
Placoderms bony dermal plates had paired pectoral and pelvic fins
Arthrodires heavy bony shields covered the head and gill and another covered trunk; active predators
Antiarchs small armored placoderms with eyes on top if head and flat belly (bottom feeders)
Phylogenetic relationships of placoderms to other jawed fishes have been a matter of conjecture. Evidence suggests that
placoderms are a basal gnathostome group sister group to other gnathostomes
Placoderms disappeared leaving no descendants

Chondrichthyes (Cartilaginous Fishes)

Elasmobranchii - sharks, rays, skates and sawfishes and Holocephali (chimaeras or ratfishes)
No bone in their body other than their scales and teeth
Mouth is on ventral surface rather than terminal except in Paleozoic sharks like Cladoselache
Placoid dermal scales composed of basal plate and bony spine of dentin that protrudes through epidermis = sandpaper
Skeleton of pelvic fins of males modified to form claspers for transferring sperm
Macrolecithal eggs and in oviparous species often encased in horny/leathery shell with tendrils that entwine it
Elasmobranchs (Elasmobranchii)
o Sharks, skates, rays, sawfishes
o Exposed gill slits (naked) vs covered by operculum usually 5 pairs (Hexanchus 6, Heptanchus 7)
o 7 slits may have been primitive because that was the number in Cladoselache (largest # in any jawed fish)
o Spiracle has miniature gill-like surface (pseudobranch)
o Heterocercal caudal fin
o Shark
Squaliformes as dissection species because of generalized anatomy
Spiny Atlantic dogfish Squalus acanthias usually studied named for prominent spine associated with
each dorsal fin; Squalous suckleyi is the Pacific spiny dogfish
o Rajiformes (rays, skates and sawfishes) compressed dorsoventrally
5 ventral gill slits but spiracle (chief incurrent route for repiratory water is dorsal)
Most rays and skates subsist on mollusks; location of spiracle removes it from mud and debris
Manta rays are filter feeders
Tail of Rajiformes changed from muscular locomotor organ to lean whiplike organ of defense/offense
Electric rays (torpedos) houses electroc organ that immobilizes prey
Stingrays rows of spines along tails with poison
Sawfishes have rajiform shape to lesser degree (sword for impaling small fishes and poking bottom)
o Chimaerans lack scales on most of the surface, with fleshy operculum, closed spiracle, upper jaw is solidly fused with
cartilaginous braincase and instead of teeth, hard flat bony plates on jaws crush molluscan shells

Acanthodians and Osteichtheyans (spiny and bony fishes) => united in Teleostomi (represents sister group to chondrichthyans)
o Extinct spiny fishes
o Stout hollow spines associated with median and paired fins and sometimes paired spines
o Body was covered by bony armor of small scales and dermal plate on the head
o Skeleton consisted of bone and cartilage
o With large operculum (primitively consisting of elongate scales with ancillary gill covers followed by single opercular
o Acanthodian spines found in early Silurian; fossils abundant from Devonian and Carboniferous, then disappear
o Include all bony fishes and their tetrapod descendants
o Characterized by an air sac (lung/swim bladder) that may be secondarily lost and presence of large units of dermal
bone on head and shoulder girdle
o Subdivided based on structure of paired appendages: Actinopterygii (ray fin) and Sarcopterygii (lobe fin+tetrapod)

Actinopterygii (ray finned)

Ancient & modern bony fishes - membranous fins supported by slender fin rays from basal skeletal elements within body wall
Bony operculum; air sac (swim bladder/lung); internal nares lacking
During Paleozoic, bony dermal armor and scales overlain with form of enamel called ganoin; caudal fins: heterocercal. Both
traits have all but disappeared in more recent species
Basal Actinopterygians
o Oldest known ray finned fishes paleoniscoids (Devonian to beginning of Mesozoic)
o African freshwater fishes Polypterus and Calamoichthys
large ganoid scales; ossified endoskeleton and air sacs connected to pharynx by air duct (aerial respiration)
sometimes called African lungfishes but shouldnt be confused with Dipnoans
o Sturgeons and paddlefishes
Cartilaginous endoskeleton (embryonic cartilage not replaced)
Scales lack ganoin
Paddlefish skin is naked except for small bony scales on the tail
Taxon Chondrostei (mostly extinct monophyletic assemblage where sturgeons & paddlefish are assigned)
o Includes surviving Mesozoic ray-finned fishes: gars (2 genera), bowfins (1 species: Amia calva) and vast array of
more recent ray finned fishes the teleosts
o Gars and bowfins classified as non-monophyletic holosteans are placed in distinct clades (Ginglymodi &
Freshwater fishes
Trunk and tail with ganoid scaled that are modified ganoid scales of paleoniscoid fishes
Trunk and tail of Amia have modern fish scales
Much of endoskeleton is ossified in both genera but braincase (neurocranium) remains cartilaginous
o Teleosts placed in taxon Teleostei
Most recently evolved modern
Scales in the dermis have become thin and flexible
Dermal skull bones are thinner and more numerous than in any other bony fishes
Haws and palate more independently maneuverable
Pelvic fins in many species are far forward
Body has become altered in many ways organisms that occupy all aquatic niches on palnet
96% of living fishes; species exploding since Mesozoic

Sarcopterygii (lobe finned)

Bony fishes that have a prominent fleshy lobe at base of paired fins and include their tetrapod descendants whose
appendages have been modified into limbs; lobefin contains part of the fin skeleton
Internal nares that open into the oropharyngeal cavity and retain a gas filled air sac
Gill slits covered by bony operculum that grows caudad from 2nd pharyngeal arch
Actinistia coelacanths
o All extinct except the relict coelacanth: Latimeria
Rhipidistia includes Dipnoi
o Skeletal elements in fin lobes correspond to proximal skeletal elements of early tetrapod limbs
o Skull similar to first amphibians
o With air sacs probably used at least occasionally as lungs and most had internal nares (not for breathing)
o Clouthier and Ahlberg places lungfishes within rhipidistians
o Freshwater habitat but new analyses suggest marginal marine habitat including fossil lungfish
o true lungfishes with 3 living genera
Protopterus (aftrica), Lepidosiren (brazil), Neoceratodus (Australia)
Inefficient gills P and L would suffocate if held underwater
N utilizes gills except when oxugen content of water is low
Wet season: inhabit streams and swamps; when sun comes: dig deep burrows in mud and spend hot
season in state of aestivation (lowered metabolism minimizes water loss and reduces needs)
Resemble amphibians (also seen in Amia and Polypterus), swim bladder/lungs have ducts leading to
pharynx with blood supplied by branch from 6th aortic arch instead of from forsal aorta in teleosts
Heart atrium partially divided into 2 chambers. Both usually have larval stage with external gills and both
have internal nares
o Remember that if we accept lungfishes as rhipidistians, they are among living taxa, the sister group to tetrapods. It is
not until we consider the numerous fossil taxa that we appreciate true relationship between lungfishes and tetrapods

Basal tetrapods have been subdivided although relationships among these groups remain unclear
A number of basal taxa share a unique labyrinthine folding of the dentine seen in cross section of teeth. This shared feature
was the basis for placing this diverse group in Labrythodontia (now recognized as a paraphyletic group).
By the Carboniferous, many groups are recognized including Temnospondyli, Anthracosauria and Microsauria
Hypothesis of relationship of living amphibians (lissamphibia) sister group to Temnospondyli. Alternatively, lissamphibians
may have evolved from a temnospondyl ancestor
Class amphibian is a paraphyletic group that omits its amniote descendants
o Oldest amphibians; swamp dwellers, Ichthyostega oldest (Devonian)
o Large, widely dispersed, diverse assemblage; kinships unclear from lack of fossil evidence
o Structure of labrynthodont vertebrae played prominent role in attempts to reconstruct phylogenetic lineages and
identify lanrynthodont ancestors of amniotes
o Had many features seldom seen in modern amphibians (minute bony scales in dermis, fish-like tail supported by
dermal fin rays, skulls similar to rhipidistian fishes)
o Grooves in skull bones under skin = had sensory canal system of neuromast organs that monitored aquatic
environment (todays aquatic amphibians have this system; terrestrial species lost it at metamorphosis)
o Common in the Permian with its fossil record extending back to the Mississipian
o Has skeletal similarities to modern frogs and salamanders = close relationship
o A # of lissamphibian skeletal features & smaller size explained as the retention of juvenile ancestral temnospondyl
features (paedomorphosis); condition in caecilians doesnt fit easily = independent origin from microsaurs
o Represent diverse group of fossil forms known from Pennsylvanian to Lower Permian
o Share a # of skeletal features with caecilians = close relationship/ convergence on an elongate body for burrowing
o Include 3 group of extant amphibians
Apoda (gymnophiona) limbless, burrowing caecilians
Urodela (caudate) tailed amphibians
Anura (Salientia) frogs, taids and tree toads
o Taxon also includes Triassic and Jurassic anurans whose skeletons resemble those of todays frogs and toads
o Given a monophyletic origin of amphibians, alternative hypothese suggest that lissamphibians are either
monophyletic (common temnospondyl ancestor) or diphyletic (apodans descendewd from microsaurian ancestor)
o Fossil record of apodans consists solely of vertebrate from Cenozoic and Late Cretaceous; however the best
preserved material is from the Jurassic of Arizona. This material consists of both cranial and postcranial skeleton
showing an animal with an apodian style skul but with limbs, although reduced. Supports monophyly of Lissamphibia
Apodans (caecilians)
o Circumtropical, limbless amphibians that except for aquatic species, live in burrows in swampy locations.
o Small eyes and buried beneath bones of the skull
o Some species have minute scales in their dermis; 250 vertebrae
o Vent is almost at the end of body short tail
o Terrestrial species lay yolky eggs and the larcal stage is passed in the egg envelopes. Aquatic genera viviparous
Urodeles (8 families)
o 2 families: Proteidae and Sirenidae (Necturus and Siren) are perennibranchiate (retain larval gills throughoutlife)
o Other families have few perennibranchiate populations; some perennibranchiates undergo complete metamorphosis
when administered iodinated compounds or thyroid hormone
o Necturus only genus of Proteidae in North America
Attains sexual maturity in 5 years
o Proteus European relative blind and lives in dark caves
o Siren lacks hind limbs and lives in muddy ditches and weed choked lakes or ponds
o Plethodon loses gills at metamorphosis but fails to develop lungs; lives in most sites lays eggs in damp places,
cutaneous respiration, larvae hatch with legs already formed and may never enter water
o Amphiuma eel-like urodele with tiny appendages
Amphiuma pholeter 1 digit
A. means has subspecies with 2/3 digits
o Cryptobranchus looks ferocious with broad flat head and wrinkled skin which often coneals single gill slit
o Hynobius basal urodele and Ambystoma terrestrial genera. However, A. mexicanum (Mexican axolotl) is an
aquatic perennibranchiate that can be caused to discard its gills experimentally
o Notophthalmus and Salamandra true salamanders (family Salamandridae)
Salamandra atria viviparous
Notophthalmus viridescens interesting life history
After several months of aquatic larval existence, red spotted newt loses its gills and gill slits,
sprouts legs and now an eft, leaves pond for land. Skin develops a thick cornified layer that blocks
the openings of the sensory canal system and skin glands, the body assumes a bright orange red
color, and a series of dorsolateral black bordered red sports appear. Red eft stage 1-3 years
At approach of sexual maturity, the eft, stimulated by the hormone prolactin, joins a mass migration
to freshwater ponds where mating occurs and eggs are deposited. The tail commences to change
from round to laterally compressed with dorsal and ventral keels, the thick cornified layer of
epidermis is shed, exposing the openings to mucous glands and the sensory canal system and
they body gradually becomes olive green above and light yellow below, a protective coloration
appropriate for life in a freshwater pond. The animal is now a newt. In some localities the larvae
remain in the pond, mature and retain vestigial gills throughout life
o Tailless amphibians where several caudal vertebrae are fised into 1 elongated urostyle
o Most representative frogs are the family Ranidae; toads are the family of Bufonidae (more terrestrial than frogs)
o Tree toads/tree frogs family Hylidae
o Breathe with lungs and skin and most amphibious
o A few, Rana cancrivora crab eating frog of Thailand can tolerate salt water
o Most anurans breed during/ shortly after rain
o Robber frogs lay eggs in rain filled crevices/pile of leaf mold/base of grass hummock away from waters edge and
above flood level = larval stage not in water; as adaptation tadpoles remain within jelly until metamorphosis
o Some species of tree toads lay eggs in a brood pouch under the skin of their back and fully metamorphosed treefrogs
emerge through an opening in its skin; similar condition found in aquatic frog Pipa
o Nectophrynoides vivipara viviparous; as many as 100 young develop in uterous
o Earliest known anuran triadobatrachus (=protobatrachus) from Triassic. Skull similar to todays Anurans and body
was shortened as a result of a reduction that had already taken place in the # of trunk vertebrae. The ribs were not as
foreshortened , the tibia and fibula were not fused and bony scales covered the abdomen
o Anthracosauria, small Paleozoic group is thought to be in direct line to amniotes. Fossil record extends from
Missisipian to the Triassic