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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)
Hagfishes
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)
Lampreys
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
Acanthodians and Osteichtheyans (spiny and bony fishes) => united in Teleostomi (represents sister group to chondrichthyans)
Acanthodians
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
cover)
o Acanthodian spines found in early Silurian; fossils abundant from Devonian and Carboniferous, then disappear
Osteichthyans
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)
Amphibia
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
Labrynthodonts
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)
Temnospondyls
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
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
Lissamphibians
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
Anurans
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
Anthracosaus
o Anthracosauria, small Paleozoic group is thought to be in direct line to amniotes. Fossil record extends from
Missisipian to the Triassic