SYSTEMATIC BIOLOGY

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CLASS ARCHOSAUROMORPHA

Archosauromorpha (ar-ko-SOR-o-MOR-fa) is made up of three Greek roots that mean "ancient lizard forms" [ancient- archaios (αρχαίος), lizard- savra (σαύρα), and form- morphi (μορφή)].  The reference is to the ancient lizards (i.e. dinosaurs) that make up most of the members of this class.

This group of vertebrates is remarkably diverse and has exploited environments on the land, water, and air since its appearance in the Permian.  This monophyletic group of diapsids is sister to the Lepidosauromorpha and includes crocodilians, pterosaurs, rhynchosaurs, dinosaurs, and birds.    Although the basal archosaurs were somewhat sprawling and lizard-like in their gaits, many evolved a more upright and bipedal stance.  The crocodilian line appears to have evolved from a line that was bipedal, or partly bipedal (alternating between bipedal and quadripedal stances).  Furthermore, bipedalism seems to have freed the forearms and allowed for the evolution of winged flight in two separate lines (pterosaurs and birds).  The ankles of the crocodilians could rotate while most of the other archosaurs evolved a more simple hinge-like ankle joint.  

Crocodilians as a group have returned to a semi-aquatic existence.  Thus, they have returned to a sprawling, lizard-like stance with a deep, flattened tail for propulsion in the water.  The once diverse group has been reduced to 23 species, which tend to be the top predators of tropical and subtropical wetlands.  

Pterosaurs (from two Greek roots meaning "winged lizard") evolved true flight on outstretched bat-like wings during the Triassic Period.  The skin was stretched tight and supported by an elongated fourth finger.  Although thin, the skin of the wings, which could have a span of 13-15 meters for Quetzacoatlus, was strengthened by bands of collagen fibers.  Traces of hair on the bodies of some pterosaur fossils suggests that they were endothermic.  Thus, their presumed physiology and wing structure support the view that they were strong and agile fliers.

Dinosaurs (from two Greek roots meaning "terrible lizard"), or, more appropriately, the non-avian dinosaurs, appeared during the Triassic Period as small bipedal carnivores, and they formed two large groups defined by the structure of the pelvic girdle: the Saurischia ("lizard-hipped dinosaurs") and the Ornithischia ("bird-hipped dinosaurs) that flourished in the Jurassic and Cretaceous Periods.  The saurischia include the theropods and sauropods, the most popular of the dinosaurians.  Theropods ("beast-footed dinosaurs"), which remained bipedal and carnivorous, included Tyrannosaurus, Coelophysis, Velociraptor, and Troodon.  Some of these taxa likely had feathers.  The resemblance to birds is more than superficial.  In fact, from a cladistic perspective, all birds are embedded within the theropods.  The sauropods ("lizard-footed dinosaurs") included herbivorous taxa that returned to a quadrupedal stance and included the largest terrestrial vertebrates that have ever lived (e.g. Apatosaurus, Camarasaurus, Brachiosaurus, and Diplodocus).  Typically, they had long necks, small heads, immense bodies on pillar-like legs, and long tails.   Sauropods had weak teeth that could do little more than tear away plant stems, leaves, twigs, etc., but did allow them to consume large masses of food that they processed in large fermentation chambers.

The ornithischians were entirely herbivorous and included the armored ornithischians (thyreophores - "shield-bearers"), the horned ornithischians (ceratopsians- "horn-faced dinosaurs"), and the ornithopods (bird-footed dinosaurs).  The thyreophores were quadrupedal animals with small heads, short necks, and large bodies on pillar-like legs.  Their bodies were covered by nodules of bony armor or plates and most had defensive spikes or clubs on the ends of their tails.  Some of the principle taxa included Stegasaurus and Ankylosaurus.  The ceratopsians appeared as bipedal animals in the lower Cretaceous, but most evolved to become large quadrupeds by the middle of the Cretaceous.  They were characterized by having an anterior shearing beak, the back of the skull expanded into a frill, and one or more horns on the face and/or frill.   Common taxa included Protoceratops and Triceratops.    The ornithopods remained bipedal or facultatively bipedal.  The principle feature of the head was the evolution of batteries of cheek teeth that were inset, suggesting that they had a cheek and could masticate plants material much like a cow does.  Most evolved a beak in the front of the jaw, which gave them the name, duck-billed dinosaurs.  Complex crests that may have served as resonating chambers on many taxa point to the likelihood of communication by sound, a characteristic of highly social vertebrates.  Indeed, evidence suggests that all of the dinosaurs lived in social groups.

Birds are the most successful of all tetrapods with more than 9,500 living species.  They are characterized by having a body covered with feathers, a light skull with no teeth, a highly reduced tail, and thoracic/lumbar vertebrae that are fused.  They retain the bipedal stance and foot structure characteristic of their theropod ancestors with the forelegs modified into wings that form airfoils made of relatively long flight feathers.  Within this basic body plan, living birds show remarkable diversity in which they have exploited aerial environments as well as terrestrial, arboreal, and aquatic environments.  They range in size from the ostrich (up to 3m tall and nearly 200kg) to the hummingbird that is no  longer that 7 cm and weighs less than 2-5 g.  More than 60% of all living birds belong to a single order (Passeriformes), commonly called the songbirds.  In general, birds are highly social animals with elaborate behaviors that include complex vocalizations and ornate displays which often feature brightly colored plumage.

HIERARCHICAL TAXONOMY OF THE CLASS ARCHOSAUROMORPHA. The following descriptions come from Benton (2005) and the Animal Diversity Web from the University of Michigan Museum of Zoology.  The structure of the following system is also based, in part, on the phylogenetic treatment by Mikko's Phylogeny Web (from the Finnish Museum of Natural History).     Taxa in red are extinct.
 

Photo of a crocodile in Costa Rica (left) and a painting of a pterosaur (right).

 

Painting of a hadrosaur, a member of the ornithischian dinosaurs (left) and a photo of a cockatiel, Nymphicus (right).

Image of Pterosaur from: http://www.nmnh.si.edu/paleo/PaleoArt/Historical/Highlights/pterosaur.html

Image of Hadrosaur from : http://www.copyrightexpired.com/earlyimage/prehistoriclifebeforekt/hadrosaurus01.html 

Images of the crocodile and the cockatiel are from the Systematic Biology Biodiversity Archive.

CLASS ARCHOSAUROMORPHA

This is one of the most diverse groups of extant and extinct vertebrates.  It includes crocodilians, pterodactyls, dinosaurs, and birds.  According to Benton (2005), the archosauromorphs had the following synapomorphic characters: a posterodorsal process on the premaxilla, a saggital crest, slender cervical ribs that taper, a notch on the anterior margin of the interclavicle, and an iliac blade with a small anterior process and larger posterior process.  Two different groups, the pterosaurs and the birds, independently evolved the ability of true flight.

The class can divided into the following major groups:

SUBCLASS RHYNCHOSAURIA+

SUBCLASS ARCHOSAURIA

CROCODILIANS

PTEROSAURS

DINOSAURS

SAURISCHIA

THEROPODS

SAURPODS

ORNITHISCHIA

BASAL ORNITHISCHIANS

ARMORED ORNITHISCHIANS

HORNED ORNIHISCHIANS

ORNITHOPODS

SUBCLASS AVES

EXTINCT TOOTHED BIRDS

NEORNITHES

PALEOGNATHAE

NEOGNATHAE

GALLOANSERAE CLADE

WATERBIRD ASSEMBLAGE

GRUIFORM-RALLIFORM CLADE

PELACONIFORM+CICIONIIFORM CLADE

SHOREBIRD CLADE

FALCONIFORM CLADE

ALBATROSS-LOON PENGUIN CLADE

OWLS

APODIFORM-GOATSUCKER CLADE

TURACO CLADE

COLUMBIFORM CLADE

PARROT CLADE

CUCKOO CLADE

PERCHING BIRD ASSEMBLAGE

WOODPECKER CLADE

MOUSEBIRD-TROGON-HORNBILL-KINGFISHER CLADE

SONGBIRD CLADE

SUBCLASS RHYNCHOSAURIA+

This is a paraphyletic group of very different basal archosaurs.  They appeared in the upper Permian and disappeared by the end of the Triassic.

ORDER UNNAMED BASAL ARCHOSAUR FAMILIES

FAMILY TRILOPHOSAURIDAE

Herbivores.  These animals had heavy skulls with broad, flattened cheek teeth and an anterior beak (ensheathed in horn?).  These secondarily lost the lower temporal fenestra.

Upper Triassic

Trilophosaurus

FAMILY RHYNCHOSAURIDAE

Herbivores.  They had specialized shearing teeth in the cheek region (two rows) and a beak with a large overbite.  These characters suggest that they must have fed on very tough plants.  The hind feet had specialized claws that seem to have functioned in digging.  These were very successful herbivores throughout the Triassic.  Some animals were up to 2m long. 

Triassic

Hyperodapedon

ORDER PROLACERTIFORMES

Carnivores, somewhat lizard-like with very long necks.  Some likely were aquatic and fed on fish and cephalopods.  These were among the first of the archosauromorphs. 

Upper Permian to upper Triassic

Protorosaurus, Tanystropheus, Prolacerta

SUBCLASS ARCHOSAURIA

These animals all have the following characters:

·        Antorbital fenestra

·        Laterosphenoid

·        Lateral mandibular fenestra

·        Laterally flattened, not rounded, teeth

INFRACLASS UNNAMED+

ORDER UNNAMED

FAMILY PROTEROSUCHIDAE

These were small (to 1.5m), slender and some-what sprawling carnivores. 

Lower Triassic

Proterosuchus

FAMILY ERYTHROSUCHIDAE

Very large carnivores (up to 5m).  They had evolved a 3-pronged pelvis; a fourth trochanter on the femur, and an elongate metatarsal 3 (longer than metatarsal 4).

Middle Triassic

Vjushkovia

FAMILY EUPARKERIIDAE

Members of this family seem to have been quadripedal and bipedal.  They had evolved an s-shaped femur and dermal scales (osteoderms) that went down the back. 

Middle Triassic

Euparkeria

INFRACLASS CRUROTARSI (the crocodilian line)

These animals, mainly carnivores, were characterized by an ankle joint that allowed rotation between the astragulus and the calcaneum.

Unassigned Taxa: Doswellia, Tarjadia, Parringtonia.

SUPERORDER UNNAMED+

ORDER PHYTOSAURIA

FAMILY PHYTOSAURIDAE

Piscivores, but also took tetrapods and carrion.  They had a nostril mound just before the eyes. 

Upper Triassic

Parasuchus, Paleorhinus, Belodon, Ebrachosuchus, Mesorhinosaurus, Rutiodon, Smilosuchus, Leptosuchus, Phytosaurus, Angistorhinopsis, Coburgosuchus, Nicrosaurus, Mysteriosuchus, Redondasaurus, Pseudopalatus.

FAMILY ORNITHOSUCHIDAE (INCERTAE SEDIS)

Carnivore.  These animals superficially resembled early dinosaurs in that they were bipedal and quadripedal; however, they had the crocodilian ankle joint. 

Upper Triassic

Ornithosuchus

FAMILY STAGONOLEPIDIDAE (INCERTAE SEDIS)

Herbivores.  Also called aetosaurs, they had stocky bodies that were covered with bony plates.  They had a short skull with a shovel snout that suggests rooting in the soil for tubers, etc.

Upper Triassic

Stagonolepis

FAMILY RAUISUCHIDAE (INCERTAE SEDIS)

Carnivores, some quite large (6-7m).  This may be a paraphyletic group of up to 4 families.  Some of these likely had an erect gait (pillar-like legs) and could move bipedally or quadripedally. 

Upper Triassic

Saurosuchus

FAMILY POPOSAURIDAE (INCERTAE SEDIS)

Carnivore.  Sometimes placed together with the rauchosuchids.  These had a high skull with recurved teeth, somewhat like the skulls of theropod dinosaurs. 

Upper Triassic

Postosuchus

SUPERORDER CROCODYLOMORPHA

Major wrist bones modified as rods; coracoid with a long backward-pointing spine; pelvis with an open acetabulum.  Also, specializations of the skull.

Unassigned Taxa: Barbarenasuchus+, Clarencea+, Dyoplax+, Hallopus+, Macelognathus+, Parrishia+, Trialestes+.

ORDER UNNAMED+

FAMILY SALTOPOSUCHIDAE

Carnivore. Long skull, and long hind legs, likely bipedal.  Animals small (0.5m). 

Upper Triassic

Saltoposuchus

FAMILY SPHENOSUCHIDAE

Carnivore.  This animal had a heavier build and was larger (1.4m).  It could run fast, but it was a quadruped.  It showed other specializations in the skull that are diagnostic for crocodilians. 

Upper Triassic to lower Jurassic

Sphenosuchus

ORDER CROCODYLIA

Unassigned Taxa: Eopneumatosuchus+, Microchampsa+, Artzosuchus+.

FAMILY PROTOSUCHIDAE+

The protosuchids were small quadripedal animals with hind legs longer than the forelegs (a consequence of a bipedal ancestry).  Posterior region of the skull is pneumatic.

Lower Jurassic

Protosuchus, Orthosuchus, Platyognathus, Dianosuchus, Erythrochampsa, Nothochampsa, Stegmosuchus, Tagarosuchus, Hemiprotosuchus, Gobiosuchus.

MESOEUCROCODYLIAN CLADE

FAMILY TELEOSAURIDAE+

Marine crocodilians with narrow snouts were piscivores of shallow coastal seas. 

Lower to middle Jurassic

FAMILY METRIORHYNCHIDAE+

These animals were wholly aquatic with a tail fin (the end of the tail was down-turned like the ichthyosaurs), the limbs were modified into paddles, and they had no body armor. 

Upper Jurassic

Geosaurus

FAMILY NOTOSUCHIDAE+

These were highly modified to a terrestrial existence.  They were small (a mater or less) and had modified teeth (sharp front teeth and flattened ‘molars’.  Some may even have been herbivorous.

Lower Cretaceous

Argentinosuchus, Chimerasuchus

FAMILY SEBICIDAE+

Carnivores with large skulls, high snouts, and laterally flattened teeth. 

Paleocene to Miocene

Sebecus

NEOSUCHIAN CLADE

These are all heavily armored aquatic reptiles that superficially resemble lizards.  They are all somewhat aquatic.  The teeth are thecodont.  Modern taxa have webbed feet and nostrils on top of the head which can be closed by flaps.  The eyes can be covered by a clear membrane.  They have a muscular partition (similar to a diaphragm), alveoli in the lungs, and a four-chambered heart.

FAMILY GONIOPHOLIDIDAE+

Aquatic crocodilians with long snouts.  Found in marine and freshwater deposits.

Middle Jurassic to upper Cretaceous

Coelosuchus, Dakotasuchus, Microsaurus, Pinacosuchus, Pliogonodon, Polydectes, Kansajsuchus, Oweniasuchus, Symptosuchus, Turanosuchus, Vectisuchus, Goniopholis, Sunosuchus, Calsoyasuchus, Eutretauranosuchus.

Bernissartia, Gilchristosuchus, Rugosuchus.

FAMILY DYROSAURIDAE+

Some of these were very large.  The skull of one large species was 1.8 m long and the estimated weight was 8 tons.

Early Cretaceous through the Eocene

Chenanisuchus, Sokotosuchus, Phosphatosaurus, Dyrosaurus, Arambourgisuchus, Congosaurus, Rhabdognathus, Hyposaurus, Sarcosuchus

EUSUCHIAN CLADE

Unassigned Taxa:  Hylaeochampsa+, Allodaposuchus+, Borealosuchus+, Pristichamphus+..

GAVIALOIDEA

FAMILY GAVIALIDAE

Gavials. Freshwater from Pakistan across northern India , Nepal , Bhutan , Bangladesh and Burma .  They are aquatic fish-eating crocodilians with narrow snouts and very weak limbs.  They inhabit fast-flowing streams. Oviparous.

Eocene to present

Gavialis, Eothoracosaurus+, Thoracosaurus+, Argochampsa+, Eosuchus+, Eogavialis+, Gryposuchus+, Ikanogavialis+, Sisquisiquesuchus+, Piscogavialis+, Hesperogavialis+.

CROCODYLOIDEA

FAMILY CROCODYLIDAE

Crocodiles and False Gavials. Freshwater, brackish water and marine environments; Africa , Madagascar , Iran , India through southeastern Asia to Australia .  Most can be recognized by the fourth tooth of the lower jaw which sits in a notch on the upper jaw and is visible.  Oviparous.

Late Cretaceous to present

Unassigned Taxa: Prodiplocynodon+, Asiatosuchus+, Aigialosuchus+, Charactosuchus+, Dollosuchus+, Holopsisuchus+, Lianghusuchus+, Megadontosuchus+, Necrosuchus+, Planocrania+.

Subfamily Mekosuchinae+: Pallimnarchus, Australosuchus, Kembara, Baru, Trilophosuchus, Quinkana, Volia, Mekosuchus.

Subfamily Crocodylinae: Crocodylus, Mecistops, Osteolaemus, Euthecodon+.

Subfamily Tomistominae: Tomistoma, Cavialosuchus, Kentisuchus+, Gavialosuchus+, Paratomistoma+, Thecachampsa+, Rhamphosuchus+.

ALLIGATOROIDEA

Unassigned Taxa: Leidyosuchus+, Deinosuchus+, Strangerochampsa+, Brachychampsa+, Allodaposuchus+, Dinosuchus+, Balanerodus+, Eoalligator+, Hispanochampsa+, Marrocosuchus+.

FAMILY DIPLOCYNODONTIDAE+

Tadzhikosuchus, Baryphracta, Diplocynodon.

FAMILY ALLIGATORIDAE

Alligators. Freshwater; Southern US to South America and a disjunct species in the Yangtze River .  Oviparous.

Late Cretaceous to present

Listrognathosuchus+, Brachygnathosuchus+.

Subfamily Alligatorinae: Alligator, Akantosuchus+, Albertochampsa+, Chrysochampsa+, Hassiacosuchus+, Navahosuchus+, Ceratosuchus+, Allognathosuchus+, Wannaganosuchus+.

Subfamily Caimaninae: Caiman, Melanosuchus, Paleosuchus, Necrosuchus+, Eocaiman+, Paleosuchus+, Parussaurus+.

FAMILY NETTOSUCHIDAE+

Mourasuchus, Orthogenysuchus, Nettosuchus.

INFRACLASS AVEMETATARSALIA+

SUPERORDER FOR SCLEROMOCHLUS+

This animal was the most basal of the Avemetatarsalia so far known.  That is, it is a sister to the Dinosauria+Pterosauria clade.  It was small (<0.2m) long and had several avian features.  It may have been a climber or a glider.  It might even have been able to hop. 

Upper Triassic

Scleromochlus

SUPERORDER ORNITHODIRA

The ankle joint is simplified to a hinge joint.  The animal stands on its toes in a digitigrade posture.

ORDER PTEROSAURIA

These animals flew almost as a bat with wing membrane stretched by the hand.  However, in this case, only the fourth finger is elongated.  Their bodies seem to have been covered with hair.  They also had five long toes on their feet.

SUBORDER RHAMPHORHYNCHOPHORIDA

These likely are paraphyletic and include the collection of basal taxa.  Mostly fish eaters, they had long tails. 

Upper Triassic to Lower Cretaceous

Eudimorphodon, Ramphorhynchus, Dimorphodon

SUBORDER PTERODACTYLOIDEA

These animals radiated into many ecological niches and some were as large as a small airplane.  In general, they had a reduced tail  and reduced, or highly modified dentition.  Some like Pteranodon lost its teeth all together. 

Lower to upper Cretaceous

Pteranodon, Dsungaripteris, Ornithocheirus, Pterodaustro, Ctenochasma, Quetzalcoatlus

ORDER INCERTAE SEDIS+

LAGERPETON, MARASUCHUS

These taxa appear to be basal dinosaurids and sisters to the rest of the dinosaurians (including birds).  Lagerpeton is known from fragmentary evidence while Marasuchus is more completely known.  Marasuchus was a small carnivore with an s-shaped neck, very long hind limbs and features of the pelvis and leg that suggest it was a fully bipedal animal.  Its long tail also served as a counter balance. 

Middle Triassic

Lagerpeton, Marasuchus

SUPERORDER DINOSAURIA

ORDER SAURISCHIA

The lizard-hipped dinosaurs

SUBORDER UNNAMED

FAMILY HERRERASAURIDAE

Carnivore. Represented by two genera, both bipeds.  They both had a dinosaurian leg, pelvis, ankle, and skull which was relatively narrow and high.  There was a reduction of toes in the fore and hind feet. 

Upper Triassic

Herrerasaurus, Eoraptor

SUBORDER THEROPODA

This includes all of the carnivorous dinosaurs (and technically also the birds)

INFRAORDER COELOPHYSOIDEA

This group had light bodies, long graceful necks and long tails.  It had a skull that was long and narrow.  The ischium was 2/3 or less as long as the pubis.  Well known.  Coelophysis ranges from 0.8-3.1m. 

Upper Triassic to lower Jurassic

Coelophysis

INFRAORDER CERATOSAURIA

Theropods with crests.

FAMILY CERATOSAURIDAE

These animals had crests or horn-like protuberances, which likely were for sexual display. 

Jurassic

Dilophosaurus, Ceratosaurus

FAMILY ABELISAURIDAE

These had very short skulls with expansions of bone to form protuberances. 

Middle to upper Cretaceous

Carnotaurus

INFRAORDER TETANURIDAE

These have the following features:

Large maxillary fenestra; teeth do not extend back of the antorbital fenestra; dorsal vertebrae pleurocoelus

CARNOSAURIA

These are the basal tetanurids, and they all have a distinctive maxilla, opisthocoelus cervical vertebrae, and a very strong first metatarsal.

FAMILY MEGALOSAURIDAE

Skull narrow; orbit was high and smaller than the antorbital fenestra. 

Middle to upper Jurassic

Megalosaurus, Magnosaurus

FAMILY SPINOSAURIDAE

These had spiny sails on their backs, a characteristic large claw on the forefoot, and an elongate alligator-like mouth. 

Cretaceous

Baryonyx, Suchomimus

FAMILY ALLOSAURIDAE

Larger than Megalsaurus, tall narrow orbit with heavy bony crests over them.  11-12m. 

Upper Jurassic

Allosaurus

FAMILY CARCHARODONTOSAURIDAE

These were very large with skulls 1.6m long. 

Cretaceous

Carchodontosaurus

COELOSAURIA (SISTER TO THE AVES)

The coelosaurs share the following characters: enlarged maxillary fenestra, premaxillary teeth without serrations.

FAMILY COELURIDAE

The basal coelosaurians.  These were small (0.7-1.4m) and had feathers. 

Upper Jurassic

Composgnathus, Sinosauropteryx

FAMILY TYRRANOSAURIDAE

Very large predators (scavengers?).  They had an extra joint between dentary and post-dentary elements that increased the gape and perhaps the force of the bite.  They had very small forelimbs with two or three digits.  The ankle was specialized such that all three metatarsals were locked together (the arctometatarsus).  Likely, they could go no faster than 40km/hr. 

Upper Cretaceous

Tyrannosaurus, Albertosaurus

FAMILY ORNITHOMIMIDAE

These animals had an ostrich-like body with long forelimbs, each terminating in three digits.  Early taxa had teeth, but later ones had entirely toothless jaws that were covered with a horny beak.  The beak was ridged on the inside like that of a mallard duck (function?). 

Upper Jurassic to upper Cretaceous.

Struthiomimus, Dromiceiomimus

THE MANIRAPTORA

FAMILY ALVAREZSAURIDAE

These animals were slender with oddly reduced forelimbs (Mononykus had only digit one that was modified into a powerful spike).  They had certain avian featured that included a keeled sternum.  Femur shorter than the tibia and fibula reduced to a spike (as in a chicken leg). 

Upper Cretaceous

Mononykus

FAMILY THERIZINOSAURIDAE

These animals were large (up to 5m) with a heavy pelvis, stocky legs, short tail and relatively long forelimbs with three long digits.  The skull was small and had a few peg-like teeth and a beak at the front.  Likely it was a herbivore (foliage-eaters?).

Upper Cretaceous

Alxasaurus

FAMILY DROMAEOSAURIDAE

These were relatively small animals (up to 3 m) that were characterized by having a specialized claw 2 that could swivel and slash.  Clearly predators, these animals may have hunted in packs. 

Lower Cretaceous

Deinonychus, Velociraptor

FAMILY TROODONTIDAE

These animals had slender necks and a head with very large orbits that faced forward (binocular vision?).  For a dinosaur, its braincase was relatively large.  Adapted as a runner, its middle hind toe (toe 3) was the longest one as in birds. 

Upper Cretaceous

Troodon

OVORAPTOSAURS (SEE AVES)

SUBORDER SAUROPODOMORPHA

INFRAORDER UNNAMED

These had 4 fused sacral vertebrae, a straight femur, no distal tarsals.

BASAL SAUROPOD FAMILY

This early animal shows the features common to the sauropod clade: a small head with a down-turned lower jaw, lanceolate teeth with serrated crowns, long neck with more than 10 cervical vertebrae, large thumb claw and no claws on toes 4&5, a short blade on the ilium.  This is from a single genus, a relatively small (2.5m) bipedal animal.

Upper Jurassic

Thecodontosaurus

FAMILY PLATEOSAURIDAE

The most typical of the prosauropods, the platyosaurids.  These animals could be quadrupedal or bipedal. 

Upper Triassic

Plateosaurus, Riojasaurus, Melanosaurus

FAMILY MASSOSPONDYLIDAE

INFRAORDER SAUROPODA

NEOSAUROPODS

The fenestra on the lower jaw has been lost and the lower temporal fenestra lies beneath the orbit. The legs are pillar-like.  Phalanges are lost

FAMILY CETIOSAURIDAE

These seem to be the precursors to the diplodocids and brachiosaurids, its hind legs still are longer than the front legs. 

Middle Jurassic

Cetiosaurus

FAMILY OMEISAURIDAE

These animals had unusually long necks, even for sauropods.

Upper Jurassic

Mamenchisaurus

FAMILY DIPLODOCIDAE

Long broad snout with long pencil-shaped teeth at the front of the jaws.  Their teeth angled out from the jaw, perhaps serving as a pincer to crop vegetation.  Nostrils are near the top of the head between the eyes.  There has been much debate about the way in which the diplodocids used their necks.  The structure of the neck suggests that it could not flex the neck up into an s-shape, but remained outstretched or raised only slightly.

Upper Jurassic

Diplodocus

MACRONARIA

Enlarged nasal ridge.  Front legs longer than the hind legs.  Contains the most massive terrestrial vertebrates known.

FAMILY CAMARASAURIDAE

They had hollow chambers in the vertebrae and forked neural spines.  The animals had a deep body and relatively long cervical ribs. 

Camarasaurus, Aragosaurus, Lourinhasaurus.

FAMILY BRACHIOSAURIDAE

This animal was a dinosaurian giraffe.  It could flex its neck upwards. 

Upper Jurassic

Brachiosaurus

FAMILY EUHELOPODIDAE

The jaws of these animals suggest that they ate tough vegetation.

Middle Jurassic

Shunosaurus

FAMILY TITANOSAURIDAE

These were huge animals that reached the theoretical limits  of a terrestrial vertebrate.  The skull was similar to that of Diplodocus.  Some had armor plates embedded in their skin.

Lower to upper Cretaceous

Argentinosaurus, Nemegtosaurus, Quesitosaurus

ORDER ORNITHISCHIA

These animals had a pubis that pointed backwards and all were herbivores.

SUBORDER UNNAMED The basal Ornithischia

FAMILY PISANOSAURIDAE

This is known from fragmentary evidence of the dentary and maxilla.  The cheek teeth had a deep root and blunt serrations.  The teeth are set back from the outer jaw suggesting that they had a cheek for chewing. 

Upper Triassic

Pisanosaurus

FAMILY FABROSAURIDAE

These were small animals (less than a meter) with the ornithischian pelvis.  It had typical ornithiscian characters of the skull including a predentary, the tip of which was toothless, in the lower jaw. 

Lower Jurassic

Lesothosaurus

SUBORDER THYREOPHORA

These were the armored ornithiscians.

INFRAORDER UNNAMED

FAMILY SCELIDOSAURIDAE

These were bipedal with rows of keeled scutes on the back and in rows on the sides.

Lower to middle Jurassic

Scelidosaurus, Scutellosaurus

INFRAORDER STEGOSAURIA

They were quadripeds with low, tube-like skulls.  The back had two rows of alternating triangular plates and a tail that terminated in two rows of spikes. 

Upper Jurassic

Stegosaurus

INFRAORDER ANKYLOSAURIA

FAMILY NODOSAURIDAE

These had rows of spiny plates on the back and bony armor over the pelvis.

Middle Jurassic to Early Cretaceous

Polocanthus

FAMILY ANKYLOSAURIDAE

They became heavily armored with armored skulls and bodies.  Their tail terminated in a large bony club. 

Lower to upper Cretaceous

Ankylosaurus, Euoplocephalus

SUBORDER CERAPODA

INFRAORDER PACHYCEPHALOSAURIA

These bipedal animals had unusually thickened skull roofs and likely engaged in head butting like modern sheep.  Also, their skeletons seem to have evolved to dissipate the force of a heavy blow to the top of the head.

Upper Cretaceous

Pachycephalosaurus, Prenocephale, Stegoceras, Homalocephale

INFRAORDER CERATOPSIA

These animals had a beak-like rostral bone, which was covered with a horny beak, at the tip of the upper jaw. 

FAMILY PSITTACOSAURIDAE

These were bipedal with an ornithopod body, but a clearly ceratopsian head. 

Lower Cretaceous

Psittacosaurus

FAMILY PROTOCERATOPSIDAE

These were quadripedal and had a characteristic frill and an incipient horn on the nose.

Middle Cretaceous

Protoceratops

FAMILY CERATOPSIDAE

These had limbs adapted for running and galloping.  They had tall neural spines on the back to which powerful neck muscled were attached.  The taxa varied according to the frill and number and placement of horns.  The frills and horns likely served multiple purposes, among them, species signaling and display, and defense. 

Upper Cretaceous

Triceratops, Centrosaurus, Torosaurus

SUBORDER ORNITHOPODA

FAMILY HETERODONTOSAURIDAE

These animals had differentiated teeth: incisors, canines, and cheek teeth that were inset suggesting a cheek.  The canines seem to have been part of their sexual dimorphism (some specimens, presumably female, lacked them).

Lower Jurassic

Heterodontosaurus

FAMILY HYPSILOPHODONTIDAE

These were similar to the heterodontosaurids, but they lacked tusks and their forelimbs were relatively longer.  They had stiff tails and the proportions of the hind limbs suggest that they were very fast runners.

Upper Jurassic to upper Cretaceous.

Hypsilophodon

FAMILY IGUANODONTIDAE

These animals had a tail stiffened by ossified tendons and forelegs terminating in a “hand” with small hooves and a thumb spike.  They could be bipeds or quadipeds.

Lower Cretaceous

Iguanodon, Ouranosaurus

FAMILY HADROSAURIDAE

These were the duck-billed dinosaurs.  Their jaws were toothless in the front which was sheathed by a horny beak.  Their cheek teeth grew in batteries of 5-6 rows with 45-60 teeth each, clear adaptations to chewing tough plant material.  Some have a variety of crests, likely for display (visual and sound).

Upper Cretaceous

Edmontosaurus, Kritosaurus, Anatosaurus, Brachylophosaurus, Saurolophus, Parasauroolophus, Lambeosaurus, Corythosaurus.

SUBCLASS AVES

INFRACLASS UNNAMED

These animals had long bony tails and teeth.

ORDER UNNAMED

FAMILY ARCHAEOPTERYGIDAE

In most details like a small theropod.  In addition, they had at least 3 of the 5 air sacs found in birds.  The pubis seems to have been vertical (also seen in some theropods).  Its toe 1 is reflexed to lie at the back of the foot (reflexed hallux).  No evidence of a beak.

Upper Jurassic

Archaeopteryx

RAHONAVIS

This bird appears to be basal in structure but not in time.  It was a flyer with an enlarged 2nd toe.  Likely a member of the Archaeopterygidae but with some advanced features. 

Upper Cretaceous

Rahonavis

JEHOLORNIS

This, too appears to be close to Archaeopteryx, but it shows significantly advanced hand.

INFRACLASS PYGOSTYLIA

Animals with a tail reduced to a pygostyle.

SUPERORDER UNNAMED+

ORDER UNNAMED

FAMILY CONFUCIUSORNITHIDAE

They have no teeth, but do have a horny beak.  The sacrum is fused to form the synsacrum characteristic of modern birds and the tail has been reduced to 7 fused caudal vertebrae. 

Lower Cretaceous

Confuciusornis, Changchengornis

ORDER OVORAPTOROSAURIA

These animals are tentatively placed in the birds.  They were flightless, had a highly modified toothless skull, and a pygostyle. 

Upper Cretaceous

Ovoraptor

SUPERORDER ORNITHOTHORACES+

ORDER ENANTIORNITHES

Birds with teeth and short backs (<13 thoracic vertebrae) and other features of modern birds.  Characterized by the presence of a tarsometatarsus (fused distal tarsals and metatarsals). 

Lower to upper Cretaceous

Gobipteryx, Sinornis, Otogornis, Bolouchia, Liaoxiornis, Longipteryx, Iberomesornis, Concornis, Eoalulavis

ORDER INCERTAE SEDIS

PATAGONOPTERYX

This was a flightless bird with adaptations for running, an ecological equivalent to the ratites (but not directly related to them). 

Upper Cretaceous

Patagonopteryx

VORONA

Known only from a hind leg.  This animal showed more complete fusion of the metatarsals than the eantionithes. 

Upper Cretaceous

Vorona

ORDER HESPERORNITHIFORMES

These were toothed flightless diving birds whose wings were almost entirely reduced. 

Upper Cretaceous

Hesperornis, Baptornis

ORDER ICHTHYORNIFORMES

These were diving birds that were good fliers.  They had large beaks lined with small sharp teeth.  Likely, the fished as terns do today. 

Upper Cretaceous

Ichthyornis

Other upper Cretaceous Ornithurine birds of uncertain affinities include: Ambiortus, Gansus , Apsaravis, Limenavis

SUPERORDER NEORNITHES

PALAEOGNATHAE

These birds have a characteristic palate (extensive vomer-pterygoid joint)

ORDER LITHORNITHIFORMES+

Variable in size (hen to crane-sized).  Some could fly, others were flightless. 

Paleocene to Eocene

Lithornis, Palaeotis

RATITES

These are represented by the tinnamous and ostriches-rheas-cassowaries-emus.  They have become complete ground-dwelling animals and have lost the sternal keel.

ORDER STRUTHIONIFORMES

Typically, they are large to very large flightless birds that are adapted to running.  They have a southern continental distribution.

FAMILY APTERYGIDAE

Kiwis.  Flightless birds of forests and open woodlands of New Zealand .  Kiwis are medium-sized birds (.35-.65m high).  They are unusual for many reasons.  Their wings are vestigial, the plumage is hair-like, they are nocturnal, and the nostrils occur at the tips of their beaks.  They are not adapted to running; their legs are relatively short and stout.

Apteryx.

FAMILY CASUARIIDAE

Cassowaries.  Flightless birds of the rainforests and savannah woodlands of Papua-New Guinea and northern Australia .  Cassowaries are very large (1-1.7m high).  The wings are highly reduced, almost vestigial.  The legs are powerful and terminate in three-toed feet.  They can run at speeds of 50km/hr and jump 1.5m high.  Their plumage is almost like coarse hair.  They have a distinctive crest on the tops of their heads

Casuarius.

FAMILY DINORNITHIDAE+

Moas.

Anomalopteryx, Dinornis, Emeus, Euryapteryx, Megalapteryx, Pachyornis.

FAMILY DROMAIIDAE

Emus.  Flightless birds of the savannahs and semi-arid areas of Australia .  Emus are very large (1.5-1.9m high).  They have reduced wings and long necks.  They have powerful legs and feet with three toes that allow them to reach speeds of 48km/hr.  Reduced now to a single species.

Pleistocene to the present.

Dromaius.

FAMILY RHEIDAE

Rheas.  Flightless birds of the pampas and scrub lands of tropical and temperate South America .  Rheas are very large (0.9-1.5m).  They have reduced wings and long necks.  Their legs are powerful and adapted for running (>60km/hr).  The feet have three toes.

Pterocnemia, Rhea.

FAMILY STRUTHIONIDAE

Ostriches.  Flightless birds of savannas and semi-arid regions of Africa .  Ostriches are very large 1.75-1.75m high.  Its powerful legs are adapted for high speed running (>50-70km/hr) and feet with only two toes, one of which has a large claw.  They have a long neck and reduced wings.

Struthio.

ORDER TINAMIFORMES

FAMILY TINAMIDAE

Tinamous.  They are weak fliers of forests and savannah of most of Souh America and southern Central America .  The are compact birds with reduced tails and strong legs.  They stand 0.15-0.50m high.

Miocene to the present

Subfamily Rhynchotinae (Steppe Tinamous): Eudromia, Nothoprocta, Nothura, Rhynchotus, Taoniscus, Tinamotis.

Subfamily Tinaminae ( Forest Tinamous): Crypturellus, Nothocercus, Tinamus.

NEOGNATHAE

The largest group of extant birds.  They also have a characteristic ankle in which the calcaneum, not the astragulus (as in the ratites and theropods) forms the ascending process on the tibia.

GALLOANSERAE Ducks and Fowl make up a clade of birds that is the sister group of all other neognaths.  They have a fossil record that may go back to the upper Cretaceous.  Diatryma was large flightless bird with a large beak (a predator?).

ORDER ANSERIFORMES

FAMILY ANATIDAE

Swans, Geese, and Ducks.  Aquatic birds of freshwater, brackish water, and marine environments of all continents, oceanic islands, and continental margins.  They are medium to moderatedly large birds (0.30-1.80m) with wingspans of 0.43 to 2m.  They have flattened bills that generally are adapted to filter feeding.  They have long necks, large bodies, and short tails.  Their diets include plants, fish, animal foods like fish eggs and invertebrates.  Many are migratory.

Subfamily Dendrocyninae (Whistling Ducks): Dendrocygna.

Subfamily Anserinae (Swans and Geese): Cyngus, Anser, Branta, Tadorna, Plectopteris.

SubfamilyAnatinae (True Ducks): Alopochen, Nettapus, Aix, , Anas, Marmaronetta, Netta, Aythya, Somateria, Polysticta, Histrionicus, Melanitta, Bucephala, Clangula, Mergus, Oxyura.

FAMILY ANHIMIDAE

Screamers.

Anhima, Chauna.

FAMILY ANSERANATIDAE

Magpie Geese.

Anseranas.

ORDER GALLIFORMES fowl

FAMILY PHASIANIDAE

Pheasants, Partridges, Quails.  Terrestrial birds of forest and open habitats through Eurasia , Australia , and Africa .  They range from small to large birds (0.14-2.5m).  Typically, they have stout bodies, short legs, a stout beak, and rounded wings.  They can have tails that are short or long and modified for display.  Similarly, they can be drab or brightly colored.  They feed mainly on the ground where they can take a wide variety of plants, invertebrates, and small vertebrates.  Usually not migratory.

Lower Eocene to the present. 

Subfamily Perdicinae (Partridges): Alectoris, Ammoperdix, Aneurophasis, Arborophilia, Bambusicola, Caloperdix, Coturnix, Francolinus, Galloperdix, Haematortyx, Lerwa, Margaroperdix, Melanoperdix, Ophrysia, Perdicula, Perdix, Ptilopachus, Rhizothera, Rollulus, Tetraogallus Tetraophasis, Xenoperdix

Subfamily Phasianinae (Pheasants): Afropavo, Argusianus, Catreus, Chrysolophus, Crossoptilon, Gallus, Ithiginis, Lophophorus, Lophura, Phasianus, Pavo, Polyplectron, Pucrasia, Rheinardia, Syrmaticus, Tragopan.

FAMILY MEAGRIDIDAE (SOME SYSTEMS PLACE THESE INTO THE PHASIANIDAE)

Turkeys .  These birds occur in open woodland habitats from northern Mexico to southern Canada .  Turkeys are moderately large birds (0.71-1.10m).  Their heads and necks are naked.  They have strong legs and usually prefer to run, but they can fly for short distances, followed by a long glide.  Turkeys are omnivores.  The males are quite distinctive with their large fan-like tail, waddle, and beard.  Not migratory.

Miocene to the present.

Meleagris.

FAMILY TETRAONIDAE (SOME SYSTEMS PLACE THESE INTO THE PHASIANIDAE)

Grouse.  Terrestrial birds of forests in the cold temperate and subarctic habitats of the Northern Hemisphere.  Grouse have compact bodies and medium-sized to moderately large birds (0.30-0.90m) with short legs.  They generally are terrestrial but can burst into a short flight.  Not migratory.

Bonasa, Centrocercus, Dendragapus, Falcipennis, Lagopus, Tetrao, Tympanuchus.

FAMILY ODONTOPHORIDAE

New World Quails.  Terrestrial birds of forests and open habitats from the northern US to northern and central South America .  Quails are small to medium-sized birds (0.17-0.37m).  They have compact bodies and short, strong legs and short wings.  They are generally herbovores.

Lower Oligocene to the present.

Callipepla, Colinus, Cyrtonyx, Dactylortyx, Dendrortyx, Odontophorus, Oreortyx, Philortyx, Rhynchortyx.

FAMILY NUMIDIDAE

Guinea Fowls.  Birds of raiforest to semi-arid habitats in Africa .  Guinea Fowl are medium-sized birds (0.40-0.72m).  They are characterized by having a rounded body, long neck and small head.  Many have a crest on the head and have large naked patches on the head and neck.  They are omnivores. 

Acryllium, Agelastes, Guttera, Numida.

FAMILT CRACIDAE

Chachalacas, Curassows, and Guans.  Arboreal birds of tropical America (South America north to southern Texas ).  They are medium-sized to moderately large birds (0.43-090m).  They are adapted to perching, and thus have an elongate hind toe.  They are generally herbivores and territorial.

Middle Eocene to the present.

Subfamily Cracinae (Curassows): Crax, Mitu, Nothocrax, Pauxi.

Subfamily Penelopinae: Aburria, Chamaepetes, Oreophasis, Ortalis, Penelope, Penelopina, Pipile.

FAMILY MEGAPODIIDAE

Megapodes.  Birds of tropical forests and semi-arid woodlands in Australia , Papua New Guinea , Philippines , and other islands east of the Wallace Line.  One occurs in the Orient.  Megapodes are medium-sized birds (0.28-0.70m) with naked heads and necks.  They have powerful legs and sharp claws with which they construct remarkable incubation mounds in which the eggs are incubated by the heat from rotting vegetation.

Aepypodius, Alectura, Eulipoa, Leipoa, Macrocephalon, Megapodius, Talegalla.

FAMILY TURNICIDAE

Button Quails

Turnix

‘WATERBIRD ASSEMBLAGE’ The waterbirds make up another large clade of birds. 

GRUIFORM-RALLIFORM CLADE

Eocene to present. 

ORDER GRUIFORMES

FAMILY EURYPYGIDAE

Sunbittern.

Eurypyga.

FAMILY HELIORNITHIDAE

Finfoot.

Heliopais, Heliornis, Podica.

FAMILY ARAMIDAE

Limpkin.  Wading bird found in freshwater wetlands of temperate South America to Georgia .  Limpkins are medium-sized birds (0.56-0.70m) that resemble ibises with straight bills.  They have a wingspan of about 1m.  They are secretive brds with long legs, long neck, and long drooping beak.  They feed mainly on aquatic mollusks, but will eath other invertebrates and small vertebrates.  They do not migrate.

Aramus.

FAMILY CARIAMIDAE

Seriemas.

Cariama, Chunga.

FAMILY GRUIDAE

Cranes.  Wading birds of all continents except South America and Antarctica.  They are moderately large to large (0.90-1.76m) and have wingspans of 1.8-2.2m.  They have long legs and long necks.  Most of them have elaborate displays.  Cranes are opportunistic omnivores.  Some species are migratory.

Subfamily Balearicinae: Balearica.

Subfamily Gruinae: Anthropoides, Bugeranu, Grus.

FAMILY PSOPHIIDAE

Trumpeters.

Psophia.

ORDER RALLIFORMES

FAMILY RALLIDAE

Rails, Crakes, and Coots.  Birds of wetlands, but aquatic environments, or terrestrial habitats and with global distribution; absent only from the Arctic and the Antarctic.  Small to medium-sized birds (0.13-0.63m).  Rails are shy birds with short tails, short wings, and laterally-compressed bodies.  Coots have more rounded bodies and tend to be more aquatic.  They