| SYSTEMATIC BIOLOGY | |
| CLASS BATRACHOMORPHA |
Batrachomorpha (ba-TRAK-o-MORF-a) comes from two Greek roots meaning "frog form" [frog- batrachos (βάτραχος) and form- morphi (μορφή)]. The reference is to the frog-like nature of the animals in this class.
The batrachomorphs, also called the amphibians, are made of two major groups: the Temnospondyli (from two Greek roots meaning "cut vertebrae") and the Lissamphibia (from two Greek roots meaning "smooth amphibians"). All of the extant amphibians are carnivores or insectivores. The temnospondyls were amphibious with gilled larvae and adults that were aquatic, semi-terrestrial or fully terrestrial. Some evolved to become sexually mature as larvae. They had a long evolutionary history from the Pennsylvanian to the lower Cretaceous. One of the most important taxa was Eryops, a semi-aquatic to terrestrial predator of the lower Permian. It had a sprawling stance, was 1.8-2 m long, and weighed nearly 100 kg.
All of the living amphibians are in the Lissamphibia, which includes three major groups: Gymnophiona (from two Greek roots meaning "naked snake") commonly called the caecilians, Urodela (from a Latin and a Greek root meaning "visible tail"), and the Anura (from a Greek and a Latin root meaning "no tail"). The caecilians are blind, legless, worm-like amphibians that live a fossorial or aquatic lifestyle. Their annular scales make them appear to be segmented like giant earthworms. Even though they are worm or snake-like, they have tails that are short or absent with a very elongate trunk (up to 200 vertebrae).
Urodela (also called the Caudata) are the salamanders and newts. They have four sprawling legs, a short body, and a long tail, though some taxa have lost their limbs entirely. Salamanders undergo a metamorphosis in which the larvae have external gills but otherwise resemble the adults. However, some purely aquatic taxa (e.g. Necturus) retain larval characters as adults. The largest salamanders are the Hellbenders (Cryptobranchus,), which are nearly 2 meters long, are lungless and gilless, but respire entirely through the skin in cold well-oxygenated upland streams.
The Anura (also known as Salientia) are the frogs and toads. These animals are tailless as adults (Anura is from two Greek roots meaning without tails) and a highly modified axial skeleton that provides for rigid support of the torso. The shortened forelegs and elongate, muscular hind legs are adapted to leaping. Most exhibit complete metamorphosis; however, a few groups have direct development. Frogs and toads are among the most successful vertebrates with about 4,000 described species worldwide.
| HIERARCHICAL TAXONOMY OF THE CLASS BATRACHOMORPHA. 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. |
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Image of Eryops, an extinct temnospondyl (left) and Xenopus, the extant African Clawed Frog. |
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Image of Eryops from: http://www.bertsgeschiedenissite.nl/geschiedenis%20aarde/amfibieen.htm Image of Xenopus from the Systematic Biology Biodiversity Archive. CLASS BATRACHOMORPHA (AMPHIBIA) Benton (2004) uses the term Batrachomorpha to define a monophyletic group that includes the extinct Temnospondyls and the living amphibians (Lissamphibia: caecilians, frogs, and salamanders). These groups are united by several synapomorphies including four digits on the forefeet, a flattened skull with fused skull elements, stalked or pedicillate teeth, 26 or fewer presacral vertebrae, large orbits, paired occipital condyles, and the maxilla articulates with the quadratojugal. They are aquatic or tied to an aquatic environment through reproduction. Most lay eggs in water and require water for an aquatic larval stage. Skull frog-like, triangular to parabolic with polygonal pattern. Many had a lateral line. Some known to have gilled larvae (tadpoles). A large opening in the palate and several large fangs attached to the palate, as well. Though most were adapted to an aquatic existence, the later taxa had several lines that were primarily terrestrial. They are characterized by having complex, primitive vertebrae made of separate unfused components that include the pleurocentra, intercentra, neural arches, and zygopophyses. Some evolved a weaker (more aquatic) structure in which the pleurocentra were lost or highly reduced. Carboniferous
to Triassic (lower Cretaceous). ORDER
UNNAMED FAMILY
TRIMERORACHIDAE THE DVINOSAURIA Broad skulls and lateral line. Permian to Triassic FAMILY ARCHEGOSAURIDAE FAMILY RHINESUCHIDAE Tupilakosaurus THE CAPITOSAURIA Broad flat skulls of animals that were semi-aquatic to fully aquatic and common denizens of freshwater environments. Some very large (3-4m). Triassic FAMILY
MASTODONSAURIDAE FAMILY
CAPITOSAURIDAE Benthosuchus THE
TREMATOSAURIA Mostly
aquatic and mostly restricted to the Triassic FAMILY
TREMATOSAURIDAE FAMILY
METAPOSAURIDAE FAMILY
PLAGIOSAURIDAE Short
broad skulls and reduced limbs, clearly aquatic. Triassic Plagioscutum FAMILY
RHYTIDOSTEIDAE FAMILY
BRACHYOPIDAE To
the upper Jurassic FAMILY
CHIGUTISAURIDAE To
the lower Cretaceous Koolasuchus THE
TEMNOSPONDYLS FAMILY
DENDRERPETONTIDAE Large
skulls with rounded anterior. Well
developed middle ear apparatus suggests a terrestrial existence. Carboniferous Balanerpeton,
Denderpeton FAMILY
ERYOPIDAE Massive
skeleton, terrestrial form. Lower
Permian Eryops FAMILY
DISSOROPHIDAE Animals
likely terrestrial; short skulls with very large eyes and large ear drums. Lower
Permian Doleserpeton FAMILY
BRANCHIOSAURIDAE These
seem to have become paedomorphic, sexually mature as larvae. Upper
Carboniferous (Pennsylvanian) to Lower Permian Branchiosaurus FAMILY
AMPHIBAMIDAE These animals have teeth that are bicuspid and pedicillate. They are sisters to the temnospondils or part of the same clade. Pough et al. (1998) summarize evidence that suggests the Lissamphibia arose as paedomorphic temnospondyls. Evidence cited refers to similarities between larval temnospondyls and lissamphibia relative to reductions in skull elements, similar kinds of teeth, and large eyes. ORDER
UNNAMED?+ FAMILY
ALBANERPETONTIDAE The
body was long and salamander-like. The
tail was short and mostly unusually thin. Also,
their neck vertebrae were reduced to an atlas and axis, which were fused to the
first thoracic vertebrae. These
likely were features that were adaptations to burrowing. Middle
Jurassic to Miocene Albanerpeton,
Celtedens The caecilians look like earthworms (also called apoda). They are legless and burrow or swim (tropical taxa). Their vertebral column may have more than 200 vertebrae. Lower Jurassic to present Eocaecilia+ FAMILY RHINATREMATIDAE American
Tailed Caecilians. Wet terrestrial
environments (seeps, wet leaf litter, underground) of northern Epicrionops,
Rhinatrema. FAMILY ICHTHYOPHIIDAE Asiatic Tailed
Caecilians. Semi-aquatic animals of
tropical Caudacaecilia,
Ichthyophis. FAMILY URAEOTYPHLIDAE Indian
Caecilians. Terrestrial animals of Uraeotyphlus. FAMILY SCOLEOMORPHIDAE African
Caecilians. Fossorial animals of
Camaroon and Crotaphatrema,
Scoleocomorphus. FAMILY CAECILIIDAE Caecilians.
Mainly terrestrial animals of tropical Boulengerula,
Brasilotyphlus, Caecilia, Dermophis, Gegeneophis, Geotrypetes, Gymnopis, Herpele,
Hypogeophis, Idiocranium, Indotyphlus, Luetkenotyphlus, Microcaecilia,
Mimosiphonops, Oscaecilia, Parvicaecilia, Prasilinia, Schistometopum, Siphonops,
Sylvacaecilia. FAMILY TYPHIONECTIDAE Aquatic
Caecilians. Freshwater (secondarily
aquatic) in northern Atretochoana,
Chthonerpeton, Nectocaecilia, Potomotyphlus, Typhlonectes. ORDER URODELA (CAUDATA) The salamanders and newts (also called caudata) show few specialized features relative to the non-amniote tetrapods. Living salamanders have four legs, a short body, and a long tail, though some taxa have lost their limbs entirely. They do not have middle ear cavities or external ears (they hear by means of a structure called the opercular apparatus). Salamanders range in size from 3cm to 2m. Middle Jurassic to the present Karaurus+ (Jurassic of FAMILY SIRENIDAE Sirens.
Freshwater and fully aquatic; in swamps, lakes, and marshes of the
southeastern US to northern Siren,
Pseudobranchus. FAMILY CRYPTOBRANCHIDAE Hellbenders.
Freshwater; cold mountain or rocky streams; disjunct distribution Andrias,
Cryptobranchus. FAMILY HYNOBIIDAE Asiatic
Salamanders. Freshwater and
terrestrial; mountain streams and ponds from the Batrachuperus,
Hynobius, Onychodactylus, Ranodon. FAMILY AMPHIUMIDAE Amphiumas or
Cretaceous to the present Amphiuma,
Proamphiuma+. FAMILY PLETHODONTIDAE Lungless
Salamanders. They occur from the
northeastern and northwaestern Miocene to present Aneides,
Batrachoseps, Bolitoglossa, Chiropterotriton, Dendrotriton, Desmognathus,
Ensatina, Gyrinophylus, Haideotriton, Hemidactylium, Hydromantes, Ixalotriton,
Leurognathus, Lineatriton, Nototriton, Nyctanolis, Oedipina, Parvimolge,
Phaeognathus, Plethodon, Pseudoeurycea, Pseudotriton, Stereochilus, Thorius,
Typhlotriton. FAMILY RHYACOTRITONIDAE Torrent Salamanders. They are found in fast-moving streams of the northwestern US, particularly in the Cascades. They unusual squared glands near the vent. They have no operculum, and very reduced lungs. They have complete metamorphosis (but the larval development is very slow, up to 3 years). Rhyacotriton. FAMILY PROTEIDAE Mudpuppies
and Waterdogs. Two species of
aquatic salamanders: one in northeastern US and the other in limestone caves of
southeastern Miocene to the present Necturus,
Proteus, Mioproteus+, Orthophyla+. FAMILY SALIMANDRIDAE Newts and
Salamanders. These animals occur in
four disjunct locations: Eocene to present Chioglossa,
Cynops, Euproctus, Mertensiella, Neurergus, Notophthalmus, Pachytriton,
Paramesotriton, Pleurodeles, Salamandra, Taricha, Triturus, Tylototriton. FAMILY AMBYSTOMATIDAE Mole
Salamanders. Burrowers, they occur
from southeastern Upper Pliocene to the present. Ambystoma,
Amphitriton+. FAMILY DICAMPTODONTIDAE Pacific
Giant Salamanders. They occur in
conifer forests of the Paleocene to the present. Dicamptodon. The frogs and toads (also called salientia) are highly modified to a jumping locomotion. They have lost their tails and have reduced the ribs and vertebrae. Lower Jurassic to present FAMILY ASCAPHIDAE Tailess
frogs. Frogs of cold mountain
streams of Jurassic to
the present (Paragonia, Ascaphus. FAMILY LEIOPELMATIDAE Jurassic to the present. Leiopelma. FAMILY BOMBINATORIDAE FAMILY DISCOGLOSSIDAE Fire Belly
Frogs. These are small toads of Upper Jurassic to the present Alytes, Barbourula, Bombina, Discoglossus. FAMILY PELOBATIDAE Spadefoot
Toads. These fossorial toads occur
in dry areas of Oligocene to the present. Ophryophryne,
Pelobates, Scaphiopus, Scutiger, Spea. FAMILY PELODYTIDAE Parsley
Frogs. Fossorial frogs from western Europe and the Eocene to the present. Pelodytes,
Miopelodytes+, Propelodytes+. FAMILY RHINOPHRYNIDAE Burrowing
Toads. Fossorial toad that occurs
from southern Paleocene to the present. Rhinophrynus FAMILY PIPIDAE Tongueless
Frogs. Aquatic frogs of tropical Lower Cretaceous to the present. Hymenochirus,
Pipa, Pseudohymenochirus, Xenopus/ FAMILY ALLOPHRYNIDAE FAMILY BRACHYCEPHALIDAE Gold Frogs
and Shield Toads. Frogs of the humid
forests of southeastern Brachycephalus,
Psyllophryne. FAMILY BUFONIDAE True Toads.
Toads are found throughout the temperate and tropical areas of the world except Upper Paleocene to present Ansonia,
Atelopus, Bufo, Capensibufo, Crepidophryne, Dendrophryniscus, Didynamipus,
Frostius, Laurentophryne, Leptophryne, Melanophryniscus, Mertensophryne,
Nectophryne, Nectophrynoides, Nimbaphrynoides, Oreophrynella, Osornophryne,
Pedostibes, Pelophryne, Peltophryne, Pseudobufo, Rhamphophryne, Werneria,
Wolterstorffina. FAMILY HELEOPHRYNIDAE Ghost Frogs.
Frogs of mountain streams in southern No fossil record. Heleophryne. FAMILY LEPTODACTYLIDAE (INCERTAE SEDIS) Neotropical
Frogs. Mainly, these are frogs that
occur in the tropics of the Common in
Cenozoic of Adenomera,
Alsodes, Atelognathus, Barycholos, Batrachophrynus, Batrachyla, Caudiverbera,
Ceratophrys, Chaophrys, Crossodactylodes, Crossodactylus, Cyclorhamphus,
Dischidodactylus, Edalorhina, Eleutherodactylus, Euparkerella, Eupsophus,
Geobatrachus, Holoaden, Hydrolaetare, Hylactophryne, Hylodes, Hylorina,
Ishnocnema, Lepidobatrachus, Leptodactylus, Limnomedusa, Lithodytes,
Macrogenioglottus, Megaelosia, Odontophrynus, Paratelmatobius, Phrynocerus,
Phrynopus, Physalaemus, Pleurodema, Proceratophrys, Pseudopaludicola,
Scythrophrys, Somuncuria, Syrrhophus, Telmatobius, Telmatobufo, Thoropa,
Vanzolinius, Zachaenus. FAMILY MYOBATRACHIDAE Australian
Frogs. Frogs of Australia with a few
taxa in Miocene to te present Adelotus,
Arenophryne, Assa, Crinia, Heleioporus, Kyarranus, Lechriodus, Limnodynastes,
Megistolotis, Mixophynes, Myobatrachus, Neobatrachus, Notaden, Philoria,
Pseudophryne, Rheobatrachus, Taudactylus,Uperoleia. FAMILY RHINODERMATIDAE Darwin’s
Frogs. Frogs of the temperate
forests of southern No fossils are known. Rhinoderma. FAMILY HYLIDAE Miocene to the present Acris,
Allophryne, Anotheca, Aparasphenodon, Corythomantis, Cryptobatrachus, Cyclorana,
Duellmanohyla, Flectonotus, Gastrotheca, Hemiphractis, Hyla, Nyctimantis,
Osteocephalus, Osteopilus, Phrynohyas, Phyllodytes, Plectrohyla, Pseudacris,
Pternohyla, Ptychohyla, Scinax, Smilisca, Sphaenorhynchus, Stephania,
Trachycephalus, Triprion. Agalychnis,
Hylomantis, Pachymedusa, Phrynomedusa, Phyllomedusa. FAMILY MEGOPHRYIDAE South Asian
Frogs. Generally frogs of the forest
floor from west of the Wallace Line to No fossil record. Leptobrachella,
Leptobrachium, Megophrys, Vibrissaphora. FAMILY HYPEROLIIDAE African Tree
Frogs. Mainly arboreal frogs of
sub-Saharan No fossil record. Acanthixalus,
Afrixalus, Chrysobatrachus, Heterixalus, Hyperolius, Kassina, Leptopelis,
Opistholhylax, Phlyctimantis, Tachycnemis. FAMILY PSEUDIDAE Paradox
Frogs. Aquatic frogs of tropical
lowlands east of the No fossils are known. Lysapsus,
Pseudis. FAMILY CENTROLENIDAE Glass Frogs
and Leaf Frogs. Arboreal frogs of
the humid tropics of Central and Centrolene,
Centrolenella, Cochranella, Hyalinobatrachium. FAMILY MICROHYLIDAE Microhylid
Frogs. Frogs with a wide
distribution in the tropics and temperate zones of the world.
They are absent from most of Miocene to the present Anodonthyla,
Arcovomer, Asterophrys, Baragenys, Breviceps, Calluella, Callulina, Chaperina,
Chiasmocleis, Cophixalus, Cophyla, Ctenophryne, Dasypops, Dermatonotus,
Dyscophus, Elachistocleis, Gastrophryne, Gastrophrynoides, Gastrophyrne,
Glyphoglossus, Hamptophryne, Hoplophryne, Hylophorbus, Hyophryne, Kalophrynus,
Kaloula, Melanobatrachus, Metaphrynella, Microhyla, Myersiella, Nelsonophryne,
Oreophryne, Otophryne, Parhoplophryne, Phrynella, Phrynomantis, Platypelis,
Plethodontohyla, Probreviceps,
Ramanella, Relictivomer, Rhombophryne, Scaphiophryne, Spelaeophryne,
Sphenophryne, Stereocyclops, Stumpffia, Synapturanus, Uperodon, Xenobatrachus.
FAMILY DENDROBATIDAE Poison Dart
Frogs. These brightly-colored
arboreal frogs are found from Nicaraugua to Colostethus,
Dendrobates, Mannophryne, Phyllobates. FAMILY HEMISOTIDAE Shovel-Nosed
Frogs. Burrowing frogs of tropical
and subtropical savannah and scrub forests No known fossil history. Hemisus. FAMILY ARTHROLEPTIDAE Schreeching
Frogs. Rainforests of sub-Saharan Arthroleptis,
Astylosternus, Cardioglossa, Leptodactylodon, Scotobleps, Trichobatrachus. FAMILY SOOGLOSSIDAE Seychelle
Frogs. These small terrestrial frogs
occur on only 2 small islands of the Nesomantis,
Sooglossus. FAMILY RANIDAE Riparian
Frogs and True Frogs. These frogs
occur throughout the tropics and temperate zones of all continents except for
most of Miocene to the present. Amolops,
Anhydrophryne, Arthroleptella, Arthroleptides, Batrachylodes, Cacosternum,
Ceratobatrachus, Conraua, Dendrobatorana, Dimorphognathus, Discodeles,
Elachyglossa, Hildebrandtia, Hoplobatrachus, Micrixalus, Microbatrachella,
Nannophrys, Nanorana, Natalobatrachus, Nothophryne, Nyctibatrachus, Occidozyga,
Palmatorappia, Petropedetes, Phrynobatrachus, Phrynodon, Platymantis,
Pyxicephalus, Rana, Staurois. FAMILY RHACOPHORIDAE No fossils are known. Boophis,
Callixalus, Chiromantis, Cryptothylax, Mantella, Mantidactylus, Philautus,
Rhacophorus, Theloderma. Prosalirus+, Triadobatrachus+ |