cribrosa'', a
reef-forming productid brachiopod (Middle Permian, Glass Mountains, Texas)
Marine biota Permian marine deposits are rich in
fossil mollusks,
brachiopods, and
echinoderms. Brachiopods were highly diverse during the Permian. The extinct order
Productida was the predominant group of Permian brachiopods, accounting for up to about half of all Permian brachiopod genera. Brachiopods also served as important ecosystem engineers in Permian reef complexes. Amongst
ammonoids,
Goniatitida were a major group during the Early-Mid Permian, but declined during the Late Permian. Members of the order
Prolecanitida were less diverse. The
Ceratitida originated from the family
Daraelitidae within Prolecanitida during the mid-Permian, and extensively diversified during the Late Permian. Only three families of
trilobite are known from the Permian,
Proetidae, Brachymetopidae and
Phillipsiidae. Diversity, origination and extinction rates during the Early Permian were low. Trilobites underwent a diversification during the Kungurian-Wordian, the last in their evolutionary history, before declining during the Late Permian. By the Changhsingian, only a handful (4–6) genera remained. Corals exhibited a decline in diversity over the course of the Middle and Late Permian.
Terrestrial biota Terrestrial life in the Permian included diverse plants,
fungi,
arthropods, and various types of
tetrapods. The period saw a massive desert covering the interior of
Pangaea. The warm zone spread in the northern hemisphere, where extensive dry desert appeared. The rocks formed at that time were stained red by iron oxides, the result of intense heating by the sun of a surface devoid of vegetation cover. A number of older types of plants and animals died out or became marginal elements. The Permian began with the Carboniferous flora still flourishing. About the middle of the Permian a major transition in vegetation began. The
swamp-loving
lycopod trees of the Carboniferous, such as
Lepidodendron and
Sigillaria, were progressively replaced in the continental interior by the more advanced
seed ferns and early
conifers as a result of the
Carboniferous rainforest collapse. At the close of the Permian, lycopod and
equisete swamps reminiscent of Carboniferous flora survived only in
Cathaysia, a series of equatorial islands in the
Paleo-Tethys Ocean that later would become
South China. The Permian saw the radiation of many important conifer groups, including the ancestors of many present-day families. Rich forests were present in many areas, with a diverse mix of plant groups. The southern continent saw extensive seed fern forests of the
Glossopteris flora. Oxygen levels were probably high there. The
ginkgos and
cycads also appeared during this period.
Insects '', a
permocupedid beetle from the Middle Permian of Russia Insects, which had first appeared and become abundant during the preceding Carboniferous, experienced a dramatic increase in diversification during the Early Permian. Towards the end of the Permian, there was a substantial drop in both origination and extinction rates. By the start of the Permian, there was already an active coevolutionary arms race between insects and plant reproductive structures, evidenced by both insect-caused damage in plants and defensive structures in plants aimed at minimising predation by insects. The dominant insects during the Permian Period were early representatives of
Paleoptera,
Polyneoptera, and
Paraneoptera.
Palaeodictyopteroidea, which had represented the dominant group of insects during the Carboniferous, declined during the Permian. This is likely due to
competition by
Hemiptera, due to their similar mouthparts and therefore ecology. Primitive relatives of
damselflies and
dragonflies (
Meganisoptera), which include the largest flying insects of all time, also declined during the Permian.
Holometabola, the largest group of modern insects, also diversified during this time.
Mecoptera (sometimes known as scorpionflies) first appeared during the Early Permian, going on to become diverse during the Late Permian. Some Permian mecopterans, like
Mesopsychidae have long proboscis that suggest they may have pollinated gymnosperms. The earliest known
beetles appeared at the beginning of the Permian. Early beetles such as members of
Permocupedidae were likely
xylophagous, feeding on decaying wood. Several lineages such as Schizophoridae expanded into aquatic habitats by the Late Permian. Members of the modern orders
Archostemata and
Adephaga are known from the Late Permian. Complex wood boring traces found in the Late Permian of China suggest that members of
Polyphaga, the most diverse group of modern beetles, were also present by the Late Permian.
Tetrapods '', a
weigeltisaurid from the Late Permian of Europe. Weigeltisaurids represent the oldest known gliding vertebrates. The terrestrial fossil record of the Permian is patchy and temporally discontinuous. Early Permian records are dominated by equatorial Europe and North America, while those of the Middle and Late Permian are dominated by temperate
Karoo Supergroup sediments of South Africa and the Ural region of European Russia. Early Permian terrestrial faunas of North America and Europe were dominated by primitive
pelycosaur synapsids including the herbivorous
edaphosaurids, and carnivorous
sphenacodontids,
diadectids and
amphibians. Early Permian reptiles, such as
acleistorhinids, were mostly small insectivores.
Amniotes Synapsids (the group that would later include mammals) thrived and diversified greatly during the Cisuralian. Permian synapsids included some large members such as
Dimetrodon. The special adaptations of synapsids enabled them to flourish in the drier climate of the Permian and they grew to dominate the vertebrates. Predator-prey interactions among terrestrial synapsids became more dynamic. If terrestrial deposition ended around the end of the Cisuralian in North America and began in Russia during the early Guadalupian, a continuous record of the transition is not preserved. Uncertain dating has led to suggestions that there is a global hiatus in the terrestrial fossil record during the late Kungurian and early
Roadian, referred to as "Olson's Gap" that obscures the nature of the transition. Other proposals have suggested that the North American and Russian records overlap, with the latest terrestrial North American deposition occurring during the Roadian, suggesting that there was an extinction event, dubbed "
Olson's Extinction". The Middle Permian faunas of South Africa and Russia are dominated by therapsids, most abundantly by the diverse
Dinocephalia. Dinocephalians become extinct at the end of the Middle Permian, during the
Capitanian mass extinction event. Late Permian faunas are dominated by advanced therapsids such as the predatory sabertoothed
gorgonopsians and herbivorous beaked
dicynodonts, alongside large herbivorous
pareiasaur parareptiles. The
Archosauromorpha, the group of reptiles that would give rise to the
pseudosuchians,
dinosaurs, and
pterosaurs in the following Triassic, first appeared and diversified during the Late Permian, including the first appearance of the
Archosauriformes during the latest Permian.
Cynodonts, the group of therapsids ancestral to modern
mammals, first appeared and gained a worldwide distribution during the Late Permian. Another group of therapsids, the
therocephalians (such as
Lycosuchus), arose in the Middle Permian. There were no flying vertebrates, though the extinct lizard-like reptile family
Weigeltisauridae from the Late Permian had extendable wings like modern
gliding lizards, and are the oldest known gliding vertebrates. File:EdaphosaurusDB.jpg|
Edaphosaurus pogonias and
Platyhystrix – Early Permian, North America and Europe File:Dimetr eryopsDB.jpg|
Dimetrodon grandis and
Eryops – Early Permian, North America File:Ocher fauna DB.jpg|Ocher fauna,
Estemmenosuchus uralensis and
Eotitanosuchus – Middle Permian, Ural Region File:Titanophoneus 3.jpg|
Titanophoneus and
Ulemosaurus – Ural Region File:Inostrancevia 4DB.jpg|
Inostrancevia alexandri and
Scutosaurus – Late Permian, North European Russia (Northern Dvina)
Amphibians Permian stem-amniotes consisted of
lepospondyli and
batrachosaurs, according to some phylogenies; according to others, stem-amniotes are represented only by
diadectomorphs. Temnospondyls reached a peak of diversity in the Cisuralian, with a substantial decline during the Guadalupian-Lopingian following Olson's extinction, with the family diversity dropping below Carboniferous levels.
Embolomeres, a group of aquatic crocodile-like limbed vertebrates that are
reptilliomorphs under some phylogenies. They previously had their last records in the Cisuralian, are now known to have persisted into the Lopingian in China. Modern amphibians (
lissamphibians) are suggested to have originated during Permian, descending from a lineage of
dissorophoid temnospondyls or
lepospondyls.
Fish The diversity of fish during the Permian is relatively low compared to the following Triassic. The dominant group of
bony fishes during the Permian were the "
Paleopterygii" a
paraphyletic grouping of
Actinopterygii that lie outside of
Neopterygii. The diversity of
coelacanths is relatively low throughout the Permian in comparison to other marine fishes, though there is an increase in diversity during the terminal Permian (Changhsingian), corresponding with the highest diversity in their evolutionary history during the Early Triassic. The last common ancestor of all living lungfish is thought to have existed during the Early Permian. Though the fossil record is fragmentary, lungfish appear to have undergone an evolutionary diversification and size increase in freshwater habitats during the Early Permian, but subsequently declined during the middle and late Permian. Conodonts experienced their lowest diversity of their entire evolutionary history during the Permian. Permian chondrichthyan faunas are poorly known. Members of the chondrichthyan clade
Holocephali, which contains living
chimaeras, reached their apex of diversity during the Carboniferous-Permian, the most famous Permian representative being the "buzz-saw shark"
Helicoprion, known for its unusual spiral shaped spiral tooth whorl in the lower jaw.
Hybodonts, a group of shark-like chondrichthyans, were widespread and abundant members of marine and freshwater faunas throughout the Permian.
Xenacanthiformes, another extinct group of shark-like chondrichthyans, were common in freshwater habitats, and represented the
apex predators of freshwater ecosystems.
Flora Four
floristic provinces in the Permian are recognised, the
Angaran, Euramerican, Gondwanan, and Cathaysian realms. The
Carboniferous Rainforest Collapse would result in the replacement of
lycopsid-dominated forests with
tree-fern dominated ones during the late Carboniferous in Euramerica, and result in the differentiation of the Cathaysian floras from those of Euramerica. The tree-like
calamites, distant relatives of modern
horsetails, lived in coal swamps and grew in
bamboo-like vertical thickets. A mostly complete specimen of
Arthropitys from the Early Permian
Chemnitz petrified forest of Germany demonstrates that they had complex branching patterns similar to modern
angiosperm trees. By the Late Permian, high thin forests had become widespread across the globe, as evidenced by the global distribution of weigeltisaurids. of Permian
wetland environment, showing an
Eryops The oldest likely record of
Ginkgoales (the group containing
Ginkgo and its close relatives) is
Trichopitys heteromorpha from the earliest Permian of France. The oldest known fossils definitively assignable to modern
cycads are known from the Late Permian. In Cathaysia, where a wet tropical frost-free climate prevailed, the
Noeggerathiales, an extinct group of tree fern-like
progymnosperms were a common component of the flora The earliest Permian (~ 298 million years ago) Cathyasian Wuda Tuff flora, representing a coal swamp community, has an upper canopy consisting of
lycopsid tree
Sigillaria, with a lower canopy consisting of
Marattialean tree ferns, and Noeggerathiales.
Bennettitales, which would go on to become in widespread the Mesozoic, first appeared during the Cisuralian in China.
Lyginopterids, which had declined in the late Pennsylvanian and subsequently have a patchy fossil record, survived into the Late Permian in Cathaysia and equatorial east Gondwana. == Permian–Triassic extinction event ==