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Clade of tetrapods

Synapsida is one of the two major clades of vertebrate animals in the group Amniota, the other being the Sauropsida (which includes reptiles and birds). The synapsids were the dominant land animals in the late Paleozoic and early Mesozoic, but the only group that survived into the Cenozoic are mammals. Unlike other amniotes, synapsids have a single temporal fenestra, an opening low in the skull roof behind each eye orbit, leaving a bony arch beneath each; this accounts for their name. The distinctive temporal fenestra developed about 318 million years ago during the Late Carboniferous period, Therefore, calling synapsids "mammal like reptiles" is incorrect under the new definition of "reptile", so they are now referred to as stem mammals, and sometimes as proto mammals, or paramammals. Synapsids were the largest terrestrial vertebrates in the Permian period, 299 to 251 million years ago, equalled only by some large pareiasaurs at the end of the Permian. Most lineages of pelycosaur grade synapsids were replaced at the end of the Early Permian by the more advanced therapsids. Synapsid numbers and variety were severely reduced by the Permian–Triassic extinction event. Only the therapsid dicynodonts and eutheriodonts (consisting of Therocephalia and Cynodontia) are known to have continued into the Triassic period. The cynodont group Probainognathia, which includes Mammaliaformes (mammals and their closer ancestors), were the only synapsids to survive beyond the Triassic. During the Triassic, the sauropsid archosaurs became some of the largest and most numerous land vertebrates, only rivaled in size by kannemeyeriiform dicynodonts, and gave rise to the dinosaurs. When all non avian dinosaurs were wiped out by the Cretaceous–Paleogene extinction event, the mammalian synapsids diversified again to become the largest land and marine animals on Earth.

At the turn of the 20th century synapsids were thought to be one of the four main subclasses of reptiles. However, this notion was disproved upon closer inspection of skeletal remains, as synapsids are differentiated from reptiles by their distinctive temporal openings. These openings in the skull bones allowed the attachment of larger jaw muscles, hence a more efficient bite. Synapsids were subsequently considered to be a later reptilian lineage that became mammals by gradually evolving increasingly mammalian features, hence the name "mammal like reptiles" (also known as pelycosaurs). These became the traditional terms for all Paleozoic (early) synapsids. More recent studies have debunked this notion as well, and reptiles are now classified within Sauropsida (sauropsids), the sister group to synapsids, thus making synapsids their own taxonomic group. The monophyly of Synapsida is not in doubt, and the expressions such as "Synapsida contains the mammals" and "synapsids gave rise to the mammals" both express the same phylogenetic hypothesis. This terminology reflects the modern cladistic approach to animal relationships, according to which the only valid groups are those that include all of the descendants of a common ancestor: these are known as monophyletic groups, or clades. Additionally, Reptilia (reptiles) has been revised into a monophyletic group and is considered entirely distinct from Synapsida, falling within Sauropsida, the sister group of Synapsida within Amniota.

The synapsids are traditionally divided for convenience, into therapsids, an advanced group of synapsids and the branch within which mammals evolved, and stem mammals, (previously known as pelycosaurs), comprising the other six more primitive families of synapsids. Stem mammals were all rather lizard like, with sprawling gait and possibly horny scutes, while therapsids tended to have a more erect pose and possibly hair, at least in some forms. In traditional taxonomy, the Synapsida encompasses two distinct grades: the low slung stem mammals have given rise to the more erect therapsids, who in their turn have given rise to the mammals. In traditional vertebrate classification, the stem mammals and therapsids were both considered orders of the subclass Synapsida. The trend towards differentiation is found in some labyrinthodonts and early anapsid reptilians in the form of enlargement of the first teeth on the maxilla, forming a form of protocanines. This trait was subsequently lost in the diapsid line, but developed further in the synapsids. Early synapsids could have two or even three enlarged "canines", but in the therapsids, the pattern had settled to one canine in each upper jaw half. The lower canines developed later.

The jaw transition is a good classification tool, as most other fossilized features that make a chronological progression from a reptile like to a mammalian condition follow the progression of the jaw transition. The mandible, or lower jaw, consists of a single, tooth bearing bone in mammals (the dentary), whereas the lower jaw of modern and prehistoric reptiles consists of a conglomeration of smaller bones (including the dentary, articular, and others). As they evolved in synapsids, these jaw bones were reduced in size and either lost or, in the case of the articular, gradually moved into the ear, forming one of the middle ear bones: while modern mammals possess the malleus, incus and stapes, basal synapsids (like all other tetrapods) possess only a stapes. The malleus is derived from the articular (a lower jaw bone), while the incus is derived from the quadrate (a cranial bone). Mammalian jaw structures are also set apart by the dentary squamosal jaw joint. In this form of jaw joint, the dentary forms a connection with a depression in the squamosal known as the glenoid cavity. In contrast, all other jawed vertebrates, including reptiles and nonmammalian synapsids, possess a jaw joint in which one of the smaller bones of the lower jaw, the articular, makes a connection with a bone of the cranium called the quadrate bone to form the articular quadrate jaw joint. In forms transitional to mammals, the jaw joint is composed of a large, lower jaw bone (similar to the dentary found in mammals) that does not connect to the squamosal, but connects to the quadrate with a receding articular bone.

Over time, as synapsids became more mammalian and less 'reptilian', they began to develop a secondary palate, separating the mouth and nasal cavity. In early synapsids, a secondary palate began to form on the sides of the maxilla, still leaving the mouth and nostril connected. Eventually, the two sides of the palate began to curve together, forming a U shape instead of a C shape. The palate also began to extend back toward the throat, securing the entire mouth and creating a full palatine bone. The maxilla is also closed completely. In fossils of one of the first eutheriodonts, the beginnings of a palate are clearly visible. The later Thrinaxodon has a full and completely closed palate, forming a clear progression.

In addition to the glandular skin covered in fur found in most modern mammals, modern and extinct synapsids possess a variety of modified skin coverings, including osteoderms (bony armor embedded in the skin), scutes (protective structures of the dermis often with a horny covering), hair or fur, and scale like structures (often formed from modified hair, as in pangolins and some rodents). While the skin of reptiles is rather thin, that of mammals has a thick dermal layer. The ancestral skin type of synapsids has been subject to discussion. Among the early synapsids, only two species of small varanopids have been found to possess osteoderms; fossilized rows of osteoderms indicate bony armour on the neck and back. However, some recent studies have cast doubt on the placement of Varanopidae in Synapsida, while others have countered and lean towards this traditional placement. Skin impressions indicate some early synapsids basal possessed rectangular scutes on their undersides and tails. The pelycosaur scutes probably were nonoverlapping dermal structures with a horny overlay, like those found in modern crocodiles and turtles. These differed in structure from the scales of lizards and snakes, which are an epidermal feature (like mammalian hair or avian feathers). Recently, skin impressions from the genus Ascendonanus suggest that at least varanopsids developed scales similar to those of squamates. It is currently unknown exactly when mammalian characteristics such as body hair and mammary glands first appeared, as the fossils only rarely provide direct evidence for soft tissues. An exceptionally well preserved skull of Estemmenosuchus, a therapsid from the Upper Permian, preserves smooth skin with what appear to be glandular depressions, an animal noted as being semi aquatic. The oldest known fossil showing unambiguous imprints of hair is the Callovian (late middle Jurassic) Castorocauda and several contemporary haramiyidans, both non mammalian mammaliaform (see below, however). More primitive members of the Cynodontia are also hypothesized to have had fur or a fur like covering based on their inferred warm blooded metabolism. The oldest examples of nocturnality in synapsids is believed to have been in species that lived more than 300 million years ago. However, Permian coprolites from Russia showcase that at least some synapsids did already have fur in this epoch. These are the oldest impressions of hair on synapsids.

Early synapsids, as far back as their known evolutionary debut in the Late Carboniferous period, Because they were vulnerable to desiccation, secretions from apocrine like glands may have helped keep the eggs moist. ), though this would limit the mobility of the parent. The latter may have been the primitive form of egg care in synapsids rather than simply burying the eggs, and the constraint on the parent's mobility would have been solved by having the eggs "parked" in nests during foraging or other activities and periodically be hydrated, allowing higher clutch sizes than could fit inside a pouch (or pouches) at once, and large eggs, which would be cumbersome to carry in a pouch, would be easier to care for. The basis of Oftedal's speculation is the fact that many species of anurans can carry eggs or tadpoles attached to the skin, or embedded within cutaneous "pouches" and how most salamanders curl around their eggs to keep them moist, both groups also having glandular skin.

Aerial locomotion first began in non mammalian haramiyidan cynodonts, with Arboroharamiya, Xianshou, Maiopatagium and Vilevolodon bearing exquisitely preserved, fur covered wing membranes that stretch across the limbs and tail. Their fingers are elongated, similar to those of bats and colugos and likely sharing similar roles both as wing supports and to hang on tree branches. Within true mammals, aerial locomotion first occurs in volaticotherian eutriconodonts. A fossil Volaticotherium has an exquisitely preserved furry patagium with delicate wrinkles and that is very extensive, "sandwiching" the poorly preserved hands and feet and extending to the base of the tail. Argentoconodon, a close relative, shares a similar femur adapted for flight stresses, indicating a similar lifestyle. Therian mammals would only achieve powered flight and gliding long after these early aeronauts became extinct, with the earliest known gliding metatherians and bats evolving in the Paleocene.

Recently, it has been found that endothermy was developed as early as Ophiacodon in the late Carboniferous. The presence of fibrolamellar, a specialised type of bone that can grow quickly while maintaining a stable structure, shows that Ophiacodon would have used its high internal body temperature to fuel a fast growth comparable to modern endotherms.

Over the course of synapsid evolution, progenitor taxa at the start of adaptive radiations have tended to be derived carnivores. Synapsid adaptive radiations have generally occurred after extinction events that depleted the biosphere and left vacant niches open to be filled by newly evolved taxa. In non mammaliaform synapsids, those taxa that gave rise to rapidly diversifying lineages have been both small and large in body size, although after the Late Triassic, progenitors of new synapsid lineages have generally been small, unspecialised generalists. Asaphestera, Archaeothyris and Clepsydrops, the earliest known synapsids, lived in the Pennsylvanian subperiod (323–299 mya) of the Carboniferous period and were one of many types of primitive synapsids that are now informally grouped together as stem mammals or sometimes as protomammals (previously known as pelycosaurs). The early synapsids spread and diversified, becoming the largest terrestrial animals in the latest Carboniferous and Early Permian periods, ranging up to in length. They were sprawling, bulky, possibly cold blooded, and had small brains. Some, such as Dimetrodon, had large sails that might have helped raise their body temperature. A few relict groups lasted into the later Permian but, by the middle of the Late Permian, all had either died off or evolved into their successors, the therapsids. The therapsids, a more advanced group of synapsids, appeared during the Middle Permian and included the largest terrestrial animals in the Middle and Late Permian. They included herbivores and carnivores, ranging from small animals the size of a rat (e. g.: Robertia), to large, bulky herbivores a ton or more in weight (e. g.: Moschops). After flourishing for many millions of years, these successful animals were all but wiped out by the Permian–Triassic mass extinction about 250 mya, the largest known extinction in Earth's history, possibly related to the Siberian Traps volcanic event. Only a few therapsids went on to be successful in the new early Triassic landscape; they include Lystrosaurus and Cynognathus, the latter of which appeared later in the early Triassic. However, they were accompanied by the early archosaurs (soon to give rise to the dinosaurs). Some of these archosaurs, such as Euparkeria, were small and lightly built, while others, such as Erythrosuchus, were as big as or bigger than the largest therapsids. After the Permian extinction, the synapsids did not count more than three surviving clades. The first comprised the therocephalians, which only lasted the first 20 million years of the Triassic period. The second were specialised, beaked herbivores known as dicynodonts (such as the Kannemeyeriidae), which contained some members that reached large size (up to a tonne or more). And finally there were the increasingly mammal like carnivorous, herbivorous, and insectivorous cynodonts, including the eucynodonts from the Olenekian age, an early representative of which was Cynognathus. Unlike the dicynodonts, which were large, the cynodonts became progressively smaller and more mammal like as the Triassic progressed, though some forms like Trucidocynodon remained large. The first mammaliaforms evolved from the cynodonts during the early Norian age of the Late Triassic, about 225 mya. During the evolutionary succession from early therapsid to cynodont to eucynodont to mammal, the main lower jaw bone, the dentary, replaced the adjacent bones. Thus, the lower jaw gradually became just one large bone, with several of the smaller jaw bones migrating into the inner ear and allowing sophisticated hearing. Whether through climate change, vegetation change, ecological competition, or a combination of factors, most of the remaining large cynodonts (belonging to the Traversodontidae) and dicynodonts (of the family Kannemeyeriidae) had disappeared by the Rhaetian age, even before the Triassic–Jurassic extinction event that killed off most of the large non dinosaurian archosaurs. The remaining Mesozoic synapsids were small, ranging from the size of a shrew to the badger like mammal Repenomamus. During the Jurassic and Cretaceous, the remaining non mammalian cynodonts were small, such as Tritylodon. No cynodont grew larger than a cat. Most Jurassic and Cretaceous cynodonts were herbivorous, though some were carnivorous. The family Tritheledontidae, which first appeared near the end of the Triassic, was carnivorous and persisted well into the Middle Jurassic. The other, Tritylodontidae, first appeared at the same time as the tritheledonts, but was herbivorous. This group became extinct at the end of the Early Cretaceous epoch. Dicynodonts are generally thought to have become extinct near the end of the Triassic period, but there was evidence this group survived, in the form of six fragments of fossil bone that were found in Cretaceous rocks of Queensland, Australia. If true, it would mean there is a significant ghost lineage of Dicynodonts in Gondwana. However, these fossils were re described in 2019 as being Pleistocene in age, and possibly belonging to a diprotodontid marsupial. Today, the 5,500 species of living synapsids, known as the mammals, include both aquatic (whales) and flying (bats) species, and the largest animal ever known to have existed (the blue whale). Humans are synapsids, as well. Most mammals are viviparous and give birth to live young rather than laying eggs with the exception being the monotremes. Triassic and Jurassic ancestors of living mammals, along with their close relatives, had high metabolic rates. This meant consuming food (generally thought to be insects) in much greater quantity. To facilitate rapid digestion, these synapsids evolved mastication (chewing) and specialized teeth that aided chewing. Limbs also evolved to move under the body instead of to the side, allowing them to breathe more efficiently during locomotion. This helped make it possible to support their higher metabolic demands.