Synapsid

Synapsids Temporal range: Late Mississippian - Early Cretaceous, 320–100 Ma PreЄ Є O S D C P T J K Pg N Descendant taxon Mammalia survives to present
Dimetrodon grandis skeleton, National Museum of Natural History
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
clade:	Amniota
Class:	Synapsida Osborn, 1903
Subgroups
†Caseasauria †Caseidae †Eothyrididae Eupelycosauria †Edaphosauridae Sphenacodontia †Varanopidae
Synonyms
Theropsida

Synapsids ('fused arch') are a group of animals that includes mammals and everything more closely related to mammals than to other living amniotes.^[1] They are easily separated from other amniotes by having an opening low in the skull roof behind each eye, leaving a bony arch beneath each, accounting for their name.^[2] Primitive synapsids are usually called pelycosaurs; more advanced mammal-like ones, therapsids. The non-mammalian members are described as mammal-like reptiles in classical systematics,^[3][4] but are referred to as "stem-mammals" (or sometimes "proto-mammals") under cladistic terminology.^[5] Synapsids evolved from basal amniotes and are one of the two major groups of the later amniotes, the other major group being the sauropsids (reptiles and birds). They are distinguished from other amniotes by having a single opening (temporal fenestra) in their skull behind each eye, which developed in the ancestral synapsid about 324 million years ago (mya) during the late Carboniferous Period.

Synapsids were the dominant terrestrial animals in the middle to late Permian period, 299 to 251 million years ago (mya). As with almost all groups then extant, their numbers and variety were severely reduced by the Permian extinction. Some species survived into the Triassic period, but archosaurs quickly became the dominant animals and few of the non-mammalian synapsids outlasted the Triassic, although survivors persisted into the Cretaceous. However, as a phylogenetic unit they included the mammal descendants, and in this sense synapsids are still very much a living group of vertebrates. In the form of mammals, Synapsids (most recently and notably humans) again became the dominant land animals after they outcompeted birds following the K-T extinction event.

Linnaean and cladistic classifications

Synapsids as a reptilian subclass

Synapsids were originally defined at the turn of the 20th century, as one of the four main subclasses of reptiles, on the basis of their distinctive temporal openings. These openings in the cheek bones allowed attachment of larger jaw muscles, hence a more efficient bite. Synapsids were considered to be the reptilian lineage that led to mammals via gradually evolved, increasingly mammalian features, hence the name "mammal-like reptiles" which became a broad, traditional description for all non-mammalian synapsids.^[3][4]

The "mammal-like reptiles"

The traditional classification of synapsids as reptiles is continued by a number of palaeontologists (e.g. Carroll 1988, Colbert & Morales 2001) and in general biology. In the 1990s this approach was complemented by a cladistic one, according to which the only valid groups are those that include common ancestors and all of their descendants: these are known as monophyletic groups, or clades. Because mammals are directly descended from the synapsids, mammals are included under Synapsida as a clade, though in formal classification mammals are treated as a separate class that has evolved from within Synapsida.

Phylogenetically synapsids are the entire synapsid/mammal branch of the tree of life, though practically the term is most often used when referring to the reptile-grade synapsids. The term "mammal-like reptiles" represents a paraphyletic grade, but is commonly used both colloquially and in the technical literature to refer to all non-mammalian synapsids. The actual monophyly of Synapsida is not in doubt however, and the expressions "Synapsida contains the mammals" and "synapsids gave rise to the mammals" both express the same phylogenetic hypothesis.

Primitive and advanced synapsids

The mammal-like reptiles are traditionally divided into a primitive group and an advanced group, known respectively as 'pelycosaurs' and therapsids. 'Pelycosaurs' make up the six most primitive families of synapsids.^[6] They were all rather lizard-like with sprawling gait and possibly horny scutes. The therapsids contain the more advanced synapsids, having a more erect pose and possibly hair, at least in some forms. In traditional taxonomy the Synapsida encompasses two distinct grades successively closer to mammals: The low-slung pelycosaurs have given rise to the more erect therapsids, who in their turn have given rise to the mammals. In traditional vertebrate classefication the Pelycosauria and Therapsida were both considered orders of the subclass Synapsida.^[2][3] At least one paleontologist, Benton, still follows such traditional classification; Benton classifies the synapsids as a paraphyletic class.^[4]

In phylogenetic nomenclature the terms are used somewhat different as the daughter clades are included. Most papers published during the 21st century have treated "Pelycosauria" as an informal grouping of primitive members, if it is used at all, being virtually synonymous with Synapsida itself. Therapsida has remained in use as a clade containing both the traditional therapsid families and mammals. However, in practical usage, the terms are used almost exclusively when referring to the more basal members that lie outside of Mammaliaformes.

Characteristics

Temporal openings

The synapsids are distinguished by a single hole behind each eye.

Synapsids evolved a temporal fenestra behind each eye orbit on the lateral surface of the skull. It may have evolved to provide new attachment sites for jaw muscles. A similar development took place in the Diapsids, who evolved two rather than one opening behind each eye. Originally, the opening in the skull left the inner cranium only covered by the jaw muscles, but in higher therapsids and mammals the sphenoid bone has expanded to close the opening. This has left the lower margin of the opening as an arch extending from the lower edges of the braincase.

Teeth

Eothyris, an early synapsid with multiple canines.

Synapsids are characterized by having differentiated teeth. These include the canines, molars, and incisors. 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 proto-canines. This trait was subsequently lost in the Sauropsid line, but developed further in the synapsids. Early synapsids could have 2 or even 3 enlarged "canines", but in the therapsids, the pattern had settled to one canine in each upper jaw half. The lower canines developed later.

Jaw

Most paleontologists hold fossilized jaw remains to be the distinguishing feature used to classify synapsids and reptiles. 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, 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 mammals possess the malleus, incus and stapes, mammal-like reptiles (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 skull bone).^[7]

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 skull 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.

Palate

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.^[8]

Skin

Synapsid combination of hair and scutes, detail of a rat's tail.

The actual skin of the synapsids has been subject to some discussion. Basal reptilian skin is rather thin, and lack the thick dermal layer that produces leather in mammals.^[9] Exposed parts of reptiles are protected by horny scales or scutes. Mammal hide has a thick, fibrous dermis and rarely exhibits scutes. A hallmark of mammals is the presence of copious glands and hair follicles.

When the change from reptilian to mammalian type skin took place is not known, though fossilized rows of osteoderms indicate horny armour on the neck and back of pelycosaurs, and skin impressions indicate that some retained rectangular scutes on their undersides.^[10] The pelycosaur scutes probably were non-overlapping 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).^[11] The remaining upper surface of the pelycosaurs may have borne scutes too, or may have been glandular and leathery like that of a mammal.

It is currently unknown at what stage the synapsids acquired mammalian characteristics such as body hair and mammary glands, as the fossils only rarely provide direct evidence for soft tissues. An exceptionally well preserved skull of Estemmenosuchus, a therapsid from the Upper Permian show smooth hairless skin with what appears to be glandular depressions.^[12] The oldest known fossil showing unambiguous imprints of hair is the Callovian (late middle Jurassic) Castorocauda, an early mammal.^[13] The more advanced therapsids could have had a combination of naked skin, scutes and hair, a combination still found in some modern mammals like rodents and the opossum.

Metabolism

Sail-back pelycosaurs like Edaphosaurus indicate an early trend toward temperature regulation in synapsids.

The first pelycosaurs had the usual reptilian cold-blooded metabolism by all indications, including sprawling gait and a low slung body.^[2] However, there appears to have been an early trend towards a form of temperature regulation in several Pelycosaur lines, as indicated by the large "sails" in both edaphosaurids and sphenacodontids (e.g. Dimetrodon).

The sphenacodontids gave rise to the therapsids, who may have inherited the temperature regulation. The legs and feet of the early therapsid groups point to a more erect posture, traditionally interpreted as a sign of more efficient metabolism.^[14] None of them show any sign of a sail, indicating any temperature regulation would have relied on the creatures own metabolism rather than external heat. In the later cynodonts, the presence of a secondary palate, erect posture and other indicators of high metabolic rate suggests that many mammalian features, including an effective insulating layer of body hair, had evolved by this stage. This is now confirmed by impressions of fur in rocks directly underlying some fossil therapsids.^[15]

Evolutionary history

Main article: Evolution of mammals

Archaeothyris, one of the oldest synapsids found.

Archaeothyris and Clepsydrops are the earliest known synapsids.^[16] They lived in the Pennsylvanian subperiod of the Carboniferous Period and belonged to the series of primitive synapsids which are conventionally grouped as pelycosaurs. The pelycosaurs were the first successful group of amniotes, spreading and diversifying until they became the dominant large terrestrial animals in the latest Carboniferous and Early Permian Periods. They were sprawling, bulky, cold-blooded and had small brains. They were the largest land animals of their time, ranging up to 3 m (10 ft) in length. Many, like Dimetrodon, had large sails that may have helped raise their body temperature. A few relict groups lasted into the later Permian, but most of the pelycosaurs became extinct before the end of Permian.

Sphenacodon was a carnivorous pelycosaur that was closely related to Dimetrodon and the therapsids.

The therapsids, a more advanced group of synapsids, appeared during the first half of the Permian and went on to become the dominant large terrestrial animals during the latter half. They were by far the most diverse and abundant animals of the Middle and Late Permian and 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 extinction in Earth's history, which may have been related to the Siberian Traps volcanic event.

Nikkasaurus - an enigmatic synapsid from the Middle Permian of Russia.

Lystrosaurus was the most common synapsid shortly after the Permian–Triassic extinction event.

Only a few therapsids (and some relict 'pelycosaur' taxa) survived the Permian extinction and 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. Now, however, they were accompanied by the early archosaurs (soon to give rise to the dinosaurs). Some of these, like Euparkeria, were small and lightly built, while others, like Erythrosuchus, were as big as or bigger than the largest therapsids.

Triassic therapsids included three groups. Specialised, beaked herbivores known as dicynodonts (such as Lystrosaurus and its descendants, the Kannemeyeriidae), contained some members which reached large size (up to a tonne or more). The increasingly mammal-like carnivorous, herbivorous, and insectivorous cynodonts included the eucynodonts from the Olenekian age, an early representative of which was Cynognathus. Finally, there were the therocephalians, which only lasted into the early part of the Triassic.

Cynognathus was the largest predatory cynodont of the Triassic.

Unlike the dicynodonts, which remained large, the cynodonts became progressively smaller and more mammal-like as the Triassic progressed. From the most advanced and tiny cynodonts, which were only the size of a shrew, came the first mammal precursors, during the Carnian age of the Late Triassic, about 220 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 Norian 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 first appeared near the end of the Triassic. They were carnivorous and persisted well into the Middle Jurassic. The other, Tritylodontidae, first appeared at the same time as the tritheledonts, but they were herbivorous. This group became extinct at the end of the Early Cretaceous epoch. Dicynodonts are thought to have become extinct near the end of the Triassic period, but there is evidence that this group survived. New fossil finds have been found in the Cretaceous rocks of Gondwana.

Today, there are 5,400 species of living synapsids known as the mammals, including both aquatic (whales) and flying (bats) species, and the largest animal ever known to have existed (the blue whale). Humans are synapsids as well. Uniquely among the synapsids, however, most mammals are viviparous and give birth to live young rather than laying eggs, the exception being the monotremes.

Synapsids' evolution into mammals is believed to have been triggered by moving to a nocturnal niche. Proto-mammals with higher metabolic rates were able to keep their bodies warm at night, and were more likely to survive. This meant consuming food (generally thought to be insects) in much greater quantity. To facilitate rapid digestion, proto-mammals evolved mastication (chewing) and specialized teeth that aided chewing. Limbs also evolved to move under the body instead of to the side, allowing proto-mammals to breathe more efficiently during locomotion^[17] and also to be able to change direction more quickly in order to catch small prey at a faster rate. This helped make it possible to support their higher metabolic demands. It is believed that, rather than out-running predators, proto-mammals adapted the strategy of outmaneuvering predators using their improved locomotor capabilities.^[15]

Taxonomy

Classification

• Series Amniota
  • CLASS SYNAPSIDA *
    • Order Pelycosauria *
      • Suborder Caseasauria
        • Family Caseaidae
        • Family Eothyrididae
      • Suborder Eupelycosauria *
        • Family Edaphosauridae
        • Family Lupeosauridae
        • Family Ophiacodontidae
        • Family Sphenacodontidae
        • Family Varanopidae
    • Order Therapsida *
      • Suborder Biarmosuchia
      • (unranked) Eutherapsida
        • Suborder Dinocephalia
          • Infraorder Anteosauria
          •  ?Infraorder Tapinocephalia
      • (unranked) Neotherapsida
        • Suborder Anomodontia
        • Superfamily Venyukoviamorpha
          • Family Venyukoviidae
        •  ?Infraorder Dicynodonta
      • (unranked) Theriodontia
        • Suborder Gorgonopsia
      • (unranked) Eutheriodontia
      • Suborder Therocephalia
      • Suborder Cynodontia *
  • CLASS MAMMALIA

Phylogeny

Synapsida

†Caseasauria Eupelycosauria †Varanopidae †Ophiacodontidae †Edaphosauridae Sphenacodontia †Sphenacodontidae Therapsida †Biarmosuchia †Eotitanosuchus Eutherapsida †Dinocephalia Neotherapsida †Anomodontia Theriodontia †Gorgonopsia Eutheriodontia †Therocephalia Cynodontia †Dvinia †Procynosuchidae Epicynodontia †Thrinaxodon Eucynodontia †Cynognathus Probainognathia †Tritheledontidae †Chiniquodontidae †Prozostrodon Mammaliaformes Mammalia

Notes

1. ^ Laurin and Reisz 2007.
2. ^ ^{a b c} Romer, A.S. & Parsons, T.S. (1985): The Vertebrate Body. (6th ed.) Saunders, Philadelphia.
3. ^ ^{a b c} Carroll 1988: 397.
4. ^ ^{a b c} Benton 2005: 122.
5. ^ Donoghue, Philip (2005), "Matters of the Record: Saving the stem group---a contradiction in terms?". Page 555 (pdf document page 3 of 6)
6. ^ Benton 2005: 120.
7. ^ Salentijn, L. Biology of Mineralized Tissues: Prenatal Skull Development, Columbia University College of Dental Medicine post-graduate dental lecture series, 2007
8. ^ Hopson 1987.
9. ^ Hildebran, M. & Goslow, G. (2001): Analysis of Vertebrate Structure. 5th edition. John Wiley & sons inc, New York. 635 pages ISBN 0-471-29505-1
10. ^ Botha-Brink, J. and Modesto, S.P. (2007). "A mixed-age classed ‘pelycosaur’ aggregation from South Africa: earliest evidence of parental care in amniotes?" Proceedings of the Royal Society B, 274(1627): 2829–2834. doi:10.1098/rspb.2007.0803
11. ^ Carroll, R.L. (1969). "Problems of the origin of reptiles." Biological Reviews, 44: 393-432.
12. ^ Kardong, K.V. (2002): Vertebrates: Comparative anatomy, function, evolution. 3rd Edition. McGraw-Hill, New York
13. ^ Ji, Q.; Luo, Z-X, Yuan, C-X, and Tabrum, A.R. (February 2006). "A Swimming Mammaliaform from the Middle Jurassic and Ecomorphological Diversification of Early Mammals". Science 311 (5764): 1123. doi:10.1126/science.1123026. PMID 16497926. http://www.sciencemag.org/cgi/content/abstract/311/5764/1123.  See also the news item at "Jurassic "Beaver" Found; Rewrites History of Mammals". http://news.nationalgeographic.com/news/2006/02/0223_060223_beaver.html. 
14. ^ Carroll, R. L. (1988), Vertebrate Paleontology and Evolution, WH Freeman & Co.
15. ^ ^{a b} Hoyt 1997.
16. ^ Lambert 2001: 68-69.
17. ^ Bramble and Jenkins 1994.

See also

• Anapsida
• Diapsida
• Euryapsida
• List of synapsids
• Mammal classification
• Prehistoric mammal
• Vertebrate paleontology
• Timeline of evolution

References

• Benton, Michael J. (2005). Vertebrate Paleontology, 3rd ed. Oxford: Blackwell Science Ltd. ISBN 0-632-05637-1.
• Bramble, D. M.; Jenkins, F. A. (1993). "Mammalian locomotor-respiratory integration: Implications for diaphragmatic and pulmonary design". Science 262 (5131): 235–240. doi:10.1126/science.8211141. PMID 8211141. 
• Carroll, Robert L. (1988). Vertebrate Paleontology and Evolution. New York: W.H. Freeman & Co. ISBN 0-7167-1822-7.
• Hopson, James A. (1987). "The Mammal-Like Reptiles: A Study of Transitional Fossils". The American Biology Teacher 49 (1): 16–26. 
• Hoyt, Donald F. (1997). Synapsid Reptiles. Zoo 138 Vertebrate Zoology Home Page.
• Lambert, David (2001). Dinosaur Encyclopedia. ISBN 0-7894-7935-4.
• Laurin, Michel, and Robert R. Reisz (2007). Synapsida: Mammals and their extinct relatives. Version 6 April 2007. The Tree of Life Web Project.

Further reading

• Colbert, E. H. (1969). Evolution of the Vertebrates (2nd ed.). New York: John Wiley & Sons Inc. ISBN 0471164666. 

External links

• Synapsids - at Macroevolution.net
• Synapsida - Pelycosauria - at Palaeos
• Transitional Vertebrate Fossils - includes description of important transitional genera in the evolutionary sequence linking primitive synapsids to mammals