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Synapsids

Synapses [1] [2] [3] ( Latin Synapsida ) , also teromorphs or beast-like [4] [5] [6] ( Latin Theromorpha , from other Greek θήρ , θηρίον - beast + μορφή - form) Is a taxonomic group of the amniotic clade that arose about 318 million years ago [7] . One of the synapsid groups - cynodonts (Cynodontia) - was the ancestor of mammals [8] [9] .

† Synapsids
Dimetrodon.jpg
Skeleton Dimetrodon grandis
Scientific classification
Domain:Eukaryotes
Kingdom:Animals
Kingdom :Eumetazoi
No rank :Bilateral symmetrical
No rank :Secondary
Type of:Chordate
Subtype :Vertebrates
Infratype :Maxillary
Overclass :Tetrapods
Grade:† Synapsids
International scientific name

Synapsida Osborn , 1903

For most of the 20th century, synapsids were considered one of the subclasses of the reptile class. So, in R. Carroll’s monograph [10], synapsids acted precisely as a subclass of the Reptilia class, which included 2 orders: pelicosaurs (Pelycosauria) and terapsid (Therapsida).

Later, the widespread dissemination of ideas characteristic of phylogenetic taxonomy led to a revision of the classification of synapsids. They were removed from the reptile class, having received at the end of the 20th century the status of an independent class [11] [12] .

According to modern concepts, synapsids together with mammals form the clade Synapsidomorpha ( synapsid-like ), which, together with the sister group Sauropsida ( zavropsida ), forms, in turn, the clade of amniotes - the largest group in the superclass of tetrapods [7] [13] [14] . Moreover, supporters of evolutionary taxonomy distinguish two classes in the synapsidoid structure: the paraphyletic class Synapsida and the daughter monophyletic class Mammalia [11] [12] . Proponents of cladistics (which is characterized by the desire to get rid of non-monophyletic taxa) usually do not distinguish the synapsid class, treating the groups included in it independently, and the term “Synapsida” is often used instead of the term “Synapsidomorpha”. With this approach, the concept of “synapsid” covers all mammals from the platypus to humans [15] .

Content

Study History

First Discoveries

 
S. S. Kutorg (1805-1861)
 
R. Owen (1804-1892)

The history of the study of synapsids dates back to 1838 . This year, Russian zoologist S. S. Kutorg , exploring finds made by Russian mining engineers in copper sandstones in the western slopes of the Ural Mountains in the 1770s, gave the first scientific descriptions of the fossils of synapsids, highlighting the species , and Syodon biarmicum . The scientist considered these species to be mammals; according to modern ideas, they belong to the group of deinocephals [16] [17] [18] . In 1841, another Russian zoologist, G. I. Fischer von Waldheim , in the course of studying fossil remains from the same region, described the species Rhopalodon wangenheimi (also nowadays referred to as deiencephalus) [19] [20] .

In 1845, the English zoologist Richard Owen , studying finds made in the Karoo region in southern Africa, , described the species Dicynodon lacerticeps - the first representative of the dicynodont group, and attributed this species to reptiles [21] . In 1854, the American zoologist Joseph Lady described the species of Bathygnathus borealis (reclassified as Dimetrodon borealis [22] ), the first pelicosaurus [23], found on the Prince Edward Island on the Canadian island . Gradually new discoveries appeared; summarizing the results of the first studies, R. Owen established the Anomodontia reptile detachment in 1859, and in 1876 isolated a separate Theriodontia detachment (which included forms with numerous well-differentiated teeth). In 1878, the American paleontologist Edward Cope introduced the Theromorpha detachment with its division into suborders Pelycosauria and Anomodontia (the latter, in turn, included the Dicynodontia and Theriodontia groups), pointing out the systematic proximity of Theromorpha and mammals [24] [25] . Soon, Cope, however, expanded the scope of Theromorpha to include cotylosaurs [26] ; at the same time, he assigned the Tritylodontidae family in 1884 not to Theromorpha, but to mammals (only in 1956 did the German paleontologist Walter Kühne attribute this family to cynodonts ) [27] .

End of XIX - beginning of XX century

 
E. Cope (1840–1897)

During the nineteenth and early twentieth centuries, many finds of synapsid fossils were made - mainly in three regions remote from each other: South Africa , North America and Eastern Europe (the situation did not fundamentally change even later: although the remains of synapsids were found in various places on earth of the Shara, the leading sources of information about the synapsid fauna were just the three mentioned regions) [28] . The study of the Karu region continued, through the efforts of paleontologists such as R. Owen, G. Seeley , R. Broome , D. Watson , and as a result, well-known representatives of therapies such as gorgonops , , trinaxodone , diademodone , trirachodone were described , cynognatus , cannumeria , bauria , moschops , , arctops [29] [30] . In the 2nd half of the 1870s, an effective search for fossil remains in North America began: E. Cope described species of Pelicosaurus ( 1875 ) from finds in Illinois and Dimetrodon limbatus ( 1878 ) from finds in Texas , and the opponent Kopa O. C. Marsh in the same 1878 described species of pelicosaurs Sphenacodon ferox and Ophiacodon mirus using fossil remains from New Mexico [31] . The most important stage of the paleontological study of Eastern Europe was the annual expeditions of V.P. Amalitsky to the Northern Dvina basin ( 1895 - 1914 ), during which, in particular, foreigners , dvins , venyukovies were described [32] [33] .

 
G. Osborne (1857-1935)

The American paleontologist Henry Osborne in 1903 introduced the division of the Reptilia class into two subclasses: Diapsida and Synapsida . The scope of the synapsid subclass in Osbourne’s interpretation, however, significantly differed from that currently accepted, since Osbourne included, along with the superorder Anomodontia, also cotylosaurs , turtles and saurotertergy (i.e., groups with anapsid and euryapsid skull structures), and the detachment Pelycosauria diapsids (considering the combination proposed by Cope with Anomodontia in one taxon and the term Theromorpha itself to be illegal). Other paleontologists introduced refinements into this classification, but the name Synapsida persisted and over time almost completely replaced the name Theromorpha [24] [34] .

As a result, the “extra” groups were withdrawn from the Synapsida volume, and already in 1925, in the fundamental monograph of Samuel Williston 's The Osteology of the Reptiles, this taxon appears, as in most subsequent works of the 20th century, as a subclass consisting of two orders: Pelycosauria Cope, 1878 (with more primitive forms) and Therapsida Broom, 1905 (to which more advanced forms were assigned). Williston himself, however, the first of these groups called Theromorpha, and the volume of Pelycosauria interpreted more narrowly; in 1940, and in their study “Review of the Pelycosauria” [35] adopted the name Pelycosauria [36] [37] for this unit. On the other hand, the volume of Anomodontia was gradually limited only by dicynodonts and closely related groups [38] .

XX century and beginning of XXI century

Throughout the 20th century and at the beginning of the 21st century, the study of synapsids continued. New genera and species were described, and based on an analysis of the features of their skeleton, microstructure of bones and the dental system, new information was obtained on the morphology, physiology and lifestyle of synapsids [39] [40] . The division of pelicosaurs into 6 families ( Ophiacodontidae , Varanopidae , Eothyrididae , Caseidae , Edaphosauridae , Sphenacodontidae ) [41] [42] and terapsid into 6 main groups ( , Dinocephia , Anomodontia , Corgon , Gorgon 38] , and basically a consensus has been established regarding the phylogenetic relationships between these groups of synapsids (see section “Classification of synapsids”) A special place in the synapsid system was occupied by the Early Permian genera tetraceratops ( Tetraceratops , discovered in 1897 and described in 1908) and raranimus ( Raranimus , discovered in 1998 and described in 2009), which combine the characteristics characteristic of pelicosaurs and terapsids; these two genera apparently form, respectively, the two earliest (sequentially separated) branches of therapids [38] [43] [44] .

 
Tetraceratops insignis (possibly the oldest terapsid)

At the end of the 20th century, the interpretation of the Synapsida taxon as an independent class (and not a subclass of the Reptilia class as before) became widespread [11] [12] ; note that Adrian Brink in 1963 proposed [45] to raise the taxonomic rank of Synapsida to a class with two subclasses: Pelycosauria and Therapsida [46] . At the same time, the spread of views characteristic of radical representatives of cladistics has led many modern authors to use the name Synapsida in an expanded sense and apply it to the entire clade of Synapsidomorpha, which includes not only the synapsid in the classical sense, but also mammals (in relation to In this clade, in addition to the name Synapsidomorpha [13] , the names Theropsida [14] [47] [48] or Pan-Mammalia [49] are also sometimes used. Moreover, mammals are also introduced into Therapsida, and the term Pelycosauria is not used as referring to the paraphyletic group [38] [50] .

For other paleontologists, this approach is notorious, because to refer to the taxa Synapsida or Therapsida in their traditional volume, the bulky expressions “non-mammalian synapsids” ( English nonmammalian Synapsida ) and “non-mammalian therapsids” ( English nonmammalian Therapsida) ) (pelicosaurs are called “basal synapsids” [51] ). In this regard, Tom Kemp noted that the paraphyleticity of Synapsida and Therapsida is of a "technical" character; at the same time, pelicosaurs significantly differ from terapsids in many ways (and therapsids from mammals), and in similar situations, when the species forming the paraphyletic group have enough similarity to each other in phenotype , the interpretation of this group as a taxon is quite legal [28] . Proponents of this point of view continue to regard Synapsida and Mammalia as separate classes [11] [12] .

Structural Features

 
Typical synapsid structure of the skull

A characteristic feature of synapsids is the presence of one temporal foramen on each side of their skull , bounded from above by the temporal arch of the orbital and scaly bones (in sauropertigia there is also one temporal foramen, but lying above the orbital bone, and in diapsids there are two temporal foramen). The figure on the right shows the structure of the skull typical of synapsids and bones are indicated: p - parietal , po - orbital , sq - scaly , j - zygomatic , qj - square-zygomatic , q - square [52] . The temporal holes serve to exit the jaw muscles, and their presence allowed those amniotes who had them, more effectively - compared to amphibians and turtles (whose skull does not have temporal holes) - bite their prey.

The lower jaw of the synapsids has a complete set of bones, characteristic of early amniotes. With the skull, it is articulated by the jaw joint formed (like in other amniotes, but not in mammals) by square and articular bones. In mammals, such a joint is replaced by a secondary jaw joint , which is already formed by scaly and dental bones [53] [54] . Traditionally, it is the formation of the secondary jaw joint that is considered as a leading feature that allows to draw a line between the classes of mammals and synapsids; The revised wording of the diagnostic character of mammals, proposed in 2002 by Luo Cheshey , Z. Kelyan-Yavorovskaya and R. Cifelli , reads as follows: “articulation of the lower jaw with the skull, in which the articular head of the tooth bone enters the glenoid (articular cavity) on the scaly bone” (it should be noted here that the synapsids from the family also have scaly and tooth bones, although there is no articular head or pronounced glenoid region) [55] . However, other views are also known (see Classification of Mammals ) on the ratio of the volumes of the classes of mammals and synapsids [56] [57] .

Already in some early synapsids (pelicosaurs), teeth show the initial stages of differentiation: the teeth located on the premaxillary bones resemble the shape of incisors , followed by fangs and cheek teeth (the latter retain their original conical shape) [58] . Differentiation of teeth acquires a more pronounced character in advanced synapsids (terapsid); it is clearly expressed in the representatives of the theriodont group (“beast-toothed”), and if the gorgonops’s cheek teeth remain simple conical (and sometimes even disappear), then they have a complex chewing surface in cynodonts [59] .

An important evolutionary conquest of advanced theriodonts was the formation of a secondary bone palate (it occurred independently in the Triassic terocephals and cynodonts). The secondary palate significantly strengthened the upper jaw and reliably isolated the nasal and oral cavities, allowing the animal to breathe continuously while chewing food [60] [61] .

Apparently, synapsids, unlike zavropsids , initially had smooth, skinless skin, in which there were numerous glands . More advanced synapsids probably had a hairline that provided thermal insulation (it is believed that hair could initially appear as a means of touch - such as vibrissae [62] ). Synapsids propagated by laying eggs [63] .

Synapsid classification

The synapsid class includes two orders - Pelycosauria ( pelicosaurs , Middle Carboniferous - Late Permian ) and Therapsida ( therapsids , Middle Permian - Early Cretaceous ); the latter includes the following main groups: ( biarmosuchia ), Dinocephalia ( dinocephalus ), Anomodontia ( anomodonts ) and Theriodontia ( theriodonts , or animal’s teeth ) [64] [65] . Some authors raise the rank of therapid to a subclass containing the units Biarmosuchia, Dinocephalia, Anomodontia and the superorder Theriodontia; the latter, in turn, includes the detachments of Gorgonopsia ( gorgonops ) and ( real theriodonts ) [66] . According to current data [42] [67] , the classification of synapsids can be represented in the form of the following two cladograms :

Pelycosauria
Caseasauria

† Eothyrididae



† Caseidae



Eupelycosauria

† Varanopidae




† Ophiacodontidae




† Edaphosauridae


Sphenacodontia

† Haptodus garnettensis




† Palaeohatteria




† Pantelosaurus




† Cutleria


Sphenacodontoidea

† Sphenacodontidae



Therapsida











Therapsida

† Biarmosuchia




† Eotitanosuchidae


Eutherapsida

† Dinocephalia


Neotherapsida
Anomodontia

† Venjukoviidae




† Galeopsidae



† Dicynodontia




Theriodontia

† Gorgonopsia


Eutheriodontia

† Therocephalia




Cynodontia (incl. Mammalia )









From these cladograms it can be seen that both orders (like the entire Synapsida class) are paraphyletic groups. Moreover, the order Pelycosauria ( pelicosaurs ) includes taxa represented by the terminal nodes of the first cladogram, excluding the last.

For a detailed tracking of the origin of mammals, we present one more cladogram revealing the internal structure of the suborder Cynodontia ( cynodonts ) and compiled taking into account the results of a study by Liu Jun and P. Olsen (2010) [68] :

Cynodontia


† Dviniidae



† Procynosuchidae





† Madysauridae


Epicynodontia

† Galesauridae




† Thrinaxodontidae


Eucynodontia

Cynognathia

† Cynognathidae


Homphodontia

† Diademodontidae




† Trirachodontidae



† Traversodontidae





Probainognathia

† Chiniquodontidae




† Probainognathidae


Prozostrodontia

† Prozostrodon




† Therioherpetidae




† Tritheledontidae




† Tritylodontidae




† Brasilodontidae




Mammalia
















Evolutionary History

 
Archaeothyris florensis , one of the oldest representatives of synapsids

Separation of the synapsid branch of amniot from its branch of zawropside occurs, according to modern data, 310 million years ago (medium carbon ) or somewhat earlier; the upper limit for the separation time is estimated at 333 million years ago (lower carbon ) [69] [70] .

As one of the oldest representatives of synapsids, archaeothyris ( ) from the family Ophiacodontidae , who lived 306 million years ago (Middle Carboniferous ) and described in 1972 from fossils found in Nova Scotia, is usually considered. Synapsids also include the early amniot , which lived somewhat earlier (314 million years ago) and was described in 1964 by fragmentary remains (also from Nova Scotia); it is also (presumably) classified as Ophiacodontidae, however, the systematic position of this genus remains unclear [48] [71] . Outwardly, the early synapsids resembled large modern lizards such as iguanas or monitor lizards , but with relatively shorter limbs (an example is the lizard - , an active medium-sized predator living at the end of the Early Permian from the same family Ophiacodontidae [72] ) [64] .

 
Dimetrodon incisivus with a “sail” on its back, tormenting a lizard

By the beginning of Perm, synapsids accounted for up to 70% of the known genera of amniotes; by this time, they had already diverged into a number of independent families (including fish-eating, herbivorous and carnivorous animals [42] ). Moreover, in the early Permian, representatives of the Sphenacodontidae family occupied dominant positions as terrestrial predators . The most famous among them was Dimetrodon , on the back of which there was a “sail” from the skin membrane and numerous spinous processes supporting it, which most likely served the purpose of thermoregulation (some Edaphosauridae also had a similar “sail”; representatives of this family occupied the trophic niche of large specialized phytophages ) [73] [74] . According to the calculations of C. Bramwell and P. Fellgett, a 200 kg dimetrodon would be heated without a sail from 26 ° C to 32 ° C in 205 minutes, and with a sail - in 80 minutes [75] .

At the turn of the early and middle Permian, the pelicosaurs are replaced by therapsids, which were distinguished by a significantly more active lifestyle and a high metabolic rate and occupied dominant positions among terrestrial vertebrates during the middle and late Permian [42] [76] . They quickly replaced the pelicosaurs, and only a few of the latter survived to the end of the Permian period. A certain exception is caseids (Caseidae), which experienced their heyday precisely in the Middle Permian and occupied the trophic niche of large specialized phytophages, which at the beginning of the period were occupied by edaphosaurs [42] [74] .

 
Inostrancevia alexandri attacking scutosaurs

Therapsids quickly diverged into a number of groups; Among them, , therocephals (Therocephalia), and early cynodonts (Cynodontia) led predatory lifestyles, and deinocephals (Dinocephalia) included both predatory and herbivorous forms [77] . Phytophages were also dicynodonts (Dicynodontia), various in size and body structure. The role of the dominant land predators was now played by the Gorgonopsia , hunting large prey; they clung to it with large saber-shaped fangs, and then tore the victim with sharp longitudinal jerks [78] . One of the largest gorgonops was foreigners ( Inostrancevia ), whose skull length could reach 60 cm [79] [80] .

At the end of the Permian period (about 252 million years ago [81] ), a catastrophic Permian extinction occurred , which resulted in the disappearance of approximately 90% of marine animal species and about 70% of terrestrial animal families. Extinction also affected the synapsid: the last pelicosaurs disappeared, and among the synapsids only three groups survived the catastrophe — large herbivorous dicynodonts and small (probably insectivorous or omnivorous ) terocephals and cynodonts [82] .

 
Herbivorous cynodont
Tritylodon longaevus

In the Triassic, terocephals, however, were relatively few and disappeared towards the end of the middle part of the period [83] . Dicynodonts were more widespread throughout the Triassic, although only one (of the treasure) out of four of their groups that survived Permian extinction underwent significant diversification during the period; by the end of the Triassic, dicynodonts also disappear (although, according to some reports, they existed in Australia until the end of the Early Cretaceous [84] ) [85] . But the true blossoming in the Triassic is experienced by the cynodonts, and most of their families belong to this period. Most cynodonts remained predatory or insectivorous, but specialized herbivorous groups also appeared: (Gomphodontia) and tritylodonts (Tritylodontydae). Triassic cynodonts have a significant number of signs characteristic of mammals [86] .

In the Late Triassic, the first mammals appear. Most evolutionary lines of cynodonts cease to exist by the end of the Triassic. Representatives of a few families met in the Jurassic : the family from the homophodont group (represented by the Scalenodontoides in the early Jurassic) and the phylogenetic families of the (ended in the Middle Jurassic) and Tritylodontydae (its representatives survived to the end Cretaceous, where it is represented by the genus Xenocretosuchus [87] ) [88] [89] .

Notes

  1. ↑ GRAMOTA.RU - reference and information Internet portal "Russian Language" | Dictionaries | Verification of the word (unspecified) . gramota.ru. Date of treatment July 6, 2018.
  2. ↑ synapsid (s) (Russian) , Gufo.me. Date of treatment July 6, 2018.
  3. ↑ where-stress.rf (neopr.) .
  4. ↑ Bestial / A.K. Rozhdestvensky // Euclid - Ibsen. - M .: Soviet Encyclopedia, 1972. - ( Great Soviet Encyclopedia : [in 30 vol.] / Ch. Ed. A. M. Prokhorov ; 1969-1978, vol. 9).
  5. ↑ Biological Encyclopedic Dictionary / Ch. ed. M.S. Gilyarov ; Editorial: A.A. Baev , G.G. Vinberg, G.A. Zavarzin et al. - M .: Sov. Encyclopedia , 1986.- S. 211. - 831 p. - 100,000 copies.
  6. ↑ Cherepanov, Ivanov, 2007 , p. 35, 250.
  7. ↑ 1 2 White A. T. Reptilomorpha: Cotylosauria (section Amniota) (neopr.) . // Website "PALAEOS: The history of life on Earth". Date of treatment May 3, 2016. Archived December 20, 2010.
  8. ↑ Vaughan, Ryan, Czaplewski, 2011 , p. 47-48.
  9. ↑ Diversity of Mammals / O. L. Rossolimo, I. Ya. Pavlinov , S. V. Kruskop, A. A. Lisovsky, N. N. Spasskaya, A. V. Borisenko, A. A. Panyutina. - M .: KMK Publishing House, 2004. - Part I. - 366 p. - (variety of animals). - ISBN 5-87317-098-3 . - S. 64.
  10. ↑ Carroll, vol. 3, 1993 , p. 203-205.
  11. ↑ 1 2 3 4 Benton, 2005 , p. 394.
  12. ↑ 1 2 3 4 Vaughan, Ryan, Czaplewski, 2011 , p. 47.
  13. ↑ 1 2 Synapsidomorpha (neopr.) . // Website " Wikispecies : The free species directory". Date of treatment May 3, 2016.
  14. ↑ 1 2 Rieppel O. Monophyly, Paraphyly, and Natural Kinds // Biology and Philosophy , 2005, 20 (2). - P. 465-487. - DOI : 10.1007 / s10539-004-0679-z .
  15. ↑ Lee M. S. Y., Reeder T. W., Slowinski J. B., Lawson R. Resolving Reptile Relationships: Molecular and Morphological Markers // Assembling the Tree of Life / Ed. by J. Cracraft and M. J. Donoghue. - Oxford: Oxford University Press, 2004 .-- xiii + 576 p. - ISBN 0-19-517234-5 . - P. 451-467.
  16. ↑ Palmer, 2005 , p. 153.
  17. ↑ Ochev, Surkov, 2003 , p. one.
  18. ↑ Ivakhnenko, 2008 , p. 101, 118-125.
  19. ↑ Palmer, 2005 , p. 154.
  20. ↑ Ochev, Surkov, 2003 , p. 2.
  21. ↑ Fröbisch J. Anomodontia: Introduction // Early Evolutionary History of the Synapsida / Ed. by C. F. Kammerer, K. D. Angielczyk, J. Fröbisch. - Dordrecht: Springer Science & Business Media , 2013 .-- xi + 337 p. - (Vertebrate Paleobiology and Paleoanthropology Series). - ISBN 978-94-007-6841-3 . - P. 89-92.
  22. ↑ Brink K. S., Maddin H. C., Evans D. C., Reisz R. R. Re-evaluation of the historic Canadian fossil Bathygnathus borealis from the Early Permian of Prince Edward Island // Canadian Journal of Earth Sciences , 2015, 52 (12). - P. 1109-1120. - DOI : 10.1139 / cjes-2015-0100 .
  23. ↑ Reisz R. R. “Pelycosaur” -Grade Synapsids: Introduction // Early Evolutionary History of the Synapsida / Ed. by C. F. Kammerer, K. D. Angielczyk, J. Fröbisch. - Dordrecht: Springer Science & Business Media , 2013 .-- xi + 337 p. - (Vertebrate Paleobiology and Paleoanthropology Series). - ISBN 978-94-007-6841-3 . - P. 3-5.
  24. ↑ 1 2 Ivakhnenko, 2008 , p. 101.
  25. ↑ Zittel K. A. History of Geology and Palæontology to the End of the Nineteenth Century . - London: Walter Scott, 1901. - xiii + 561 p. - P. 517.
  26. ↑ Hay O. P. Bibliography and Catalog of the Fossil Vertebrata of North America . - Washington: U. S. Government Printing Office, 1902. - 868 p. - (Bulletin of the United States Geological Survey, no. 179). - P. 426.
  27. ↑ Hu Yaoming, Meng Jin, Clark J. M. A new tritylodontid from the Upper Jurassic of Xinjiang, China // Acta Palaeontologica Polonica , 2009, 54 (12). - P. 385-391. - DOI : 10.4202 / app.2008.0053 .
  28. ↑ 1 2 Kemp T. S. The origin and early radiation of the therapsid mammal-like reptiles: a palaeobiological hypothesis // Journal of Evolutionary Biology , 2006, 19 (4). - P. 1231-1247. - DOI : 10.1111 / j.1420-9101.2005.01076.x .
  29. ↑ Gow C. E. The importance of Karoo fossils in the search for mammal origins // Palaeontologia africana , 1997, 33 . - P. 23-27.
  30. ↑ Carroll, vol. 2, 1993 , p. 188, 194-197, 200-201, 208-213, 219.
  31. ↑ Baur G. , The History of the Pelycosauria, with a Description of the Genus Dimetrodon , Cope // Transactions of the American Philosophical Society. New Series , 1889, 20 (1). - P. 5-62. - DOI : 10.2307 / 1005488 .
  32. ↑ Ivakhnenko M.F. Tetrapods of the East European Placard - Late Paleozoic territorial-natural complex . - Perm, 2001 .-- 200 p. - (Proceedings of the Paleontological Institute of the Russian Academy of Sciences, vol. 283). - ISBN 5-88345-064-4 . - S. 19-20.
  33. ↑ Ochev, Surkov, 2003 , p. 2-5.
  34. ↑ Osborn H. F. The Reptilian Subclasses Diapsida and Synapsida and the Early History of the Diaptosauria // Memoirs of the AMNH , 1903, 1 (8). - P. 1109-1120.
  35. ↑ Romer A. S., Price L. I. Review of the Pelycosauria. - New York: Geological Society of America, 1940. - x + 538 p. - (Geological Society of America. Special papers, no. 28).
  36. ↑ Williston S. W. The Osteology of the Reptiles . - Cambridge: Harvard University Press , 1925. - xiii + 300 p. - P. 211, 228, 236-237.
  37. ↑ Spindler F., Scott D., Reisz R. R. New Information on the Cranial and Postcranial Anatomy of the Early Synapsid Ianthodon schultzei (Sphenacomorpha: Sphenacodontia), and its Evolutionary Significance // Fossil Record , 2015, 18 . - P. 17-30. - DOI : 10.5194 / fr-18-17-2015 .
  38. ↑ 1 2 3 4 Rubidge B. S., Sidor C. A. Evolutionary Patterns among Permo-Triassic Therapsids // Annual Review of Ecology and Systematics , 2001, 32 . - P. 449-480. - DOI : 10.1146 / annurev.ecolsys.32.08.081501.114113 .
  39. ↑ Carroll, vol. 2, 1993 , p. 222-224.
  40. ↑ Huttenlocker A. K., Rega E. The Paleobiology and Bone Microstructure of Pelycosaurian-Grade Synapsids // Forerunners of Mammals: Radiation, Histology, Biology / Ed. by A. Chinsamy-Turan. - Bloomington, Indiana: Indiana University Press, 2011 .-- xv + 330 p. - ISBN 978-0-253-35697-0 . - P. 91-120.
  41. ↑ Benton, 2005 , p. 120-124.
  42. ↑ 1 2 3 4 5 Kemp T. S. The Origin and Radiation of Therapsids // Forerunners of Mammals: Radiation, Histology, Biology / Ed. by A. Chinsamy-Turan. - Bloomington, Indiana: Indiana University Press, 2011 .-- xv + 330 p. - ISBN 978-0-253-35697-0 . - P. 3-30.
  43. ↑ Amson E., Laurin M. On the Affinities of Tetraceratops insignis , an Early Permian Synapsid // Acta Palaeontologica Polonica , 2011, 56 (2). - P. 301-312. - DOI : 10.4202 / app.2010.0063 .
  44. ↑ Liu Jun, Rubidge B. S., Li Jinling. New Basal Synapsid Supports Laurasian Origin for Therapsids // Acta Palaeontologica Polonica , 2009, 54 (3). - P. 393-400. - DOI : 10.4202 / app.2008.0071 .
  45. ↑ Brink A. S. The taxonomic position of the Synapsida // South African Journal of Science , 1963, 59 (5). - P. 153-159.
  46. ↑ Ivakhnenko, 2008 , p. 102.
  47. ↑ Marjanović D., Laurin M. Fossils, Molecules, Divergence Times, and the Origin of Lissamphibians // Systematic Biology , 2007, 56 (3). - P. 369-388. - DOI : 10.1080 / 10635150701397635 .
  48. ↑ 1 2 Kermack D. M., Kermack K. A. The Evolution of Mammalian Characters . - Dordrecht: Springer Science & Business Media , 2013 .-- x + 149 p. - ISBN 978-1-4214-0040-2 . - P. 10-13.
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Links

  • Laurin M., Reisz R. R. Tree of Life - Synapsida . 1997.
  • Synapsida - Pelycosauria
  • Transitional Vertebrate Fossils
  • Synapsids - Extinct animal wiki (rus.)
Source - https://ru.wikipedia.org/w/index.php?title= Synapsids&oldid = 101530658


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