В данной статье приведён примерный перечень ископаемых переходных форм, то есть останков существ, которые демонстрируют примитивные черты по сравнению с наиболее развитыми представителями своего таксона. Окаменелости перечислены в определённой последовательности, показывая переход от одной таксономической группы к другой и представляя значительные шаги в эволюции основных черт живых организмов в различных эволюционных линиях. Эти изменения часто представляют собой серьезные изменения в анатомии, связанные с образом жизни, например, приобретение оперённых крыльев птицами или ног тетраподами. Как отмечал ещё Дарвин, палеонтологическая летопись страдает опеределённой неполнотой[1].
В идеале этот список должен включать в себя только «истинные» переходные окаменелости, представляющие прямых предков, из которых далее развились известные таксономические группы. Однако большинство представленных здесь находок являются вымершими боковыми ветвями, более или менее тесно связанными с настоящими предковыми видами[2]. Все они также включают уникальные черты, характерные для своих эволюционных линий. Ископаемые, демонстрирующие небольшое количество таких уникальных черт, называются «переходными». Если же ископаемое имеет множество уникальных черт, не наблюдаемых ни в предковых, ни в производных группах, оно определяется как «промежуточное». Поскольку все виды всегда подвергаются естественному отбору, сам термин «переходное ископаемое» не является корректным. Однако это часто используемый термин и полезная концепция в эволюционной биологии. Окаменелости, представленные в списке, отражают важные шаги в развитии основных функций организмов в различных эволюционных линиях и в этом смысле соответствуют общепринятой интерпретации этого термина.
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Это неполный список, который, возможно, никогда не будет удовлетворять определённым стандартам полноты. Вы можете дополнить его из авторитетных источников.
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An early member of the Tetrapodomorpha, the piscine line leading to tetrapods, Osteolepis?! is generalised enough to give a fair approximation of the common ancestor of tetrapods and lungfish.[6]
Fish
A small to medium sized sarcopterygian fish with internal nostrils and pectoral fins stiffened by bony components broadly homologous to the humerus and radius/ulna found in tetrapods.[6]
Though not on the evolutionary path to tetrapods, Eusthenopteron is of fairly general build and is very well known, serving as an iconic model organism in tetrapod evolution.[7]
A medium sized,inly pelagic fish, Eusthenopteron[англ.]*inly use the pectoral and pelvic fins for navigation, and the tail for propulsion.[7] The fin was of diphycercal, foreshadowing the straightening of the spine and the evolution of a contiguous fin in fish like Panderichthys[англ.]
A large, predatory shallow water fish. As common in shallow water fish, the pectoral and pelvic fins were flexible and paddle-like for propulsion.[8] The dorsal and anal fins are lost, the tail fin contiguous.[9] The spiracles were short and wide, indication large amount of oxygen were taken up by the lungs rather than through the gills.[10]
"Fish" with stout, fleshy pectoral fins with a joint between the innermost and the two next bony elements, corresponding to the elbow in higher tetrapods. The cleithrum bone was free of the skull, functioning as anchoring for the pectoral fins, and at the same time allowing for movement of the neck.[12][13]
Though fragmentary, the find includes a shoulder blade (Cleitrum bone) as well as elements of the limbs, which shows it had comparable limbs Ichthyostega[англ.]* and Hynerpeton[англ.]*, indicating feet rather than fins.
Possibly oldest animal to have feet rather than fins.[16]
A large, dorso-ventrally flattened predatory fish with a well armoured labyrinthodont-like skull. While the fins themselves has not been found, the shoulder girdle is essentially similar to that of Acanthostega, indicating it too had feet rather than fins.[16]
Together with Ichthyostega[англ.]* the sole early labyrinthodont known from fairly complete skeletons. It is the oldest animal known to have feet rather than fins, thusking it a true tetrapod and Древнейший известный unquestionable ichthyostegalian.[17]
First known animal with toes rather than fins. The feet were broad and paddle-like, adapted for movement in water.[18] It retained functional gills in adulthood, behind a fleshy operculum.
Fairly closely related to Acanthostega[англ.]*. It possibly represent an early (and ultimately unsuccessful) line adapted to moving on land by inchworm-like movements.
Together with Acanthostega the sole early labyrinthodont known from fairly complete skeletons.
Early labyrinthodont with polydactylous, paddle-like feet and reinforced vertebrae and neural spines. It probably spent time on land, yet retained gills and a tail with fin rayes.
A large animal with paddle-like six-toed feet. It did however not have gills in adulthood, and is thus the oldest labyrinthodont known to depend entirely on breathing with its lungs.[21]
Intermediate between the earlier Ichthyostegalian and the later, more advanced labyrinthodonts.
Despite an extra toe on the forelimbs, Pederpes had limbs that terminated in feet adapted primarely for walking rather than paddles for combined swimming and walking like the earlier groups.[22]
The Limnoscelis was originally described as a "cotylosaur" (early reptiles) together with the other diadectomorphans. Today the large-bodied diadectomorphs are thought to have had a larval stage, falling close to, but just outside the amphibian/reptile divide.
A large, predatory reptile-like amphibian. The limbs are extremely heavily built, indicating it fed on slow moving prey.
An early anapsid reptile, considered to be ancestral to both the synapsid and sauropsid lines, and thus the oldest representative of the crown group amniotes.
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Древнейший известный archosaur, Proterosuchus was one of the largest land reptile during the Early Triassic, about the size of to today's Komodo dragons. It looked somewhat crocodile-like, with sprawling legs, long jaws, powerful neck muscles and a long tail. A distinct proterosuchid trait is the peculiar hook-shaped mouth.
Known from a somewhat fragmentary find, Spondylosoma was possibly an early dinosaur, or near dinosaur.[34] It has however also been classified as a rauisuchian.[35]
A small (1 meter, ~ 10 kg) bipedal carnivore with numerous sharp teeth. It was a swift digigrade runner. The forelimbs were half the length of the hindlimbs and the hands had five fingers
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The find is represented only by a hind leg, but one that is very bird-like. It belonged to a small maniraptoran dinosaur with long, pennaceous feathers on its hind legs and (in all likelihood) arms.
Known for its mosaic of avian and theropod characteristics Archaeopteryx is both the first primitive bird in the fossil record and Идин из первых transitional fossils discovered.
Traditionally seen as the first proper bird, though it is not directly ancestral to modern birds.[39] An excellent intermediate form between dinosaurs and birds. Capable of gliding, but lacking alula and keel, it could likely not sustain powered flight.
With its short tail and toothless beak, Confuciusornis is very modern looking compared to Archaeopteryx[англ.]*. The toothless beak is however a case of convergent evolution, as more advanced birds retained teeth, illustration the sometimes confusing mosaic evolution of the dinosaur-bird transition.
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An advanced member of the family Sphenacodontidae, from which the therapsids (advanced synapsids) evolved
A pelycosaur-grade synapsid. At up to 4 meters, Dimetrodon was one of the largest animals of its time. The distinct sail of the backkes it the most recognized synapsid known
Primitive traits
Cold blooded metabolism dependent of external heat source (hence the "sail")[44]
Sprawling gait
No secondary palate
No enlarged side teeth in the lower jaw
Derived traits
Distinctly elongated 2nd and 3rd tooth on the maxilla, corresponding to the canine inmmals. The first canine generally longer than the second.[45]
A European ape which is considered to be the predecessor of the great apes.
Some objections have been raised to this fossils status due to its location in Spain, but Pierolapithecus is likely a transitional taxon between generalized apes and the lineage which led to great apes.
Pleisomorphic traits
Relatively short fingers and walked in a similar quadrupedal fashion like baboons
Lacks adaptations for both gibbon-style brachaition as well as derived knuckle-walking like in chimpanzee's and gorilla's
Derived traits
Flat, wider rib cage like great apes for tree-climbing
The clavicle is large and similar to modern chimps suggesting a dorsally positioned scapula
Intermediate between extinct quadrupedal and bipedal apes. While the relationship between some species are being revised, Australopithecus afarensis[англ.]* is considered to be, by most experts, the ancestor to all later hominids.
Very successful hominid, which was probably ancestral to both modern humans and neanderthals. Probably the first hominid to leave and successfully colonize territories outside of Africa.
Homo rhodesiensis was the immediate ancestor of modern humans which evidently displaced the neanderthals in Europe and the island 'hobbits' of southeast Asia. H. rhodesiensis evolved from Homo erectus[англ.]* about half a million years ago but still retains some primitive characteristics such as relatively thick bones and molars larger than modern humans.
Ancestral to modern humans.
Primitive traits
Large teeth
Heavy brow ridge
Extremely robust build in most groups
Derived traits
Rounder, less broad based cranium
Larger brain size, approaching (and sometimes exceeding) modern values
↑Darwin, C (1859) On the Origin of Species. Chapter 10: On the Imperfection of the Geological Record.
↑Weishampel D.B. (1996). Fossils, Phylogeny, and Discovery: A Cladistic Study of the History of Tree Topologies and Ghost Lineage Durations. Journal of Vertebrate Paleontology. 16 (2): 191–197. doi:10.1080/02724634.1996.10011307. JSTOR4523710.
↑ 12R. Cloutier. Taxonomic review of Eusthenopteron foordi. // Devonian Fishes and Plants of Miguasha, Quebec, Canada. — Dr. Friedrich Pfeil, München, 1996. — P. 487–502.
↑ 12Anderson, J. S.; Reisz, R. R.; Scott, D.; Fröbisch, N. B.; Sumida, S. S. (2008). A stem batrachian from the Early Permian of Texas and the origin of frogs and salamanders. Nature. 453: 515–518. doi:10.1038/nature06865. PMID18497824. {{cite journal}}: Неизвестный параметр |author-separator= игнорируется (справка); Шаблон цитирования имеет пустые неизвестные параметры: |author-name-separator= (справка)
↑Estes, R., and O. A. Reig. (1973): The early fossil record of frogs: a review of the evidence. Pp. 11-63 In J. L. Vial (Ed.), Evolutionary Biology of the Anurans: Contemporary Research onjor Problems. University of Missouri Press, Columbia.
↑Moss J.L. (1972). The Morphology and phylogenetic relationship of the Lower Permian tetrapod Tseajaia campi Vaughn (Amphibia: Seymouriamorpha). University of California Publications in Geological Sciences. 98: 1–72.
↑Berman, D.S.; Sumida, S.S.; Lombard, R.E. (1992). Reinterpretation of the temporal and occipital regions in Diadectes and the relationship of diadectomorphs. Journal of Paleontology. 66: 481–499.
↑ 12Gauthier J., Kluge, A.G., & Rowe, T. (1988) The early evolution of the Amniota. In: M. J. Benton (ed.) The phylogeny and classification of the tetrapods, Volume 1: amphibians, reptiles, birds (1): pp 103-155. Oxford: Clarendon Press.
↑R. L. Paton, T. R. Smithson and J. A. Clack, "An amniote-like skeleton from the Early Carboniferous of Scotland", (abstract), Nature 398, 508-513 (8 April 1999)
↑Langer, M.C. (2004). Basal Saurischia. In: Weishampel, D.B., Dodson, P., and Osmólska, H. (eds.). The Dinosauria (second edition). University of California Press:Berkeley, 25-46. ISBN 0-520-24209-2
↑Galton, P.M. (2000). Are Spondylosoma and Staurikosaurus (Santaria Formation, Middle-Upper Triassic, Brasil) the oldest saurischian dinosaurs?. Palaontologische Zeitschrift. 74 (3): 393–423.
↑R.N.rtinez et al. A basal dinosaur from the dawn of the dinosaur era in southwestern Pangaea. Science, Vol. 331, 14 January 2011, p. 206.
↑Chinsamy A.,rtin L.D., Dobson P. (1998). Bone microstructure of the diving Hesperornis and the volant Ichthyornis from the Niobrara Chalk of western Kansas. Cretaceous Research. 19 (2): 225–235. doi:10.1006/cres.1997.0102.{{cite journal}}: Википедия:Обслуживание CS1 (множественные имена: authors list) (ссылка)
↑Romer, A.S. (1940). Review of the Pelycosauria. Special Papers of the Geological Siciety of America. 28: 1–538. {{cite journal}}: Неизвестный параметр |coauthors= игнорируется (|author= предлагается) (справка)
↑GA Floridesa, Kalogiroua SA; SA; Wrobelb, L Tassoub (2001). Natural environment and thermal behavior of Dimetrodon limbatus". Journal of Thermal Biology. 26 (1): 15–20. {{cite journal}}: Неизвестный параметр |author-separator= игнорируется (справка); Шаблон цитирования имеет пустые неизвестные параметры: |author-name-separator= (справка)
↑Ruben, J.A. The Evolution ofmmalian Endothermy // Forerunners ofmmals. — Bloomington : Indiana University Press, 2011. — P. 272–286. — ISBN 0-253-35697-0.
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