Largest prehistoric animals

The largest prehistoric animals include both vertebrate and invertebrate species. Many of them are described below, along with their typical range of size (for the general dates of extinction, see the link to each). Many species mentioned might not actually be the largest representative of their clade due to the incompleteness of the fossil record and many of the sizes given are merely estimates since no complete specimen have been found. Their body mass, especially, is largely conjecture because soft tissue was rarely fossilized. Generally the size of extinct species was subject to energetic[1] and biomechanical constraints.[2]

From left to right: a polar bear (Ursus maritimus), a giraffe (Giraffa camelopardalis), †Paraceratherium transouralicum, †Patagotitan mayorum, two humans (Homo sapiens), †Palaeoloxodon recki, an African bush elephant (Loxodonta africana) and a white rhino (Ceratotherium simum)

Non-mammalian synapsids (Synapsida)

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Caseasaurs (Caseasauria)

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The herbivorous Alierasaurus was the largest caseid and the largest amniote to have lived at the time, with an estimated length around 6–7 m (20–23 ft).[3] Cotylorhynchus hancocki is also large, with an estimated length and weight of at least 6 m (20 ft)[4] and more than 500 kg (1,100 lb).[5]

Edaphosaurids (Edaphosauridae)

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Size comparison of some species of Edaphosaurus

The largest edaphosaurids were Lupeosaurus at 3 m (9.8 ft) long[6] and Edaphosaurus, which could reach even more than 3 m (9.8 ft) in length.[7]

Sphenacodontids (Sphenacodontidae)

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The biggest carnivorous synapsid of Early Permian was Dimetrodon, which could reach 4.6 m (15 ft) and 250 kg (550 lb).[8] The largest members of the genus Dimetrodon were also the world's first fully terrestrial apex predators.[9]

Tappenosauridae

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The Middle Permian Tappenosaurus was estimated at 5.5 m (18 ft) in length, nearly as large as the largest dinocephalians.[10]

Therapsids (Therapsida)

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Anomodonts (Anomodontia)

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Lisowicia compared to a human

The plant-eating dicynodont Lisowicia bojani is the largest-known of all non-mammalian synapsids, at about 4.5 m (15 ft) long, 2.6 m (8 ft 6 in) tall, and 9,000 kg (20,000 lb) in body mass.[11][12][13] However, in 2019 its weight was later more reliably estimated by modelling its mass from the estimated total volume of its body. These estimates varied depending on the girth of its rib cage and the amount of soft tissue modelled around the skeleton, with an overall average weight of 5.9 metric tons (6.5 short tons), and a lowermost estimate with minimal body fat and other tissues at 4.9 metric tons (5.4 short tons) and a maximum of 7 metric tons (7.7 short tons) at its bulkiest.[14]

Biarmosuchians (Biarmosuchia)

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Adult specimens of the Late Permian Eotitanosuchus (a possible synonym to Biarmosuchus[15]) could reach 6 m (20 ft) in length and over 600 kg (1,300 lb) in weight.[15]

Dinocephalians (Dinocephalia)

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Anteosaurus overviewing the landscape

The largest carnivorous non-mammalian synapsids was the dinocephalian Anteosaurus, which was 5–6 m (16–20 ft) long, and weighed 500–600 kg (1,100–1,300 lb).[16][17] Fully grown Titanophoneus from the same family Anteosauridae likely had a skull of 1 m (3.3 ft) long.[17]

Gorgonopsians (Gorgonopsia)

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Photo-reconstruction of Inostrancevia

Inostrancevia latifrons is the largest known gorgonopsian, with a skull length of more than 60 cm (24 in), a total length approaching 3.5 m (11 ft) and a mass of 300 kg (660 lb).[18] Rubidgea atrox is the largest African gorgonopsian, with skull of nearly 45 cm (18 in) long.[19] Other large gorgonopsians include Dinogorgon with skull of ~40 cm (16 in) long,[20] Leontosaurus with skull of almost 40 cm (16 in) long,[19] and Sycosaurus with skull of ~38 cm (15 in) long.[19]

Therocephalians (Therocephalia)

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The largest of therocephalians is Scymnosaurus,[21][22] which reached a size of the modern hyena.[23]

Non-mammalian cynodonts (Cynodontia)

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Mammals (Mammalia)

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Non-therian mammals

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Gobiconodonts (Gobiconodonta)

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A reconstruction of Repenomamus

The largest gobiconodont and the largest well-known Mesozoic mammal was Repenomamus.[27][28][29][30][31][32] The known adult of Repenomamus giganticus reached a total length of around 1 m (3 ft 3 in) and an estimated mass of 12–14 kg (26–31 lb).[29] With such parameters it surpassed in size several small theropod dinosaurs of the Early Cretaceous.[33] Gobiconodon was also a large mammal,[31][32] it weighed 5.4 kilograms (12 lb),[29] had a skull of 10 cm (3.9 in) in length, and had 35 cm (14 in) in presacral body length.[34]

Multituberculates (Multituberculata)

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The largest multituberculate,[35] Taeniolabis taoensis is the largest non-therian mammal known, at a weight possibly exceeding 100 kg (220 lb).[36]

Monotremes (Monotremata)

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Photo-reconstruction of Murrayglossus hacketti (Zaglossus hacketti) by paleoartist Roman Uchytel

Metatherians (Metatheria)

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Thylacosmilus compared to a human

Marsupials (Marsupialia)

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  • The largest known marsupial, and the largest metatherian, is the extinct Diprotodon, about 3 m (9.8 ft) long, standing 2 m (6 ft 7 in) tall and weighing up to 2,786 kg (6,142 lb).[48] Fellow vombatiform Palorchestes azael was similar in length being around 2.5 m (8.2 ft), with body mass estimates indicating it could exceed 1,000 kg (2,200 lb).[49]
  • The largest known carnivorous marsupial was Thylacoleo carnifex. Measurements taken from a number of specimens show they averaged 101 to 164 kg (223 to 362 lb) in weight.[50][51]
  • The largest known kangaroo was an as yet unnamed species of Macropus, estimated to weigh 274 kg (604 lb),[52] larger than the largest known specimen of Procoptodon, which could grow up to 2 m (6 ft 7 in) and weigh 230 kg (510 lb).[53] Some species from the genus Sthenurus were similar in size or a bit larger than the extant grey kangaroo (Macropus giganteus).[54]
  • The largest potoroid ever recorded was Borungaboodie, which was nearly 30% bigger than the largest living species and weighted up to 10 kg (22 lb).[55]

Non-placental eutherians

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Restoration of Coryphodons

Cimolestans (Cimolesta)

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The largest known cimolestan is Coryphodon, 1 m (3 ft 3 in) high at the shoulder, 2.5 m (8 ft 2 in) long[56][57] and up to 700 kg (1,500 lb) of mass.[58] Barylambda was also a huge mammal, at 650 kg (1,430 lb).[59] Wortmania and Psittacotherium from the group Taeniodonta were among the largest mammals of the Early Paleocene.[60] Lived as soon as half a million years after K–Pg boundary, Wortmania reached 20 kg (44 lb) in body mass. Psittacotherium, which appeared two million years later, reached 50 kg (110 lb).[60]

Leptictids (Leptictida)

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The largest leptictid ever discovered is Leptictidium tobieni from the Middle Eocene of Germany. It had a skull 101 mm (4.0 in) long, head with trunk 375 mm (14.8 in) long, and tail 500 mm (20 in) long.[61] Close European relatives from the same family Pseudorhyncocyonidae had skulls of 67–101 mm (2.6–4.0 in) in length.[61]

Tenrecs and allies (Afroscida)

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The larger of the two species of bibymalagasy (Plesiorycteropus madagascariensis), extinct tenrec relatives from Madagascar, is estimated to have weighed from 10 to 18 kilograms (21 to 40 lb).[62]

Even-toed ungulates (Artiodactyla)

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The extinct Hippopotamus gorgops is the most massive of the fossil even-toed ungulates
 
A skeleton of Bison latifrons
  • The largest of Bovinae as well as the largest bovid was Bison latifrons. It reached a weight from 1,250 kg (2,760 lb)[69][70] to 2,000 kg (4,400 lb),[71] 4.75 m (15.6 ft) in length, shoulder height of 2.31 m (7.6 ft),[72] and had horns that spanned 2.13 m (7 ft 0 in).[73] The North American Bison antiquus reached up to 4.6 m (15 ft) long, 2.27 m (7.4 ft) tall, weight of 1,588 kg (3,501 lb),[74] and horn span of 1 m (3.3 ft).[72] The African Pelorovis reached 2 t (2.2 short tons) in weight and had bony cores of the horns about 1 m (3 ft 3 in) long.[75] Another enormous bovid, the african giant buffalo (Syncerus antiquus) reached 3 m (9.8 ft) in length from muzzle to the end of the tail, 1.85 m (6.1 ft) in height at the withers, 1.7 m (5.6 ft) in height at the hindquarters,[76][77] and the distance between the tips of its horns was as large as 2.4 m (7 ft 10 in).[76] Aside from local populations and subspecies of extant species, such as the gaur population in Sri Lanka, European bison in British Isles, Caucasian wisent and Carpathian wisent, the largest modern extinct bovid is aurochs (Bos primigenius) with an average height at the shoulders of 155–180 cm (61–71 in) in bulls and 135–155 cm (53–61 in) in cows, while aurochs populations in Hungary had bulls reaching 155–160 cm (61–63 in).[78] The kouprey (Bos sauveli), reaching 1.7–1.9 m (5 ft 7 in – 6 ft 3 in) in shoulder height,[79][80] has existed since the Middle Pleistocene[81] and is also considered to be possibly extinct.[82][83]
  • The long-legged Megalotragus is possibly the largest known alcelaphine bovid,[84] bigger than the extant wildebeest.[85] The tips of horns of M. priscus were located at a distance of about 1.2 m (3 ft 11 in) from each other.[86]
 
Megaloceros giganteus
  • The extinct cervid Irish elk (Megaloceros giganteus) reached over 2.1 m (7 ft) in height, 680 kg (1,500 lb) in mass and could have antlers spanning up to 4.3 m (14 ft) across, about twice the maximum span for a moose's antlers.[87][88] The giant moose (Cervalces latifrons) reached 2.1 to 2.4 m (6.9 to 7.9 ft) high[89] and was twice as heavy as the Irish elk but its antler span at 2.5 m (8.2 ft) was smaller than that of Megaloceros.[90][91] North American stag-moose (Cervalces scotti) reached 2.5 metres (8.2 ft) in length and a weight of 708.5 kilograms (1,562 lb).[92][93]
  • The largest known giraffid, aside from the extant giraffe, is Sivatherium, with a body weight of 1,250 kg (2,760 lb).[94]
  • The largest protoceratid was Synthetoceras, it reached 2 m (6 ft 7 in) long and 150–200 kg (330–440 lb) in mass.[95][96]
  • The largest known wild suid to ever exist was Kubanochoerus gigas, having measured up to 500 kg (1,100 lb) and stood around 1 m (3 ft 3 in) tall at the shoulder.[97] Megalochoerus could be similar in size, possibly weighing 303 kg (668 lb) or 526 kg (1,160 lb).[98]
  • The largest tayassuid extinct Platygonus species were similar in size to modern peccaries especially giant peccary, at around 1 m (3.3 ft) in body length, and had long legs, allowing them to run well. They also had a pig-like snout and long tusks which were probably used to fend off predators.[99]
  • The largest camelid was Titanotylopus from the Miocene of North America. It possibly reached 2,485.6 kg (5,480 lb) and a shoulder height of over 3.4 m (11 ft).[100][101] The Syrian camel (Camelus moreli) was twice as big as the modern camels.[102] It was 3 m (9.8 ft) at the shoulder[103] and 4 m (13 ft) tall.[102] Camelops had legs to be 20% longer than that of Dromedary, and was about 2.3 m (7 ft 7 in) tall at the shoulder and weighed about 1,000 kg (2,200 lb).[104]
  • The anoplotheriid Anoplotherium is thought to have been capable of reaching up to 271 kg (597 lb) in the case of A. commune and 229 kg (505 lb) in the case of A. latipes.[105] A. latipes in particular could have measured more than 2.5 m (8 ft 2 in) in length and 1.25 m (4 ft 1 in) in shoulder height. Because it was probably capable of facultative bipedalism, it could have been capable of standing over 3 m (9.8 ft) tall.[106]

Cetaceans (Cetacea)

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Size comparison between a human and two species of Basilosaurus, B. cetiodes (dark blue) and B. isis
  • The heaviest archeocete, and possibly the heaviest known mammal was Perucetus, with weight estimated at 85–340 t (84–335 long tons; 94–375 short tons), while length is estimated at 17.0–20.1 meters (55.8–65.9 ft),[107] possibly rivalling the Blue Whale in mass. However, Motani and Pyenson in 2024 argued that it is extremely difficult for Perucetus to rival or exceed the blue whale in weight. They discussed that since Perucetus is much shorter than the blue whale in length, it should be at least 3.375 times denser or 1.83 times fatter to weigh heavier, which is impossible for vertebrates whose whole-body density range from 0.75 to 1.2. Motani and Pyenson tested the hypotheses of Bianucci and colleagues by performing various body mass estimation methods: the regression-based and volumetric mass estimation resulted in 60–114 t (59–112 long tons; 66–126 short tons) for a length range of 17–20 m (56–66 ft), though the likely body mass range would fall within 60–70 t (59–69 long tons; 66–77 short tons) . They also claimed that the previous estimation is inflated by assumed isometry, and that the effect from pachyostosis on the estimation of body mass is not negligible as it resulted in underestimation.[108] The longest of known Eocene archeocete whales was Basilosaurus at 17–20 m (56–66 ft) in length.[109][110][111]
  • The largest squalodelphinid was Macrosqualodelphis at 3.5 m (11 ft) in length.[112]
  • Some Neogene rorquals were comparable in size to modern huge relatives. Parabalaenoptera was estimated to be about the size of the modern gray whale,[113] about 16 m (52 ft) long. Some balaenopterids perhaps rivaled the blue whale in terms of size,[113] though other studies disagree that any baleen whale grew that large in the Miocene.[114]

Odd-toed ungulates (Perissodactyla)

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Relative sizes of †Paraceratherium, †Elasmotherium, white rhino, Indian rhino, black rhino and Sumatran rhino compared to a human
 
Life restoration of Moropus elatus
  • One of the largest known perissodactyls, and the second largest land mammal (see Palaeoloxodon namadicus) of all time was the hornless rhino Paraceratherium. The largest individual known was estimated at 4.8 m (15.7 ft) tall at the shoulders, 7.4 m (24.3 ft) in length from nose to rump, and 17 t (18.7 short tons) in weight.[115][116]
  • A large specimen of an unnamed species of the related Dzungariotherium has been estimated to be around 20.6 metric tonnes.[117]
  • Some prehistoric horned rhinos also grew to large sizes. The biggest Elasmotherium reached up to 5–5.2 m (16–17 ft) long,[118] 2.5 m (8 ft 2 in) high[119] and weighed 3.5–5 t (3.9–5.5 short tons).[120][118][119] Such parameters make it the largest rhino of the Quaternary.[120] Woolly rhinoceros (Coelodonta antiquitatis) of the same time reached 1,100–1,500 kg (2,400–3,300 lb)[121] or 2,000 kg (4,400 lb),[122][123] 1.93 m (6 ft 4 in) at the shoulder height and 4.6 m (15 ft) in length.[124]
  • Metamynodon, an amynodontid, reached 4 m (13 ft) in length, comparable to Hippopotamus in measurement and shape.[125]
  • The giant tapir (Tapirus augustus) was the largest tapir ever, at about 623 kg (1,373 lb)[126] and 1 m (3.3 ft) tall at the shoulders.[127] Earlier, this mammal was estimated even bigger, at 1.5 m (4.9 ft) tall, and assigned to the separate genus Megatapirus.[127]
  • The largest known lophiodont is Lophiodon, with L. lautricense being estimated to reach more than 2,000 kg (4,400 lb) in weight.[128]
  • One of the biggest chalicotheres was Moropus.[129] It stood about 2.4 metres (8 ft) tall at the shoulder.[130]
  • Late Eocene perissodactyls from the family Brontotheriidae attained huge sizes. The North American Megacerops (also known as Brontotherium[131]) reached 2.5 m (8 ft 2 in) tall at the shoulders,[132] 5 m (16 ft) in length,[131] and 3 t (6,600 lb) in weight.[133] Embolotherium from Asia was equal in size.[134]
  • The largest prehistoric horse was Equus giganteus of North America. It was estimated to grow to more than 1,250 kg (1.38 short tons) and 2 m (6 ft 7 in) at the shoulders.[135] The largest anchitherine equid was Hypohippus at 403 to 600 kg (888 to 1,323 lb), comparable to large modern domestic horses.[136][137] Megahippus is another large anchitheriine. With the body mass of 266.2 kg (587 lb) it was much heavier than most of its close relatives.[136]
  • Among the largest-sized genera of palaeotheres, close relatives of horses, is Palaeotherium, with P. giganteum being estimated to reach weights of more than 700 kg (1,500 lb).[138] Previously until the naming of P. giganteum in 1994, P. magnum was considered the largest species of Palaeotherium,[139] potentially reaching 1.3 m (4 ft 3 in) in shoulder height and 2.52 m (8 ft 3 in) in length.[140] Another palaeothere Cantabrotherium is estimated to have weighed about 600 kg (1,300 lb).[138]

Phenacodontids (Phenacodontidae)

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The largest known phenacodontid is Phenacodus. It was 1.5 m (4 ft 11 in) long[141] and weighed up to 56 kg (123 lb).[142]

Dinoceratans (Dinocerata)

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The largest known dinoceratan was Eobasileus with skull length of 102 cm (40 in), 2.1 m (6 ft 11 in) tall at the back and 1.5 m (4 ft 11 in) tall at the shoulder.[143] Another huge animal of this group was Uintatherium, with skull length of 76 cm (30 in), 1.5 m (4 ft 11 in) tall at the shoulder,[143] 4 m (13 ft) in length and 2.25 t (2.48 short tons), the size of a rhinoceros.[144] Despite their large size, Eobasileus as well as Uintatherium had a very small brain.[143][144]

Carnivores (Carnivora)

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Caniformia

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Arctodus simus reconstruction
 
Chapalmalania, the giant procyonid
 
Skeletal mount of Epicyon haydeni
  • The largest terrestrial mammalian carnivore and the largest known bear, as well as the largest known mammalian land predator of all time, was Arctotherium angustidens, the South American short-faced bear. A humerus of A. angustidens from Buenos Aires indicates that the males of the species could have weighed 1,588–1,749 kg (3,501–3,856 lb) and stood at least 3.4 m (11 ft) tall on their hind-limbs.[145][146] Another huge bear was the giant short-faced bear (Arctodus simus), with the average weight of 625 kg (1,378 lb) and the maximum estimated at 957 kg (2,110 lb).[147] There is a guess that the largest individuals of this species could reached even larger mass, up to 1,200 kg (2,600 lb).[145] The extinct cave bear (Ursus spelaeus) was also heavier than many recent bears. Largest males weighed as much as 1,000 kg (2,200 lb).[148] Ailuropoda baconi from the Pleistocene was larger than the modern giant panda (Ailuropoda melanoleuca).[149]
  • The biggest odobenid and one of the biggest pinnipeds to have ever existed is Pontolis magnus, with a skull length of 60 cm (24 in) (twice as large as the skulls of modern male walruses)[150] and having a total body length of more than 4 m (13 ft).[151][152] Only the modern male elephant seals (Mirounga) reach similar sizes.[151] The second largest prehistoric pinniped is Gomphotaria pugnax with a skull length of nearly 47 cm (19 in).[150]
  • One of the largest of prehistoric otariids is Thalassoleon, comparable in size to the biggest extant fur seals. An estimated weight of T. mexicanus is no less than 295–318 kg (650–701 lb).[153]
  • The biggest known mustelid to ever exist was likely the giant otter, Enhydriodon. It exceeded 3 m (9.8 ft) in length, and would have weighed in at around 200 kg (440 lb), much larger than any other known mustelid, living or extinct.[154][155][156] There were other giant otters, like Siamogale, at around 50 kg (110 lb)[157] and Megalenhydris, which was larger than a modern-day giant river otter.[158] Megalictis was the largest purely terrestrial mustelid[159] (although Enhydriodon had recently been mentioned as the largest mustelid that also happens to be a terrestrial predator[154]). Similar in size to the jaguar, Megalictis ferox had even wider skull, almost as wide as of the black bear.[159] Megalictis had a powerful bite force, allowing it to eat large prey and crush bones, as modern hyenas and jaguars can.[159] Another large-bodied mustelid was the superficially cat-like Ekorus from the Miocene of Africa. At almost 44 kg (97 lb), the long-legged Ekorus was about the size of a wolf[160] and filling a similar to leopards ecological niche before big cats came to the continent.[161] Other huge mustelids include Perunium[162] and hypercarnivorous Eomellivora, both from the Late Miocene.[163]
  • The heaviest procyonid was possibly South American Chapalmalania. It reached 1.5 metres (4.9 ft) in body length with a short tail and 150 kilograms (330 lb), comparable in size to an American black bear (Ursus americanus).[164] Another huge procyonid was Cyonasua, which weighted about 15–25 kg (33–55 lb), about the same size as a medium-sized dog.[165]
  • The largest canid of all time was Epicyon haydeni, which stood 90 cm (35 in) tall at the shoulder, had a body length of 2.4 m (7.9 ft) and weighed 100–125 kg (220–276 lb),[166][167][168] with the heaviest known specimen weighing up to 170 kg (370 lb).[42] The extinct dire wolf (Aenocyon dirus) reached 1.5 m (4.9 ft) in length and weighed between 50 and 110 kg (110 and 243 lb).[42][169] The largest wolf (Canis lupus) subspecies ever existed in Europe is the Canis lupus maximus from the Late Pleistocene of France. Its long bones are 10% larger than those of extant European wolves and 20% longer than those of C. l. lunellensis.[170] The Late Pleistocene Italian wolf was morphometrically close to C. l. maximus.[171]
  • The largest bear-dog was a species of Pseudocyon weighing around 773 kg (1,704 lb), representing a very large individual.[172]

Feliformia

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Comparison between Amphimachairodus giganteus and the modern domestic cat
 
Skeleton of Panthera atrox at the La Brea Tar Pits Museum

Hyaenodonts (Hyaenodonta)

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The largest hyaenodont was Simbakubwa at 1,500 kg (3,300 lb).[196] Another giant hyaenodont, Megistotherium reached 500 kg (1,100 lb)[42] and had a skull of 66.4 cm (26.1 in) in length.[197]

Oxyaenids (Oxyaenidae)

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Sarkastodon

The largest known oxyaenid was Sarkastodon weighing in at 800 kg (1,800 lb).[42]

Mesonychians (Mesonychia)

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Some mesonychians reached a size of a bear. Such large were Mongolonyx from Asia[198] and Ankalagon from North America.[199][200] Another large mesonychian is Harpagolestes with a skull length of a half a meter in some species.[198]

Bats (Chiroptera)

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Found in Quaternary deposits of South and Central Americas, Desmodus draculae had a wingspan of 0.5 m (20 in) and a body mass of up to 60 g (2.1 oz). Such proportions make it the largest vampire bat that ever evolved.[201]

Hedgehogs, gymnures, shrews, and moles (Eulipotyphla)

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Deinogalerix skeleton

The largest known animal of the group Eulipotyphla was Deinogalerix,[202] measuring up to 60 cm (24 in) in total length, with a skull up to 21 cm (8.3 in) long.[203]

Rodents (Rodentia)

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The giant beaver (Castoroides ohioensis)
  • Several of the extinct South American dinomyids were much bigger than the modern rodents. Josephoartigasia monesi was the largest-known rodent of all time, approximately weighing an estimated 480–500 kg (1,060–1,100 lb).[204] Phoberomys pattersoni weighed 125–150 kg (276–331 lb).[204] Both Josephoartigasia and Phoberomys reached about 1.5 m (4 ft 11 in) tall at the shoulder.[205] Another huge dinomyid, Telicomys gigantissimus had a minimal weight of 200 kg (440 lb).[205]
  • Amblyrhiza inundata from the family Heptaxodontidae was a massive animal, it weighed 50–200 kg (110–440 lb).[206][205]
  • The largest beaver was the giant beaver (Castoroides) of North America. It grew over 2 m in length and weighed roughly 90 to 125 kg (198 to 276 lb), also making it one of the largest rodents to ever exist.[207]
  • The largest old world porcupine are the Hystrix refossa was larger than living porcupines. It was approximately 20% larger than its closest relative, the living Indian porcupine (H. indica), reaching lengths of over 115 cm (45 in).

Rabbits, hares, and pikas (Lagomorpha)

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The biggest known prehistoric lagomorph is Minorcan giant lagomorph Nuralagus rex at 12 kg (26 lb).[208]

Pangolins (Pholidota)

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The largest pangolin was the extinct Manis palaeojavanica[209] Its total length is measured up to 2.5 m (8.2 ft).[210]

Primates (Primates)

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Reconstruction of Gigantopithecus blackii

Elephants, mammoths, and mastodons (Proboscidea)

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Steppe mammoth skeletal mount
 
Mounted Deinotherium skeleton
  • The elephant Palaeoloxodon namadicus has been suggested to have been the largest land mammal ever, based on a particularly large partial femur which was estimated to have belonged to an individual 22 t (24.3 short tons) in weight and about 5.2 m (17.1 ft) tall at the shoulder, though the author of the estimate said that this was speculative and should be treated with caution.[115] In 2023, a publication by Gregory S. Paul and Larramendi estimated that another specimen identified as cf. P. namadicus, also only known from a partial femur, would have weighed 18–19 tonnes (40,000–42,000 lb).Other authors have noted that weight estimates for proboscideans based on single bones can lead to estimates that are "highly improbable" compared to accurate estimates from complete skeletons.[228] In 2024, Biswas, Chang and Tsai estimated a maximum shoulder height of over 4.5 metres (15 ft) and suggested that the body mass for 5 measured specimens ranged from 13.2 to 18.5 tonnes (29,000 to 41,000 lb) from specimens from Taiwan.[229] The largest individual reported individual of the steppe mammoth of Eurasia (Mammuthus trogontherii) was estimated to reach 4.5 m (14.8 ft) at the shoulders and 14.3 t (15.8 short tons) in weight.[115][230] Stegodon zdanskyi, the biggest species of Stegodon, was 13 t (14.3 short tons) in body mass.[115] Another enormous proboscidean is Stegotetrabelodon syrticus, over 4 m (13 ft) in height and 11 to 12 t (12.1 to 13.2 short tons) in weight.[115] The Columbian mammoth (Mammuthus columbi) was about 4 m (13.1 ft) tall at the shoulder but didn't weigh as much as other huge mammoths. Its average mass was 9.5 t (10.5 short tons) with one unusually large specimen about 12.5 t (13.8 short tons).[115] Columbian mammoths had very long tusks. The largest known mammoth tusk, 4.9 m (16 ft) long, belonged to this species.[231]
  • The mammutid "Mammut" borsoni is one of the largest known proboscideans and land mammals. The average fully-grown male is estimated to have been 4.1 m (13 ft) tall and weighed about 16 t (17.6 short tons), with very large males possibly rivalling the estimated size of the largest Palaeoloxodon namadicus.[115] This species also had the longest tusks of any animals with the largest recorded specimen being 5.02 m (16.5 ft) long from basis to tip along the curve.[232]
  • Deinotherium was the largest proboscidean in Deinotheriidae family. Bones retrieved in Crete confirm the existence of specimen 4.1 m (13 ft) tall at the shoulders and more than 14 t (15.4 short tons) in weight.[115]

Sea cows (Sirenia)

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According to reports, Steller's sea cows have grown to 8 to 9 m (26 to 30 ft) long as adults, much larger than any extant sirenians.[233] The weight of Steller's sea cows is estimated to be 8–10 t (8.8–11.0 short tons).[234]

With its direct ancestor the Cuesta sea cow being around 9 m (30 ft) long and possibly 10 tonnes (11 short tonnes) in weight.[235]

Arsinoitheres (Arsinoitheriidae)

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Skeleton of Arsinoitherium

The largest known arsinoitheriid was Arsinoitherium. A. zitteli would have been 1.75 m (5 ft 9 in) tall at the shoulders, and 3 m (9.8 ft) long.[236][237] A. giganteum reached even larger size than A. zitteli.[238]

Hyraxes (Hyracoidea)

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Some of the prehistoric hyraxes were extremely large compared to modern small relatives. The largest hyracoid ever evolved is Titanohyrax ultimus.[239] With the mass estimation in rage of 600 kg (1,300 lb) to over 1,300 kg (2,900 lb) it was close in size to Sumatran rhinoceros.[240] Another enormous hyrax is Megalohyrax which had skull of 391 mm (15.4 in) in length[241] and reached the size of tapir.[239][242] More recent Gigantohyrax was three times as large as the extant relative Procavia capensis,[243] although it is noticeably smaller than earlier Megalohyrax and Titanohyrax.[244]

Desmostylians (Desmostylia)

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Desmostylus skeletal diagram

The largest known desmostylian was a species of Desmostylus, with skull length of 81.8 cm (32.2 in) and comparable in size to the Steller's sea cow.[245]

Paleoparadoxia is also known as one of the largest desmostylians, with body length of 3.03 m (9.9 ft).[246]

Armadillos, glyptodonts and pampatheres (Cingulata)

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The largest cingulate known is Doedicurus, at 4 m (13 ft) long, 1.5 m (4 ft 11 in) high[144] and reaching a mass of approximately 1,910 to 2,370 kg (2.11 to 2.61 short tons).[citation needed] The largest species of Glyptodon, Glyptodon clavipes, reached 3–3.3 m (9.8–10.8 ft) in length[247][144] and 2 t (2.2 short tons) in weight.[citation needed]

Anteaters and sloths (Pilosa)

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Skeleton and illustration of Megatherium

The largest known pilosan is Eremotherium, a ground sloth with an estimated weight of up to 6.55 t (7.22 short tons) and a length of up to 6 m (20 ft),[248] which is as big as a bull African bush elephant. The closely related ground sloth Megatherium attained similarly large dimensions.[249]

Astrapotherians (Astrapotheria)

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Some of the largest known astrapotherians weighed about 3–4 t (3.3–4.4 short tons), including the genus Granastrapotherium[250] and some species of Parastrapotherium (P. martiale).[251] The skeleton remains suggests that the species Hilarcotherium miyou was even larger, with a weight of 6.456 t (7.117 short tons).[252]

Litopterns (Litopterna)

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The largest known litoptern was Macrauchenia, which had three hoofs per foot. It was a relatively large animal, with a body length of around 3 m (9.8 ft).[253]

Notoungulates (Notoungulata)

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The largest notoungulate known of complete remains is Toxodon. It was about 2.7 m (8 ft 10 in) in body length, and about 1.5 m (4 ft 11 in) high at the shoulder and resembled a heavy rhinoceros. Although incomplete, the preserved fossils suggests that Mixotoxodon were the most massive member of the group, with a weight about 3.8 t (4.2 short tons).[254]

Pyrotherians (Pyrotheria)

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The largest mammal of the South American order Pyrotheria was Pyrotherium at 2.9–3.6 m (9 ft 6 in – 11 ft 10 in) in length and 1.8–3.5 t (4,000–7,700 lb) in weight.[255]

Reptiles (Reptilia)

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Lizards and snakes (Squamata)

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Megalania skeletal reconstruction on Melbourne Museum steps
  • Mosasaurs are the largest-known squamates. The largest-known mosasaur is likely Mosasaurus hoffmanni, estimated at more than 17 m (56 ft) in length,[256][257] however these estimations are based on heads and total body length ratio 1:10, which is unlikely for Mosasaurus, and probably that ratio is about 1:7.[258] Another giant mosasaur is Tylosaurus, estimated at 10–14 m (33–46 ft) in length.[259][260] Another mosasaur, Prognathodon reached similar sizes.
  • The largest known prehistoric snake is Titanoboa cerrejonensis, estimated at 12.8 m (42 ft) or even 14.3 m (47 ft)[261] in length and 1,135 kg (2,502 lb) in weight,[262] and madtsoiid Vasuki indicus which is estimated to reach between 11–15 m (36–49 ft).[263] A close rival in size to those snakes is palaeophiid marine snake Palaeophis colossaeus, which may have been around 9 m (30 ft) in length[262][264][265] or even up to 12.3 m (40 ft).[266] Another known very large fossil snake is Gigantophis garstini, estimated at 9.3–10.7 m (31–35 ft) in length,[267][268] although later study shows smaller estimation about 6.6–7.2 m (22–24 ft).[269] The largest fossil python is Liasis dubudingala with length roughly 9 m (30 ft).[270] The largest viper as well as the largest venomous snake ever recorded is Laophis crotaloides from the Early Pliocene of Greece. This snake reached over 3 m (9.8 ft) in length and 26 kg (57 lb) in weight.[271][272] Another huge fossil viper is indeterminate species of Vipera. With a length of around 2 m (6 ft 7 in) it was one of the biggest predators of Mallorca during the Early Pliocene.[273] The largest known blind snake is Boipeba tayasuensis with estimated total length of 1.1 m (3 ft 7 in).[274]
  • The largest known land lizard is probably megalania (Varanus priscus) at 7 m (23 ft) in length.[275] As extant relatives, megalania could have been venomous and in that case this lizard was also the largest venomous vertebrate ever evolved.[276] However, maximum size of this animal is subject to debate.[277] Recent studies have estimated it at 5.5 m (18 ft) long.[278]

Turtles, tortoises and close relatives (Pantestudines)

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Cryptodira

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  • The largest known turtle ever was Archelon ischyros at 5 m (16 ft) long and 2,200 kg (4,900 lb).[279] Possible second-largest sea turtle was Protostega at 3.9 m (13 ft) in total body length.[280][281] There is even a larger specimen of this genus from Texas estimated at 4.2 m (14 ft) in total length.[282][280] Partially known Cratochelone is estimated to reach 4 m (13 ft) in total length.[283] Another huge prehistoric sea turtle is the Late Cretaceous Gigantatypus, estimated at over 3.5 m (11 ft) in length.[284] Psephophorus terrypratchetti from the Eocene attained 2.3–2.5 m (7.5–8.2 ft) in body length.[285]
  • The largest tortoise was Megalochelys atlas at up to 2 m (6.6 ft) in shell length[286] and weighing 0.8–1.0 t (1,800–2,200 lb).[133] M. margae had carapace of 1.4–2 m (4.6–6.6 ft) long; an unnamed species from Java reached at least 1.75 m (5.7 ft) in carapace length.[287] The Cenozoic Titanochelon were also larger than extant giant tortoises, with a shell length of up to 2 m (6 ft 7 in).[288][289] Other giant tortoises include Centrochelys marocana at 1.8–2 m (5.9–6.6 ft) in carapace length and Mesoamerican Hesperotestudo sp. at 1.5 m (4.9 ft) in carapace length.[287]
  • The largest trionychid ever recorded is indeterminate specimen GSP-UM 3019 from the Middle Eocene of Pakistan. Bony carapace of GSP-UM 3019 is 120 cm (3.9 ft) long and 110 cm (3.6 ft) wide indicates the total carapace diameter (with soft margin) about 2 m (6.6 ft).[290] Drazinderetes tethyensis from the same formation had a bony carapace 80 cm (2.6 ft) long and 70 cm (2.3 ft) wide.[290] Another huge trionychid is North American Axestemys byssinus at over 2 m (6.6 ft) in total length.[291]

Side-necked turtles (Pleurodira)

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The fossil of carapace of Stupendemys geographicus

The largest freshwater turtle of all time was the Miocene podocnemid Stupendemys, with an estimated parasagittal carapace length of 2.86 m (9 ft 5 in) and weight of up to 1,145 kg (2,524 lb).[292] Carbonemys cofrinii from the same family had a shell that measured about 1.72 m (5 ft 8 in),[293][294][295] complete shell was estimated at 1.8 m (5.9 ft).[296]

Macrobaenids (Macrobaenidae)

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The largest macrobaenids were the Early Cretaceous Yakemys, Late Cretaceous Anatolemys, and Paleocene Judithemys. All reached 70 cm (2.3 ft) in carapace length.[297]

Meiolaniformes

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Skeleton of Meiolania platyceps

The largest meiolaniid was Meiolania. Meiolania platyceps had a carapace 100 cm (3.3 ft) long[287] and probably reached over 3 m (9.8 ft) in total body length.[298] An unnamed Late Pleistocene species from Queensland was even larger, up to 200 cm (6.6 ft) in carapace length.[287] Ninjemys oweni reached 100 cm (3.3 ft) in carapace length[287] and 200 kg (440 lb) in weight.[299]

Sauropterygians (Sauropterygia)

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Placodonts and close relatives (Placodontiformes)

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Placodus was among the largest placodonts, with a length of up to 3 m (9.8 ft).[300]

Nothosaurs and close relatives (Nothosauroidea)

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The largest nothosaur as well as the largest Triassic sauropterygian was Nothosaurus giganteus at 7 m (23 ft) in length.[301]

Plesiosaurs (Plesiosauria)

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  • The largest known plesiosauroid was an indeterminate specimen possibly belonging to Aristonectes (identified as cf. Aristonectes sp.), with a body length of 11–11.9 metres (36–39 ft) and body mass of 10.7–13.5 metric tons (11.8–14.9 short tons).[302] Another long plesiosauroid was Albertonectes at 11.2–11.6 metres (37–38 ft).[303] Thalassomedon rivaled it in size, with its length at 10.86–11.6 m (35.6–38.1 ft).[304] Other large plesiosauroids are Styxosaurus and Elasmosaurus. Both reached some more than 10 m (33 ft) in length.[305][306] Hydralmosaurus (previously synonymized with Elasmosaurus and Styxosaurus) reached 9.44 m (31.0 ft) in total body length.[306] In past, Mauisaurus was considered to be more than 8 m (26 ft) in length,[307][306] but later it was determined as nomen dubium.[308]
 
Size estimation of three species of Pliosaurus.

Proterosuchids (Proterosuchidae)

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Proterosuchus fergusi is the largest known proterosuchid with a skull length of 47.7 cm (18.8 in) and a possible body length of 3.5–4 m (11–13 ft).[318]

Erythrosuchids (Erythrosuchidae)

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Life reconstruction of Erythrosuchus africanus

The largest erythrosuchid was Erythrosuchus africanus with a maximum length of 4.75–5 m (15.6–16.4 ft).[319]

Phytosaurs (Phytosauria)

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Some of the largest known phytosaurs include Redondasaurus with a length of 6.4 m (21 ft)[320] and Smilosuchus with a length of more than 7 m (23 ft).[321]

Non-crocodylomorph pseudosuchians (Pseudosuchia)

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Size comparison of Sillosuchus to a human

Crocodiles and close relatives (Crocodylomorpha)

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Large crocodylomorphs (†Deinosuchus, †Purussaurus, †Gryposuchus, †Euthecodon, †Sarcosuchus, and modern Crocodylus porosus) compared to a human
 
Skull size comparison of large crocodylomorphs (Left to right, †Razanandrongobe, †Machimosaurus, †Barinasuchus, †Sarcosuchus, modern Crocodylus porosus, and †Purussaurus)

Aegyptosuchids (Aegyptosuchidae)

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The Late Cretaceous Aegisuchus was originally estimated to reach 15 m (49 ft) in length by the lower estimate and as much as 22 m (72 ft) by the upper estimate although a length of over 15 m is likely a significant overestimate.[331] However, this estimation is likely to be a result of miscalculation, and its length would be only around 3.9 m (13 ft).[332]

Crocodylians (Crocodylia)

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Paralligatorids (Paralligatoridae)

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The largest paralligatorid was likely Kansajsuchus, estimated at up to 8 m (26 ft) long.[353]

Tethysuchians (Tethysuchia)

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  • Some extinct pholidosaurids reached giant sizes. In the past, Sarcosuchus imperator was believed to be the largest crocodylomorph, with initial estimates proposing a length of 12 m (39 ft) and a weight of 8 t (8.8 short tons).[354] However, recent estimates have now shrunk to a length of 9 to 9.5 m (29.5 to 31.2 ft) and a weight of 3.5 to 4.3 metric tons (3.9 to 4.7 short tons).[355] Related to Sarcosuchus, Chalawan thailandicus could have reached more than 10 m (33 ft) in length,[356] although other estimates suggest 7–8 m (23–26 ft).[341]
  • The largest dyrosaurid was Phosphatosaurus gavialoides, estimated at 9 m (30 ft) in length.[357][341]

Stomatosuchids (Stomatosuchidae)

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Stomatosuchus, a stomatosuchid, was estimated at 10 m (33 ft) in length.[358]

Notosuchians (Notosuchia)

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  • Some of largest terrestrial notosuchian crocodylomorphs were the Miocene sebecid Barinasuchus, with a skull of 95–110 cm (37–43 in) long, and Eocene sebecid Dentaneosuchus with estimated mandible length of 1 m (3.3 ft).[359][360] Various estimates suggest a possible length of these animals between 3–10 m (9.8–32.8 ft). Using proportion of Stratiotosuchus which is also large to have 47 cm (19 in) long skull,[361] Barinasuchus is estimated to have length at least 6.3 m (21 ft).[359][360]
  • Other huge notosuchian, although only known from fragmentary material, is an early member Razanandrongobe, which skull size may exceeded that of Barinasuchus and overall length may be around 7 m (23 ft).[362][363]

Thalattosuchians (Thalattosuchia)

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Plesiosuchus compared to a human

Basal crocodylomorphs

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Redondavenator was the largest Triassic crocodylomorph ever recorded,[370] with a skull of at least 60 cm (2.0 ft) in length.[371][372] Another huge basal crocodylomorph was Carnufex[370] at 3 m (9.8 ft) long even through that is immature.[373]

Pterosaurs (Pterosauria)

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Hatzegopteryx (A-B), Arambourgiania (C) and Quetzalcoatlus sp. (D-E)

Choristoderes (Choristodera)

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The largest known choristoderan, Kosmodraco dakotensis (previously known as Simoedosaurus dakotensis[385]) is estimated to have had a total length of around 5 m (16 ft).[386][385]

Tanystropheids (Tanystropheidae)

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Reconstruction of Tanystropheus, note that anatomical features based on smaller species T. longobardicus, while size is based on T. hydroides

Tanystropheus, the largest of all tanystropheids, reached up to 5 m (16 ft) in length.[387]

Thalattosaurs (Thalattosauria)

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The largest species of thalattosaur, Miodentosaurus brevis grew to more than 4 m (13 ft) in length.[388] The second largest member of this group is Concavispina with a length of 3.64 m (11.9 ft).[389]

Ichthyosaurs (Ichthyosauria)

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Shonisaurus popularis (green) and Shastasaurus sikanniensis (red) compared with a human

In April 2018, paleontologists announced the discovery of a previously unknown ichthyosaur that may have reached lengths of 26 m (85 ft) making it one of the largest animals known, rivaling some blue whales in size.[390][391] These remains were later named Ichthyotitan and it has been estimated to reach up to 25 m (82 ft), which makes it the largest ichthyosaur and the largest marine reptile ever.[392] Another large ichthyosaur was the Late Triassic Shastasaurus sikanniensis at 21 m (69 ft) in length[393][394] and 81.5 t (180,000 lb) in weight.[395] Another, larger ichthyosaur was found in 1850 in Aust.[396] Its remains seemed to surpass the measurements of the other ichthyosaur, but the researchers commented that the remains were too fragmentary for a size estimate to be made.[396] Another huge ichthyosaur was Shonisaurus popularis at 15 m (49 ft) in length and 29.7 t (65,000 lb) in weight.[394] The largest Middle Triassic ichthyosaur as well as the largest animal of that time was Cymbospondylus youngorum at 17.65 m (57.9 ft) in length[395] and 44.7 t (99,000 lb) in weight.[395]

Tangasaurids (Tangasauridae)

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The largest tangasaurid was Hovasaurus with an estimated snout-vent length of 30–35 cm (12–14 in) and a tail of 60 cm (24 in).[397]

Pareiasaurs (Pareiasauria)

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Largest pareiasaurs reached up to 3 m (9.8 ft) in length. Such sizes had Middle Permian Bradysaurus, Embrithosaurus, and Nochelesaurus from South Africa,[398] and the Late Permian Scutosaurus from Russia.[398] The most robust Scutosaurus had 1.16 t (2,600 lb) in body mass.[398]

Captorhinids (Captorhinidae)

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The heavy built Moradisaurus grandis, with a length of 2 m (6 ft 7 in),[399] is the largest known captorhinid.[400] The second largest captorhinid was Labidosaurikos with the largest adult skull specimen 28 cm (11 in) long.[401]

Non-avian dinosaurs (Dinosauria)

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Sauropodomorphs (Sauropodomorpha)

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The largest of non-sauropod sauropodomorphs ("prosauropod") was Euskelosaurus. It reached 12.2 m (40 ft) in length and 2 t (2.2 short tons) in weight.[402] Another huge sauropodomorph Yunnanosaurus youngi reached 13 m (43 ft) long.[403]

Sauropods (Sauropoda)

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Size comparison of selected giant sauropod dinosaurs (from left to right): Supersaurus, Argentinosaurus, Diplodocus, Mamenchisaurus, and Sauroposeidon
  • A mega-sauropod, Maraapunisaurus fragillimus (previously known as Amphicoelias fragillimus), is a contender for the largest-known dinosaur in history. It has been estimated at 58–60 m (190–197 ft) in maximum length and 122,400 kg (269,800 lb) in weight.[404] Unfortunately, the fossil remains of this dinosaur have been lost.[404] More recently, it was estimated at 35–40 m (115–131 ft) in length and 80–120 t (180,000–260,000 lb) in weight.[405]
  • Known from the incomplete and now disintegrated remains, the Late Cretaceous Bruhathkayosaurus matleyi was an anomalously large sauropod.[406] Informal estimations suggested as huge parameters as 45 m (148 ft) in length and 139–220 t (306,000–485,000 lb) in weight.[407] Some estimation however, suggests 37 m (121 ft) and 95 t (209,000 lb) but it still much heavier than most other sauropods.[407] More recent estimations by Gregory Paul in 2023 has placed its weight range around 110 t (240,000 lb) to a 170 t (370,000 lb). If true, it would make Bruhathkayosaurus the single largest terrestrial animal to have walked the earth and would have rivalled the largest blue whale recorded.[408]
  • BYU 9024, a massive cervical vertebra found in Utah,[409] may belong to a Barosaurus lentus[410][411] or Supersaurus vivianae[412] of a huge size, possibly 45–48 m (148–157 ft) in length and 60–66 t (132,000–146,000 lb) in body mass.[410][413] Supersaurus vivianae itself may have been the longest dinosaur yet discovered as a study of 3 specimens suggested length of 39 m (128 ft) or over 40 m (130 ft).[412]
 
Mounted skeleton of Mamenchisaurus sinocanadorum
 
Reconstructed skeleton of Argentinosaurus

Other huge sauropods include Argentinosaurus, Alamosaurus, and Puertasaurus with estimated lengths of 30–33 m (98–108 ft) and weights of 50–80 t (55–88 short tons).[422] Patagotitan was estimated at 37 m (121 ft) in length[423] and 57 t (63 short tons) in average weight,[424] and was similar in size to Argentinosaurus and Puertasaurus.[425] Giant sauropods like Supersaurus, Sauroposeidon, and Diplodocus probably rivaled them in length but not in weight.[404] Dreadnoughtus was estimated at 49 t (108,000 lb) in weight[424] and 26 m (85 ft) in length, but the most complete individual was immature when it died.[426] Turiasaurus is considered the largest dinosaur from Europe,[427][428] with an estimated length of 30 m (98 ft) and a weight of 50 t (55 short tons).[422][428] However, lower estimates at 21 m (69 ft) and 30 t (66,000 lb) would make it smaller than the Portuguese Lusotitan, which reached 24 m (79 ft) in length and 34 t (75,000 lb) in weight.[429]

Many large sauropods are still unnamed and may rival the current record holders:

  • The "Archbishop", a large brachiosaur that was discovered in 1930. As of October 2023, a scientific paper on the specimen is still in progress.[430]
  • Brachiosaurus nougaredi is yet another large brachiosaur from Early Cretaceous North Africa. The remains have been lost, but the sacrum drawing remains. It suggests a sacrum of almost 1.3 m (4.3 ft) long,[431] making it the largest dinosaur sacrum discovered so far, except those of Argentinosaurus and Apatosaurus.[432]
  • In 2010, the femur of a large sauropod was discovered in France. The femur suggests an animal that grew to immense sizes.[433]

Non-avian theropods (Theropoda)

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Size comparison of selected giant theropod dinosaurs (from left to right): Spinosaurus , Giganotosaurus, Tyrannosaurus, Mapusaurus, and Carcharodontosaurus

Armoured dinosaurs (Thyreophora)

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The largest-known thyreophoran was Ankylosaurus at 9 m (30 ft) in length and 6 tonnes (6.6 short tons) in weight.[444][445] Stegosaurus was also 9 m (30 ft) long[428] but around 5 tonnes (5.5 short tons) tonnes in weight.[citation needed]

Pachycephalosaurs (Pachycephalosauria)

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The largest pachycephalosaur was the eponymous Pachycephalosaurus. Previously claimed to be at 7 m (23 ft) in length,[428] it was later estimated about 4.5 metres (14.8 ft) long and a weight of about 450 kilograms (990 lb).[446]

Ceratopsians (Ceratopsia)

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Size comparison of several members of Ceratopsidae (from left to right): Nasutoceratops, Styracosaurus, Centrosaurus, Pachyrhinosaurus, Triceratops, Pentaceratops, Chasmosaurus, and Anchiceratops

The largest ceratopsian known is Triceratops horridus, along with the closely related Eotriceratops xerinsularis both with estimated lengths of 9 m (30 ft). Pentaceratops and several other ceratopsians rival them in size.[447] Titanoceratops had one of the longest skull of any land animal, at 2.65 m (8.7 ft) long.[448]

Ornithopods (Ornithopoda)

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From left to right: Shantungosaurus giganteus, Magnapaulia laticaudus, Edmontosaurus annectens, Saurolophus angustirostris, Hypsibema missouriensis, Charonosaurus jiayinensis, Iguanodon bernissartensis

Birds (Aves)

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From left to right: a human, †Anomalopteryx didiformus, †Megalapteryx didinus, †Euryapteryx curtus, Casuarius casuarius, †Pachyornis elephantopus, Rhea americana, Struthio camelus, †Dinornis novaezealandiae, †Aepyornis maximus

The largest bird in the fossil record may be the extinct elephant bird species Aepyornis maximus of Madagascar, whose closest living relative is the kiwi. Giant elephant birds exceeded 2.3 metres (7.5 ft) in height, and average a mass of 850 kg (1,870 lb)[468]

The largest fowl was the mihirung Dromornis stirtoni of Australia. It exceeded 2.7 m (8.9 ft) in height, and average a mass of 500 kilograms (1,100 lb)[469]

Another contender is Brontornis burmeisteri, an extinct flightless bird from South America which reached a weight of 319 kg (703 lb) and a height of approximately 2.8 metres (9.2 ft).[470]

The tallest recorded bird was Pachystruthio dmanisensis, a relative of the ostrich. This particular species of bird stood at 3.5 metres (11.5 ft) tall and average a mass of 450 kg (990 lb)[471]

The largest known flightless neoave was the terror bird Paraphysornis brasiliensis of South America, the Brazilian terror bird exceeded 240 kg (530 lb) in mass,[472]

Table of heaviest extinct bird species

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Rank Common Name Binomial Name Average mass
kg (lb)
1 Aepyornis maximus 850 kg (1,870 lb)[468]
2 Dromornis stirtoni 500 kg (1,100 lb)[469]
3 Pachystruthio dmanisensis 450 kg (990 lb)[471]
4 Brontornis burmeisteri 319 kg (703 lb)[470]
5 Dromornis planei 300 kg (660 lb)[469]
6 Genyornis newtoni 275 kg (606 lb)[469]
7 East Asian ostrich Struthio anderssoni 270 kg (600 lb)[473]
8 Dromornis murrayi 250 kg (550 lb)[474]
9 Paraphysornis brasilienis 240 kg (530 lb)[472]
10 Aepyornis hildebrandti 235 kg (518 lb)[468]
11 Dromornis australis 220 kg (490 lb)[469]
12 Gastornis gigantea 200 kg (440 lb)[469]
13 Ilbandornis lawsoni 175 kg (386 lb)[469]
14 Devincenzia pozzi 161 kg (355 lb)[475]
15 Ilbandornis woodburnei 150 kg (330 lb)[469]
16 Titanis walleri 150 kg (330 lb)[476]
17 Gastornis parisiensis 135 kg (298 lb)[477]
18 South Island giant moa Dinornis robustus 125 kg (276 lb)[478]
19 Gargantuavis philoinos 120 kg (260 lb)[479]
20 Palaeeudyptes klekowskii 116 kg (256 lb)[480]
21 North Island giant moa Dinornis novaezealandiae 100 kg (220 lb)[478]
22 Phorusrhacos longissmus 94 kg (207 lb)[470]
23 Mullerornis modestus 80 kg (180 lb)[468]
24 Heavy-footed moa Pachyornis elephantopus 80 kg (180 lb)[478]
25 Giant Teratorn Argentavis magnificens 71 kg (157 lb)[481]
26 Barawertornis tedfordi 70 kg (150 lb)[469]
27 Mesembriornis incertus 70 kg (150 lb)[482]
28 Crested moa Pachyornis australis 67 kg (148 lb)[478]
29 Eastern moa Emeus crassus 58 kg (128 lb)[478]
30 Broad-billed moa Euryapteryx curtus 47.5 kg (105 lb)[478]
31 Upland moa Megalapteryx didinus 40 kg (88 lb)[478]
32 Bush moa Anomalopteryx didiformis 40 kg (88 lb)[478]
33 Sylviornis neocaledoniae 30.5 kg (67 lb)[482]
34 Eremopezus eocaenus 30 kg (66 lb)[483]
35 Mantell's moa Pachyornis geranoides 27 kg (60 lb)[478]
36 Patagornis marshi 23 kg (51 lb)[482]
37 Teratornis merriami 22.5 kg (50 lb)[484]
38 Pelagornis sandersi 21.7 kg (48 lb)[485]
39 Llallawavis scagliai 18 kg (40 lb)[486]
40 Giant darter Giganhinga kiyuensis 17.7 kg (39 lb)[487]
41 Giant swan Cygnus falconeri 16 kg (35 lb)[488]
42 Leptoptilos robustus 16 kg (35 lb)[489]
43 Haast's eagle Hieraaetus moorei 12 kg (26 lb)[478]
44 Dodo Raphus cucullatus 10.2 kg (22 lb)[490]
45 South Island adzebill Aptornis defossor 10 kg (22 lb)[478]
46 South Island goose Cnemiornis calcitrans 10 kg (22 lb)[478]
47 North Island adzebill Aptornis otidiformis 8 kg (18 lb)[478]
48 North Island goose Cnemiornis gracilis 8 kg (18 lb)[478]
49 Hercules parrot Heracles inexpectatus 7 kg (15 lb)[491]
50 Spectacled cormorant Phalacrocorax perspicillatus 6.4 kg (14 lb)[492]

Enantiornitheans (Enantiornithes)

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One of the largest enantiornitheans was Enantiornis,[493] with a length in life of around 78.5 cm (30.9 in), hip height of 34 cm (13 in), weight of 6.75 kg (14.9 lb),[494] and wingspan comparable to some of the modern gulls, around 1.2 m (3 ft 11 in).[493] Gurilynia was the largest Mesozoic bird from Mongolia, with a length of 53 cm (21 in), hip height of 23.2 cm (9.1 in), and weight of 2.1 kg (4.6 lb).[494]

Avisauridae

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Two Mirarce sitting on a head of ceratopsian dinosaur

The Late Cretaceous Avisaurus was almost as large as Enantiornis. It had a wingspan around 1.2 m (3 ft 11 in),[493] a length of 72 cm (28 in), hip height of 31.5 cm (12.4 in), and weight of 5.1 kg (11 lb).[494] Even larger could be the Soroavisaurus. One tibiotarsus (PVL-4033) indicates an animal with a length of 80 cm (31 in), hip height of 35 cm (14 in), and weight of 7.25 kg (16.0 lb).[494] However, according to Walker and Dyke (2009) which considered PVL-4033 as Martinavis sp., its tibiotarsus length is 85.6 mm (3.37 in),[495] much shorter than that of Lectavis (156 mm (6.1 in) tibiotarsus)[496] which the same book estimated a length of 41 cm (16 in), hip height of 30 cm (12 in), and weight of 1.15 kg (2.5 lb).[494] Mirarce was comparable in size to a turkey, much larger than most of other enantiornitheans.[497]

Pengornithidae

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One of the biggest Early Cretaceous enantiornithine bird was Pengornis at 50 cm (1.6 ft) in length[428] and skull length of 54.7 mm (2.15 in).[498]

Gargantuaviidae

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Gargantuavis is the largest known bird of the Mesozoic, a size ranging between the cassowary and the ostrich, and a mass of 140 kg (310 lb) like modern ostriches.[499] In 2019 specimens MDE A-08 and IVPP-V12325 were measured at 1.8 m (5 ft 11 in) in length, 1.3 m (4 ft 3 in) in hip height, and 120 kg (260 lb) in weight.[479]

Dromornithiformes

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A cast of Dromornis stirtoni from Australia

The largest dromornithid was Dromornis stirtoni over 3 m (9.8 ft) tall[500] and 528–584 kg (1,164–1,287 lb) in mass for males.[501]

Gastornid (Gastornithiformes)

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Large individuals of Gastornis reached up to 2 m (6 ft 7 in) in height.[502] Weight of Gastornis ranges from 100 kg (220 lb) to 156 kg (344 lb) and sometimes to 180 kg (400 lb) for European specimens and from 160 kg (350 lb) to 229 kg (505 lb) for North American.[503][477][504]

Waterfowl (Anseriformes)

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Reconstruction of Garganornis ballmanni

Possibly flightless, the Miocene Garganornis ballmanni was larger than any extant members of Anseriformes, with 15.3–22.3 kg (34–49 lb) in body mass.[505] Another huge anseriform was the flightless New Zealand goose (Cnemiornis). It reached 15–18 kg (33–40 lb), approaching in size to small species of moa.[506]

Swans (Cygnini)

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The largest known swan was the Pleistocene giant swan (Cygnus falconeri), which reached a bill-to-tail length of about 190–210 cm (75–83 in),[507] a weight of around 16 kg (35 lb), and a wingspan of 3 m (9.8 ft).[488][508][509] The New Zealand swan (Cygnus sumnerensis) weighed up to 10 kg (22 lb), compared to the related extant black swan at only 6 kg (13 lb).[510] The large marine swan Annakacygna yoshiiensis from the Miocene of Japan far exceeded the extant mute swan in both size and weight.[511]

Anatinae

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Finsch's duck (Chenonetta finschi) reached 1–2 kg (2.2–4.4 lb) in weight, surpassing related modern Australian wood duck (800 g (1.8 lb)).[512]

Pelicans, ibises and allies (Pelecaniformes)

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Storks and allies (Ciconiiformes)

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Leptoptilos robustus compared in size to a human

The largest known of Ciconiiformes was Leptoptilos robustus, standing 1.8 m (5 ft 11 in) tall and weighing an estimated 16 kg (35 lb).[515][489] Ciconia maltha is a relatively large species of Ciconia, with a height of over 5 feet (1.5 meters) and a wingspan up to 10 feet (3.0 meters) across.[516]

Cranes (Gruiformes)

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A large true crane (Gruinae) from the late Miocene (Tortonian) of Germany was equal in size to the biggest extant cranes and resembled the long-beaked Siberian crane (Leucogeranus leucogeranus).[517]

Shorebirds (Charadriiformes)

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Miomancalla howardi was the largest known charadriiform of all time, weighing approximately 0.6 kg (1.3 lb) more than the second-largest member, the great auk (Pinguinus impennis).[518]

Hesperornithines (Hesperornithes)

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The largest known of the hesperornithines was Canadaga arctica at 2.2 m (7 ft 3 in) long.[519]

New World vultures (Cathartiformes)

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A skeleton of Teratornis

One of the heaviest flying birds of all time was Argentavis, a Miocene teratornithid. The immense bird had a wingspan estimated up to 5.09–6.5 m (16.7–21.3 ft)[481][520] and a weight up to 70 to 72 kg (154 to 159 lb).[521][481] Argentavis's humerus was only slightly shorter than an entire human arm.[522] Another huge teratorn was Aiolornis, with a wingspan of around 5 m (16 ft).[523] The Pleistocene Teratornis merriami reached 13.7 kg (30 lb) and 2.94–3.38 m (9.6–11.1 ft) in wingspan, with lower size estimates still exceeding the largest specimens of California condor (Gymnogyps californianus).[524]

Seriemas and allies (Cariamiformes)

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Size comparison of Kelenken and a human

The largest known-ever Cariamiforme and largest phorusrhacid or "terror bird" (highly predatory, flightless birds of America) was Brontornis, which was about 175 cm (69 in) tall at the shoulder, could raise its head 2.8 m (9 ft 2 in) above the ground and could have weighed as much as 400 kg (880 lb).[525] The immense phorusrhacid Kelenken stood 3 m (9.8 ft) tall[526][527] with a skull 716 mm (28.2 in) long (460 mm (18 in) of which was beak), had the largest head of any known bird.[526] South American Phorusrhacos stood 2.4-2.7 m (7.9-8.8 ft) tall, and weighed nearly 130 kilograms (290 lb), as much as a male ostrich.[528][529] The largest North American phorusrhacid was Titanis, which reached a height of approximately 2.5 m (8.2 ft),[530] slightly taller than an African forest elephant.

Accipitriforms (Accipitriformes)

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Haast's eagle, the largest bird of prey, attacking moa

The largest known bird of prey ever was the enormous Haast's eagle (Hieraaetus moorei), with a wingspan of 2.6 to 3 m (8 ft 6 in to 9 ft 10 in), relatively short for their size.[531][532] Total length was probably up to 1.4 m (4 ft 7 in) in female[533] and they weighed about 10 to 15 kg (22 to 33 lb).[534] Another giant extinct hawk was Titanohierax about 7.3 kg (16 lb) that lived in the Antilles and The Bahamas, where it was among the top predators.[535] An unnamed late Quaternary eagle from Hispaniola could be 15–30% larger than the modern golden eagle (Aquila chrysaetos).[536] Some extinct species of Buteogallus surpassed their extant relatives in size. Buteogallus borrasi was about 33% larger than the modern great black hawk (B. urubitinga).[537] B. daggetti, also known as "walking eagle", was around 40% larger than the savanna hawk (B. meridionalis).[538] Eyles's harrier (Circus eylesi) from the Pleistocene-Holocene of New Zealand was more than twice heavier than the extant C. approximans.[539]

Moa (Dinornithiformes)

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The tallest known bird was the South Island giant moa (Dinornis robustus), part of the moa family of New Zealand that went extinct about 500 years ago. It stood up to 3.7 m (12 ft) tall,[540] and weighed approximately half as much as a large elephant bird due to its comparatively slender frame.[541]

Tinamous (Tinamiformes)

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MPLK-03, a tinamou specimen that existed during the Late Pleistocene in Argentina, possibly belongs to the modern genus Eudromia and surpacces extant E. elegans and E. formosa in size by 2.2–8% and 6–14%, respectively.[542]

Elephant birds (Aepyornithiformes)

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The largest bird in the fossil record may be the extinct elephant birds (Vorombe, Aepyornis) of Madagascar, which were related to the ostrich. They exceeded 3 m (9.8 ft) in height and 500 kilograms (1,100 lb) in weight.[541]

Ostriches (Struthioniformes)

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With 450 kg (990 lb) in body mass, Pachystruthio dmanisensis from the lower Pleistocene of Crimea was the largest bird ever recorded in Europe. Despite its giant size, it was a good runner.[543] A possible specimen of Pachystruthio from the lower Pleistocene of Hebei Province (China) was about 300 kg (660 lb) in weight, twice heavier than the common ostrich (Struthio camelus).[544] Remains of the massive Asian ostrich (Struthio asiaticus) from the Pliocene[545] indicate a size 20% bigger than adult male of the extant Struthio camelus.[546]

Pigeons and doves (Columbiformes)

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A painting of a live Dodo from the early 1600s.

The largest pigeon relative known was the dodo (Raphus cucullatus), possibly exceeding 1 m (3.3 ft) in height and weighing as much as 28 kg (62 lb), although recent estimates have indicated that an average wild dodo weighed much less at approximately 10.2 kg (22 lb).[547][548]

Pheasants, turkeys, gamebirds and allies (Galliformes)

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The largest known of the Galliformes was likely the giant malleefowl, which could reach 7 kg (15 lb) in weight.[549]

Songbirds (Passeriformes)

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The largest known songbird is the extinct giant grosbeak (Chloridops regiskongi) at 280 mm (11 in) long.[citation needed]

Cormorants and allies (Suliformes)

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The spectacled cormorant or Pallas's Cormorant (Phalacrocorax perspicillatus)
  • The largest known cormorant was the spectacled cormorant of the North Pacific (Phalacrocorax perspicillatus), which became extinct around 1850 and averaged around 6.4 kg (14 lb) and 1.15 m (3 ft 9 in).[492]
  • The largest known darter was Giganhinga with estimated weight about 17.7 kg (39 lb),[487] earlier study even claims 25.7 kg (57 lb).[550]
  • The largest known plotopterid, penguin-like flightless bird was Copepteryx titan that is known from 22 cm (8.7 in) long femur, almost twice as long as that of emperor penguin.[551]

Grebes (Podicipediformes)

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The largest known grebe, the Atitlán grebe (Podylimbus gigas), reached a length of about 46–50 centimetres (18–20 in).[552]

Bony-toothed birds (Odontopterygiformes)

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The largest known of the Odontopterygiformes— a group which has been variously allied with Procellariiformes, Pelecaniformes and Anseriformes and the largest flying birds of all time other than Argentavis were the huge Pelagornis, Cyphornis, Dasornis, Gigantornis and Osteodontornis.[citation needed] They had a wingspan of 5.5–6 m (18–20 ft) and stood about 1.2 m (3 ft 11 in) tall.[citation needed] Exact size estimates and judging which one was largest are not yet possible for these birds, as their bones were extremely thin-walled, light and fragile, and thus most are only known from very incomplete remains.[citation needed]

Woodpeckers and allies (Piciformes)

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The largest known woodpecker is the possibly extinct imperial woodpecker (Campephilus imperialis) with a total length of about 56–60 cm (22–24 in).[553]

Parrots (Psittaciformes)

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The largest known parrot is the extinct Heracles inexpectatus with a length of about 1 meter (3.3 feet).[554]

Penguins (Sphenisciformes)

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Size comparison of the giant penguin Anthropornis nordenskjoeldi

One of the heaviest penguins ever known is Kumimanu fordycei, with a body mass estimate of 148 to 159.7 kg (326 to 352 lb), derived from humerus measurements.[555] Another example is Palaeeudyptes klekowskii of Antarctica, with a bill-to-tail length estimated at 2.02 m (6 ft 8 in) and an estimated body weight of 84.2 kg (186 lb), slightly smaller than previous estimates.[555][556] The Eocene Anthropornis nordenskjoeldi is comparable in size, and was once estimated to reach lengths of 2.05 m (6 ft 9 in) and a weight of 108 kg (238 lb).[557] However, recent estimation from humerus measurements put A. nordenskjoeldi more in the range of 67 kg (148 lb) in weight.[555] Other large penguins include the New Zealand giant penguin (Pachydyptes pondeorsus) weighing around 65.4 to 94.6 kg (144 to 209 lb), and Icadyptes salasi at 52.8 to 73.0 kg (116.4 to 160.9 lb).[558][555]

Owls (Strigiformes)

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The largest known owl of all time was the Cuban Ornimegalonyx at 1,100 mm (43.3 in) tall probably exceeding 9 kg (20 lb).[559]

Amphibians (Amphibia)

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The largest known amphibian of all time was the 9.1 m (30 ft) long temnospondyl Prionosuchus.[560]

Lissamphibians (Lissamphibia)

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Frogs and toads (Anura)

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Size comparison of Beelzebufo

The largest known frog ever was an as yet unnamed Eocene species that was about 58–59-centimetre-long (22.8–23.2 in).[561] The Late Cretaceous Beelzebufo grew to at least 23.2 cm (9.1 in) (snout-vent length), which is around the size of a modern African bullfrog.[562]

Salamanders, newts and allies (Urodela)

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Andrias matthewi size comparison

Diadectomorphs (Diadectomorpha)

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Size comparison of Diadectes

The largest known diacectid, herbivorous Diadectes, was a heavily built animal, up to 3 m (9.8 ft) long, with thick vertebrae and ribs.[565][566]

Anthracosauria

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The largest known anthracosaur was Anthracosaurus, with skull about 40 cm (16 in) in length.[567]

Embolomeri

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Restoration of Pholiderpeton

The longest member of this group was Eogyrinus attheyi, species sometimes placed under genus Pholiderpeton.[568] Its skull had length about 41 cm (16 in).[569]

Temnospondyls (Temnospondyli)

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Scale diagram of small and large specimens of Prionosuchus

The largest known temnospondyl amphibian is Prionosuchus, which grew to lengths of 9 m (30 ft).[560] Another huge temnospondyl was Mastodonsaurus giganteus at 6 m (20 ft) long.[570] Unnamed species of temnospondyl from Lesotho is partial, but possible body length estimation is 7 m (23 ft).[571]

Fishes (Pisces)

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Fishes are a paraphyletic group of non-tetrapod vertebrates.

Jawless fish (Agnatha)

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Conodonts (Conodonta)

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Iowagnathus grandis is estimated to have length over 50 cm (1.6 ft).[572]

Heterostracans (Heterostraci)

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Some members of Psammosteidae such as Obruchevia and Tartuosteus are estimated to reached up to 2 m (6.6 ft).[573]

Thelodonts (Thelodonti)

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Although known from partial materials, Thelodus parvidens (=T. macintoshi) is estimated to reached up to 1 m (3.3 ft).[574]

Cephalaspidomorphs (Cephalaspidomorphi)

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A species of Parameteoraspis reached up to 1 m (3.3 ft).[575]

Spiny sharks (Acanthodii)

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The largest of the now-extinct Acanthodii was Xylacanthus grandis, an ischnacanthiform based on a ~35 cm (14 in) long jaw bone. Based on the proportions of its relative Ischnacanthus, X. grandis had an estimated total length of 2.5 m (8 ft 2 in).[576]

Placoderms (Placodermi)

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Cast of a Dunkleosteus skull

The largest known placoderm was the giant predatory Dunkleosteus. The largest and most well known species was D. terrelli, various estimate put its length around 4.1–10 m (13.5–32.8 ft) in length and 1–4 t (1.1–4.4 short tons) in weight.[577] Another large placoderm, Titanichthys, may have rivaled it in size.[578] Titanichthys is estimated to have a length around 4.1–7.5 m (13–25 ft)[577][579][580][581]

Cartilaginous fish (Chondrichthyes)

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Mackerel sharks (Lamniformes)

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How estimates for the size of Megalodon using different assumptions (brown) compare with the whale shark (blue), great white shark (yellow), and human (black) for scale
  • Species in the extinct genus Otodus were huge. A giant shark, Otodus megalodon[582][583][584] is by far the biggest mackerel shark ever known.[585] Most estimates of megalodon's size extrapolate from teeth, with maximum length estimates up to 10.6–20 m (35–66 ft)[583][584][586] and average length estimates of 10.5 m (34 ft).[587][588] Due to fragmentary remains, there have been many contradictory size estimates for megalodon, as they can only be drawn from fossil teeth and vertebrae.[589]: 87 [590] Mature male megalodon may have had a body mass of 12.6 to 33.9 metric tons (13.9 to 37.4 short tons), and mature females may have been 27.4 to 59.4 metric tons (30.2 to 65.5 short tons), assuming that males could range in length from 10.5 to 14.3 m (34 to 47 ft) and females 13.3 to 17 m (44 to 56 ft).[591] Related to megalodon, Otodus angustidens and O. chubutensis reached the large sizes too. Each was estimated at 9.3 m (31 ft)[592] and 12.2 m (40 ft),[593] respectively.
  • Other giant mackerel sharks were Pseudoscapanorhynchidae from the Cretaceous period. Cretodus had a size range of 9–11 m (30–36 ft) (for C. crassidens),[594] Leptostyrax reached lengths of 6.3–8.3 m (21–27 ft).[595]
  • The Cenozoic Parotodus reached up to 7.6 m (25 ft) in length.[596]
  • The heaviest thresher shark was likely Alopias grandis. It was similar in size or even larger than the extant great white shark and probably did not have an elongated dorsal tail, characteristic of modern relatives.[597]

Ground sharks (Carcharhiniformes)

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The Cenozoic Hemipristis serra was considerably larger than its modern-day relatives and had much larger teeth. Its total length is estimated to be at 6 metres (20 ft) long.[598]

Hybodonts (Hybodontiformes)

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One of the largest hybodontiforms was the Jurassic Asteracanthus with body length of up to 3 m (9.8 ft).[599] Crassodus reifi is known from less materials, however it is estimated that reached over 3 m (9.8 ft).[600]

Ctenacanthiformes

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The largest member of ctenacanthiformes is Saivodus striatus with estimated length around 6–9 m (20–30 ft).[601][577]

Skates and allies (Rajiformes)

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The giant sclerorhynchid Onchopristis reached about 4.25 m (13.9 ft) in length.[602]

Eugeneodont (Eugeneodontida)

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Size comparation of Helicoprion

The largest known eugeneodont is an as-yet unnamed species of Helicoprion discovered in Idaho. The specimens suggest an animal that possibly exceeded 12 m (39 ft) in length.[603] Another fairly large eugeneodont is Parahelicoprion. Being more slimmer than Helicoprion, it reached nearly the same size,[603] possibly up to 12 m (39 ft) in length.[604] Both had the largest sizes among the animals of Paleozoic era.[605][604]

Lobe-finned fish (Sarcopterygii)

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Coelacanths (Actinistia)

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Size estimation of Mawsonia gigas

The largest coelacanth is Cretaceous Mawsonia gigas with estimated total length up to 5.3 m (17 ft). Jurassic Trachymetopon may have reached size close to that, about 5 m (16 ft).[606] An undetermined mawsoniid from the Maastrichtian deposits of Morocco probably reached 3.65–5.52 m (12.0–18.1 ft) in length.[607][606]

Lungfish (Dipnoi)

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Cretaceous Ceratodus sp. from Western Interior is estimated to had a length around 4 m (13 ft).[608]

Stem-tetrapods (Tetrapodomorpha)

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Reconstruction of Rhizodus
 
Reconstruction of Hyneria

Ray-finned fish (Actinopterygii)

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Acipenseriformes

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Pachycormiformes

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Largest specimen of Leedsichthys compared to human and other pachycormid fish

The largest known ray-finned fish and largest bony fish of all time was the pachycormid, Leedsichthys problematicus, at around 16.5 m (54 ft) long.[615] Earlier estimates have had claims of larger individuals with lengths over 27 m (89 ft).[616][617]

Ichthyodectiformes

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Comparation of some ichthyodectiforms: Xiphactinus (1), Ichthyodectes (2), Cladocyclus (3), Chirocentrites (4)

The largest known of ichthyodectiform fish was Xiphactinus, which measured up to 6.1 m (20 ft) long.[618] Ichthyodectes reached 3 m (9.8 ft) long, twice lesser than Xiphactinus.[619]

Pycnodontiformes

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The largest known pycnodontiform was Gyrodus circularis, with length up to 2 m (6 ft 7 in).[620]

Bichirs (Polypteriformes)

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The Late Cretaceous Bawitius was likely the largest bichir of all time. It reached up to 3 m (9.8 ft) in length.[621]

Opahes, ribbonfishes, oarfishes and allies (Lampriformes)

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Megalampris was likely the largest fossil opah. This fish was around 4 m (13 ft) in length when alive, which is twice the length of the largest living opah species, Lampris guttatus.[622]

Salmon and trout (Salmoniformes)

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The largest salmon was Oncorhynchus rastrosus, varying in size from 1.9 m (6 ft 3 in) and 177 kg (390 lb)[623] to 2.4 m (7 ft 10 in) and 200 kg (440 lb).[624][623]

Pufferfishes, boxfishes, triggerfishes, ocean sunfishes and allies (Tetraodontiformes)

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Lizardfishes (Aulopiformes)

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The largest lizardfish was Stratodus which could reach length of 5 m (16 ft).[628]

Echinoderms (Echinodermata)

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Crinozoa

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Fossil of Seirocrinus subangularis

Sea lilies (Crinoidea)

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The longest stem of Seirocrinus subangularis reached over 26 m (85 ft).[629]

Asterozoa

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Starfish (Asteroidea)

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Helianthaster from Hunsrück Slate had a radius of about 25 cm (9.8 in).[630]

Graptolites (Graptolithina)

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The longest known graptoloid graptolite is Stimulograptus halli at 1.45 m (4.8 ft). It was found in Silurian deposits of the United Kingdom.[631]

Kinorhynchs (Kinorhyncha)

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Cambrian kinorhynchs from Qingjiang biota, also known as "mud dragons", reached 4 cm (1.6 in) in length, much larger than extant relatives that grow only a few millimeters in length.[632][633]

Arthropods (Arthropoda)

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Dinocaridida

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Gilled lobopodians

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Size estimation of Omnidens.

Based on the findings of mouthparts, the Cambrian gilled lobopodian Omnidens amplus is estimated to have been 1.5 metres (4.9 ft).[634] It is also known as the largest Cambrian animal known to exist.[634]

Radiodont (Radiodonta)

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Scaled diagram of Aegirocassis

The largest known radiodont is Aegirocassis benmoulai, estimated to have been at least 2 m (6 ft 7 in) long.[635][636]

Chelicerata

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Sea spiders (Pycnogonida)

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The largest fossil sea spider is Palaeoisopus problematicus with legspan about 32 cm (13 in).[637]

Horseshoe crabs and allies (Xiphosura)

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Chasmataspidids (Chasmataspidida)

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Size comparison of the chasmataspidids

The largest chasmataspidids were the Ordovician Hoplitaspis at 29 cm (11 in) in length and similar in size range Chasmataspis.[642]

Eurypterids (Eurypterida)

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Size comparison of the largest known eurypterids

Arachnids (Arachnida)

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Artiopods (Artiopoda)

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Retifacies probably reached up to 55 cm (22 in).[657] Tegopelte is another one example of large non-trilobite artiopod, reached 280 mm (11 in) long[658] and was the largest of the Burgess Shale bilaterians, surpassing all other benthic organisms by at least twice.[658]

Trilobites (Trilobita)

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Some of trilobites exceeded 60 cm (24 in) in length. A nearly complete specimen of Isotelus rex from Manitoba attained a length over 70 cm (28 in), and an Ogyginus forteyi from Portugal was almost as long. Fragments of trilobites suggest even larger record sizes. An isolated pygidium of Hungioides bohemicus implies that the full animal was 90 cm (35 in) long.[659]

Myriapods (Myriapoda)

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A life-size reconstruction of Arthropleura

The largest known myriapod by far was Arthropleura. Measuring 2.5 metres (8 ft 2 in) long[660] and 50 centimetres (20 in) wide.[661] Some specimens could have been even larger, up to 2.63 metres (8 ft 8 in) in length and 50 kilograms (110 lb) in weight.[662][663]

Non-hexapod crustaceans (Crustacea)

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Cycloids (Cyclida)

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The largest cyclid was Opolanka decorosa, the Late Triassic Halicyne-like cycloid which reached over 6 cm (2.4 in) across the carapace.[664]

Remipedes (Remipedia)

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Tesnusocaris had body length at least 9.5 cm (3.7 in),[665] larger than every living remipedes which could reach up to 4.5 cm (1.8 in).[666]

Insects (Insecta)

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Sawflies, wasps, bees, ants and allies (Hymenoptera)

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Titanomyrma with rufous hummingbird for scale

Fleas (Siphonaptera)

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The largest known in Siphonaptera was probably Pseudopulex magnus, growing to 0.90 in (22.8 mm) in length.[671]

Earwigs (Dermaptera)

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Labidura herculeana (St. Helena earwig) specimen

Extinct as recently as after 1967[672][673] and also submitted as the Holocene subfossils,[674] the Saint Helena giant earwig (Labidura herculeana, with synonym Labidura loveridgei) reached 84 mm (3.3 in) in length including forceps 34 mm (1.3 in) long.[672]

Chresmodidae

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Chresmodidae had long specialized legs like of the modern Gerridae family. One of the Chresmodidae, Chresmoda obscura, could have reached a size of about 19 centimetres (7.5 in).[675]

Beetles (Coleoptera)

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One of the largest known fossil beetles in the superfamily Scarabaeoidea is Protognathinus spielbergi. It had total length including mandibles about 5.5 centimetres (2.2 in).[676] The largest fossil scarabaeid was Oryctoantiquus borealis with an estimated body length of 5 centimetres (2.0 in).[677]

Titanopterans (Titanoptera)

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Reconstruction of Gigatitan vulgaris

Related to modern orthopterans, titanopterans from the Triassic period were much larger. The wingspan of Gigatitan vulgaris was up to 40 centimetres (16 in).[678] Clatrotitan andersoni also reached a huge size, having a forewing 13.8 centimetres (5.4 in) long.[679]

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Makarkinia adamsi from the Crato Formation is estimated to have the longest forewings of any neuropteran species, estimated at 160 mm (6.3 in).[680]

Cockroaches, termites, mantises and allies (Dictyoptera)

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Dragonflies, damselflies and griffinflies (Odonatoptera)

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Reconstruction of Meganeura

Mayflies (Ephemeroptera)

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Palaeodictyoptera

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The largest known palaeodictyopteran was Mazothairos, with an estimated wingspan of up to 560 mm (22 in).[693] If a subcircular wing known from Piesberg Quarry belongs to a palaeodictyopteran, it possibly had single wing length at least 30 cm (12 in).[694]

Archaeognatha (jumping bristletails) and other wingless primitive insects

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  • The largest known machilid is Triassic Gigamachilis, with 40 millimetres (1.6 in) body length not counting the length of the filament, and estimated total length about 80 millimetres (3.1 in).[695]
  • The largest specimens of the extinct suborder Monura reached 30 millimetres (1.2 in) or more, not counting the length of the filament.[696]
  • Although Ramsdelepidion was once considered as a 60 millimetres (2.4 in)-long silverfish,[697] it was later considered that classification is uncertain and just treated as stem group insect.[698]
  • The wingless early insect Carbotriplura had body length about 103 millimetres (4.1 in) without tail filaments.[699]

Arrow worms (Chaetognatha)

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The Cambrian stem-chaetognathan Timorebestia koprii had reached up to 20 cm (7.9 in) in body length and 30 cm (12 in) including the antennae.[700] Capinatator had a third of length, about 10 cm (3.9 in), but it is not considered as stem member and still had length similar to the largest modern arrow worms.[700][701]

Ringed worms (Annelida)

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Websteroprion is the largest known fossil eunicidan annelid, with estimated length ranges 0.42–8.3 m (1 ft 5 in – 27 ft 3 in), however comparison with closely related extant taxa indicates length around 1–2 m (3 ft 3 in – 6 ft 7 in).[702] It also had the biggest scolecodonts of any prehistoric polychaete, up to 13.2 mm (0.52 in) in length and possibly larger.[702]

Molluscs (Mollusca)

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Snails and slugs (Gastropoda)

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Campanile giganteum shell

Bivalves (Bivalvia)

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Tusk shells (Scaphopoda)

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Cephalopods (Cephalopoda)

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Parapuzosia seppenradensis shell

Nautiloids (Nautiloidea)

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The largest and longest known of nautiloids was Endoceras giganteum with a shell length of 5.73 m (18.8 ft). There is a record of individual whose shell length had reached 9.14 m (30.0 ft), but it is doubtful.[714]

Ammonites (Ammonoidea)

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The largest known ammonite was Parapuzosia seppenradensis.[715] A partial fossil specimen found in Germany had a shell diameter of 1.95 m (6 ft 5 in), but the living chamber was incomplete, so the estimated shell diameter was probably about 3.5 m (11 ft) and weighed about 705 kg (1,554 lb) when it was alive.[716] However, a later study estimates shell diameter up to around 2 m (6 ft 7 in).[717]

Belemnites (Belemnoidea)

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The largest known belemnite was Megateuthis gigantea, reaching about 50 and 700 mm (2.0 and 27.6 in) in maximum diameter and length of rostrum, respectively.[718]

Squids, octopuses, cuttlefishes and allies (Neocoleoidea)

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Brachiopods (Brachiopoda)

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Gigantoproductus giganteus

The largest brachiopod ever evolved was Striatifera striata from Akkermanovka Quarry, Russia, with height up to 0.5 metres (1 ft 8 in).[722] Another huge brachiopod was the Carboniferous Gigantoproductus giganteus, with shell width from 30 cm (12 in)[723] to over 35 centimetres (14 in).[712][724] Titanaria costellata had large and long shell 35–36 cm (14–14 in) in width, nearly as large as Gigantoproductus.[725]

Hyoliths (Hyolitha)

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The largest hyolith is Macrotheca almgreeni, with length about 50 centimetres (20 in).[712][726]

Cnidarians (Cnidaria)

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Jellyfishes and allies (Medusozoa)

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The largest fossil jellyfish is Cambrian Cordubia gigantea, with diameter of 88 centimetres (35 in).[727] Specimens from the Cambrian of Wisconsin reached 70 cm (28 in) in length.[728]

Vendobionts (Vendobionta)

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Petalonamids (Petalonamae)

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A large specimen of Trepassia wardae

Longest specimens of Trepassia wardae (also known as Charnia wardi) reached 185 cm (73 in) in length.[729] Charnia masoni is known from specimens as small as only 1 cm (0.39 in), up to the largest specimens of 66 cm (26 in) in length.[730]

Proarticulata

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Dickinsonia tenuis reached 1.4 m (4.6 ft) in length, that makes it one of the largest precambrian organisms.[731][732]

Sponges (Porifera)

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The largest known Permian sponge Gigantospongia had diameter up to 2.5 metres (8 ft 2 in).[733]

See also

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References

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