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]
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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)
Further information: Synapsida
Caseasaurs (Caseasauria)
The herbivorous Alierasaurus was the largest caseid and the largest amniote to have lived at the time, with an estimated length around 6–7m (20–23ft).[3]Cotylorhynchus hancocki is also large, with an estimated length and weight of at least 6m (20ft)[4] and more than 500kg (1,100lb).[5]
Edaphosaurids (Edaphosauridae)
Size comparison of some species of Edaphosaurus
The largest edaphosaurids were Lupeosaurus at 3m (9.8ft) long[6] and Edaphosaurus, which could reach even more than 3m (9.8ft) in length.[7]
Sphenacodontids (Sphenacodontidae)
The biggest carnivorous synapsid of Early Permian was Dimetrodon, which could reach 4.6m (15ft) and 250kg (550lb).[8] The largest members of the genus Dimetrodon were also the world's first fully terrestrial apex predators.[9]
Tappenosauridae
The Middle Permian Tappenosaurus was estimated at 5.5m (18ft) in length, nearly as large as the largest dinocephalians.[10]
Therapsids (Therapsida)
Anomodonts (Anomodontia)
Lisowicia compared to a human
The plant-eating dicynodont Lisowicia bojani is the largest-known of all non-mammalian synapsids, at 4.5m (15ft) long, 2.6m (8ft 6in) tall and 9,000kg (20,000lb) in body mass.[11][12][13]
Dinocephalians (Dinocephalia)
Anteosaurus overviewing the landscape
Among the largest carnivorous non-mammalian synapsids was the dinocephalian Anteosaurus, which was 5–6m (16–20ft) long, and weighed 500–600kg (1,100–1,300lb).[14][15] Fully grown Titanophoneus from the same family Anteosauridae likely had a skull of 1m (3.3ft) long.[15] Another enormous dinocephalian was the Late Permian Eotitanosuchus (a possible synonym to Biarmosuchus[16]). Adult specimens could reach 6m (20ft) in length and over 600kg (1,300lb) in weight.[16]
Gorgonopsians (Gorgonopsia)
Photo-reconstruction of Inostrancevia
Inostrancevia latifrons is the largest known gorgonopsian, with a skull length of more than 60cm (24in), a total length approaching 3.5m (11ft) and a mass of 300kg (660lb).[17]Rubidgea atrox is the largest African gorgonopsian, with skull of nearly 45cm (18in) long.[18] Other large gorgonopsians include Dinogorgon with skull of ~40cm (16in) long,[19]Leontosaurus with skull of almost 40cm (16in) long,[18] and Sycosaurus with skull of ~38cm (15in) long.[18]
Therocephalians (Therocephalia)
The largest of therocephalians is Scymnosaurus,[20][21] which reached a size of the modern hyena.[22]
Non-mammalian cynodonts (Cynodontia)
The largest known non-mammalian cynodont is Scalenodontoides, a traversodontid, which had a maximum skull length of approximately 617 millimetres (24.3in) based on a fragmentary specimen.[23]
Paceyodon davidi was the largest of morganucodontans, cynodonts close to mammals. It is known by a right lower molariform 3.3mm (0.13in) in length, which is bigger than molariforms of all other morganucodontans.[24]
The largest known docodont was Castorocauda, almost 50cm (20in) in length.[25]
Mammals (Mammalia)
Further information: Mammal
Non-therian mammals
Gobiconodonts (Gobiconodonta)
A reconstruction of Repenomamus
The largest gobiconodont and the largest well-known Mesozoic mammal was Repenomamus.[26][27][28][29][30][31] The known adult of Repenomamus giganticus reached a total length of around 1m (3ft 3in) and an estimated mass of 12–14kg (26–31lb).[28] With such parameters it surpassed in size several small theropod dinosaurs of the Early Cretaceous.[32]Gobiconodon was also a large mammal,[30][31] it weighed 5.4 kilograms (12lb),[28] had a skull of 10cm (3.9in) in length, and had 35cm (14in) in presacral body length.[33]
Multituberculates (Multituberculata)
The largest multituberculate[34]Taeniolabis taoensis is the largest non-therian mammal known, at a weight possibly exceeding 100kg (220lb).[35]
Monotremes (Monotremata)
Photo-reconstruction of Murrayglossus hacketti (Zaglossus hacketti) by paleoartist Roman Uchytel
The largest known monotreme (egg-laying mammal) ever was the extinct long-beaked echidna species known as Murrayglossus, known from a couple of bones found in Western Australia. It was the size of a sheep, weighing probably up to 30kg (66lb).[36]
The largest known ornithorhynchid is Obdurodon tharalkooschild, it was even larger than 70cm (28in)-long Monotrematum sudamericanum.[37]
Kollikodon was likely the largest monotreme in Mesozoic. Its body length could be up to a 1m (3ft 3in).[38]
Metatherians (Metatheria)
A model of Thylacosmilus
The largest non-marsupial metatherian was Thylacosmilus, weigh 80 to 120 kilograms (180 to 260lb),[39][40] one estimate suggesting even 150kg (330lb).[41] Proborhyaenid Proborhyaena gigantea is estimated to weigh over 50kg (110lb) and possibly reached 150kg (330lb).[42]Australohyaena is another large metatherian, weighing up to 70 kilograms (150lb).[43]
Stagodontid mammal Didelphodon was one of the largest Mesozoic metatherians and all Cretaceous mammals.[44] Its skull could reached over 10 centimetres (3.9in) in length[45] and a weight of complete animal was 5.2 kilograms (11lb).[46]
Marsupials (Marsupialia)
The largest known marsupial, and the largest metatherian, is the extinct Diprotodon, about 3m (9.8ft) long, standing 2m (6ft 7in) tall and weighing up to 2,786kg (6,142lb).[47] Fellow vombatiform Palorchestes azael was similar in length being around 2.5m (8.2ft), with body mass estimates indicating it could exceed 1,000kg (2,200lb).[48]
The largest known carnivorous marsupial was Thylacoleo carnifex. Measurements taken from a number of specimens show they averaged 101 to 164kg (223 to 362lb) in weight.[49][50]
The largest known kangaroo was an as yet unnamed species of Macropus, estimated to weigh 274kg (604lb),[51] larger than the largest known specimen of Procoptodon, which could grow up to 2m (6ft 7in) and weigh 230kg (510lb).[52] Some species from the genus Sthenurus were similar in size or a bit larger than the extant grey kangaroo (Macropus giganteus).[53]
The largest potoroid ever recorded was Borungaboodie, which was nearly 30% bigger than the largest living species and weighted up to 10kg (22lb).[54]
Non-placental eutherians
Restoration of Coryphodons
Cimolestans (Cimolesta)
The largest known cimolestan is Coryphodon, 1m (3ft 3in) high at the shoulder, 2.5m (8ft 2in) long[55][56] and up to 700kg (1,500lb) of mass.[57]Barylambda was also a huge mammal, at 650kg (1,430lb).[58]Wortmania and Psittacotherium from the group Taeniodonta were among the largest mammals of the Early Paleocene.[59] Lived as soon as half a million years after K–Pg boundary, Wortmania reached 20kg (44lb) in body mass. Psittacotherium, which appeared two million years later, reached 50kg (110lb).[59]
Leptictids (Leptictida)
The largest leptictid ever discovered is Leptictidium tobieni from the Middle Eocene of Germany. It had a skull 101mm (4.0in) long, head with trunk 375mm (14.8in) long, and tail 500mm (20in) long.[60] Close European relatives from the same family Pseudorhyncocyonidae had skulls of 67–101mm (2.6–4.0in) in length.[60]
Even-toed ungulates (Artiodactyla)
The extinct Hippopotamus gorgops is the most massive of the fossil even-toed ungulates
The largest known land-dwelling artiodactyl was Hippopotamus gorgops with a length of 4.3m (14ft), a height of 2.1m (6ft 11in), and a weight of 5t (11,000lb).[61]
Daeodon and similar in size and morphology Paraentelodon[62] were the largest-known entelodonts that ever lived, at 3.7m (12ft) long and 1.77m (5.8ft) high at the shoulder.[63] The huge Andrewsarchus from the Eocene of Inner Mongolia had skull 83.4cm (32.8in) long[64] though the taxonomy of this genus is disputive.[65][66]
A skeleton of Bison latifrons
The largest of Bovinae as well as the largest bovid was Bison latifrons. It reached a weight from 1,250kg (2,760lb)[67][68] to 2,000kg (4,400lb),[69]4.75m (15.6ft) in length, shoulder height of 2.31m (7.6ft),[70] and had horns that spanned 2.13m (7ft 0in).[71] The North American Bison antiquus reached up to 4.6m (15ft) long, 2.27m (7.4ft) tall, weight of 1,588kg (3,501lb),[72] and horn span of 1m (3.3ft).[70] The African Pelorovis reached 2t (2.2 short tons) in weight and had bony cores of the horns about 1m (3ft 3in) long.[73] Another enormous bovid, the african giant buffalo (Syncerus antiquus) reached 3m (9.8ft) in length from muzzle to the end of the tail, 1.85m (6.1ft) in height at the withers, 1.7m (5.6ft) in height at the hindquarters,[74][75] and the distance between the tips of its horns was as large as 2.4m (7ft 10in).[74] 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 155–180cm (61–71in) in bulls and 135–155cm (53–61in) in cows, while aurochs populations in Hungary had bulls reaching 155–160cm (61–63in).[76] The kouprey (Bos sauveli), reaching 1.7–1.9m (5ft 7in– 6ft 3in) in shoulder height,[77][78] has existed since the Middle Pleistocene[79] and is also considered to be possibly extinct.[80][81]
The long-legged Megalotragus is possibly the largest known alcelaphine bovid,[82] bigger than the extant wildebeest.[83] The tips of horns of M. priscus were located at a distance of about 1.2m (3ft 11in) from each other.[84]
Megaloceros giganteus
The extinct cervid Irish elk (Megaloceros giganteus) reached over 2.1m (7ft) in height, 680kg (1,500lb) in mass and could have antlers spanning up to 4.3m (14ft) across, about twice the maximum span for a moose's antlers.[85][86] The giant moose (Cervalces latifrons) reached 2.1 to 2.4m (6.9 to 7.9ft) high[87] and was twice as heavy as the Irish elk but its antler span at 2.5m (8.2ft) was smaller than that of Megaloceros.[88][89] North American stag-moose (Cervalces scotti) reached 2.5 metres (8.2ft) in length and a weight of 708.5 kilograms (1,562lb).[90][91]
The largest known giraffid, aside from the extant giraffe, is Sivatherium, with a body weight of 1,250kg (2,760lb).[92]
The largest protoceratid was Synthetoceras, it reached 2m (6ft 7in) long and 150–200kg (330–440lb) in mass.[93][94]
The largest known wild suid to ever exist was Kubanochoerus gigas, having measured up to 500kg (1,100lb) and stood around 1m (3ft 3in) tall at the shoulder.[95]Megalochoerus could be similar in size, possibly weighing 303kg (668lb) or 526kg (1,160lb).[96]
The largest camelid was Titanotylopus from the Miocene of North America. It possibly reached 2,485.6kg (5,480lb) and a shoulder height of over 3.4m (11ft).[97][98] The Syrian camel (Camelus moreli) was twice as big as the modern camels.[99] It was 3m (9.8ft) at the shoulder[100] and 4m (13ft) tall.[99]Camelops had legs to be 20% longer than that of Dromedary, and was about 2.3m (7ft 7in) tall at the shoulder and weighed about 1,000kg (2,200lb).[101]
Cetaceans (Cetacea)
Further information: Cetacean
Size comparison between a human and two species of Basilosaurus, B. cetiodes (dark blue) and B. isis
The largest of known Eocene archeocete whales was Basilosaurus at 17–20m (56–66ft) in length.[102][103][104]
The largest prehistoric sperm whale, or toothed whale was Livyatan melvillei weighing in at about 57 tonnes (63 short tons).[105][106]
The largest squalodelphinid was Macrosqualodelphis at 3.5m (11ft) in length.[107]
Some Neogene rorquals were comparable in size to modern huge relatives. Parabalaenoptera was estimated to be about the size of the modern gray whale,[108] about 16m (52ft) long. Some balaenopterids perhaps rivaled the blue whale in terms of size,[108] though other studies disagree that any baleen whale grew that large in the Miocene.[109]
Odd-toed ungulates (Perissodactyla)
Relative sizes of †Paraceratherium, †Elasmotherium, white rhino, Indian rhino, black rhino and Sumatran rhino compared to a humanLife restoration of Moropus elatus
The largest known perissodactyl, and the second largest land mammal (see Palaeoloxodon namadicus) of all time was the hornless rhino Paraceratherium. The largest individual known was estimated at 4.8m (15.7ft) tall at the shoulders, 7.4m (24.3ft) in length from nose to rump, and 17t (18.7 short tons) in weight.[110][111]
Some prehistoric horned rhinos also grew to large sizes. The biggest Elasmotherium reached up to 5–5.2m (16–17ft) long,[112]2.5m (8ft 2in) high[113] and weighed 3.5–5t (3.9–5.5 short tons).[114][112][113] Such parameters make it the largest rhino of the Quaternary.[114] Woolly rhinoceros (Coelodonta antiquitatis) of the same time reached 1,100–1,500kg (2,400–3,300lb)[115] or 2,000kg (4,400lb),[116][117]1.93m (6ft 4in) at the shoulder height and 4.6m (15ft) in length.[118]
Metamynodon, an amynodontid, reached 4m (13ft) in length, comparable to Hippopotamus in measurement and shape.[119]
The giant tapir (Tapirus augustus) was the largest tapir ever, at about 623kg (1,373lb)[120] and 1m (3.3ft) tall at the shoulders.[121] Earlier, this mammal was estimated even bigger, at 1.5m (4.9ft) tall, and assigned to the separate genus Megatapirus.[121]
One of the biggest chalicotheres was Moropus.[122] It stood about 2.4 metres (8ft) tall at the shoulder.[123]
Late Eocene perissodactyls from the family Brontotheriidae attained huge sizes. The North American Megacerops (also known as Brontotherium[124]) reached 2.5m (8ft 2in) tall at the shoulders,[125]5m (16ft) in length,[124] and 3t (6,600lb) in weight.[126]Embolotherium from Asia was equal in size.[127]
The largest prehistoric horse was Equus giganteus of North America. It was estimated to grow to more than 1,250kg (1.38 short tons) and 2m (6ft 7in) at the shoulders.[128] The largest anchitherine equid was Hypohippus at 403 to 600kg (888 to 1,323lb), comparable to large modern domestic horses.[129][130]Megahippus is another large anchitheriine. With the body mass of 266.2kg (587lb) it was much heavier than most of its close relatives.[129]
Phenacodontids (Phenacodontidae)
The largest known phenacodontid is Phenacodus. It was 1.5m (4ft 11in) long[131] and weighed up to 56kg (123lb).[132]
Dinoceratans (Dinocerata)
The largest known dinoceratan was Eobasileus with skull length of 102cm (40in), 2.1m (6ft 11in) tall at the back and 1.5m (4ft 11in) tall at the shoulder.[133] Another huge animal of this group was Uintatherium, with skull length of 76cm (30in), 1.5m (4ft 11in) tall at the shoulder,[133]4m (13ft) in length and 2.25t (2.48 short tons), the size of a rhinoceros.[134] Despite their large size, Eobasileus as well as Uintatherium had a very small brain.[133][134]
Carnivores (Carnivora)
Caniformia
Arctodus simus reconstructionChapalmalania, the giant procyonidSkeletal 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 1,588–1,749kg (3,501–3,856lb) and stood at least 3.4m (11ft) tall on their hind-limbs.[135][136] Another huge bear was the giant short-faced bear (Arctodus simus), with the average weight of 625kg (1,378lb) and the maximum recorded at 957kg (2,110lb).[137] There is a guess that the largest individuals of this species could reached even larger mass, up to 1,200kg (2,600lb).[138] The extinct cave bear (Ursus spelaeus) was also heavier than many recent bears. Largest males weighed as much as 1,000kg (2,200lb).[139]Ailuropoda baconi from the Pleistocene was larger than the modern giant panda (Ailuropoda melanoleuca).[140]
The biggest odobenid and one of the biggest pinnipeds to have ever existed is Pontolis magnus, with skull length of 60cm (24in) (twice as large as the skulls of modern male walruses)[141] and more than 4m (13ft) in total body length.[142][143] Only the modern males of elephant seals (Mirounga) reaches the similar sizes.[142] The second largest prehistoric pinniped is Gomphotaria pugnax with the skull length of nearly 47cm (19in).[141]
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 295–318kg (650–701lb).[144]
The biggest known mustelid to ever exist was likely the giant otter, Enhydriodon. It exceeded 3m (9.8ft) in length, and would have weighed in at around 200kg (440lb), much larger than any other known mustelid, living or extinct.[145][146][147] There were other giant otters, like Siamogale, at around 50kg (110lb)[148] and Megalenhydris, which was larger than a modern-day giant river otter.[149]Megalictis was the largest purely terrestrial mustelid[150] (although Enhydriodon had recently been mentioned as the largest mustelid that also happens to be a terrestrial predator[145]). Similar in size to the jaguar, Megalictis ferox had even wider skull, almost as wide as of the black bear.[150]Megalictis had a powerful bite force, allowing it to eat large prey and crush bones, as modern hyenas and jaguars can.[150] Another large-bodied mustelid was the superficially cat-like Ekorus from the Miocene of Africa. At almost 44kg (97lb), the long-legged Ekorus was about the size of a wolf[151] and filling a similar to leopards ecological niche before big cats came to the continent.[152] Other huge mustelids include Perunium[153] and hypercarnivorous Eomellivora, both from the Late Miocene.[154]
The heaviest procyonid was possibly South American Chapalmalania. It reached 1.5 metres (4.9ft) in body length with a short tail and 150 kilograms (330lb), comparable in size to an American black bear (Ursus americanus).[155] Another huge procyonid was Cyonasua, which weighted about 15–25kg (33–55lb), about the same size as a medium-sized dog.[156]
The largest canid of all time was Epicyon haydeni, which stood 90cm (35in) tall at the shoulder, had a body length of 2.4m (7.9ft) and weighed 100–125kg (220–276lb),[157][158][159] with the heaviest known specimen weighing up to 170kg (370lb).[41] The extinct dire wolf (Aenocyon dirus) reached 1.5m (4ft 11in) in length and weighed between 50 and 110kg (110 and 243lb).[41][160] 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.[161] The Late Pleistocene Italian wolf was morphometrically close to C. l. maximus.[162]
The largest bear-dog was a species of Pseudocyon weighing around 773kg (1,704lb), representing a very large individual.[163]
Feliformia
Comparison between Amphimachairodus giganteus and the modern domestic catSkeleton of Panthera atrox at the La Brea Tar Pits Museum
The largest nimravid was probably Quercylurus major as its fossils suggest it was similar in size to the modern-day brown bear and was scansorial.[164] In 2021, Eusmilus was declared as the largest of the holplophonine nimravids, reaching the weight of nearly 111kg (245lb), comparable to a small African lion.[165] However, the largest Hoplophoneus was estimated at 160kg (350lb).[41]
The biggest saber-toothed cats are Amphimachairodus kabir and Smilodon populator, with the males possibly reaching 350–490kg (770–1,080lb) and 220–450kg (490–990lb) respectively.[41][166][167] Another contender for the largest felid of all time is Machairodus. M. horribilis from China was estimated at 405kg (893lb)[168] while the North American M. lahayishupup weighed up to 410kg (900lb).[169][170][171]Xenosmilus was also a huge cat. It reached around 2m (6.6ft) long[172] and weighed around 300–350kg (660–770lb).[168]
The heaviest known pantherine felids are the Ngangdong tiger (Panthera tigris soloensis), which are estimated to have weighed up to 486kg (1,071lb),[167] the American lion (Panthera atrox), weighing up to 363kg (800lb)[173][174] and the Eurasian cave lion (Panthera spelaea), weighing up to 339kg (747lb).[167] Being the ancestor of the modern jaguar,[175]Panthera gombaszoegensis was much larger, up to 150kg (330lb) in maximum weight.[175]
Some extinct feline felids also surpassed their modern relatives in size. The Eurasian giant cheetah (Acinonyx pardinensis) reached 60–121kg (132–267lb), approximately twice as large as the modern cheetah.[176] The North American Pratifelis was larger than the extant cougar.[177]
The largest barbourofelid was Barbourofelis fricki, with the shoulder height of 90cm (35in).[178]
The largest viverrid known to have existed is Viverra leakeyi, which was around the size of a wolf or small leopard at 41kg (90lb).[179]
The largest known fossil hyena is Pachycrocuta, estimated at 90–100cm (35–39in) at the shoulder[180] and 190kg (420lb) weight.[41] Another huge hyena with mass over 100kg (220lb) is the cave hyena. It is actually a subspecies of the African spotted hyena, which is at 10% smaller than the extinct cave hyena.[181]
The percrocutid feliform, Dinocrocuta, was two or even three times as large as the extant spotted hyena, 160 or 240kg (350 or 530lb).[182]
The extinct giant fossa (Cryptoprocta spelea) had a body mass in range from 17kg (37lb)[183] to 20kg (44lb),[184] much larger than the modern fossa weighs (up to 8.6kg (19lb) for adult males[185]).
Hyaenodonts (Hyaenodonta)
The largest hyaenodont was Simbakubwa at 1,500kg (3,300lb).[186] Another giant hyaenodont, Megistotherium reached 500kg (1,100lb)[41] and had a skull of 66.4cm (26.1in) in length.[187]
Oxyaenids (Oxyaenidae)
Sarkastodon
The largest known oxyaenid was Sarkastodon weighing in at 800kg (1,800lb).[41]
Mesonychians (Mesonychia)
Some mesonychians reached a size of a bear. Such large were Mongolonyx from Asia[188] and Ankalagon from North America.[189][190] Another large mesonychian is Harpagolestes with a skull length of a half a meter in some species.[188]
Bats (Chiroptera)
Found in Quaternary deposits of South and Central Americas, Desmodus draculae had a wingspan of 0.5m (20in) and a body mass of up to 60g (2.1oz). Such proportions make it the largest vampire bat that ever evolved.[191]
Hedgehogs, gymnures, shrews, and moles (Eulipotyphla)
Deinogalerix skeleton
The largest known animal of the group Eulipotyphla was Deinogalerix,[192] measuring up to 60cm (24in) in total length, with a skull up to 21cm (8.3in) long.[193]
Rodents (Rodentia)
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 480–500kg (1,060–1,100lb).[194]Phoberomys pattersoni weighed 125–150kg (276–331lb).[194] Both Josephoartigasia and Phoberomys reached about 1.5m (4ft 11in) tall at the shoulder.[195] Another huge dinomyid, Telicomys gigantissimus had a minimal weight of 200kg (440lb).[195]
Amblyrhiza inundata from the family Heptaxodontidae was a massive animal, it weighed 50–200kg (110–440lb).[196][195]
The largest beaver was the giant beaver (Castoroides) of North America. It grew over 2 m in length and weighed roughly 90 to 125kg (198 to 276lb), also making it one of the largest rodents to ever exist.[197]
Rabbits, hares, and pikas (Lagomorpha)
The biggest known prehistoric lagomorph is Minorcan giant lagomorph Nuralagus rex at 12kg (26lb).[198]
Primates (Primates)
Reconstruction of Gigantopithecus blackii
The largest known primate as well as the largest hominid of all time was Gigantopithecus blackii, standing 3m (9.8ft) tall and weighing 540kg (1,200lb).[199][200] However In 2017, new studies suggested a body mass of 200–300kg (440–660lb) for this primate.[201] Another giant hominid was Meganthropus palaeojavanicus at 2.4m (7ft 10in) in body height,[202] although it is known from very poor remains.[203]
During the Pleistocene, some archaic humans were close in sizes or even larger than early modern humans. Neanderthals (Homo neanderthalensis) reached 77.6kg (171lb) and 66.4kg (146lb) in average weight for males and females, respectively, larger than the parameters of modern humans (Homo sapiens) (68.5kg (151lb) and 59.2kg (131lb) for males and females, respectively).[204] A tibia from Kabwe (Zambia) indicates an indeterminate Homo individual of possibly 181.2cm (71.3in) in height. It was one of the tallest humans of the Middle Pleistocene and noticeably large even compared to recent humans.[205] The tallest Homo sapiens individuals from the Middle Pleistocene of Spain reached 194cm (76in) and 174cm (69in) for males and females, respectively.[205] Some Homo erectus could be as large as 185cm (73in) tall and 68kg (150lb) in weight.[206][207]
The largest known Old World monkey is the prehistoric baboon, with a male specimen of Dinopithecus projected to weigh an average of 46kg (101lb) and up to 57kg (126lb).[208] It exceeds the maximum weight record of the chacma baboon, the largest extant baboon. One source projects a specimen of Theropithecus oswaldi to have weighed 72kg (159lb).[209]
The largest known New World monkey was Cartelles, which is studied as specimen of Protopithecus, weighing up to 34.27kg (75.6lb). Caipora bambuiorum is another large species, weighing up to 27.74kg (61.2lb).[210]
The largest omomyids were Macrotarsius and Ourayia from the Middle Eocene. Both reached 1.5–2kg (3.3–4.4lb) in weight.[211]
Some prehistoric lemuriform primates grew to huge sizes as well. Archaeoindris was a 1.5-metre-long (4.9ft) sloth lemur that lived in Madagascar and weighed 150–187.8kg (331–414lb),[212] as large as an adult male gorilla.[213]Palaeopropithecus from the same family was also heavier than most modern lemurs, at 25.8–45.8kg (57–101lb).[214]Megaladapis is another large extinct lemur at 1.3 to 1.5m (4ft 3in to 4ft 11in) in length[citation needed] and an average body mass of around 140kg (310lb).[215] Other estimates suggest 46.5–85.1kg (103–188lb) but its still much larger than any extant lemur.[214]
The largest known land mammal ever was a proboscidean called Palaeoloxodon namadicus which weighed about 22t (24.3 short tons) and measured about 5.2m (17.1ft) tall at the shoulder.[110] The largest individuals of the steppe mammoth of Eurasia (Mammuthus trogontherii) estimated to reach 4.5m (14.8ft) at the shoulders and 14.3t (15.8 short tons) in weight.[110][216]Stegodon zdanskyi, the biggest species of Stegodon, was 13t (14.3 short tons) in body mass.[110] Another one enormous proboscidean is Stegotetrabelodon syrticus, over 4m (13ft) in height and 11 to 12t (12.1 to 13.2 short tons) in weight.[110] The Columbian mammoth (Mammuthus columbi) was about 4m (13.1ft) tall at the shoulder but didn't weigh as much as other huge mammoths. Its average mass was 9.5t (10.5 short tons) with one unusually large specimen about 12.5t (13.8 short tons).[110] Columbian mammoths had very long tusks. The largest known mammoth tusk, 4.9m (16ft) long, belonged to this species.[217]
The largest mammutid was the Neogene Mammut borsoni. The biggest specimen reached 4.1m (13ft) tall and weighed about 16t (17.6 short tons).[110] This species also had the longest tusks, 5.02m (16.5ft) long from basis to tip along the curve.[218]
Deinotherium was the largest proboscidean in Deinotheriidae family. Bones retrieved in Crete confirm the existence of specimen 4.1m (13ft) tall at the shoulders and more than 14t (15.4 short tons) in weight.[110]
Sea cows (Sirenia)
According to reports, Steller's sea cows have grown to 8 to 9m (26 to 30ft) long as adults, much larger than any extant sirenians.[219] The weight of Steller's sea cows is estimated to be 8–10t (8.8–11.0 short tons).[220]
Arsinoitheres (Arsinoitheriidae)
Skeleton of Arsinoitherium
The largest known arsinoitheriid was Arsinoitherium. A. zitteli would have been 1.75m (5ft 9in) tall at the shoulders, and 3m (9.8ft) long.[221][222]A. giganteum reached even larger size than A. zitteli.[223]
Hyraxes (Hyracoidea)
Some of the prehistoric hyraxes were extremely large compared to modern small relatives. The largest hyracoid ever evolved is Titanohyrax ultimus.[224] With the mass estimation in rage of 600kg (1,300lb) to over 1,300kg (2,900lb) it was close in size to Sumatran rhinoceros.[225] Another enormous hyrax is Megalohyrax which had skull of 391mm (15.4in) in length[226] and reached the size of tapir.[227][224] More recent Gigantohyrax was three times as large as the extant relative Procavia capensis,[228] although it is noticeably smaller than earlier Megalohyrax and Titanohyrax.[229]
Desmostylians (Desmostylia)
Desmostylus skeletal diagram
The largest known desmostylian was a species of Desmostylus, with skull length of 81.8cm (32.2in) and comparable in size to the Steller's sea cow.[230]
Paleoparadoxia is also known as one of the largest desmostylians, with body length of 3.03m (9.9ft).[231]
Armadillos, glyptodonts and pampatheres (Cingulata)
The largest cingulate known is Doedicurus, at 4m (13ft) long, 1.5m (4ft 11in) high[134] and reaching a mass of approximately 1,910 to 2,370kg (2.11 to 2.61 short tons).[citation needed] The largest species of Glyptodon, Glyptodon clavipes, reached 3–3.3m (9.8–10.8ft) in length[232][134] and 2t (2.2 short tons) in weight.[citation needed]
Anteaters and sloths (Pilosa)
Skeleton and illustration of Megatherium
The largest known pilosan ever was Megatherium, a ground sloth with an estimated average weight of 3.8t (4.2 short tons)[233] and a height of 6m (20ft)[233] which is almost as big as the African bush elephant. Several other sloths grew to large sizes as well, such as Eremotherium, but none as large as Megatherium.
Astrapotherians (Astrapotheria)
Some of the largest known astrapotherians weighed about 3–4t (3.3–4.4 short tons), including the genus Granastrapotherium[234] and some species of Parastrapotherium (P. martiale).[235] The skeleton remains suggests that the species Hilarcotherium miyou was even larger, with a weight of 6.456t (7.117 short tons).[236]
Litopterns (Litopterna)
The largest known litoptern was Macrauchenia, which had three hoofs per foot. It was a relatively large animal, with a body length of around 3m (9.8ft).[237]
Notoungulates (Notoungulata)
The largest notoungulate known of complete remains is Toxodon. It was about 2.7m (8ft 10in) in body length, and about 1.5m (4ft 11in) high at the shoulder and resembled a heavy rhinoceros. Although is not complete, the preserved fossils suggests that Mixotoxodon were the most massive member of the group, with a weight about 3.8t (4.2 short tons).[238]
Pyrotherians (Pyrotheria)
The largest mammal of the South American order Pyrotheria was Pyrotherium at 2.9–3.6m (9ft 6in– 11ft 10in) in length and 1.8–3.5t (4,000–7,700lb) in weight.[239]
Megalania skeletal reconstruction on Melbourne Museum steps
Giant mosasaurs are the largest-known animals within the Squamata. The largest-known mosasaur is likely Mosasaurus hoffmanni, estimated at more than 17m (56ft) in length,[240][241] 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.[242] Another giant mosasaur is Tylosaurus, estimated at 10–14m (33–46ft) in length.[243][244] Another large mosasaur is Hainosaurus bernardi (could be synonymous to Tylosaurus). It was once estimated at 17 and 15m (56 and 49ft) in length,[245][246] but later estimates put it at around 12.2m (40ft).[247]
The largest known prehistoric snake is Titanoboa cerrejonensis, estimated at 12.8m (42ft) in length and 1,135kg (2,502lb) in weight.[248] Another known very large fossil snake is Gigantophis garstini, estimated at 9.3–10.7m (31–35ft) in length,[249][250] although later study shows smaller estimation about 6.6–7.2m (22–24ft).[251] A close rival in size to Gigantophis is a fossil snake, Palaeophis colossaeus, which may have been around 9m (30ft) in length.[248][252][253] Later studies speculate that it reached a maximum length of 12.3m (40ft).[254] The largest fossil python is Liasis dubudingala with length roughly 9m (30ft).[255] 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 3m (9.8ft) in length and 26kg (57lb) in weight.[256][257] Another huge fossil viper is indeterminate species of Vipera. With a length of around 2m (6ft 7in) it was one of the biggest predators of Mallorca during the Early Pliocene.[258] The largest known blind snake is Boipeba tayasuensis with estimated total length of 1.1m (3ft 7in).[259]
The largest known land lizard is probably megalania (Varanus priscus) at 7m (23ft) in length.[260] As extant relatives, megalania could have been venomous and in that case this lizard was also the largest venomous vertebrate ever evolved.[261] However, maximum size of this animal is subject to debate.[262]
Turtles, tortoises and close relatives (Pantestudines)
Further information: Pantestudines
Size comparison between Protostegidae family: Notochelone (lightest blue), Protostega (darkest blue), and Archelon
Cryptodira
The largest known turtle ever was Archelon ischyros at 5m (16ft) long and 2,200kg (4,900lb).[263] Possible second-largest sea turtle was Protostega at 3.9m (13ft) in total body length.[264][265] There is even a larger specimen of this genus from Texas estimated at 4.2m (14ft) in total length.[266][264] Another huge prehistoric sea turtle is the Late Cretaceous Gigantatypus, estimated at over 3.5m (11ft) in length.[267]Psephophorus terrypratchetti from the Eocene attained 2.3–2.5m (7.5–8.2ft) in body length.[268]
The largest tortoise was Megalochelys atlas at up to 2.7m (9ft) in shell length[269] and weighing 0.8–1.0t (1,800–2,200lb).[126]M. margae had carapace of 1.4–2m (4.6–6.6ft) long; an unnamed species from Java reached at least 1.75m (5.7ft) in carapace length.[270] The Cenozoic Titanochelon were also larger than extant giant tortoises, with a shell length of up to 2m (6ft 7in).[271][272] Other giant tortoises include Centrochelys marocana at 1.8–2m (5.9–6.6ft) in carapace length and Mesoamerican Hesperotestudo sp. at 1.5m (4.9ft) in carapace length.[270]
The largest trionychid ever recorded is indeterminate specimen GSP-UM 3019 from the Middle Eocene of Pakistan. Bony carapace of GSP-UM 3019 is 120cm (3.9ft) long and 110cm (3.6ft) wide indicates the total carapace diameter (with soft margin) about 2m (6.6ft).[273]Drazinderetes tethyensis from the same formation had a bony carapace 80cm (2.6ft) long and 70cm (2.3ft) wide.[273] Another huge trionychid is North American Axestemys byssinus at over 2m (6.6ft) in total length.[274]
Side-necked turtles (Pleurodira)
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 2.86m (9ft 5in) and weight of up to 1,145kg (2,524lb).[275]Carbonemys cofrinii from the same family had a shell that measured about 1.72m (5ft 8in),[276][277][278] complete shell was estimated at 1.8m (5.9ft).[279]
Macrobaenids (Macrobaenidae)
The largest macrobaenids were the Early Cretaceous Yakemys, Late Cretaceous Anatolemys, and Paleocene Judithemys. All reached 70cm (2.3ft) in carapace length.[280]
Meiolaniformes
Skeleton of Meiolania platyceps
The largest meiolaniid was Meiolania. Meiolania platyceps had a carapace 100cm (3.3ft) long[270] and probably reached over 3m (9.8ft) in total body length.[281] An unnamed Late Pleistocene species from Queensland was even larger, up to 200cm (6.6ft) in carapace length.[270]Ninjemys oweni reached 100cm (3.3ft) in carapace length[270] and 200kg (440lb) in weight.[282]
Sauropterygians (Sauropterygia)
Placodonts and close relatives (Placodontiformes)
Placodus was among the largest placodonts, with a length of up to 3m (9.8ft).[283]
Nothosaurs and close relatives (Nothosauroidea)
The largest nothosaur as well as the largest Triassic sauropterygian was Nothosaurus giganteus at 7m (23ft) in length.[284]
Plesiosaurs (Plesiosauria)
Main article: Plesiosaur size
The largest known plesiosauroid was Aristonectes, with a body length of 10–11.86 metres (32.8–38.9ft) and body mass of 4t (4.4 short tons)[285] or even 10.7–13.5t (11.8–14.9 short tons).[286] Another long plesiosauroid was Albertonectes at 11.2–11.6 metres (37–38ft).[287]Thalassomedon rivaled it in size, with its length at 10.86–11.6m (35.6–38.1ft).[288] Other large plesiosauroids are Styxosaurus and Elasmosaurus. Both reached some more than 10m (33ft) in length.[285][289]Hydralmosaurus (previously synonymized with Elasmosaurus and Styxosaurus) reached 9.44m (31.0ft) in total body length.[289] In past, Mauisaurus was considered to be more than 8m (26ft) in length,[290][289] but later it was determined as nomen dubium.[291]
Size estimation of three species of Pliosaurus.
There is much controversy over the largest-known of the Pliosauroidea. Pliosaurus funkei, fossil remains of a pliosaur nicknamed as "Predator X" have been discovered and excavated from Norway in 2008. This pliosaur has been estimated at 10–13m (33–43ft) in length.[292] However, in 2002, a team of paleontologists in Mexico discovered the remains of a pliosaur nicknamed as "Monster of Aramberri", which is also estimated at 15m (49ft) in length,[293] with shorter estimation about 11.5m (38ft).[294] This species is, however, claimed to be a juvenile and has been attacked by a larger pliosaur.[295] Some media sources claimed that Monster of Aramberri was a Liopleurodon but its species is unconfirmed thus far.[293] Another very large pliosaur was Pliosaurus macromerus, known from a single 2.8-metre-long (9.2ft) incomplete mandible.[296] The Early Cretaceous Kronosaurus queenslandicus is estimated at 9–10.9m (30–36ft) in length and 10.6–12.1t (11.7–13.3 short tons) in weight.[297][298] The Late Jurassic Megalneusaurus rex could reach lengths of 7.6–9.1 metres (25–30ft).[299][300] Close contender in size was the Late Cretaceous Megacephalosaurus eulerti with a length in range of 6–9m (20–30ft).[301]
Proterosuchids (Proterosuchidae)
Proterosuchus fergusi is the largest known proterosuchid with a skull length of 47.7cm (18.8in) and a possible body length of 3.5–4m (11–13ft).[302]
Erythrosuchids (Erythrosuchidae)
Life reconstruction of Erythrosuchus africanus
The largest erythrosuchid was Erythrosuchus africanus with a maximum length of 4.75–5m (15.6–16.4ft).[303]
Phytosaurs (Phytosauria)
Some of the largest known phytosaurs include Redondasaurus with a length of 6.4m (21ft)[304] and Smilosuchus with a length of more than 7m (23ft).[305]
Non-crocodylomorph pseudosuchians (Pseudosuchia)
Further information: Pseudosuchia
Size comparison of Sillosuchus to a human
The largest shuvosaurid and one of the largest pseudosuchian from the Triassic period was Sillosuchus. Biggest specimens could have reached 9–10m (30–33ft) in length.[306][307]
The largest known carnivorous pseudosuchian of the Triassic is loricatan Fasolasuchus tenax, which measured an estimated of 8 to 10m (26 to 33ft).[308][306][307] It is both the largest "rauisuchian" known to science, and the largest non-dinosaurian terrestrial predator ever discovered.[citation needed] Biggest individuals of Postosuchus[309] and Saurosuchus[310] had a body length of around 7m (23ft). A specimen of Prestosuchus discovered in 2010 suggest that this animal also reached lengths of nearly 7m (23ft) making it one of the largest Triassic pseudosuchians.[311]
Desmatosuchus was likely one of the largest known aetosaurs, about 4–6m (13–20ft) in length and 280kg (620lb) in weight.[312][313][314]
Crocodiles and close relatives (Crocodylomorpha)
Further information: Crocodylomorpha
Large crocodylomorphs (†Deinosuchus, †Purussaurus,
†Gryposuchus, †Euthecodon,
†Sarcosuchus, and modern Crocodylus porosus) compared to a human
Aegyptosuchids (Aegyptosuchidae)
The Late Cretaceous Aegisuchus is the main contender for the title of the largest crocodylomorph ever recorded. It reached 15m (49ft) in length by the lower estimate and as much as 22m (72ft) by the upper estimate,[315] although a length of over 15 m is likely an overestimate.[315]
Crocodylians (Crocodylia)
The largest caiman and likely the largest crocodylian was Purussaurus brasiliensis estimated at 11–13m (36–43ft).[316] According to another information, maximum estimate measure 11.4m (37ft) and almost 7.8t (8.6 short tons) in length and in weight respectively.[317] Another giant caiman was Mourasuchus. Various estimates suggest the biggest specimens reached 9.47m (31.1ft) in length and 8.5t (9.4 short tons) in weight[318] or 4.7–5.98m (15.4–19.6ft) in body length.[319]
The largest alligatoroid is likely Deinosuchus riograndensis at 12m (39ft) long and weighing 8.5t (9.4 short tons).[320][321]
The largest extinct species of the genus Alligator was the Haile alligator (Alligator hailensis), which had a skull 52.5cm (20.7in) long and was similar in size to the extant American alligator (Alligator mississippiensis).[322]
The largest gavialids were Asian Rhamphosuchus at 8–11m (26–36ft)[323][324][317] and South American Gryposuchus at 10.15m (33.3ft) in length.[325][324]
The basal crocodyloidean Astorgosuchus bugtiensis from the Oligocene was large. It estimated at 8m (26ft) in length.[324]
The largest known true crocodile was Euthecodon which estimated to have reached 6.4–8.6m (21–28ft) or even 10m (33ft) long.[326][317] The largest species of the modern Crocodylus were Kenyan Crocodylus thorbjarnarsoni at 7.56m (24.8ft) in length,[327][317] Tanzanian Crocodylus anthropophagus at 7.5m (25ft) in length[328][329] and indeterminate species from Kali Gedeh (Java) at 6–7m (20–23ft) in length.[330]
Unnamed Pliocene species of Quinkana known from partial remains may have reached up to 6m (20ft) in length, although other species (known from Oligocene to Pleistocene) are smaller with length just about 3m (9.8ft). It is not only the largest mekosuchian (some studies reject it from this group[331]) but also it could have been Australia's largest Pliocene predator.[260]Paludirex is another large mekosuchian with length over 4m (13ft).[332]
Paralligatorids (Paralligatoridae)
The largest paralligatorid was likely Kansajsuchus estimated at up to 8m (26ft) long.[333]
Tethysuchians (Tethysuchia)
Some extinct pholidosaurids reached giant sizes. In the past, the Sarcosuchus imperator was believed to be the largest crocodylomorph, with initial estimates proposing a length of 12m (39ft) and a weight of 8t (8.8 short tons).[334] However, recent estimates have now shrunk to a length of 9 to 9.5m (29.5 to 31.2ft) and a weight of 3.5 to 4.3 metric tons (3.9 to 4.7 short tons).[335] Related to Sarcosuchus, Chalawan thailandicus could reached more than 10m (33ft) in length,[336] although other estimates suggest 7–8m (23–26ft).[324]
The largest dyrosaurid was Phosphatosaurus gavialoides estimated at 9m (30ft) in length.[337][324]
Stomatosuchids (Stomatosuchidae)
Stomatosuchus, a stomatosuchid, estimated at 10m (33ft) in length.[338]
Notosuchians (Notosuchia)
Skull of Barinasuchus
The largest terrestrial notosuchian crocodylomorph was very likely the Miocene sebecid Barinasuchus, with a skull of 95–110cm (37–43in) long, comparable in size to the 104cm (41in)-long skull of Daspletosaurus.[339] Various estimates suggest a possible length of Barinasuchus at 6–7.5m (20–25ft).[339]
Other huge notosuchians are Brazilian Stratiotosuchus at 4m (13ft) long,[340] and Baurusuchus at 3.5–4m (11–13ft) long,[341] both from the family Baurusuchidae.[340]
Thalattosuchians (Thalattosuchia)
Restoration of MachimosaurusPlesiosuchus compared to a human
The largest thalattosuchian as well as the largest teleosauroid was the Early Cretaceous Machimosaurus rex estimated at 7.15m (23.5ft) in length.[342]Neosteneosaurus edwardsi (previously known as Steneosaurus edwardsi[343]) was the biggest Middle Jurassic crocodylomorph, it reached 6.6m (22ft) long.[342]
Plesiosuchus was very large metriorhynchid. With the length of 6.83m (22.4ft) it exseeded even some pliosaurids of the same time and locality such as Liopleurodon.[344] Other huge metriorhynchids include Tyrannoneustes at 5m (16ft) in length[345] and Torvoneustes at 4.7m (15ft) in length.[346]
Basal crocodylomorphs
Redondavenator was the largest Triassic crocodylomorph ever recorded,[347] with a skull of at least 60cm (2.0ft) in length.[348][349] Another huge basal crocodylomorph was Carnufex[347] at 3m (9.8ft) long even through that is immature.[350]
Pterosaurs (Pterosauria)
Main article: Pterosaur size
Hatzegopteryx (A-B), Arambourgiania (C) and Quetzalcoatlus sp. (D-E)
The largest known pterosaur was Quetzalcoatlus northropi, at 127kg (280lb) and with a wingspan of 10–12m (33–39ft).[351] Another close contender is Hatzegopteryx, also with a wingspan of 12m (39ft) or more.[351] This estimate is based on a skull 3m (9.8ft) long.[352] Yet another possible contender for the title is Cryodrakon which had a 10-metre (33ft) wingspan.[353] An unnamed pterodactyloid pterosaur from the Nemegt Formation could reach a wingspan of nearly 10m (33ft).[354][355] According to various assumptions, the wingspan of Arambourgiania philadelphiae reached from 8m (26ft) to more than 10m (33ft).[354][353] South American Tropeognathus reached the maximum wingspan of 8.7m (29ft).[356][357]
The largest of non-pterodactyloid pterosaurs as well as the largest Jurassic pterosaur[358] was Dearc, with an estimated wingspan between 2.2m (7ft 3in) and 3.8m (12ft).[359] Only a fragmentary rhamphorhynchid specimen from Germany could be larger (184% the size of the biggest Rhamphorhynchus).[360] Other large non-pterodactyloid pterosaurs were Sericipterus, Campylognathoides and Harpactognathus, with the wingspan of 1.73m (5ft 8in),[361]1.75m (5ft 9in),[361] and 2.5m (8ft 2in),[360] respectively.
Choristoderes (Choristodera)
The largest known choristoderan, Kosmodraco dakotensis (previously known as Simoedosaurus dakotensis[362]) is estimated to have had a total length of around 5m (16ft).[363][362]
Tanystropheids (Tanystropheidae)
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 5m (16ft) in length.[364]
Thalattosaurs (Thalattosauria)
The largest species of thalattosaur, Miodentosaurus brevis grew to more than 4m (13ft) in length.[365] The second largest member of this group is Concavispina with a length of 3.64m (11.9ft).[366]
Ichthyosaurs (Ichthyosauria)
Shonisaurus popularis (green) and Shastasaurus sikanniensis (red) compared with a human
The largest known ichthyosaur and the largest marine reptile was the Late Triassic Shastasaurus sikanniensis at 21m (69ft) in length[367][368] and 81.5t (180,000lb) in weight.[369] In April 2018, paleontologists announced the discovery of a previously unknown ichthyosaur that may have reached lengths of 26m (85ft) making it one of the largest animals known, rivaling some blue whales in size.[370][371] Another, larger ichthyosaur was found in 1850 in Aust.[372] 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.[372] Another huge ichthyosaur was Shonisaurus popularis at 15m (49ft) in length and 29.7t (65,000lb) in weight.[368] The largest Middle Triassic ichthyosaur as well as the largest animal of that time was Cymbospondylus youngorum at 14 to 17.65m (45.9 to 57.9ft) in length[373][369] and 14.7 to 135.8t (32,000 to 299,000lb) in weight.[369]
Tangasaurids (Tangasauridae)
The largest tangasaurid was Hovasaurus with an estimated snout-vent length of 30–35cm (12–14in) and a tail of 60cm (24in).[374]
Pareiasaurs (Pareiasauria)
Largest pareiasaurs reached up to 3m (9.8ft) in length. Such sizes had Middle Permian Bradysaurus, Embrithosaurus, and Nochelesaurus from South Africa,[375] and the Late Permian Scutosaurus from Russia.[375] The most robust Scutosaurus had 1.16t (2,600lb) in body mass.[375]
Captorhinids (Captorhinidae)
The heavy built Moradisaurus grandis, with a length of 2m (6ft 7in),[376] is the largest known captorhinid.[377] The second largest captorhinid was Labidosaurikos with the largest adult skull specimen 28cm (11in) long.[378]
Non-avian dinosaurs (Dinosauria)
Further information: Dinosaur
Further information: Dinosaur size
Sauropodomorphs (Sauropodomorpha)
The largest of non-sauropod sauropodomorphs ("prosauropod") was Euskelosaurus. It reached 12.2m (40ft) in length and 2t (2.2 short tons) in weight.[379] Another huge sauropodomorph Yunnanosaurus youngi reached 13m (43ft) long.[380]
Sauropods (Sauropoda)
Size comparison of selected giant sauropod dinosaurs (from left to right): Supersaurus, Argentinosaurus, Diplodocus, Xinjiangtitan, 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 58–60m (190–197ft) in maximum length and 122,400kg (269,800lb) in weight.[381] Unfortunately, the fossil remains of this dinosaur have been lost.[381] More recently, it was estimated at 35–40m (115–131ft) in length and 80–120t (180,000–260,000lb) in weight.[382]
Known from the incomplete and now disintegrated remains, the Late Cretaceous Bruhathkayosaurus matleyi was an anomalously large sauropod.[383] Informal estimations suggested as huge parameters as 45m (148ft) in length and 139–220t (306,000–485,000lb) in weight.[384] More accurate estimation suggests 37m (121ft) and 95t (209,000lb) but it still much heavier than most other sauropods.[384]
BYU 9024, a massive cervical vertebra found in Utah,[385] may belong to Barosaurus lentus[386][387] or Supersaurus vivianae[388] of a huge size, possibly 45–48m (148–157ft) in length and 60–66t (132,000–146,000lb) in body mass.[389][387]Supersaurus vivianae itself may have been the longest dinosaur yet discovered as a study of 3 specimens suggested length of 39m (128ft) or over 40m (130ft).[388]
Mounted skeleton of Mamenchisaurus sinocanadorum
Mamenchisaurus sinocanadorum was likely the largest mamenchisaurid, reaching nearly 35m (115ft) in length and 60–80t (130,000–180,000lb) in weight.[382]Xinjiangtitan shanshanesis from the same family had 15m (49ft)-long neck, about 55% of its total length that could be at least 27m (89ft).[390]
The Middle Jurassic Breviparopus taghbaloutensis was mentioned in The Guinness Book of Records as the longest dinosaur at 48m (157ft) although this animal is known only from fossil tracks.[391][392] Originally thought to be a brachiosaurid, it was later identified as a huge diplodocoid, possibly 33.5m (110ft) in length and 62t (137,000lb) in weight.[393]
Reconstructed skeleton of Argentinosaurus
The tallest sauropod was Sauroposeidon proteles with estimated height at 16.5–18m (54–59ft).[394][395][396]Asiatosaurus could reach 17.5m (57ft) in height but this animal is known only from teeth.[396]Giraffatitan was estimated at 16m (52ft) in height.[397]
Other huge sauropods include Argentinosaurus, Alamosaurus, and Puertasaurus with estimated lengths of 30–33m (98–108ft) and weights of 50–80t (55–88 short tons).[398]Patagotitan was estimated at 37m (121ft) in length[399] and 57t (63 short tons) in average weight,[400] and was similar in size to Argentinosaurus and Puertasaurus.[401] Giant sauropods like Supersaurus, Sauroposeidon, and Diplodocus probably rivaled them in length but not in weight.[381]Dreadnoughtus was estimated at 49t (108,000lb) in weight[400] and 26m (85ft) in length but the most complete individual was immature when it died.[402]Turiasaurus is considered of being the largest dinosaur from Europe,[403][404] with an estimated length of 30m (98ft) and a weight of 50t (55 short tons).[398][404] However, with lower estimate at 21m (69ft) and 30t (66,000lb) it was smaller than Portuguese Lusotitan that reached 24m (79ft) in length and 34t (75,000lb) in weight.[405]
Many large sauropods are still unnamed and may rival the current record holders:
The "Archbishop", a large brachiosaur that was discovered in 1930. The animal was reported to get a scientific paper published by the end of 2016.[406]
Brachiosaurus nougaredi is yet another large brachiosaur from Early Cretaceous North Africa. The remains have been lost, but the sacrum drawing remains. They suggest a sacrum of almost 1.3m (4.3ft) long,[407] making it the largest dinosaur sacrum discovered so far, except those of Argentinosaurus and Apatosaurus.[408]
In 2010, the femur of a large sauropod was discovered in France. The femur suggests an animal that grew to immense sizes.[409]
Non-avian theropods (Theropoda)
Size comparison of selected giant theropod dinosaurs (from left to right): Spinosaurus , Giganotosaurus, Tyrannosaurus, Mapusaurus, and Carcharodontosaurus
The largest theropod as well as the largest terrestrial (or possibly semi-aquatic)[410] predator yet known is Spinosaurus aegyptiacus, with the largest specimen known estimated at 12.6–18m (41–59ft) in length and around 7–20.9t (8–23 short tons) in weight.[411][412] New estimates published in 2014 and 2018, based on a more complete specimen supported that Spinosaurus could reach lengths of 15 to 16 meters (49 to 52ft).[413][414][415] The latest estimates suggest a weight of 6.4 to 7.5 metric tons (7.1 to 8.3 short tons).[414][415] The White Rock spinosaurid had vertebrae comparable in dimensions to Spinosaurus, it was likely a huge theropod with a length over 10m (33ft).[416]
Other large theropods were Giganotosaurus carolinii, and Tyrannosaurus rex, whose largest specimens known estimated at 13.2m (43ft)[417] and 12.3m (40ft)[418] in length, and weigh between 4.2 to 13.8t (4.6 to 15.2 short tons)[419][420][421][411] and 4.5 metric tons (5.0 short tons)[422][423] to over 7.2 metric tons (7.9 short tons),[418] respectively. Some other notable giant theropods (e.g. Carcharodontosaurus, Acrocanthosaurus, and Mapusaurus) may also have rivaled them in size.
Macroelongatoolithus, ranging from 34–61cm (1.12–2.00ft) in length,[424] is the largest known type of dinosaur egg.[425] It is assigned to oviraptorosaurs like Beibeilong.[425]
Armoured dinosaurs (Thyreophora)
The largest-known thyreophoran was Ankylosaurus at 9m (30ft) in length and 6 tonnes (6.6 short tons) in weight.[426][427]Stegosaurus was also 9m (30ft) long[404] but around 5 tonnes (5.5 short tons) tonnes in weight.[citation needed]
Pachycephalosaurs (Pachycephalosauria)
The largest pachycephalosaur was the nominate Pachycephalosaurus. Previously claimed to be at 7m (23ft) in length,[404] it was later estimated about 4.5 metres (14.8ft) long and a weight of about 450 kilograms (990lb).[428]
Ceratopsians (Ceratopsia)
Size comparison of several members of Ceratopsidae (from left to right): Nasutoceratops, Styracosaurus, Centrosaurus, Pachyrhinosaurus, Eotriceratops, Triceratops, Pentaceratops, and Chasmosaurus
The largest ceratopsian known is Triceratops horridus, along with the closely related Eotriceratops xerinsularis both with estimated lengths of 9m (30ft). Pentaceratops and several other ceratopsians rival them in size.[429]Titanoceratops had one of the longest skull of any land animal, at 2.65m (8.7ft) long.[430]
Ornithopods (Ornithopoda)
From left to right: Shantungosaurus giganteus, Magnapaulia laticaudus, Edmontosaurus annectens, Saurolophus angustirostris, Hypsibema missouriensis, Charonosaurus jiayinensis, Iguanodon bernissartensis
The very largest known ornithopods, like Shantungosaurus were as heavy as medium-sized sauropods at up to 23t (25 short tons),[431][432] and 16.6m (54ft) in length.[431]Magnapaulia reached 12.5m (41ft) in length,[433] or, according to original description, even 15m (49ft).[434][404] The Mongolian Saurolophus, S. angustirostris, reached 13m (43ft) long and possibly more.[435] Such animal could weighed up to 11t (12 short tons).[435] The largest Edmontosaurus reached 12m (39ft) in length and around 6t (6.6 short tons) in body mass.[436] An estimated maximum length of Brachylophosaurus is 11m (36ft) resulting in weight of 7t (7.7 short tons).[437] PASAC-1, informally named "Sabinosaurus", is the largest well-known North American saurolophine,[438] around 11m (36ft) long, that is about 20% larger than other known specimens.[439]Hypsibema missouriensis was up to 10.7m (35ft) long.[440][441] The Late Cretaceous Charonosaurus was estimated around 10m (33ft) in length and 5t (5.5 short tons) in weight.[404][442]
The largest ornithopod outside of Hadrosauroidea was likely the Iguanodon. Biggest specimens reached 11m (36ft) in length[443][444] and weighed around 4.5t (5.0 short tons).[445] Another large ornithopod is Iguanacolossus, with 9m (30ft) in length and 5t (5.5 short tons) in weight.[446][447]
The largest rhabdodontid was Matheronodon, estimated at 4.8m (16ft) in length.[448]Rhabdodon reached approximately 4m (13ft) and 250kg (550lb) according to 2016 estimates.[449]
Birds (Aves)
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 known birds of all time might have been the elephant birds of Madagascar. Both were about 3m (9.8ft) tall and 500 kilograms (1,100lb) in weight.[450] Nearly the same size was the Australian Dromornis stirtoni (see below). The tallest bird ever was the giant moa at 3.6m (12ft) tall.[451]
The widest known wingspan of any flight-capable bird was Pelagornis sandersi with a wingspan of 7.3m (24ft), and a body weight of 21.7kg (48lb). The heaviest flight-capable bird was the giant teratorn, Argentavis magnificens which had a somewhat-smaller wingspan at around 5.09–6.5m (16.7–21.3ft)[452][453] but was far heavier, with accepted maximums around 70–72kg (154–159lb).[454]
Enantiornitheans (Enantiornithes)
One of the largest enantiornitheans was Enantiornis,[455] with a length in life of around 78.5cm (30.9in), hip height of 34cm (13in), weight of 6.75kg (14.9lb),[456] and wingspan comparable to some of the modern gulls, around 1.2m (3ft 11in).[455]Gurilynia was the largest Mesozoic bird from Mongolia, with a length of 53cm (21in), hip height of 23.2cm (9.1in), and weight of 2.1kg (4.6lb).[456]
Avisauridae
Two Mirarce sitting on a head of ceratopsian dinosaur
The Late Cretaceous Avisaurus was almost as large as Enantiornis. It had a wingspan around 1.2m (3ft 11in),[455] a length of 72cm (28in), hip height of 31.5cm (12.4in), and weight of 5.1kg (11lb).[456] Even larger could be the Soroavisaurus. One tibiotarsus (PVL-4033) indicates an animal with a length of 80cm (31in), hip height of 35cm (14in), and weight of 7.25kg (16.0lb).[456]Mirarce was comparable in size to a turkey, much larger than most of other enantiornitheans.[457]
Pengornithidae
One of the biggest Early Cretaceous enantiornithine bird was Pengornis at 50cm (1.6ft) in length[404] and skull length of 54.7mm (2.15in).[458]
Gargantuaviidae
Gargantuavis is the largest known bird of the Mesozoic, a size ranging between the cassowary and the ostrich, and a mass of 140kg (310lb) like modern ostriches.[459] In 2019 specimens MDE A-08 and IVPP-V12325 were measured at 1.8m (5ft 11in) in length, 1.3m (4ft 3in) in hip height, and 120kg (260lb) in weight.[460]
Dromornithiformes
A cast of Dromornis stirtoni from Australia
The largest dromornithid was Dromornis stirtoni over 3m (9.8ft) tall[461] and 528–584kg (1,164–1,287lb) in mass for males.[462]
Gastornid (Gastornithiformes)
Large individuals of Gastornis (also known as Diatryma) reaged up to 2m (6ft 7in) in height.[463] Weight of Gastornis ranges from 100kg (220lb) to 156kg (344lb) and sometimes to 180kg (400lb) for European specimens and from 160kg (350lb) to 229kg (505lb) for North American.[464][465][466]
Waterfowl (Anseriformes)
Reconstruction of Garganornis ballmanni
Possibly flightless, the Miocene Garganornis ballmanni was larger than any extant members of Anseriformes, with 15.3–22.3kg (34–49lb) in body mass.[467] Another huge anseriform was the flightless New Zealand goose (Cnemiornis). It reached 15–18kg (33–40lb), approaching in size to small species of moa.[468]
Swans (Cygnini)
The largest swan of ever evolved was the Pleistocene giant swan (Cygnus falconeri), it reached bill-to-tail length of about 190–210cm (75–83in),[469] weighed around 16kg (35lb) and had a wingspan of about 3m (9.8ft).[470][471][472] The New Zealand swan (Cygnus sumnerensis) weighed up to 10kg (22lb), much more than related black swan at only 6kg (13lb).[473] The giant Annakacygna yoshiiensis from the Miocene of Japan was much bigger than the extant mute swan.[474]
Anatinae
Finsch's duck (Chenonetta finschi) reached 1–2kg (2.2–4.4lb) in weight, surpassing related modern Australian wood duck (800g (1.8lb)).[475]
Pelicans, ibises and allies (Pelecaniformes)
The Early Pliocene Pelecanus schreiberi was larger than most extant pelicans. Pelecanus odessanus from the Late Miocene was probably the same size as P. schreiberi, its tarsometatarsus is 150mm (5.9in) long.[476]
Storks and allies (Ciconiiformes)
Leptoptilos robustus compared in size to a human
The largest known of Ciconiiformes was Leptoptilos robustus, standing 1.8m (5ft 11in) tall and weighing an estimated 16kg (35lb).[477][478]
Cranes (Gruiformes)
A huge 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).[479]
Shorebirds (Charadriiformes)
Miomancalla howardi was the largest charadriiform of all time, weighing approximately 1.5ft (0.46m)(?) more than the great auk with a height of approximately 1m (3.3ft).[480]
Hesperornithines (Hesperornithes)
The largest known of the hesperornithines was Canadaga arctica at 2.2m (7ft 3in) long.[481]
New World vultures (Cathartiformes)
A skeleton of Teratornis
One of the heaviest flying bird ever was Argentavis from the family Teratornithidae. The immense bird had a wingspan estimated up to 5.09–6.5m (16.7–21.3ft)[452][453] and a weight up to 70 to 72kg (154 to 159lb).[482][452]Argentavis's humerus was only slightly shorter than an entire human arm.[483] Another huge teratorn was Aiolornis, it had a wingspan around 5m (16ft).[484] The Pleistocene Teratornis merriami reached 13.7kg (30lb) and 2.94–3.38m (9.6–11.1ft) in wingspan.[485] Even with lower estimates, it was larger than the observed California condor (Gymnogyps californianus) of nowadays.[485]
Seriemas and allies (Cariamiformes)
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 175cm (69in) tall at the shoulder, could raise its head 2.8m (9ft 2in) above the ground and could have weighed as much as 400kg (880lb).[486] The immense phorusrhacid Kelenken stood 3m (9.8ft) tall[487][488] with a skull 716mm (28.2in) long (460mm (18in) of which was beak), had the largest head of any known bird.[487] South American Phorusrhacos stood nearly 2.4 to 2.7 meters (7ft 10in to 8ft 10in) tall, and probably weighed nearly 130 kilograms (290lb), as much as a male ostrich.[489][490] The largest North American phorusrhacid is Titanis, which is about 2.5m (8ft 2in) tall,[491] as tall as a forest elephant.
Accipitriforms (Accipitriformes)
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 3m (8ft 6in to 9ft 10in), relatively short for their size.[492][493] Total length was probably up to 1.4m (4ft 7in) in female[494] and they weighed about 10 to 15kg (22 to 33lb).[495] Another giant extinct hawk was Titanohierax about 7.3kg (16lb) that lived in the Antilles and The Bahamas, where it was among the top predators.[496] An unnamed late Quaternary eagle from Hispaniola could be 15–30% larger than the modern golden eagle (Aquila chrysaetos).[497] 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).[498]B. daggetti, also known as "walking eagle", was around 40% larger than the savanna hawk (B. meridionalis).[499] Eyles's harrier (Circus eylesi) from the Pleistocene-Holocene of New Zealand was more than twice heavier than the extant C. approximans.[500]
Moa (Dinornithiformes)
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 3.7m (12ft) tall,[451] and weighed approximately half as much as a large elephant bird due to its comparatively slender frame.[450]
Tinamous (Tinamiformes)
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.[501]
Elephant birds (Aepyornithiformes)
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 3m (9.8ft) in height and 500 kilograms (1,100lb) in weight.[450]
Ostriches (Struthioniformes)
With 450kg (990lb) 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.[502] A possible specimen of Pachystruthio from the lower Pleistocene of Hebei Province (China) was about 300kg (660lb) in weight, twice heavier than the common ostrich (Struthio camelus).[503] Remains of the massive asian ostrich (Struthio asiaticus) from the Pliocene[504] indicate a size 20% bigger than adult male of the extant Struthio camelus.[505]
Pigeons and doves (Columbiformes)
Dodo (Raphus cucullatus)
The largest pigeon relative known was the dodo (Raphus cucullatus), possibly exceeding 1m (3.3ft) in height and weighing as much as 28kg (62lb), although recent estimates have indicated that an average wild dodo weighed much less at approximately 10.2kg (22lb).[506][507]
Pheasants, turkeys, gamebirds and allies (Galliformes)
The largest known of the Galliformes was likely the giant malleefowl, which could reach 7kg (15lb) in weight.[508]
Songbirds (Passeriformes)
The largest known songbird is the extinct giant grosbeak (Chloridops regiskongi) at 280mm (11in) long.[citation needed]
Cormorants and allies (Suliformes)
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 6.4kg (14lb) and 1.15m (3ft 9in).[509]
The largest known darter was Giganhinga with estimated weight about 17.7kg (39lb),[510] earlier study even claims 25.7kg (57lb).[511]
The largest known plotopterid, penguin-like flightless bird was Copepteryx titan that is known from 22cm (8.7in) long femur, almost twice as long as that of emperor penguin.[512]
Grebes (Podicipediformes)
The largest known grebe, the Atitlán grebe (Podylimbus gigas), reached a length of about 46–50 centimetres (18–20in).[513]
Bony-toothed birds (Odontopterygiformes)
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 5.5–6m (18–20ft) and stood about 1.2m (3ft 11in) 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)
The largest known woodpecker is the possibly extinct imperial woodpecker (Campephilus imperialis) with a total length of about 56–60cm (22–24in).[514]
Parrots (Psittaciformes)
The largest known parrot is the extinct Heracles inexpectatus with a length of about 1 meter (3.3 feet).[515]
Penguins (Sphenisciformes)
Size comparison of the giant penguin Anthropornis nordenskjoeldi
The largest known penguin of all time was Palaeeudyptes klekowskii of Antarctica, its body length (tip of the bill to tip of the tail) is estimated about 2.02m (6ft 8in) and body weight is estimated about 116.21kg (256.2lb).[516] Another large penguin is Anthropornis nordenskjoeldi of New Zealand and Antarctica. Its body length is estimated 1.99m (6ft 6in) and was 97.8kg (216lb) in weight. There is also an estimate that one remain of Anthropornis can reach that body length of 2.05m (6ft 9in) and 108kg (238lb) in weight.[517] Similar in size were the New Zealand giant penguin (Pachydyptes pondeorsus) with a height of 1.4 to 1.6m (4ft 7in to 5ft 3in) and weighing possibly around 80 to 100kg (180 to 220lb) and over, and Icadyptes salasi at 1.5m (4ft 11in) tall.[518]
Owls (Strigiformes)
The largest known owl of all time was the Cuban Ornimegalonyx at 1,100mm (43.3in) tall probably exceeding 9kg (20lb).[519]
Amphibians (Amphibia)
The largest known amphibian of all time was the 9.1m (30ft) long temnospondyl Prionosuchus.[520]
Lissamphibians (Lissamphibia)
Frogs and toads (Anura)
Size comparison of Beelzebufo
The largest known frog ever was an as yet unnamed Eocene species that was about 58–59.1-centimetre-long (22.8–23.3in).[521] The Late Cretaceous Beelzebufo grew to at least 23.2cm (9.1in) (snout-vent length), which is around the size of a modern African bullfrog.[522]
Salamanders, newts and allies (Urodela)
Andrias matthewi size comparison
Andrias matthewi was the largest lissamphibian ever known, with total length up to 2.3m (7ft 7in).[523]
Habrosaurus was the largest sirenid. It reached 1.6m (5ft 3in) long.[524]
Diadectomorphs (Diadectomorpha)
Size comparison of Diadectes
The largest known diacectid, herbivorous Diadectes, was a heavily built animal, up to 3m (9.8ft) long, with thick vertebrae and ribs.[525][526]
Anthracosauria
The largest known anthracosaur was Anthracosaurus, with skull about 40cm (16in) in length.[527]
Embolomeri
Restoration of Pholiderpeton
The longest member of this group was Eogyrinus attheyi, species sometimes placed under genus Pholiderpeton.[528] Its skull had length about 41cm (16in).[529]
Temnospondyls (Temnospondyli)
Scale diagram of small and large specimens of Prionosuchus
The largest known temnospondyl amphibian is Prionosuchus, which grew to lengths of 9m (30ft).[520] Another huge temnospondyl was Mastodonsaurus giganteus at 6m (20ft) long.[530] Unnamed species of temnospondyl from Lesotho is partial, but possible body length estimation is 7m (23ft).[531]
Fishes (Pisces)
Fishes are a paraphyletic group of non-tetrapod vertebrates.
Jawless fish (Agnatha)
Further information: Agnatha
Conodonts (Conodonta)
Iowagnathus grandis is estimated to have length over 50cm (1.6ft).[532]
Heterostracans (Heterostraci)
Some members of Psammosteidae such as Obruchevia and Tartuosteus are estimated to reached up to 2m (6.6ft).[533]
Thelodonts (Thelodonti)
Although known from partial materials, Thelodus parvidens (=T. macintoshi) is estimated to reached up to 1m (3.3ft).[534]
Cephalaspidomorphs (Cephalaspidomorphi)
A species of Parameteoraspis reached up to 1m (3.3ft).[535]
Spiny sharks (Acanthodii)
The largest of the now-extinct Acanthodii was Xylacanthus grandis, an ischnacanthiform based on a ~35cm (14in) long jaw bone. Based on the proportions of its relative Ischnacanthus, X. grandis had an estimated total length of 2.5m (8ft 2in).[536]
Placoderms (Placodermi)
Cast of a Dunkleosteus skull
The largest known placoderm was the giant predatory Dunkleosteus. The largest and most well known species was D. terrelli, which grew almost 9m (29.5ft) in length[537] and 4t (4.4 short tons)[538] in weight. Its filter feeding relative, Titanichthys, may have rivaled it in size.[539]Titanichthys reached a length of 7m (23ft)[540][541] though in older paper it was estimated at 7.5m (25ft).[542]
Cartilaginous fish (Chondrichthyes)
Further information: Cartilaginous fish
Mackerel sharks (Lamniformes)
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[543][544][545] is by far the biggest mackerel shark ever known.[546] Most estimates of megalodon's size extrapolate from teeth, with maximum length estimates up to 10–20.3m (33–67ft)[544][545][547] and average length estimates of 10.5m (34ft).[548][549] Due to fragmentary remains, there have been many contradictory size estimates for megalodon, as they can only be drawn from fossil teeth and vertebrae.[550]:87[551] 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 14.3m (34 to 47ft) and females 13.3 to 17m (44 to 56ft).[552] Related to megalodon, Otodus angustidens and O. chubutensis reached the large sizes too. Each was estimated at 9.3m (31ft)[553] and 12.2m (40ft),[554] respectively.
Other giant mackerel sharks were Pseudoscapanorhynchidae from the Cretaceous period. Cretodus had a size range of 9–11m (30–36ft) (for C. crassidens),[555]Leptostyrax reached lengths of 6.3–8.3m (21–27ft).[556]
The Cenozoic Parotodus reached up to 7.6m (25ft) in length.[557]
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.[558]
Ground sharks (Carcharhiniformes)
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 (20ft) long.[559]
Hybodonts (Hybodontiformes)
One of the largest hybodontiforms was the Jurassic Asteracanthus with body length of up to 3m (9.8ft).[560]Crassodus reifi is known from less materials, however it is estimated that reached over 3m (9.8ft).[561]
Skates and allies (Rajiformes)
The giant sclerorhynchid Onchopristis reached about 4.25m (13.9ft) in length.[562]
Eugeneodont (Eugeneodontida)
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 12m (39ft) in length.[563] Another fairly large eugeneodont is Parahelicoprion. Being more slimmer than Helicoprion, it reached nearly the same size,[563] possibly up to 12m (39ft) in length.[564] Both had the largest sizes among the animals of Paleozoic era.[565][564]
Lobe-finned fish (Sarcopterygii)
Further information: Sarcopterygii
Coelacanths (Actinistia)
Size estimation of Mawsonia gigas
The largest coelacanth is Cretaceous Mawsonia gigas with estimated total length up to 5.3m (17ft). Jurassic Trachymetopon may have reached size close to that, about 5m (16ft).[566] An undetermined mawsoniid from the Maastrichtian deposits of Morocco probably reached 3.65–5.52m (12.0–18.1ft) in length.[567][566]
Lungfish (Dipnoi)
Cretaceous Ceratodus sp. from Western Interior is estimated to had a length around 4m (13ft).[568]
Stem-tetrapods (Tetrapodomorpha)
Reconstruction of RhizodusReconstruction of Hyneria
Not only the largest known rhizodont, but also the largest lobe-finned fish was the 6–7m (20–23ft) long Rhizodus.[569] Another large rhizodonts were Strepsodus with estimated length around 3–5m (9.8–16.4ft) and Barameda estimated at 3–4m (9.8–13.1ft) in length.[570][571]
Tristichopterid Hyneria reached length up to 4m (13ft).[572]
Ray-finned fish (Actinopterygii)
Further information: Actinopterygii
Pachycormiformes
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 16.5m (54ft) long.[573] Earlier estimates have had claims of larger individuals with lengths over 27m (89ft).[574][575]
Ichthyodectiformes
Comparation of some ichthyodectiforms: Xiphactinus (1), Ichthyodectes (2), Cladocyclus (3), Chirocentrites (4)
The largest known of ichthyodectiform fish was Xiphactinus, which measured up to 6.1m (20ft) long.[576]Ichthyodectes reached 3m (9.8ft) long, twice lesser than Xiphactinus.[577]
Bichirs (Polypteriformes)
The Late Cretaceous Bawitius was likely the largest bichir of all time. It reached up to 3m (9.8ft) in length.[578]
Opahes, ribbonfishes, oarfishes and allies (Lampriformes)
Megalampris was likely the largest fossil opah. This fish was around 4m (13ft) in length when alive, which is twice the length of the largest living opah species, Lampris guttatus.[579]
Salmon and trout (Salmoniformes)
The largest salmon was Oncorhynchus rastrosus, varying in size from 1.9m (6ft 3in) and 177kg (390lb)[580] to 2.4m (7ft 10in) and 200kg (440lb).[581][580]
Pufferfishes, boxfishes, triggerfishes, ocean sunfishes and allies (Tetraodontiformes)
Austromola angerhoferi had total body length about 3.2m (10ft), and total height 4m (13ft), comparable with largest ocean sunfish.[582][583]
Some extinct species of Balistes like B. vegai and B. crassidens are estimated to have total length up to 1.8m (5ft 11in).[584]
Lizardfishes (Aulopiformes)
The largest lizardfish was Stratodus which could reach length of 5m (16ft).[585]
Echinoderms (Echinodermata)
Crinozoa
Fossil of Seirocrinus subangularis
Sea lilies (Cricoidea)
Longest stem of Seirocrinus subangularis reached over 26m (85ft).[586]
Asterozoa
Starfish (Asteroidea)
Helianthaster from Hunsrück Slate had radius about 25cm (9.8in).[587]
Graptolites (Graptolithina)
The longest known graptoloid graptolite is Stimulograptus halli at 1.45m (4.8ft). It found in Silurian deposits of the United Kingdom.[588]
Kinorhynchs (Kinorhyncha)
Cambrian kinorhynchs from Qingjiang biota, also known as "mud dragons", reached 4cm (1.6in) in length, much larger than extant relatives that grow only a few millimeters in length.[589][590]
Arthropods (Arthropoda)
Dinocaridida
Gilled lobopodians
Size estimation of Omnidens.
Based on the findings of mouthparts, the Cambrian gilled lobopodian Omnidens amplus is estimated to have been 1.5 metres (4.9ft).[591] It is also known as the largest Cambrian animal known to exist.[591]
Radiodont (Radiodonta)
Scaled diagram of Aegirocassis
The largest known radiodont is Aegirocassis benmoulai, estimated to have been at least 2m (6ft 7in) long.[592][593]
Chelicerata
Sea spiders (Pycnogonida)
The largest fossil sea spider is Palaeoisopus problematicus with legspan about 32cm (13in).[594]
Horseshoe crabs and allies (Xiphosura)
Willwerathia reached 9cm (3.5in) in carapace width and was the largest species of basal ("synziphosurine") xiphosurans.[595][596] However, the Devonian Maldybulakia reached nearly 11.5cm (4.5in)[597] and was assigned to xiphosurans in 2013.[596]
Horseshoe crab trackway icnofossil Kouphichnium lithographicum from Cerin in Ain indicates length of animal 77.4–85.1cm (30.5–33.5in).[598]
Chasmataspidids (Chasmataspidida)
Size comparison of the chasmataspidids
The largest chasmataspidids were the Ordovician Hoplitaspis at 29cm (11in) in length and similar in size range Chasmataspis.[599]
Eurypterids (Eurypterida)
Size comparison of the largest known eurypterids
The largest known eurypterid was Jaekelopterus rhenaniae at 2.5m (8ft 2in) in length, which is also the largest arthropod known to exist.[600]Erettopterus grandis possibly reached this same length but this is based on an incomplete telson only. A close contender was Acutiramus bohemicus at 2.1m (6ft 11in) in length.[601] The largest megalograptid as well as the largest Ordovician eurypterid was Pentecopterus. It reached up to 1.7m (5ft 7in) in length.[602] All these were eurypterine eurypterids.[601][602]
The largest stylonurine eurypterid was Hibbertopterus, with 1.8m (5ft 11in) in length.[601]
Arachnids (Arachnida)
There are three contenders for largest-known arachnid as well as the largest scorpions of all time: Pulmonoscorpius kirktonensis, Brontoscorpio anglicus and Praearcturus gigas. Each was estimated to have been 70cm (28in),[603]90cm (35in)[604] and up to 100cm (39in),[605] respectively.
Mongolarachne jurassica is the largest described fossil spider, with the total body length of female is approximately 24.6mm (0.97in) while the front legs reach about 56.5mm (2.22in) in length.[606]Dinodiplura ambulacra had larger body length, combined length of carapace and opisthosoma reaches 26.15mm (1.030in).[607]
The largest of prehistoric whipscorpions and possibly the largest-known whipscorpion ever discovered[608] was Mesoproctus. An unnamed species M. sp. had a carapace of 32.5mm (1.28in) in length and 16mm (0.63in) in width, comparable or even larger than the extant Mastigoproctus have.[609]
The largest Ricinulei to ever exist was Curculioides bohemondi with a body length of 21.77mm (0.857in).[610]
The largest fossil acariform mite and also the largest erythraeoid mite ever recorded was Immensmaris chewbaccei with idiosoma of more than 8mm (0.31in) in length.[611]
The largest known trigonotarbid was Kreischeria with a minimal length of 51mm (2.0in).[612] The second largest was Pleophrynus at 36mm (1.4in) in length.[612]
Artiopods (Artiopoda)
Further information: Artiopoda
Retifacies probably reached up to 55cm (22in).[613]Tegopelte is another one example of large non-trilobite artiopod, reached 280mm (11in) long[614] and was the largest of the Burgess Shale bilaterians, surpassing all other benthic organisms by at least twice.[614]
Trilobites (Trilobita)
Some of these extinct marine arthropods exceeded 60cm (24in) in length. A nearly complete specimen of Isotelus rex from Manitoba attained a length over 70cm (28in), 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 90cm (35in) long.[615]
Myriapods (Myriapoda)
A life-size reconstruction of Arthropleura
The largest known myriapod by far was Arthropleura. Measuring 2.5 metres (8ft 2in) long[616] and 50 centimetres (20in) wide.[617] Some specimens could have been even larger, up to 2.63 metres (8ft 8in) in length and 50 kilograms (110lb) in weight.[618][619]
Non-hexapod crustaceans (Crustacea)
Cycloids (Cyclida)
The largest cyclid was Opolanka decorosa, the Late Triassic Halicyne-like cycloid which reached over 6cm (2.4in) across the carapace.[620]
Remipedes (Remipedia)
Tesnusocaris had body length at least 9.5cm (3.7in),[621] larger than every living remipedes which could reach up to 4.5cm (1.8in).[622]
Sawflies, wasps, bees, ants and allies (Hymenoptera)
Titanomyrma with rufous hummingbird for scale
The largest known of this group was the giant ant Titanomyrma giganteum with queens growing to 6cm (2.4in). It had a wingspan of 15cm (5.9in).[623]
Apis lithohermaea is one of the largest honey bees ever found, comparable in size to the modern Apis dorsata.[624]
The giant horntail Ypresiosirex orthosemos reached 67.9mm (2.67in) in length including the incomplete ovipositor.[625] Another example of giant sawfly is Hoplitolyda duolunica, with wingspan over 92mm (3.6in).[626]
Fleas (Siphonaptera)
The largest known in Siphonaptera was probably Pseudopulex magnus, growing to 0.90in (22.8mm) in length.[627]
Earwigs (Dermaptera)
Labidura herculeana (St. Helena earwig) specimen
Extinct as recently as after 1967[628][629] and also submitted as the Holocene subfossils,[630] the Saint Helena giant earwig (Labidura herculeana, with synonym Labidura loveridgei) reached 84mm (3.3in) in length uncluding forceps 34mm (1.3in) long.[628]
Chresmodidae
Chresmodidae had long specialized legs like of the modern Gerridae family. One of the Chresmodidae, Chresmoda obscura could reached a size of about 19 centimetres (7.5in).[631]
Beetles (Coleoptera)
One of the largest known fossil beetles in the superfamily Scarabaeoidea is Protognathinus spielbergi. It had total length including mandibles about 5.5 centimetres (2.2in).[632] The largest fossil scarabaeid was Oryctoantiquus borealis with an estimated body length of 5 centimetres (2.0in).[633]
Titanopterans (Titanoptera)
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 (16in).[634]Clatrotitan andersoni also reached a huge size, having a forewing of 13.8 centimetres (5.4in) long.[635]
Antlions and related net-winged insects (Neuroptera)
Makarkinia adamsi from the Crato Formation is estimated to have the longest forewings of any neuropteran species, estimated at 160mm (6.3in).[636]
Cockroaches, termites, mantises and allies (Dictyoptera)
Further information: Dictyoptera
Some Carboniferous cockroach-like insects grouping in Blattoptera like Archoblattina beecheri[637] and Necymylacris (Xenoblatta) scudderi[638][639] could reach around 9 centimetres in total length, which is comparable to a modern Megaloblatta longipennis.
Cretaceous cockroach Ptiloteuthis foliatus had 7.9cm (3.1in) long wing.[640]
Found in the Miocene of Austria, the giant termite Gyatermes styriensis reached 25mm (0.98in) in body length and had a wingspan of 76mm (3.0in).[641]
Dragonflies, damselflies and griffinflies (Odonatoptera)
Reconstruction of Meganeura
The largest known odonatopteran insect was Meganeuropsis permiana with single wing of 33cm (13in). Meganeura had a 32cm (13in) long wing.[642]
Triadotypid odonatan Reisia gelasii (=Triadotypus guillaumei) from Triassic had 136 millimetres (5.4in) long wing, and wingspan can be 280 millimetres (11in).[643][644]
Mayflies (Ephemeroptera)
The largest known mayfly is Permian Ponalex maximus, with 55mm (2in) long hindwing.[645] Cretaceous Epicharmeropsis quadrivenulosus had 37mm (1in) long forewing.[646]
Although Bojophlebia prokopi from the Upper Carboniferous of Moravia (Czech Republic) with a wingspan of 45cm (18in) is described as the largest mayfly,[647] later study shows that this insect is not related to mayflies.[648]
Palaeodictyoptera
The largest known palaeodictyopteran was Mazothairos, with an estimated wingspan of up to 560mm (22in).[649] If subcircular wing known from Piesberg Quarry belongs to palaeodictyopteran, it possibly had single wing length at least 30cm (12in).[650]
Archaeognatha (jumping bristletails) and other wingless primitive insects
The largest known machilid is Triassic Gigamachilis, with 40 millimetres (1.6in) body length not counting the length of the filament, and estimated total length about 80 millimetres (3.1in).[651]
The largest specimens of the extinct suborder Monura reached 30 millimetres (1.2in) or more, not counting the length of the filament.[652]
Although Ramsdelepidion was once considered as 60 millimetres (2.4in)-long silverfish,[653] it was later considered that classification is uncertain and just treated as stem group insect.[654]
Wingless early insect Carbotriplura had body length about 103 millimetres (4.1in) without tail filaments.[655]
Ringed worms (Annelida)
Websteroprion is the largest known fossil eunicidan annelid, with estimated length ranges 0.42–8.3m (1ft 5in– 27ft 3in), however comparison with closely related extant taxa indicates length around 1–2m (3ft 3in– 6ft 7in).[656] It also had the biggest scolecodonts of any prehistoric polychaete, up to 13.2mm (0.52in) in length and possibly larger.[656]
Molluscs (Mollusca)
Snails and slugs (Gastropoda)
Campanile giganteum shell
The largest known gastropods were in the genus Campanile, with the extinct Campanile giganteum having shell lengths up to 90cm (35in)[657] or even more than 120cm (47in).[658]
The largest known cowrie is Vicetia bizzottoi, with shell length of 33.5cm (13.2in).[659]
Pebasiconcha immanis is the largest land snail ever known, shell height is 25.6cm (10.1in) with a partial specimen that may exceed 30cm (12in) in height.[660]
Bivalves (Bivalvia)
The largest known bivalve ever as well as the largest inoceramid was Platyceramus platinus, a giant that usually had an axial length of 1m (3ft 3in), but some individuals could reach an axial length of up to 3m (9.8ft).[661] Another large prehistoric bivalve was Inoceramus. In 1952, 187cm (6.14ft)-long specimen of Inoceramus steenstrupi was found in the Late Cretaceous deposits of Greenland.[662]
Some Permian alatoconchid genus like Shikamaia had shell length about 1m (3ft 3in).[663] Previous estimation reconstructed length of Shikamaia around 1.6m (5ft 3in).[664]
The longest ostreid is Konbostrea, with shell height reaching up to 1.2m (3ft 11in).[665]
Rudist Titanosarcolites had overall size around 2m (6ft 7in).[666]
Tusk shells (Scaphopoda)
Complete shell length of tusk shell Prodentalium onoi is estimated to be over 30cm (12in).[667]
Cephalopods (Cephalopoda)
Main article: Cephalopod size
Parapuzosia seppenradensis shell
Nautiloids (Nautiloidea)
The largest and longest known of nautiloids was Endoceras giganteum with a shell length of 5.73m (18.8ft). There is a record of individual whose shell length had reached 9.14m (30.0ft), but it is doubtful.[668]
Ammonites (Ammonoidea)
The largest known ammonite was Parapuzosia seppenradensis.[669] A partial fossil specimen found in Germany had a shell diameter of 1.95m (6ft 5in), but the living chamber was incomplete, so the estimated shell diameter was probably about 3.5m (11ft) and weighed about 705kg (1,554lb) when it was alive.[670] However, later study estimates shell diameter up to around 2m (6ft 7in).[671]
Belemnites (Belemnoidea)
The largest known belemnite was Megateuthis gigantea, reaching about 50 and 700mm (2.0 and 27.6in) in maximum diameter and length of rostrum, respectively.[672]
Squids, octopuses, cuttlefishes and allies (Neocoleoidea)
Octopod Enchoteuthis melanae (considered as specimen of Tusoteuthis longa) had mantle length up to 2 metres (6ft 7in), comparable to the modern-day giant squid. Previously, this taxon is considered as animal like giant squid, with total length including arms over 10 metres (33ft). However, considering other fossil relatives, total length including arms is estimated to be around 3 metres (9.8ft).[673]
Both non-octopod Yezoteuthis and teuthid Haboroteuthis are estimated to be similar in size to the modern-day giant squid.[674][675]
Brachiopods (Brachiopoda)
Gigantoproductus giganteus
The largest brachiopod ever evolved was Striatifera striata from Akkermanovka Quarry, Russia, with height up to 0.5 metres (1ft 8in).[676] Another huge brachiopod was the Carboniferous Gigantoproductus giganteus, with shell width from 30cm (12in)[677] to over 35 centimetres (14in).[666][678]Titanaria costellata had large and long shell 35–36cm (14–14in) in width, nearly as large as Gigantoproductus.[679]
Hyoliths (Hyolitha)
The largest hyolith is Macrotheca almgreeni, with length about 50 centimetres (20in).[666][680]
Cnidarians (Cnidaria)
Jellyfishes and allies (Medusozoa)
The largest fossil jellyfish is Cambrian Cordubia gigantea, with diameter of 88 centimetres (35in).[681] Specimens from the Cambrian of Wisconsin reached 70cm (28in) in length.[682]
Vendobionts (Vendobionta)
Further information: Vendobionta
Petalonamids (Petalonamae)
A large specimen of Trepassia wardae
Longest specimens of Trepassia wardae (also known as Charnia wardi) reached 185cm (73in) in length.[683]Charnia masoni is known from specimens as small as only 1cm (0.39in), up to the largest specimens of 66cm (26in) in length.[684]
Proarticulata
Dickinsonia rex reached 1.4m (4.6ft) in length, that makes it one of the largest precambrian organisms.[685][686]
Sponges (Porifera)
The largest known Permian sponge Gigantospongia had diameter up to 2.5 metres (8ft 2in).[687]
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