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Therizinosaurus

Therizinosaurid genus from the Late Cretaceous period

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2014-03-25 12.46.44
Yuya Tamai from Gifu, Japan · CC BY 2.0

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Therizinosaurus was a colossal therizinosaurid that could grow up to 9–10 m (30–33 ft) long and 4–5 m (13–16 ft) tall, and weigh possibly over 5 t (5.5 short tons). Like other therizinosaurids, it would have been a slow-moving, long-necked, high browser equipped with a rhamphotheca (horny beak) and a wide torso for food processing. Its forelimbs were particularly robust and had three fingers that bore unguals which, unlike other relatives, were very stiffened, elongated, and only had significant curvatures at the tips. Therizinosaurus had the longest known manual unguals of any land animal, reaching above 50 cm (20 in) in length. Its hindlimbs ended in four functionally weight-bearing toes differing from other theropod groups in which the first toe was reduced to a dewclaw and also resembling the very distantly related sauropodomorphs.

It was one of the last and the largest representative of its unique group, the Therizinosauria (formerly known as Segnosauria; the segnosaurs). During and after its original description in 1954, Therizinosaurus had rather complex relationships due to the lack of complete specimens and relatives at the time. Maleev thought the remains of Therizinosaurus to belong to a large turtle-like reptile, and also named a separate family for the genus: Therizinosauridae. Later on, with the discovery of more complete relatives, Therizinosaurus and kin were thought to represent some kind of Late Cretaceous sauropodomorphs or transitional ornithischians, even though at some point it was suggested that it may have been a theropod. After years of taxonomic debate, nevertheless, they are now placed in one of the major dinosaur clades, Theropoda, specifically as maniraptorans. Therizinosaurus is widely recovered within Therizinosauridae by most analyses.

The unusual arms and body anatomy (extrapolated after relatives) of Therizinosaurus have been cited as an example of convergent evolution with chalicotheriines and other primarily herbivorous mammals, suggesting similar feeding habits. The elongated hand claws of Therizinosaurus were more useful when pulling vegetation within reach rather than being used for active attack or defense because of their fragility, however, they may have had some role for intimidation. Its arms also were particularly resistant to stress, which suggests a robust use of these limbs. Therizinosaurus was a very tall animal, likely having a reduced competition over the foliage in its habitat and outmatching predators like tyrannosaurid Tarbosaurus.

History of discovery

In 1948, several Mongolian Paleontological expeditions organized by the USSR Academy of Sciences were conducted in the Nemegt Formation of the Gobi Desert, Southwestern Mongolia, with the main objective of new fossils findings. The expeditions unearthed numerous dinosaur and turtle fossil remains from the stratotype locality Nemegt (also known as Nemegt Valley), but the most notable elements collected were three partial manual unguals (claw bones) of considerable size. This set of unguals was found on a subdivision of the Nemegt locality designated as Quarry V near the skeleton of a large theropod, but also in association with other elements including a metacarpal fragment and several ribs fragments. It was labelled under the specimen number PIN 551-483 and later on, these fossils were described by the Russian paleontologist Evgeny Maleev in 1954 who used them to scientifically name the new genus and type species Therizinosaurus cheloniformis, becoming the holotype specimen. The generic name, Therizinosaurus, is derived from the Greek θερίζω (therízo, meaning scythe, reap or cut) and σαῦρος (sauros, meaning lizard) in reference to the enormous manual unguals, and the specific name, cheloniformis, is taken from the Greek χελώνη (chelóni, meaning turtle) and Latin formis as the remains were thought to belong to a turtle-like reptile. Maleev also coined a separate family for this new and enigmatic taxon: Therizinosauridae. Since little was known of Therizinosaurus at the time of the original description, Maleev thought PIN 551-483 belonged to a large, 4.5 m (15 ft) long turtle-like reptile that relied on its giant hand claws to harvest seaweed.

Though it was not fully understood to what general kind of animal these fossils belonged, in 1970, the Russian paleontologist Anatoly K. Rozhdestvensky was one of the first authors to suggest that Therizinosaurus was a theropod and not a turtle. He made comparisons between Chilantaisaurus and the holotype unguals of Therizinosaurus to propose that the appendages actually came from a carnosaurian dinosaur, thereby interpreting Therizinosaurus as a theropod. Rozhdestvensky also illustrated the three holotypic manual unguals and re-identified the metacarpal fragment as a metatarsal bone, and based on the unusual shape of both metatarsal and ribs fragments he listed them as sauropod remains. These theropodan affinities were also followed by the Polish paleontologist Halszka Osmólska and co-author Ewa Roniewicz in 1970 during their naming and description of Deinocheirus—another large and enigmatic theropod from the formation that was initially known from partial arms. Similar to Rozhdestvensky, they suggested that the holotype unguals were more likely to have belonged to a carnosaurian theropod, rather than a large marine turtle.

Additional specimens

Further expeditions in the Nemegt Formation unearthed more fossils of Therizinosaurus. In 1968 prior to Rozhdestvensky, Osmólska and Roniewicz statements, the upper portion of a manual ungual was found in the Altan Uul locality and labeled as MPC-D 100/17 (formerly IGM or GIN). In 1972, another fragmented ungual (specimen MPC-D 100/16) was discovered at the Upper White Beds of the Hermiin Tsav locality, only preserving its lower portion. During the year 1973, a much more complete, larger, and articulated specimen was collected also from Hermiin Tsav. This specimen was labelled as MPC-D 100/15 and consists of both left and right arms including the scapulocoracoids, both humeri (upper arm bones), right ulna with radius and left ulna, two right carpals, the right metacarpus including a complete digit Il, and some ribs with gastralia (belly ribs). As common with fossils, some elements were not entirely preserved such as the scapulocoracoids with broken ends, and the left arm is less complete than the right one. All of these specimens were first described and referred to Therizinosaurus by the Mongolian paleontologist Rinchen Barsbold in 1976. In this new monograph, he pointed out that the rib fragments in MPC-D 100/15 were more slender than the ones from the holotype, and identified MPC-D 100/16 and 100/17 as pertaining to digits I and III, respectively. It was clear to Barsbold that MPC-D 100/15 represented Therizinosaurus as the ungual in this specimen shared the elongation and flattened morphology of all previous specimens. He concluded that Therizinosaurus was a theropod taxon since MPC-D 100/15 matched multiple theropodan characters.

Also during the year 1973, the specimen MPC-D 100/45 was discovered by the Joint Soviet-Mongolian Paleontological Expedition at the Hermiin Tsav locality. Unlike the previous findings, MPC-D 100/45 is represented by a right hindlimb composed of a very fragmented femur with the lower end of the tibia, astragalus, calcaneum, tarsal IV, a functional tetradactyl feet (four-toed) compromising four partial metatarsals, partially preserved digits I and III, and nearly complete digits II and IV. These newer remains were described by the also Mongolian paleontologist Altangerel Perle in 1982. He regarded the referral of Therizinosaurus and Therizinosauridae to Chelonia (turtles order) to be unlikely, and hypothesized Therizinosaurus and Segnosaurus—at the time of this description regarded as a theropod dinosaur—to be particularly similar based on their respective scapulocoracoid morphology, only differing in size. Perle referred MPC-D 100/45 to Therizinosaurus given that this specimen was found near the location of MPC-D 100/15 and was virtually similar to the described pes for Segnosaurus. In 1990, Barsbold and Teresa Maryanska agreed with Perle in that the hindlimb material from Hermiin Tsav he described in 1982 was therizinosaurian (then called segnosaurians) given that the metatarsus was stocky and the astragalus had a laterally arched ascending process (bony extension), but cast doubt with his referral of it to Therizinosaurus and the segnosaurian identity for this taxon since it was only known from the pectoral girdle and other forelimb elements, making direct comparisons between specimens impossible. They considered this specimen to represent a Late Cretaceous representative of the Segnosauria, but not Therizinosaurus.

In 2010 however, the North American paleontologist Lindsay E. Zanno in her large taxonomic reevaluation of Therizinosauria considered the referral of MPC-D 100/45 to Therizinosaurus to be likely based on the rationale that it was collected in the same stratigraphic context (Nemegt Formation) as the holotype, and shared the robust and four-toed morphology of other therizinosaurids such as Segnosaurus. She also excluded the rib material from the holotype as it was re-identified by Rozhdestvensky to likely have come from a sauropod dinosaur, and not Therizinosaurus itself.

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