therizinosauroid

Therizinosaurs (once called segnosaurs) were large herbivorous theropod dinosaurs whose fossils have been found across the Early to Late Cretaceous deposits in Asia and North America. Various features of the forelimbs, skull and pelvis unite these finds as both theropods and maniraptorans, close relatives to birds. The name of the representative genus, ''Therizinosaurus'', is derived from the Greek (, 'to reap' or 'scythe')Translated paper
and (, 'lizard'). The older representative, ''Segnosaurus'', is derived from the Latin ('slow') and the Greek .

History of research

Therizinosaurs were long considered an enigmatic group, whose mosaic of features resembling those of various different dinosaur groups, and scarcity of their fossils, led to controversy over their evolutionary relationships for decades after their initial discovery. The first genus, ''Therizinosaurus'', was originally identified as a turtle when described from forelimb elements in 1954. Perle noted in 1979 that the ''Segnosaurus'' fossils were possibly representative of a new family of dinosaurs, which he tentatively classified as theropods (traditionally thought of as the "meat-eating" dinosaurs). He named the family Segnosauridae, with ''Segnosaurus'' as type genus and sole member. He distinguished Segnosauridae from the theropod families Deinocheiridae and Therizinosauridae (then only known from the genera ''Deinocheirus'' and ''Therizinosaurus'', both mainly represented by large forelimbs found in Mongolia) by features of their humeri and hand claws. Later in 1979, Barsbold and Perle found the pelvic features of segnosaurids and dromaeosaurids so different from those of "true" theropods that they should be separated into three taxa of the same rank, possibly at the level of infraorder within Saurischia (one of the two main divisions of dinosaurs, the other being Ornithischia).

In 1980, Barsbold and Perle named the new theropod infraorder Segnosauria, containing only Segnosauridae. In the same article, they named the new genus ''Erlikosaurus'' (known from a well-preserved skull and partial skeleton) which they tentatively considered a segnosaurid, and reported a partial pelvis of an undetermined segnosaurian, both from the same formation as ''Segnosaurus''. Combined, the specimens provided relatively complete data on this group; they were united by their opisthopubic pelvis, slender mandible, and the toothless front of their jaws. Barsbold and Perle stated that though some of their features resembled those of ornithischians and sauropods, these similarities were superficial, and were distinct when examined in detail. While they were essentially different from other theropods (perhaps due to diverging from them relatively early), and therefore warranted a new infraorder, they did show similarities with them. Since the ''Erlikosaurus'' specimen lacked a pelvis, the authors were unsure if that of the undetermined segnosaurian could belong to it, in which case they would consider it part of a separate family. Though ''Erlikosaurus'' was difficult to compare directly to ''Segnosaurus'' due to the incompleteness of their remains, Perle stated in 1981 that there was no justification for separating it into another family.

In 1982, Perle reported hindlimb fragments similar to those of ''Segnosaurus'', and assigned them to ''Therizinosaurus'', whose forelimbs had been found in almost the same location. He concluded that Therizinosauridae, Deinocheiridae, and Segnosauridae, which all had enlarged forelimbs, represented the same taxonomic group. ''Segnosaurus'' and ''Therizinosaurus'' were particularly similar, leading Perle to suggest they belonged in a family to the exclusion of Deinocheiridae (today, ''Deinocheirus'' is recognized as an ornithomimosaur).Translated paper
/ref> Barsbold retained ''Segnosaurus'' and ''Erlikosaurus'' in the family Segnosauridae in 1983, and named the new genus ''Enigmosaurus'' based on the previously undetermined segnosaurian pelvis, which he placed in its own family, Enigmosauridae, within Segnosauria. Though the structure of the pelvis of ''Erlikosaurus'' was unknown, Barsbold considered it unlikely the ''Enigmosaurus'' pelvis belonged to it, since ''Erlikosaurus'' and ''Segnosaurus'' were so similar in other respects, while the pelvis of ''Enigmosaurus'' was very different from that of ''Segnosaurus''. Barsbold found that segnosaurids were so peculiar compared to more typical theropods that they were either a very significant deviation in theropod evolution, or that they went "beyond the borders" of this group, but opted to retain them within Theropoda.Translated paper
/ref> In the same year, Barsbold stated that the segnosaurian pelvis deviated strongly from the theropod norm, and found the configuration of their ilia generally similar to those of sauropods.

Paleontologist Gregory S. Paul concluded in 1984 that segnosaurs did not possess any theropodan features, but were instead derived, late-surviving Cretaceous prosauropods with adaptations similar to those of ornithischians. He found segnosaurs similar to prosauropods in the morphology of their snout, mandible, and hindfoot, and to ornithischians in their cheek, palate, pubis, and ankle, and similar to early dinosaurs in other respects. He proposed that ornithischians were descended from prosauropods, and that the segnosaurs were an intermediate relict of this transition, which supposedly took place during the Triassic period. In this way, he considered segnosaurians to be to herbivorous dinosaurs what monotremes are to mammals. He did not rule out that segnosaurs could be derived from theropods, or that segnosaurs, prosauropods and ornithischians were each independently derived from early dinosaurs, but found these options unlikely. He considered the common descent of these groups as support for the idea that dinosaurs were a monophyletic (natural) group, which was contested by some paleontologists at the time (who instead thought different dinosaurs groups evolved independently from thecodonts). Paleontologist David B. Norman considered Paul's idea a contentious claim "bound to provoke much argument" in 1985. In 1988, Paul maintained that segnosaurs were late surviving ornithischian-like prosauropods, and proposed a segnosaurian identity for ''Therizinosaurus''. He also placed segnosauria within Phytodinosauria, a superorder that paleontologist Robert Bakker had created in 1985 to retain all plant-eating dinosaurs. In a 1986 study of the interrelationships of saurischian dinosaurs, paleontologist Jacques Gauthier concluded that segnosaurs were prosauropods. While he conceded they had similarities with ornithischians and theropods, he proposed these featured had evolved independently. In a 1989 conference abstract about sauropodomorph interrelationships, paleontologist Paul Sereno also considered segnosaurs as prosauropods, based on skull features.

In a 1990 review article, Barsbold and paleontologist Teresa Maryańska found Segnosauria to be a rare and aberrant group of saurischians, in an unresolved position among sauropodomorphs and theropods, probably closer to the former. They therefore listed them as Saurischia ''sedis mutabilis'' ("position subject to change"). Though they agreed the hindlimbs assigned to ''Therizinosaurus'' in 1982 were segnosaurian, they did not consider this justification for ''Therizinosaurus'' itself being a segnosaur, since it was only known from forelimbs. In 1993, paleontologists Dale A. Russell and Dong Zhi-Ming described the new genus ''Alxasaurus'' from China, at the time the most complete large theropod from its time and place. While it was similar to prosauropods in some respects, the detailed morphology of its limbs linked it to ''Therizinosaurus'' and segnosaurs. Since it preserved both fore and hindlimbs, ''Alxasaurus'' showed that Perle's assignment of segnosaurian hindlimbs to ''Therizinosaurus'' was probably correct. Russell and Dong therefore proposed that Segnosauridae was a junior synonym of Therizinosauridae (since the latter name was older), with ''Alxasaurus'' being the most completely known representative so far, providing a better understanding of the group. They also named the new higher taxonomic rank Therizinosauroidea to contain ''Alxasaurus'' and Therizinosauridae (since the new genus was somewhat different from its relatives), which they placed in the group Tetanurae within Theropoda. They considered therizinosaurs most closely related to ornithomimids, troodontids, and oviraptorids, which they placed together in the group Oviraptorosauria (since they found Maniraptora, the conventional grouping of these, invalid, and the higher level taxonomy of theropods was in flux at the time).

The synonymy of Segnosauridae with Therizinosauridae was accepted by Perle himself and co-authors of a redescription of the holotype skull of ''Erlikosaurus'' in 1994, and they considered therizinosaurs maniraptoran theropods, the group that also includes modern birds (since they did find Maniraptora to be valid through their analysis). They also discussed the previous ornithischian and sauropod hypotheses for therizinosaur affinities in detail and demonstrated various faults with them. Palaeontologist Lev Alexandrovich Nessov rejected that therizinosaurs were theropods in 1995, and instead considered them a distinct group within saurischia.Nessov, L. A., 1995, Dinosaurs of northern Eurasia: New data about assemblages, ecology and paleobiogeography. Saint Petersburg In 1996, paleontologist Thomas R. Holtz Jr. found therizinosaurs to group with oviraptorosaurs in a phylogenetic analysis of coelurosauria. In 1999, paleontologist Xing Xu and colleagues described a small, basal therizinosauroid from China, ''Beipiaosaurus'', which confirmed that the group belonged among the coelurosaurian theropods, and that similarities with prosauropods had evolved independently. They published the first ever cladogram showing the evolutionary relationships of Therizinosauria, and demonstrated that ''Beipiaosaurus'' had features of more basal theropods, coelurosaurs, and therizinosaurs. Sereno found Therizinosaurs to be basal Ornithomimosaurian theropods during the year 1999.

By the early 21st century, many more therizinosaur taxa had been discovered, including outside Asia (the first being ''Nothronychus'' from North America), as well as various basal taxa that helped understanding of the early evolution of the group (such as ''Falcarius'', also from North America). Therizinosaurs were not considered as rare or aberrant anymore, but more diverse than previously thought (including in size), and their classification as maniraptoran theropods was generally accepted. The placement of Therizinosauria within Maniraptora continued to be unclear; in 2007, paleontologist Alan H. Turner and colleagues found them to group with oviraptorosaurs, while Zanno and colleagues found them to be the most basal clade within Maniraptora in 2009, bracketed by Ornithomimosauria and Alvarezsauridae. Despite the additional fossil material, the interrelations within the group were also still uncertain by 2010, when Zanno conducted the most detailed phylogenetic analysis of the Therizinosauria until that point. She cited the inaccessibility, damage, potential loss of holotype specimens, scarcity of cranial remains, and fragmentary specimens with few overlapping elements as the most significant obstacles to resolving the evolutionary relationships within the group.

Uncertain species

In 1979 Dong Zhiming named a new species of the megalosaurid ''Chilantaisaurus'', ''C. zheziangensis'', based on specimen ZhM V.001. This specimen was recovered in 1972 from the Tangshang Formation and consists of a partial tibia and partial right pes (foot) largely lacking metatarsals. Dong referred it to the genus mainly based on similarities between the unguals of this specimen and those of ''C. tashuikouensis''.Translated paper
/ref> Barsbold and Maryanska in 1990 considered ''C. zheziangensis'' as a tentative segnosaur (later known as therizinosaurs) based on its relatively short and robust pedal phalanges and enlarged, strongly curved unguals, mostly similar to ''Segnosaurus''. As this taxon may lie outside the genus ''Chilantaisaurus'', they listed this species as ''"Chilantaisaurus" zheziangensis''. Although Glut (1997) stated this specimen may have been based on part of the holotype of Nanshiungosaurus brevispinus (based on a pers. comm from Dong to Molnar in 1984), Dong in 1979 described both taxa from largely different formations and localities. Zanno in 2010 argued that the notorious side to side compression of the unguals reflect therizinosaur affinities, although examinations to the preserved tibia are required for further conclusions. In 2012 Mai-Ping Qian and colleagues placed ''"C". zheziangensis'' in the family Therizinosauridae based on its pes morphology, which is consistent to other therizinosaurids. They also illustrated most of the preserved pes. Hartman with team in 2019 added ''"C". zheziangensis'' to a phylogenetic analysis and recovered it within Therizinosauroidea in a polytomy with ''Alxasaurus'', ''Enigmosaurus'' and therizinosaurids.

In 1997 Dong Zhiming and You Hailu named and described a supposed second species of ''Nanshiungosaurus'', ''N. bohlini'', based on specimen IVPP V 11116 found in 1992 at Early Cretaceous strata from the Zhonggou Formation, Xinminbao Group. It consists of 11 cervical and 5 dorsal vertebrae with some ribs. In order to contain both ''N. brevispinus'' and ''N. bohlini'' they coined the Nanshiungosauridae family. Dong and Yu presented no clear evidence regarding the assignment of this new species to ''Nanshiungosaurus''. Li and colleagues in their 2007 description of ''Suzhousaurus'' pointed out that ''N. bohlini'' might be synonymous with the former, as both are found in the same geological group and also incompletely known. As per terms of taxonomic priority, the species name would be ''Suzhousaurus bohlini''. However, they noted that a direct comparison between specimens is difficult to near impossible because there is no overlapping material (besides dorsal vertebrae) and the holotype of ''N. bohlini'' is apparently lost. Li and team disagree in that this species belong to ''Nanshiungosaurus'' and listed it as ''"Nanshiungosaurus" bohlini''. Zanno in 2010 indicated that the anatomical traits that were originally used to characterize ''"N." bohlini'' are now known to be present in other therizinosaur taxa. Hartman with colleagues in 2019 recovered ''"N." bohlini'' as a therizinosaurid in a clade joined by ''Segnosaurus'' and ''Nothronychus''.

Around 2005 partial therizinosaur material was collected from the Laijia Formation and later used to represent "Tiantaiosaurus sifengensis" (alternatively "Tiantaisaurus"), which is a currently unpuslihed and informal therizinosaur taxon. Qian and team in 2012 noted that a whole manuscript describing the taxon was written in 2007 but never officially published.

In 1998 Zhao Xijin and Xu Xing briefly discovered a partial lower jaw with teeth (IVPP V11579) from the Early Jurassic-aged Lufeng Formation in Yunnan, and concluded that this specimen represented the oldest known record of a coelurosaurian theropod. Based mainly on teeth morphology, they indicated therizinosaur affinities. The specimen was latter described in depth in 2001 and used as the holotype specimen for the new genus and species ''Eshanosaurus deguchiianus'', named by Xing and colleagues. The team reinforced therizinosaur relationships, arguing that the teeth morphology of ''Eshanosaurus'' can be differentiated from sauropodomorphs. In the same 2001 however, James I. Kirkland and Douglas G. Wolfe noted that the holotype of ''Eshanosaurus'' preserves traits only seen in sauropodomorphs. Paul M. Barrett