Falcarius

Summary

Falcarius
Temporal range: Early Cretaceous,
~139–134 Ma
Falcarius.jpg
Reconstructed skeleton, Natural History Museum of Utah
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Clade: Maniraptora
Clade: Therizinosauria
Genus: Falcarius
Kirkland et al. 2005
Type species
Falcarius utahensis
Kirkland et al. 2005

Falcarius (meaning "sickle cutter") is a genus of therizinosaur theropod dinosaurs that lived in North America during the Early Cretaceous period in what is now the Cedar Mountain Formation. The first remains were collected in 1999 and subsequent findings were made during the 2000s. The genus is known from multiple specimens ranging from immature to fully-grown individuals.

Falcarius were 4 m (13 ft) long, bipedal herbivores with a small head and elongated neck and tail. Unlike advanced members, Falcarius had propubic pelves and three-toed feet with a reduced hallux (first digit).

The description of Falcarius, following that of the therizinosauroids Beipiaosaurus from the Early Cretaceous of China in 1999, clarifies the early evolution of the Therizinosauria and their relationship with the larger group of theropod dinosaurs, because Falcarius were transitional forms between older theropods and the much advanced Therizinosauridae. Falcarius are considered the most primitive therizinosaurs.

History of discovery

Reconstructed skull in dorsal view

The remains of Falcarius were first discovered in 1999 by commercial fossil collector Lawrence Walker at the Crystal Geyser Quarry site in Grand County. He informed paleontologist James I. Kirkland of the find, who with a team of the Utah Geological Survey from 2001 onwards began to uncover the bones that proved to be present in a two-acre (8,000-square-meter) area of Utah's Cedar Mountain Formation (Yellow Cat member). Falcarius thus lived in the Early Cretaceous period. Two extensive bone beds were discovered, including the remains of hundreds, perhaps thousands, of individuals of the new species. The minimum number of individual animals was in 2006 estimated at three hundred.[1] In 2005 over two thousand specimens had been excavated, mostly consisting of disarticulated bones.[2] These included the remains of juvenile animals.[1] In 2008 a second site was reported, the Suarez Quarry, with mainly adult individuals, but of a perhaps slightly different type. In 2010 the number of specimens from the original quarry had increased to over 2,700,[3] and was later that year reported to have risen to over three thousand.[4]

Block containing dense concentration of Falcarius bones (specimen number CEUM 8429). On display at the USU Eastern Prehistoric Museum, Price, Utah

The first remains were partially described and discussed in 2004 on several abstracts by paleontologists Lindsay E. Zanno, then a doctoral student at the University, James I. Kirkland, Scott D. Sampson, chief curator at the University of Utah's Utah Museum of Natural History, Donald D. DeBlieux, David K. Smith and R. Kent Sanders.[5][6][7] It was not formally named until a subsequent paper published in the May 2005 issue of the journal Nature.[2] Dr. Sampson was quoted as saying:

"[…] the missing link between predatory dinosaurs and the bizarre plant-eating therizinosaurs."[8]

Zanno subsequently and informally described Falcarius as "the ultimate in bizarre: a cross between an ostrich, a gorilla, and Edward Scissorhands."[8] The generic name, Falcarius, is literally taken from the Latin falcarius (meaning sickle cutter) in reference to the large hand claws. The specific name, utahensis, reflects the provenance from Utah. The holotype specimen, UMNH VP 15000, consists of a partial braincase and a large number of paratypes were assigned: specimens UMNH VP 12283, 12285, 12286, 12288, 12290, 12292, 12293, 12315, 12321-12329, 12343-12345, 12347-12349, 12360, 12363, 12366, 12369-12373, 12377, 12380, 12383-12384, 12386-12389, 12392-12394, 12396-12400, 12402-12404, 12406, 12410-12417, 12419-12425, 12427-12439, 12441-12443, 14530-14533, 14537-14538, 14542-14557, 14560-14565, 14568-14582, 14584-14656, 14658-14666, 14668-14670, 14672-14676, 14678-14680, 14682-14690, 14692-14699, 14701-14999 and 15001-15149. Apart from this extensive hypodigm, dozens of other bones were referred to the species.[2] Falcarius has since been described in detail by Zanno in a series of studies, beginning with her thesis in 2004.[9]

Description

Size comparison of an average individual

Falcarius is known from multiple specimens at different ontogenic stages. While the smallest specimen was approximately less than 1 m (3.3 ft), average adults were about 4–5 m (13–16 ft) in length weighing approximately 100 kg (220 lb).[1][3][10] Falcarius can be differentiated from other therizinosaur taxa in: on the underside of the braincase there is a well-developed depression, the recessus basisphenoidalis. At the back of the head there are deep depressions below the occipital condyle and on the subotics, each with several pneumatic grooves. On the thumb claw the raised attachment point for the tendon of the extensor muscle is bordered by deep grooves for ligaments. The large skull opening in the snout, the fenestra antorbitalis, is positioned in a depression which reaches the side of the nasal bone. There are at least five pairs of conical teeth in the front of the lower jaws. These teeth are hollowed-out at the inside. On the underside of the vertebrae of the neck there is a depression with at the mid-line a ridge. The vertebrae of the back have below their front extensions, the prezygapophyses, a groove which is divided in three smaller depressions. The middle vertebrae of the back have a second ridge extending from the base of the upper rib joint, the diapophysis, to the back of the vertebra. In the arm, the shaft of the humerus is reinforced by powerful, somewhat obliquely placed, thickenings extending towards the condyles of the lower joint, which swellings have a hollow front edge. The front edge of the hip joint has a prominent swelling, pointing sideways to the back, on the pubis.[4][3]

Skull

Dentary teeth of UMNH VP 14527 and UMNH VP 15259

The head of Falcarius is partially known. It was small and elongated. With its long neck, Falcarius could apparently reach about 1.5 m (4.9 ft) off the ground to munch leaves or fruit. The teeth numbered at least sixteen in the maxilla of the upper jaw. The dentary (lower jaw) carried twenty-eight teeth. Its small leaf-shaped and very finely serrated maxillary teeth indicate that it consumed plant material. The front-most five teeth of the lower jaw are much longer, straighter and more pointed though, and might indicate a partially omnivorous diet including meat, e.g. small animals such as lizards. In the back of the head, the braincase was relatively large. Its lower elements were moderately inflated by pneumatised, hollowed-out, bone tissue.[3][11]

Postcranial skeleton

The partitioning of the vertebral column followed that of most maniraptorans. The neck was very long with elongated cervical vertebrae. The tail was relatively long. The arm was moderately long with a somewhat robust humerus. The relatively large, and slightly recurved, pointed ten- to thirteen-centimeter (four- to five-inch) hand claws were likely used in self-defence.[12][3]

Pectoral girdle of skeletal cast

Falcarius is known from many specimens, including complete forelimb specimens. Most of the bones of the pectoral girdle and forelimb are known, although sternal bones are not preserved. Both a left and right scapula are preserved, and they are both mostly complete as well. The blades of the scapulae are about 22.5 cm (225 mm) long, and also quite slender, with barely any expansion. They are curved instead of straight. The scapula is placed in a relatively high position. Two coracoids are also preserved, although they differ more than the scapulae in morphology. The right coracoid is the better preserved of the two. The coracoid is not specially enlarged. The furcula is elongated and narrow, its branches placed at an angle of 104°. It is V-shaped and possesses a small hypocleidium at the front. The arm as a whole is not especially developed in length but shows signs of an increasing robustness. The humerus has a rather short triangular deltopectoral crest. The epicondyles are exceptionally thick, indicating a strong musculature. The ulna is lightly curved. The hand is rather long but not very robust, with in general an elongated metacarpus, although the first metacarpal is short. The first finger diverges from the second finger, the longest of the hand. The third finger is very thin. The hand claws are moderately curved, rather pointed, and moderately long.[12][3]

In the pelvis the ilium was by a moderate inclination partly adapted to a more raised position of the trunk. The pelvis was propubic or mesopubic: the pubic bone pointed more or less forwards. The leg was relatively long and adapted to running, with the lower leg longer than the thighbone. In the foot there were three weight-bearing toes. The first toe did not reach the ground nor did the first metatarsal touch the ankle.[3]

Classification

The describers in 2005 assigned Falcarius to the Therizinosauroidea sensu of Zhang, in a basal position.[2] Zanno later placed it as the basal-most taxon in the Therizinosauria, just outside a Therizinosauroidea sensu of Sereno.[4]

Because of its primitive position and relative completeness, Falcarius is a good taxon to use to compare to related groups and descendant taxa. It resembles dinosaurs belonging to the Therizinosauridae, part of the group Maniraptora. Falcarius itself probably does not belong to this former group, although it does belong to the more inclusive group Therizinosauria. The group is characterized by wider hips, a comparatively large braincase and long necks typical of plant eaters. The less basal Asian specimens were covered in quill-like feathers; this is assumed for Falcarius as well.[4] In general Falcarius can be seen as a transitional form between the original theropod build and the more derived Therizinosauridae. Falcarius shared some derived traits with the therizinosaurids: it had a long neck, small head with teeth adapted for eating plant material, a more robust arm with large hand-claws and a more vertical position of the body. On the other hand, it resembled more typical theropods in having a long tail, a propubic pelvis, a long lower leg and a foot without a fourth toe reaching the ground.[3]

The propubic pelvis of Falcarius, with the pubis pointing forwards

The cladogram below shows Hartman et al. 2019 phylogenetic analysis of Therizinosauria based on the extensive revision by Zanno in 2010. Falcarius is recovered as the primitive-most and first taxon to split off:[13]

Therizinosauria

Falcarius

Jianchangosaurus

Therizinosauroidea

Beipiaosaurus

Martharaptor

"Chilantaisaurus" zheziangensis

Enigmosaurus

Alxasaurus

Therizinosauridae

See also

References

  1. ^ a b c Zanno, L. E.; Erickson, G. M. (2006). "Ontogeny and life history of Falcarius utahensis, a primitive therizinosauroid from the Early Cretaceous of Utah". Journal of Vertebrate Paleontology. 26 (supp. 3): 143A. doi:10.1080/02724634.2006.10010069.
  2. ^ a b c d Kirkland, J. I.; Zanno, L. E.; Sampson, S. D.; Clark, J. M.; DeBlieux, D. D. (2005). "A primitive therizinosauroid dinosaur from the Early Cretaceous of Utah". Nature. 435 (7038): 84–87. Bibcode:2005Natur.435...84K. doi:10.1038/nature03468. PMID 15875020.
  3. ^ a b c d e f g h Zanno, L. E. (2010). "Osteology of Falcarius utahensis (Dinosauria: Theropoda): characterizing the anatomy of basal therizinosaurs". Zoological Journal of the Linnean Society. 158 (1): 196–230. doi:10.1111/j.1096-3642.2009.00464.x.
  4. ^ a b c d Zanno, L. E. (2010). "A taxonomic and phylogenetic re-evaluation of Therizinosauria (Dinosauria: Maniraptora)". Journal of Systematic Palaeontology. 8 (4): 503–543. doi:10.1080/14772019.2010.488045.
  5. ^ Kirkland, J. I.; Zanno, L. E.; DeBlieux, D. D.; Sampson, S. D. (2004). "A new, basal-most therizinosauroid (Theropoda: Maniraptora) from Utah demonstrates a pan-Laurasian distribution for Early Cretaceous (Barremian) therizinosauroids". Journal of Vertebrate Paleontology. 24 (supp. 3): 78A. doi:10.1080/02724634.2004.10010643.
  6. ^ Zanno, L. E. (2004). "The pectoral girdle and forelimb of a primitive therizinosauroid (Theropoda: Maniraptora): New information on the phylogenetics and evolution of therizinosaurs". Journal of Vertebrate Paleontology. 24 (supp. 3): 134A. doi:10.1080/02724634.2004.10010643.
  7. ^ Smith, D. K.; Kirkland, J. I.; Sanders, R. K.; Zanno, L. E.; DeBlieux, D. D. (2004). "A comparison of North American therizinosaur (Theropoda: Dinosauria) braincases". Journal of Vertebrate Paleontology. 24 (supp. 3): 114A. doi:10.1080/02724634.2004.10010643.
  8. ^ a b Lane, N. (2009). Life Ascending: The Ten Great Inventions of Evolution. W. W. Norton & Company. p. 230. ISBN 9780393065961. LCCN 2009005046.
  9. ^ Zanno, L. E. (2004). The Pectoral Girdle and Forelimb of a Primitive Therizinosauroid (Theropoda, Maniraptora) with Phylogenetic and Functional Implications (PhD diss.). Department of Geology and Geophysics, University of Utah.
  10. ^ Paul, G. S. (2016). The Princeton Field Guide to Dinosaurs (2nd ed.). Princeton, New Jersey: Princeton University Press. pp. 162−168. ISBN 9780691167664.
  11. ^ Smith, D. K.; Zanno, L. E.; Sanders, R. K.; Deblieux, D. D.; Kirkland, J. I. (2011). "New information on the braincase of the North American therizinosaurian (Theropoda, Maniraptora) Falcarius utahensis". Journal of Vertebrate Paleontology. 31 (2): 387−404. doi:10.1080/02724634.2011.549442. JSTOR 25835833.
  12. ^ a b Zanno, L. E. (2006). "The pectoral girdle and forelimb of the primitive therizinosauroid Falcarius Utahensis (Theropoda, Maniraptora): analyzing evolutionary trends within Therizinosauroidea". Journal of Vertebrate Paleontology. 26 (3): 636−650. doi:10.1671/0272-4634(2006)26[636:tpgafo]2.0.co;2. JSTOR 4524610.
  13. ^ Hartman, S.; Mortimer, M.; Wahl, W. R.; Lomax, D. R.; Lippincott, J.; Lovelace, D. M. (2019). "A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight". PeerJ. 7: e7247. doi:10.7717/peerj.7247. PMC 6626525. PMID 31333906.

External links

  • BBC
  • Therizinosauroidea - UCMP, Berkeley