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| ''Dreadnoughtus schrani'' || [[Titanosaur]]ia || 59,291<ref name="Lacovara et al., 2014" /> || 65.4
| ''Dreadnoughtus schrani'' || [[Titanosaur]]ia || 59,291<ref name="Lacovara et al., 2014" /> || 65.4
|-
|-
| ''[[Brachiosaurus|Brachiosaurus altithorax]]'' || [[Brachiosauridae]] || 56,000<ref name="Bensonetal2014">{{cite journal | last1 = Benson | first1 = RBJ | last2 = Campione | first2 = NE | last3 = Carrano | first3 = MT | last4 = Mannion | first4 = PD | last5 = Sullivan | first5 = C | display-authors = 5 | last6 = et al | year = 2014 | title = Rates of Dinosaur Body Mass Evolution Indicate 170 Million Years of Sustained Ecological Innovation on the Avian Stem Lineage | url = | journal = PLoS Biol | volume = 12 | issue = 5| page = e1001853 | doi = 10.1371/journal.pbio.1001853 }}</ref> || 62
| ''[[Brachiosaurus|Brachiosaurus altithorax]]'' || [[Brachiosauridae]] || 56,255<ref name="Bensonetal2014">{{cite journal | last1 = Benson | first1 = RBJ | last2 = Campione | first2 = NE | last3 = Carrano | first3 = MT | last4 = Mannion | first4 = PD | last5 = Sullivan | first5 = C | display-authors = 5 | last6 = et al | year = 2014 | title = Rates of Dinosaur Body Mass Evolution Indicate 170 Million Years of Sustained Ecological Innovation on the Avian Stem Lineage | url = | journal = PLoS Biol | volume = 12 | issue = 5| page = e1001853 | doi = 10.1371/journal.pbio.1001853 }}</ref> || 62
|-
|-
| ''[[Turiasaurus|Turiasaurus riodevensis]]'' || [[Turiasauria]] || 51,000<ref name="Bensonetal2014"/> || 56.1
| ''[[Turiasaurus|Turiasaurus riodevensis]]'' || [[Turiasauria]] || 50,923<ref name="Bensonetal2014"/> || 56.1
|-
|-
| ''[[Elaltitan|Elaltitan lilloi]]'' || [[Titanosaur]]ia || 43,000<ref name="Bensonetal2014"/> || 47.2
| ''[[Elaltitan|Elaltitan lilloi]]'' || [[Titanosaur]]ia || 42,798<ref name="Bensonetal2014"/> || 47.2
|-
|-
| ''[[Futalognkosaurus|Futalognkosaurus dukei]]'' || [[Titanosaur]]ia || 38,000<ref name="Bensonetal2014"/> || 42
| ''[[Futalognkosaurus|Futalognkosaurus dukei]]'' || [[Titanosaur]]ia || 38,139<ref name="Bensonetal2014"/> || 42
|-
|-
| ''[[Giraffatitan|Giraffatitan brancai]]'' || [[Brachiosauridae]] || 34,000<ref name="Bensonetal2014"/> || 37.5
| ''[[Giraffatitan|Giraffatitan brancai]]'' || [[Brachiosauridae]] || 34,003<ref name="Bensonetal2014"/> || 37.5
|-
|-
| ''[[Diplodocus|Diplodocus longus]]'' || [[Diplodocid]]ae || 15,000<ref name="Bensonetal2014"/> || 16.3
| ''[[Diplodocus|Diplodocus longus]]'' || [[Diplodocid]]ae || 14,813<ref name="Bensonetal2014"/> || 16.3
|}
|}


Revision as of 16:16, 26 January 2015

Dreadnoughtus
Temporal range: Late Cretaceous, 75 Ma
↓
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: †Sauropodomorpha
Clade: †Sauropoda
Clade: †Macronaria
Clade: †Titanosauria
Clade: †Lithostrotia
Genus: †Dreadnoughtus
Lacovara et al., 2014
Type species
†Dreadnoughtus schrani
Lacovara et al., 2014 (type)

Dreadnoughtus is a genus of giant titanosaurian sauropod dinosaur from the Upper Cretaceous, discovered in the (Campanian-Maastrichtian; 84–66 Ma) Cerro Fortaleza Formation of Santa Cruz province, Argentina. It is one of the largest of all known terrestrial vertebrates, possessing the greatest mass of any land animal that can be calculated with reasonable certainty,[1] using limb bone measurements.[2][3] In terms of skeletal completeness and ability to encode its anatomy into cladistic analyses, Dreadnoughtus schrani is the most complete gigantic titanosaurian sauropod dinosaur.[1]

Etymology

"Dreadnought" is an English term for the predominant type of battleship in the early 20th century, deriving from the words "dread" (fear) + "nought" (nothing). According to the research team that discovered the taxon, the genus name Dreadnoughtus "alludes to the gigantic body size of the taxon (which presumably rendered healthy adult individuals nearly impervious to attack)" and to the two dreadnought battleships of the Argentine Navy, the ARA Rivadavia and ARA Moreno. Thus, the genus name also honors the country in which Dreadnoughtus schrani was discovered. The name of the type species, schrani, was given in recognition of the American entrepreneur Adam Schran for his financial support of the project.[1]

Description

The discovery of Dreadnoughtus schrani provides insight into the size and anatomy of giant titanosaurian dinosaurs, especially of the limbs and the shoulder and hip girdles. The majority of Dreadnoughtus schrani bones are very well preserved. There is minimal deformation, especially in the limb bones. Fine features, such as locations of muscle attachment, are frequently clearly visible.

Holotype and paratype specimens

The holotype specimen, MPM-PV 1156, consists of a partial skeleton, somewhat preserved in its original layout, that comprises: a maxilla (jaw) fragment; a tooth; a posterior cervical vertebra; cervical ribs; multiple dorsal vertebrae and dorsal ribs; the sacrum; 32 caudal vertebrae and 18 haemal arches (bones from the tail) that include a sequence of 17 anterior and middle caudal vertebrae and their corresponding haemal arches found in their original layout; the left pectoral girdle and forelimb minus the front foot; both sternal plates; all pelvic elements; the left hind limb lacking a hind foot and right tibia; metatarsals I and II; and one claw from digit I.

The paratype, MPM-PV 3546, consists of a partially articulated postcranial skeleton of a slightly smaller individual whose remains were discovered in the same location as the holotype. It includes a partial anterior cervical vertebra, multiple dorsal vertebrae and ribs, the sacrum, seven caudal vertebrae and five haemal arches, a nearly complete pelvis, and the left femur.[1]

Size

Dreadnoughtus Dimensions[1]
Dimension Metric Imperial
Mass 59,300 kilograms (59.3 t) 65.4 short tons (130,800 lb)
Total length 26 m 85 ft
Head and neck length 12.2 m 40 ft
Neck-only length 11.3 m 37 ft
Torso and hip length 5.1 m 17 ft
Tail length 8.7 m 29 ft
Shoulder height ~ 2 stories (6 m) ~ 2 stories (20 ft)

At 1.74 m, the scapula of the holotype is longer than any other known Titanosaur scapula.[1] Its ilium, the top bone of the pelvis, is also larger than any previously reported, measuring 1.31 m in anterior to posterior length.[1] The forearm of the holotype is longer than any previously reported for a titanosaur; only measuring less than the elongate forearms of brachiosaurid sauropods, which exhibited a more inclined body posture.[1] Only Paralititan[4] preserves a longer humerus (upper arm bone). Although each species likely had slightly different body proportions, these measurements clearly demonstrate the massive nature of Dreadnoughtus schrani.[1]

Mass

File:Dreadnoughtus Size & Weight Comparision (Metric) JPG.jpg
Dreadnoughtus schrani was substantially more massive than any other supermassive dinosaur for which mass can be accurately calculated.

Using Equation 1 of Campione and Evans (2012),[3] which allows the inference of body mass of a quadrupedal animal solely from the minimum circumference of the shaft of its humerus and femur, the Dreadnoughtus type specimen weighed approximately 59.3 metric tons (65.4 short tons), or 59,291 kilograms.[1] A different, albeit unofficial, estimate by Matt Wedel used volumetric models that yield between 35 and 40 metric tons (39 to 44 short tons),[5] or even as low as approximately 30 metric tons (33 short tons), based on a 20% shorter torso.[6] By comparison, the animal's weight was more than eight and a half times that of a male African elephant and even exceeded the Boeing 737-900 airliner by several tons when taking the estimate from the description.[7] Linear bone measurements indicate the holotype individual was approximately 26 metres (85 ft)* long and stood about 2 stories tall.[1] Its neck is estimated to have been 11.3 m long, its torso 4 m long, and its tail 8.7 m long.

Sauropod Taxonomic Group Mass (kg) Mass (short tons)
Dreadnoughtus schrani Titanosauria 59,291[1] 65.4
Brachiosaurus altithorax Brachiosauridae 56,255[8] 62
Turiasaurus riodevensis Turiasauria 50,923[8] 56.1
Elaltitan lilloi Titanosauria 42,798[8] 47.2
Futalognkosaurus dukei Titanosauria 38,139[8] 42
Giraffatitan brancai Brachiosauridae 34,003[8] 37.5
Diplodocus longus Diplodocidae 14,813[8] 16.3

Completeness

Completeness may be assessed in different ways. Sauropod dinosaur skeletons are often recovered with little to no skull material, so completeness is often looked at in terms of postcranial completeness (i.e., the completeness of the skeleton excluding the skull). Completeness may also be assessed in terms of the numbers of bones versus the types of bones. The most important metric for understanding the anatomy of a fossil animal is the types of bones. The completeness statistics for Dreadnoughtus schrani are as follows:

  • 116 bones out of ~256 in the entire skeleton (including the skull) = 45.3% complete
  • 115 bones out of ~196 in the skeleton (excluding the skull) = 58.7% complete
  • 100 types of bones out of ~142 types in the skeleton (excluding the skull) = 70.4% complete

The completeness of Dreadnoughtus schrani compared with other extremely massive (over 40 metric tons) sauropods is as follows:[2]

Sauropod Skeletal Completeness Total Mirrored Postcranial Completeness
(i.e. types of bones)
Dreadnoughtus schrani 45.5% 70.4%
Turiasaurus riodevensis 44.1% 45.8%
Futalognkosaurus dukei 15.2% 26.8%
Paralititan stromeri 7.8% 12.7%
Argentinosaurus huinculensis 5.1% 9.2%
Antarctosaurus giganteus 2.3% 3.5%
Puertasaurus reuili 1.6% 2.8%

Thus, the skeleton of Dreadnoughtus schrani is substantially more complete than those of all other extremely massive (>40 metric tons) dinosaurs.[1]

Posture

All titanosaurs had what is called wide-gauge posture, a relative term to describe a stance in which the feet fell apart from the body midline. More derived titanosaurs had a greater degree of wide-gauge posture,[9][10] with their limbs held more widely than their ancestors and contemporaneous counterparts. The stance of Dreadnoughtus schrani was clearly wide-gauge, but not to the degree of saltasaurids because the femoral condyles are perpendicular to its shaft rather than beveled.[1] This and the fact that the head of the femur was not turned in towards the body as in saltasaurids[9] support the phylogenetic conclusion that Dreadnoughtus was not a saltasaurid. The animal’s broad sternal bones also demonstrate a wide pectoral girdle, giving it a broad-shouldered, broad-chested appearance. Paleontologist Kenneth Lacovara compared the animal's gait to an Imperial Walker's.[11]

Although the forelimbs of Dreadnoughtus schrani are longer than in any other previously known titanosaur, they are not significantly longer than the hind limbs.[1] Therefore, Lacovara et al. (2014) reconstructed its neck to have been held more horizontally, rather than anteriorly inclined in the manner of Brachiosaurus.[12]

Distinctive features

File:Dreadnoughtus tail composite image.jpg
Composite image of the tail bones of the Dreadnoughtus schrani type specimen. This individual’s tail would have been 8.7 metres (29 feet) in length.

The tail of Dreadnoughtus schrani has several characteristic features included in the diagnosis of the species. The first vertebra of the tail has a ridge on its ventral surface called a keel. In the first third of the tail, the bases of the neural spines are extensively subdivided into cavities caused by contact with air sacs (part of the dinosaur’s respiratory system). In addition, the anterior and posterior boundaries of these neural spines have distinct ridges (pre- and postspinal laminae) connecting them to the pre- and postzygapophyses (the articulation points of the neural arches). In the middle of its tail, the vertebrae have a triangular process that extends over the centrum towards each preceding vertebra.[1]

Just like modern archosaurs with tails (for example crocodilians,[13] Dreadnoughtus schrani had bones below the vertebrae called chevrons or haemel arches. These bones connect with the ventral surface of the vertebrae and are “Y” shaped when viewed anteriorly. In Dreadnoughtus schrani the bottom stem of the “Y” is broadly expanded, likely for the attachment of muscles.[1]

The shoulder girdle and forelimb of Dreadnoughtus schrani also exhibit unique features. An oblique ridge crosses the interior face of the scapular blade, extending from the top side near the far end of the blade to the bottom side near the base of the scapular blade. Finally, each end of the radius exhibits a unique form: the top, or proximal end, has a distinct concave embayment on its posterior face while the bottom, or distal end, is nearly square in shape instead of broadly expanded.[1]

Discovery and study

File:Kenneth Lacovara at dig site of Dreadnoughtus.jpg
Kenneth Lacovara surrounded by the skeleton of Dreadnoughtus schrani at the dig site in Argentina

Dr. Kenneth Lacovara, of Drexel University, discovered the remains in the Cerro Fortaleza Formation in Santa Cruz Province, Patagonia, Argentina in 2005. Due to the large size of the bones and the remote location where they were found, it took his team four austral summers to fully excavate the remains. Mules, ropes, hills, and many team members were needed to finally get the field-jacketed bones to a truck.

File:Dreadnoughtus schrani fossil skeleton at Lacovara lab Drexel University.JPG
Skeleton of Dreadnoughtus schrani in Lacovara’s fossil lab at Drexel University. Tail section (on left); Scapula (shoulder blade) (standing against divider); Cervical (neck) vertebra (center); Tibia (shinbone) and Humerus (upper arm bones) (right).

In 2009, the fossils were transported to Philadelphia via an ocean freighter for preparation and study. Fossil preparation and analysis occurred at Drexel University, the Academy of Natural Sciences of Drexel University and the Carnegie Museum of Natural History. Dreadnoughtus schrani fossils will be returned to their permanent repository at the Museo Padre Molina in Rio Gallegos, Argentina.

3D imagery

File:3D-imaging-Dreadnoughtus-femur-Lacovara-Drexel.jpg
Lacovara’s team prepared 3D laser scans of most of the fossil bones from Dreadnoughtus schrani, making them available to other researchers – or to anyone – for future study. Here, Lacovara works with doctoral candidate Anna Jaworski scanning the Dreadnoughtus femur.

The bones of both Dreadnoughtus specimens were scanned with a NextEngine 3D laser scanner.[1] Using the software Autodesk Maya, the scans of each bone were positioned in 3D space to create a digital articulated skeleton, which was then converted into 3D PDF files using the software GeoMagic. The high fidelity of these scans allowed Lacovara et al. (2014) to study the heavy fossils of Dreadnoughtus schrani in a way that was safe for the fossils and enhanced virtual and long distance collaboration.

File:Dreadnoughtus-3d-tail.jpg
Screen shot of digital reconstruction of the first 32 caudal vertebrae and associated haemal arches of the holotype of Dreadnoughtus schrani (MPM-PV 1156) in left lateral view. Downloadable, interactive 3D PDF file available at http://dx.doi.org/10.6084/m9.figshare.1119785

Phylogeny

Macronaria

Based on a cladistic analysis, Dreadnoughtus schrani appears to be a “derived” basal titanosaur that is not quite a lithostrotian.[1] Lacovara et al. (2014) note that because of the wide array of relatively “advanced” and “primitive” features in the skeleton of Dreadnoughtus schrani and the current instability of titanosaurian interrelationships, future analyses may find widely differing positions for it within Titanosauria.

Taphonomy

Based on the sedimentary deposits at the site, the two Dreadnoughtus schrani specimens appear to have been buried quickly during a fluvial avulsion event, or break in a levee resulting in a flood. This event generated a liquifacted crevasse splay deposit which entombed the two dinosaurs. Thus, rapid and relatively deep burial of the Dreadnoughtus type specimen accounts for its extraordinary completeness. Numerous small theropod teeth found amongst the bones are likely evidence of scavenging, most likely by megaraptorans,[1] perhaps Orkoraptor.

Ontogeny

The holotype specimen was likely not fully grown at the time of its death. The histology of the holotype humerus, which shows a lack of an external fundamental system (an outer layer of bone found only in fully-grown vertebrates) and abundance of fast-deposited or still-growing woven tissue in the primary fibrolamellar bone of the outer bone cortex, lead Lacovara et al. (2014) to determine that the specimen was still growing at the time of its death.[1][14] It remains unknown how large this individual would have grown had it not died when it did.

References

  1. ^ a b c d e f g h i j k l m n o p q r s t u v Lacovara, Kenneth J.; Ibiricu, L.M.; Lamanna, M.C.; Poole, J.C.; Schroeter, E.R.; Ullmann, P.V.; Voegele, K.K.; Boles, Z.M.; Egerton, V.M.; Harris, J.D.; Martínez, R.D.; Novas, F.E. (September 4, 2014). "A Gigantic, Exceptionally Complete Titanosaurian Sauropod Dinosaur from Southern Patagonia, Argentina". Scientific Reports. 4: 6196. doi:10.1038/srep06196. PMID 25186586.
  2. ^ a b Benson, Roger B. J.; Campione, Nicolás E.; Carrano, Matthew T.; Mannion, Phillip D.; Sullivan, Corwin; Upchurch, Paul; Evans, David C. (May 6, 2014). "Rates of Dinosaur Body Mass Evolution Indicate 170 Million Years of Sustained Ecological Innovation on the Avian Stem Lineage". PLOS Biology. doi:10.1371/journal.pbio.1001853.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  3. ^ a b Campione, Nicolás E.; Evans, David C. (July 10, 2012). "A universal scaling relationship between body mass and proximal limb bone dimensions in quadrupedal terrestrial tetrapods". BMC Biology: 15. doi:10.1186/1741-7007-10-60.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  4. ^ Smith, Joshua B.; Lamanna, Matthew C.; Lacovara, Kenneth J.; Dodson, Peter; Smith, Jennifer R.; Poole, Jason C.; Giegengack, Robert; Attia, Yousry (2001). "A Giant sauropod dinosaur from an Upper Cretaceous mangrove deposit in Egypt". Science. 292 (5522): 1704–1706. doi:10.1126/science.1060561. PMID 11387472. Retrieved 2014-08-31.
  5. ^ Matt Wedel (11 September 2014). "How massive was Dreadnoughtus?". svpow. Retrieved 11 September 2014.
  6. ^ Matt Wedel (15 September 2014). "How long was the torso of Dreadnoughtus?". svpow. Retrieved 15 September 2014.
  7. ^ Chang, Kenneth (4 September 2014). "Argentine Dinosaur Was an Estimated 130,000 Pounds, and Still Growing". The New York Times. Retrieved 5 September 2014.
  8. ^ a b c d e f Benson, RBJ; Campione, NE; Carrano, MT; Mannion, PD; Sullivan, C; et al. (2014). "Rates of Dinosaur Body Mass Evolution Indicate 170 Million Years of Sustained Ecological Innovation on the Avian Stem Lineage". PLoS Biol. 12 (5): e1001853. doi:10.1371/journal.pbio.1001853. {{cite journal}}: Explicit use of et al. in: |last6= (help)CS1 maint: unflagged free DOI (link)
  9. ^ a b Wilson, Jeffrey A.; Carrano, Matthew T. (June 1999). "Titanosaurs and the origin of "wide-gauge" trackways: a biomechanical and systematic perspective on sauropod locomotion". Paleobiology. 25 (2): 252–267. doi:10.1666/0094-8373-25.2.252. Retrieved 31 August 2014.
  10. ^ Wilson, J. A. (February 2006). "An Overview of Titanosaur Evolution and Phylogeny". III Jornadas Internacionales sobre Paleontología de Dinosaurios y su Entorno: 169–190.
  11. ^ "Supermassive Dinosaur Would Have 'Feared Nothing'" (mp3). Science Friday. Public Radio International. Sep 5, 2014. Retrieved Sep 6, 2014.
  12. ^ Christian; Dzemski (2011). "Neck posture in sauropods". Biology of the sauropod dinosaurs: understanding the life of giants: 251–260.
  13. ^ Wilhite, Ray (2003). "Biomechanical Reconstruction of the Appendicular Skeleton in Three North American Jurassic Sauropods". Louisiana State University Electronic Thesis & Dissertation Collection. Retrieved 31 August 2014.
  14. ^ Schroeter, Elena; Boles, Zachary; Lacovara, Kenneth (November 2011). "The Histology of a Massive Titanosaur from Argentina and Implications for Maximum Size" (PDF). Journal of Vertebrate Paleontology (Program and Abstracts Supplement): 189.
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