In defence of giraffes, they are relative latecomers in evolutionary terms: given a few tens of millions more years, it is conceivable that they might overcome some of these disadvantages to evolve longer necks. A central paradox of sauropod cervical morphology is that in the elongation of the cervical ribs, the vertebrae appear better adapted for anchoring hypaxial than epaxial musculature – even though holding the neck up was important and, due to gravity, much more difficult than drawing it down. In summary, no other clade has all of the suggested adaptations for long necks that are found in sauropods. This is within 6% of Leidy’s (1870) estimate of “almost twenty-two feet”, or about 6.7 m, and approximately equal to the 7-m neck length reported for Albertonectes by Kubo, Mitchell & Henderson (2012). A lower limit to neural spine length is imposed by the volume of muscle needed to produce the range of motion. 1560–1561). There are strong mechanical constraints on the latter: as body size increases, the eggs approach a point at which the shell cannot simultaneously be thick enough to support the egg and thin enough for the hatchling to break out of (Murray & Vickers-Rich, 2004, p. 212; Birchard & Deeming, 2009). This implies that the feeding mechanism of Diplodocus and other diplodocids was radically different from that of other sauropods. Specifically: A. M. longus colli dorsalis. Think of the word "dinosaur," and two images are likely to come to mind: a snarling Velociraptor hunting for grub, or a giant, gentle, long-necked Brachiosaurus lazily plucking the leaves off trees. The sauropods were the among largest dinosaurs and included well-known species such as the Brachiosaurus, Brontosaurus (now called Apatosaurus) and Diplodocus, which had very long necks … Dinosaurs that have long necks belong to the clade of plant-eating dinosaurs known as sauropods. ... been 2. (To be fair, though, most of this length was taken up by Diplodocus' enormously long neck and tail, not its bloated trunk.) longus colli dorsalis. 701–702). Diplodocus had an extremely long tail, composed of about 80 caudal vertebrae, which are almost double the number some of the earlier sauropods had in their tails (such as Shunosaurus with 43), and far more than contemporaneous macronarians had (such as Camarasaurus with 53). For many years, it was the longest dinosaur known. Another giant theropod, Gigantoraptor erlianensis Xu et al., 2007 belongs to another long-necked group, Oviraptorosauria. The long necks of elasmosaurid plesiosaurs were constructed very differently from those of sauropods, consisting of many very short cervicals – 76 in the neck of Albertonectes vanderveldei Kubo, Mitchell & Henderson, 2012 and 71 in Elasmosaurus platyurus Cope, 1868 (Sachs, 2005, p. 92). These trees supplanted the lush vegetation of the humid environment that preceded the volcanic event. "The only ones that survived this crisis were eusauropods. Diplodocus specie… If bifid spines conferred a great advantage, they would presumably be found throughout the neck – although the importance of stability, and the difficulty of attaining it, is greater in the posterior part of the neck, which bears greater forces than the anterior part. This is particularly clear in Fig. 10–20), in which lengths of 39, 39, 36, 29.6 and 18 cm are given for cervicals 1, 2, 4, 6 and 7, even though C2 and C7 are reported as of “size class III”. In extant lizards and crocodilians, as in basal archosaurs (Fig. As well as providing a platform for the evolution of large body size, the stability of the quadrupedal posture also enabled the evolution of longer necks: although progressive elongation displaced the centre of mass forwards from above the hindlimbs, it remained in the stable region between fore and hindlimbs. But these low spines would have reduced the lever arm with which epaxial tension members acted. It is obviously impossible for a terrestrial animal with a torso the size of a giraffe’s to carry a 10 m neck. intercristales and Mm. Because of their size, most sauropods most likely had a distinct feeding advantage over other land-dwelling dinosaurs—their extremely long necks enabled them to browse among the tops of the tallest trees. Some of the largest animals to ever walk the land belong to the sauropods, and well-known sauropods include Diplodocus and Brachiosaurus. thank you in advance for your patience and understanding. One of the best-known sauropods, Diplodocus was a very large long-necked quadrupedal animal, with a long, whip-like tail. Interpretation of sauropods as living animals is made especially difficult by the lack of good extant analogues. The skull of Diplodocus … Ayuntamiento de Cuenca, Instituto Juan de Valdés, Field Columbian Museum, Geological Series, Quarterly Journal of the Geological Society, London. The known material of the diplodocid Supersaurus Jensen, 1985 includes a cervical vertebra whose centrum is 138 cm long. 7.1 and 7.2), so that the neck of Apatosaurus must have been triangular in cross-section. In sauropods with bifid spines, then, the one- or two-segment Mm. For example, based on their relative heights, epipophyses probably dominated neural spines in Apatosaurus (Fig. This chapter asks how dinosaurs used long necks and tails. In some sauropods, the cervical neural spines are bifid (i.e., having separate left and right metapophyses and a trough between them). The single known cervical vertebra is 118 cm in total length, including overhanging prezygapophyses, and its incomplete centrum can be reconstructed after related titanosaurs as having been 105 cm long. Diplodocus is among the most easily identifiable dinosaurs, with its typical sauropod shape, long neck and tail, and four sturdy legs. At least some researchers have proposed that sauropod necks were relatively immobile, horizontal and beam-like … Dinosaur fossil from 98 million-years-ago unearthed in Argentina may have been the largest animal to ever walk the Earth, study claims The fossil remains — including tail and pelvic bones — were uncovered in 2012 The giant titanosaur sauropod dinosaur would have had a long neck and tail It was preserved in a muddy floodplain in what […] Neck statistics of some sauropods, chosen because of unusually long, short or complete necks. Here we survey the longest necked taxa in several groups of extant and extinct animals (Figs. by David Button 1. Wednesday November 18 2020, 12.01am, The Times. intercristales. 11). Second, although bone is much stiffer than tendon, it is actually not as strong in tension, so that an ossified tendon is more likely to break under load. Sauropods shared a body plan consisting of: a small head 5 and 6) indicate that in at least one specimen the vertebral column is complete. The large hypaxial muscles (M. flexor colli lateralis, M. flexor colli medialis, and M. longus colli ventralis) insert on the cervical ribs (Fig. Giraffatitan, and a comparison of the world’s largest dinosaurs. In extant animals, the mechanically significant soft tissues of the neck (muscles, tendons and ligaments) can be examined and their osteological correlates identified. In sauropods with bifid neural spines, such as Camarasaurus (Fig. Their huge size was likely a … In this respect, sauropod osteology is intermediate between the conditions of crocodilians and birds – so the widely recognized similarity of sauropod cervicals to those of birds (e.g., Wedel & Sanders, 2002; Tsuihiji, 2004), while significant, should not be accepted unreservedly. Our lack of knowledge was because sediments with dinosaurs and plants of that particular age are very uncommon.". The typical length of the neck of the ostrich is only 1.0 m (sum of vertebral lengths in Dzemski & Christian (2007), Table 1, plus 8% to allow for intervertebral cartilage – see Cobley, 2011, p. 16). B. M. cervicalis ascendens. Introduction. However, this raises as many questions as it answers since pumping blood to a height of 30 or 40 feet would strain even the biggest, most robust heart. Wh y sauropods had long necks; ... it seems likely that sauropods shared a suite of featur es that facilitated the. and will receive updates in the daily or weekly email digests if turned on. Within Theropoda, at least three lineages evolved especially long necks. One of the best-known sauropods, Diplodocus was a large long-necked four-legged animal, with a long, whip-like tail. Can that have been done by reducing the amount of muscle? Consider an ostrich neck scaled up by a linear factor of L. The weight exerted by the neck is proportional to L3 but the cross-sectional area of the bracing members is proportional to only L2. However, ligament cannot have filled the trough as envisaged by Alexander (1985, figure 4C), because pneumatic foramina are often found in the base of the troughs of presacral vertebrae, for example in the cervicals of Apatosaurus (Fig. E. M. longus colli ventralis. Puertasaurus Novas et al., 2005 is the largest titanosaur for which cervical material has been described. If the necks of sauropods were as heavily muscled as those of ostriches, then they would have appeared in cross section as shown in Fig. intercristales. However, this explanation cannot be correct, as bifid spines are not known in taxa along the line to birds – only in sauropods and a few modern birds. 1, 1, The second through seventeenth cervical vertebrae of, Blue arrows indicate epaxial muscles attaching to neural spines, red arrows indicate epaxial muscles attaching to epipophyses, and green arrows indicate hypaxial muscles attaching to cervical ribs. It is significant that all other clades of large (>10 ton) terrestrial herbivores – ceratopsians, hadrosaurs, proboscideans, and indricotheres – practiced extensive oral processing of their food, requiring massive dentition and correspondingly large heads. Heavy quadrupedal animals don’t need long tails for balance, nor do they need long necks for balancing long tails (don’t take this the wrong way, but you come across as almost implying that sauropods just randomly evolved big tails for no apparent reason). In many long-necked animals, the legs are of a similar length and so the neck elongation can be explained as a simple consequence of the need to reach down to ground level – for example in order to drink. The largest animals to ever walk the Earth were sauropods -- long-necked dinosaurs that could grow the length of three school buses. It is worth noting that the available material of Malawisaurus and Isisaurus pertains to relatively small individuals; perhaps the forces exerted by the epaxial muscles were not enough to produce distinctive scarring of the bone that we would recognize as epipophyses. In some specimens, a ligament scar and pneumatic foramen occur together in the intermetapophyseal trough (Fig. Not all muscles leave diagnostic traces on the skeleton, so the absence of epipophyses does not mean that the epaxial muscles that insert above the postzygapophyses were absent. 7.5), although the spines are higher in posterior cervicals. Unpublished MSc Thesis, University College London, London, UK, Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, Journal of Experimental Zoology (Mol Dev Evol), Physeter macrocephalus. Diplodocus is the longest dinosaur known from a complete skeleton, measuring over 35 meters long (of which the neck represents 6 meters) and 10 to 16 tonnes. Although pneumaticity was undoubtedly an important adaptation for increasing the length of the neck without greatly increasing its mass, a longer neck remains more mechanically demanding than a shorter neck of the same mass, because that mass acts further from the fulcrum of the cervicodorsal joint, increasing the moment that must be counteracted by the epaxial tension members. It was nearly complete when found, but has since been damaged and is now missing its central portion, but plaster replicas made before the damage indicate the extent of the missing portion. Aside from the factors previously discussed, the elongation of sauropod necks was made possible by the distinctive architecture of their cervical vertebrae. Figure 10 shows the cervical skeleton of Euhelopus as it actually is, and reconstructed with speculative muscle attachments that would have been more mechanically efficient: why did sauropod necks not evolve this way? In some sauropods, including Erketu and Mamenchisaurus, which were proportionally long-necked even by sauropod standards, the neural spines are strikingly low, and the epipophyses no higher – a surprising arrangement, as low spines would have reduced the lever arm with which the epaxial tension members worked. C. M. flexor colli lateralis. Most researchers, for instance, do agree that Apatosaurus and Diplodocus likely kept their necks low while grazing. Stiff cervical ribs would have helped provide lateral stabilization for the neck, which would have been especially important in taxa with epaxial tension members concentrated on the midline (i.e., those with non-bifid spines) as discussed above. 6.1 and 6.4). It is particularly notable that mamenchisaurids (Mamenchisaurus and Omeisaurus) have very low neural spines, as does Erketu in the preserved, anterior, cervicals. Brachiosaurus is one of the most famous Sauropods. Towards the middle ground of these extremes fall the cervical vertebrae of Giraffatitan, which are anteroposteriorly longer and dorsoventrally shorter than those of Apatosaurus, but not as anteroposteriorly long or as dorsoventrally short as those of Erketu. In most sauropods, the cervical vertebrae do have epipophyses, but the neural spines are as prominent or more so (Fig. In birds, ossification (or at least mineralization) of tendon has many functional effects: it (1) restricts tendon deformation; (2) reduces tendon strain at a given stress; (3) accommodates higher load bearing (to a point; see below); and (4) reduces damage to the tendon (Landis & Silver, 2002, p. 1153). The cervical architecture is rather different in crocodilians, and in non-archosaurian diapsids such as lizards, snakes, ichthyosaurs and plesiosaurs: there are no epipophyses, and the main epaxial neck muscles are the Mm. no more than one email per day or week based on your preferences. Finally we thank J Hutchinson (Royal Veterinary College) for his constructive and efficient editorial handling of the manuscript. Measured from the skeletal reconstruction of Xu et al. In many ways, the sauropods (of which Brachiosaurus was a prominent example) are more fascinating than bipedal predators like T. Rex or Velociraptor. (2007, figure 1A), it appears to have had a neck 2.15 m in length – although this is conjectural as almost no cervical material is known. Ouvrage enrichi de figures en taille-douce, The role of phylogenetic analysis in the inference of unpreserved attributes of extinct taxa, A new giant pterosaur with a robust skull from the latest Cretaceous of Romania, The first giant dinosaurs: a large sauropod from the Late Triassic of Thailand, Winning by a neck: tall giraffes avoid competing with shorter browsers, Biggest of the big: a critical re-evaluation of the mega-sauropod, First complete sauropod dinosaur skull from the Cretaceous of the Americas and the evolution of sauropod dentition, Two new oviraptorids (Theropoda: Oviraptorosauria), Upper Cretaceous Djadokhta Formation, Ukhaa Tolgod, Mongolia, Proceedings of the Academy of Natural Sciences of Philadelphia, On a gigantic saurian from the Dakota epoch of Colorado, Geology and paleontology: a new species of. This disparity is particularly evident in the cervical vertebrae (Fig. longus colli dorsalis would have had the dual function of support and lateral motion. 7.1) but neural spines may have dominated in Isisaurus Wilson & Upchurch, 2003 and Giraffatitan (Figs. They weren’t all enormous, but the big ones were extraordinary. But in some respects they seem locked into a mammalian pattern that will always prevent them from matching the necks of sauropods: extensive oral processing of food requires a large head with heavy teeth; almost no mammal has evolved more than seven cervical vertebrae; and the mammalian lung has attained a local maximum of efficiency that makes it unlikely ever to evolve into something analogous to the avian flow-through lung, so both an air-sac system and vertebral pneumaticity are precluded. Vertebrae modified from. †Diplodocus longus (nomen dubium) Marsh, 1878. The basal eusauropod Mamenchisaurus Young, 1954 is known from several species. Azdarchids are variously reported as having seven to nine cervical vertebrae, but never more; non-avian theropods do not seem to have exceeded the 13 or perhaps 14 cervicals of Neimongosaurus Zhang et al., 2001, with eleven or fewer being more typical. Unilateral branch stripping is the most likely feeding behavior of Diplodocus, as it explains the unusual wear patterns of the teeth (coming from tooth–food contact). As noted above, the epipophyses are the insertion points of the largest and longest epaxial muscles in birds, whereas in crocodilians the epipophyses are non-existent, and no major muscles insert above the postzygapophyses (Tsuihiji, 2004). On average, then, total C1–C7 neck length in known Zhejiangopterus specimens was about 3.85 times that of C5. We then examine the osteology of sauropod necks more closely, comparing their cervical anatomy with that of their nearest extant relatives, the birds and crocodilians, and discussing unusual features of sauropods’ cervical vertebrae. Bearing this in mind, the total neck length of Arambourgiania may have somewhat exceeded 3.0 m. In azhdarchids, C8 may be between 20% and 50% the length of C5 (Pereda-Suberbiola et al., 2003, p. 86), which might amount to 16–39 cm in Arambourgiania. Multiplication of cervical vertebrae obviously contributes to neck elongation. Brachiosaurus had a proportionally long neck, small skull, and large overall size, all of which are typical for sauropods. 7.3); but their presence in other titanosaurs such as Rapetosaurus Curry & Forster, 2001  (Curry & Forster, 2001, figure 3A), and Saltasaurus Bonaparte & Powell, 1980 (Powell, 1992, figure 5) and in outgroups such as Giraffatitan (Fig. Were it not for the end-Cretaceous extinction, non-avian theropods would have been the most likely candidates for evolving sauropod-like neck lengths, due to the combination of pneumaticity, small heads in some clades. In birds, the largest and mechanically most important epaxial muscles (M. longus colli dorsalis and M. cervicalis ascendens) insert on the epipophyses of the cervical vertebrae – that is, distinct dorsally projecting tubercles above the postzygapophyses. Sheer size is probably a necessary, but not sufficient, condition for evolving an absolutely long neck. Therefore, the whole-element densities of their postcranial bones will always be between 1.0 and 2.0; they cannot be more dense than bone tissue, nor can they be constructed entirely out of marrow. The sperm whale’s nose: sexual selection on a grand scale? The azhdarchid for which the most complete neck has been described is Zhejiangopterus linhaiensis Cai & Wei, 1994, so we will base our estimates on this species. 11.2), the muscles of the neural spine were presumably significant, and would have acted primarily along the midline of the neck. Sure, there were sauropods that fed on tree-tops: Brachiosaurus, for example, had wear on its teeth that matched well with a tree-munching lifestyle. For this reason, it would be reasonable to expect animals to evolve the shortest possible trachea. The long-necked theropods may not have been under the same selection pressure to evolve long necks as were sauropods. In some mature males, the trachea coils back on itself so many times that its total length exceeds 800 mm, nearly three times the total body length of approx. For many years, it was the longest dinosaur known. The air space proportion (ASP) of a bone is the proportion of its volume taken up by pneumatic cavities (Wedel, 2005). It is now well established that sauropods had an air-sac system similar to that of extant birds (Wedel, 2003), and most likely a similarly efficient flow-through lung (Wedel, 2009). Since its skull was much more robust that those of other azhdarchids (Buffetaut, Grigorescu & Csiki, 2002, p. 183), it was probably carried on a proportionally shorter and stronger neck. Their body design did not vary as much as in other dinosaur groups – they all had large bodies and small heads, and most had very long necks and tails, like this Brachiosaurus.All the sauropods were herbivorous, maintaining their massive bodies with only plant matter. Diplodocus is among the most easily identifiable dinosaurs, with its typical sauropod shape, long neck and tail, and four sturdy legs. If this is correct, then its neck was subject to a quite different biomechanical regime than those of sauropods. The reduction in head weight would have reduced the required lifting power of the necks that carried them, and therefore the muscle and ligament mass could be reduced, allowing the necks to be longer than would have been possible with heavier heads. While it could reach up to eighty feet in length, most of that length was accomplished via its long neck and whip-like tail. Its front limbs were a bit shorter than its hind limbs, which forms a horizontal stance for the most part. We thank those who provided useful feedback and helpful contributions – in particular H Mallison (Museum für Naturkunde Berlin) and the anonymous referee who also reviewed this version. However, the small absolute size of birds means that the forces acting on their necks are so different that we can’t assume that sauropod necks functioned in the same ways – just as the problems involved in flight through air for high-Reynolds number fliers such as birds are very different than than they are for low-Reynolds number fliers such as fruit-flies, whose aerodynamics are dominated by friction drag rather than form drag. If the trachea is narrow, then it is difficult to inhale sufficient air quickly – a problem exacerbated by friction of inhaled air against the tracheal wall. In sauropods with unsplit neural spines, such as Giraffatitan (Fig. 3). Even among diplodocids, it had extraordinary cervical ribs: very short, very robust, and positioned very low, far below the centra on extremely long parapophyses (Figs. The necks of sauropod dinosaurs greatly exceeded in length those of all other animals (Wedel, 2006a). In the 1990s, a nearly complete skeleton of a Camarasaurus was discovered at Howe Quarry in Wyoming, close to a 1934 dig site where an expedition from the American Museum of Natural History discovered about 4,000 dinosaur fossils. This suggests that the necks of super-giant sauropods may have been even longer than imagined: Carpenter (2006, p. 133) estimated the neck length of the apocryphal giant Amphicoelias fragillimus Cope, 1878 as 16.75 m, 2.21 times the length of 7.5 m used for Diplodocus, but if Parrish’s allometric curve pertained then the true value would have been 2.211.35 = 2.92 times as long as the neck of Diplodocus, or 21.9 m; and the longest single vertebra would have been 187 cm long. Stress is force/area, which is proportional to L3/L2 = L, so the stress on the bracing members that support the neck varies linearly with L. (The weight of the neck acts at a distance proportional to L from the torso, and the bracing members acts at a distance proportional to L above the neck-torso articulation, so these factors cancel out of the balancing moment equation.) Diplodocusmay have had thin spines lining its back, and many modern reconstructions s… intertransversarii. Alone among extant animals, birds are able to cope with such extreme tracheal elongation, due to their very efficient lungs and the large tidal volume of the whole respiratory system on account of the voluminous air-sacs. 1 and 2; and “kei” and Kevin Ryder for permission to use their photographs. It is possible that neck length was positively allometric in these clades, as in sauropods, and they may have had necks somewhat longer than isometric scaling suggests. Hutchinson ( Royal Veterinary College ) for his constructive and efficient editorial of. Accomplished via its long neck, small skull, and erect limbs reminiscent of the largest of all.. 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Speed of a giraffe ’ s nose: sexual selection on a grand scale skulls brains! To torso length of their cervical vertebrae do have epipophyses, shown in green proportionally small heads sauropods like diplodocus most likely had long necks to sauropodomorphs. … dinosaurs that could grow the length of the largest long-necked theropods lived in 2–2.5! Neck elongation, the small airspace ventral to the number of cervical vertebrae they evolve! Necks as long as those of birds suggests that sauropods spent most of the.! With a suspension bridge as living animals is made especially difficult by lack... Are available are those of sauropods modifications of the largest of all sauropods were much short! The multisegment Mm that sauropods spent most of their respective C5s this implies that feeding... Of a turtle foramina in the intermetapophyseal trough ( Fig Clench, 1978 ) ASP calculations probably dominated spines. 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Their bodies and 7.2 ), from which Fig of C5 thank you in advance for your patience and.. Carotici, which forms a horizontal stance for the large cervical muscles have varied a lot over decades!