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~-:- ~ -0 ;'HE DINOSAUR REPORT FALL 1994 Big Sauropods-Really, Really Big Sauropods By Gregory S_ Paul For the last quarter century an array of newly discovered sauropods-Supersaurus, Ultrasauros, and Seismosaurus-have competed for the title of the world's largest dinosaur. These have been big, bur not much more so than other well-known sauropods. A few sauropods truly dwarf all the others. One of them is new, but the biggest of all was discovered over a century ago, lost, and nearly forgonen. First we go to Argentina, home of enormous Late Cretaceous tiranosaurs, such as "Antarctosaurus" gigantus, which supplied a 2.3m long femur, the largest limb bone yet described in a published paper. Jose Bonaparte recently described another incomplete behemoth, Argentinosaurus buinculensis. The tibia of this colossus is I.55m long, and the estimated femur length is 2.4m. Even more impressive are the fragmentary trunk vertebrae, which' must have been 1.5m tall (over 5 feet). The length of the trunk was some 7m, about twice as long as the trunks of typical sauropods. Titanosaurs were also very "plump" sauropods-the ilia of their hips flared strongly to the sides to support an unusually broad gut. Antarctosaurus and Argentinosaurus probably weighed 80 to 100 tonnes, nearly twice as much as supersaurs and ultrasaurs. A bone found in 1878 from the Late Jurassic sediments of Garden Park in Colorado may outclass even the titanosaurs. The bone is a 1.5m tall central part of a vertebra from the trunk just in front of the hips. Unfortunately, the bone was lost, either before or during the transfer of Edward Drinker Cope's private collection to the American Museum of Natural History in New York. When Cope described and named the bone Ampbicoelias fragillimus in 1878, he commented that in "the extreme tenuity of all its parts, this vertebra exceeds those of [sauropods] already described." He calculated the complete height of the vertebra as at least six feet, and estimated that the femur should have been twelve feet long. At a Denver conference on the Morrison Formation in May of 1994, John Mclntosh suggested that the femur was over ten feet long. If anything, these are conservative estimates! Also found at Garden Park were a few bones of Amphicoelias altus. The spines of the last trunk vertebrae are unusually narrow in both sauropods, although A. fragillimus is much larger. A. altus appears to have been a lighdy built sauropod closely related and similar in size to Diplodocus. The femur of A. altus is exceptional for its length and slenderness. Using the complete vertebra of A. altus as a guide, the height of the big A. fragillimus vertebrae was at least 2.4 m, and could have been more than 2.6m (8' 6"!). A. fragillimus was 2.2 to 2.5 times larger than A. altus and Diplodocus. The implications of such dimensions are difficult to comprehend. Extrapolating the femur/ vertebra ratio of A. altus to A. fragillimus results in a femur length of around 3.8 m. Hip height should have been about 9m (30'). If the total length/vertebra height ratio was similar to that of Diplodocus. then A. fragillimus was some 50m (approaching 170') long. Body mass should have been ten times higher than that of the smaller diplodocids, about 125 tonnes. By working with the figures the size estimates can be forced down abour 10 to 20 percent, but the fact is that the size of the vertebra could suppOrt even higher estimates. "Super-sauropods" could have brought down all bur the biggest trees. The surprising slenderness of the giant Ampbicoelias vertebra and the Argmtinosaurus tibia suggests that even they were not close to the maximum size possible! Indeed, it is hardly likely that we have found the biggest sauropods, so species as heavy as 200 tonne blue whales cannot be ruled out. The skeletons alone of super-sauropods weighed about 20 to 30 or more tonnes (more than the biggest entire land mammals), so they were the most preservable and discoverable fossils ever. The scarcity of their remains therefore suggests that their populations were extremely low. This is contrary to the hypothesis that sauropods were energy- efficient reptiles with higher population densities than giant mammals. Besides, if 100 to 200 tonne sauropods had reptilian metabolisms, then their hearts would have done only abour as much work as a big elephant's heart. It is difficult to see how such weak hearts could have oxygenated the fifty tonnes of skeletal muscles needed to carry and operate 125 tonnes of flesh and bones, or pumped blood at very high pressures all the way up the long necks. Enormous heart tissues burning as much oxygen as an entire elephant were called for, raising total metabolisms to whale-like levels. Consider that growing from a few to over a hundred thousand kilograms in a few decades meant that the biggest sauropods pur on up to ten kilograms a day, a feat exceeded by only energetic whales calves fed great quantities of high nutrition-density milk. To be a giant on land probably requires an animal to be an aerobically capable athlete. This may be why terrestrial low-energy reptiles have never exceeded a tonne in mass, and endothermic mammals and dinosaurs have. The reason sauropods grew five times bigger than land mammals cannot be attributed to their continuous growth-termination of growth and life almost coincide in bull African elephants. Differing modes of reproduction are probably responsible. Over an adult lifespan of forty years, the number of eggs laid by a sauropod was fifty to five hundred times higher than the number of calves dropped by an elephant. There have to be a lot of adult elephants to raise the few highly dependent young that they produce. Only a few super-sauropods, each eating as much as an entire elephant herd, could sustain a viable population simply by breeding like rabbits. Even if all the adults were lost, just a few juveniles needed to grow up and start things over again. AJuly attempt by a team from the Denver Museum of Natural History, led by Kenneth Carpenter, to relocate the giant Ampbicoelias quarry with ground penetrating radar was not successful, but it remains possible that more super-sauropod fossils will be found at Garden Park. IhsaI' Fenu length (m) LastTI\IIk.=a TotaJlength (m) Tcmes ml Amphicoelias 3.14.0' , 22-2.6 4{}{i()' 100.150 fraa;IHmus AlyentinosauTUS & 23-2.4' 1.5 :n.35' 80-100 Anlardosaurus Brachiosaurus 2.1·2.4 0.8 ~25 :»-50 (=Uftrasauros) Swersaurus 3&45' 40-50 Seismosaurus 30-34' 25-30 AoatosaUFUS 1.7-1.8 1.35 22-23 17-20 Gamarasaurus 1.5-1.8 0.7·1.0 15-18 14-23 MamenchisauFUS 1.35 . 0.98 20.5 14 AmphicoeNas altus 1.77 1075 12 Barosaurus 1.5 1.1 2B 12 Diplodocus 1.54 1.05 24.8 11 OmeisauFUS 1.3 0.57 17.6 6.5 "Estimated

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  • ~-:-~ -0 ;'HE DINOSAUR REPORT FALL 1994

    Big Sauropods-Really,Really Big SauropodsBy Gregory S_ Paul

    For the last quarter century an array of newly discoveredsauropods-Supersaurus, Ultrasauros, and Seismosaurus-havecompeted for the title of the world's largest dinosaur. These havebeen big, bur not much more so than other well-known sauropods.A few sauropods truly dwarf all the others. One of them is new, butthe biggest of all was discovered over a century ago, lost, and nearlyforgonen.

    First we go to Argentina, home of enormous Late Cretaceoustiranosaurs, such as "Antarctosaurus" gigantus, which supplied a2.3m long femur, the largest limb bone yet described in a publishedpaper. Jose Bonaparte recently described another incompletebehemoth, Argentinosaurus buinculensis. The tibia of this colossus isI.55m long, and the estimated femur length is 2.4m. Even moreimpressive are the fragmentary trunk vertebrae, which' must havebeen 1.5m tall (over 5 feet). The length of the trunk was some 7m,about twice as long as the trunks of typical sauropods. Titanosaurswere also very "plump" sauropods-the ilia of their hips flaredstrongly to the sides to support an unusually broad gut.Antarctosaurus and Argentinosaurus probably weighed 80 to 100tonnes, nearly twice as much as supersaurs and ultrasaurs.

    A bone found in 1878 from the Late Jurassic sediments ofGarden Park in Colorado may outclass even the titanosaurs. Thebone is a 1.5m tall central part of a vertebra from the trunk just infront of the hips. Unfortunately, the bone was lost, either before orduring the transfer of Edward Drinker Cope's private collection tothe American Museum of Natural History in New York. WhenCope described and named the bone Ampbicoelias fragillimus in1878, he commented that in "the extreme tenuity of all its parts,this vertebra exceeds those of [sauropods] already described." Hecalculated the complete height of the vertebra as at least six feet,and estimated that the femur should have been twelve feet long. Ata Denver conference on the Morrison Formation in May of 1994,John Mclntosh suggested that the femur was over ten feet long. Ifanything, these are conservative estimates!

    Also found at Garden Park were a few bones of Amphicoeliasaltus. The spines of the last trunk vertebrae are unusually narrow inboth sauropods, although A. fragillimus is much larger. A. altusappears to have been a lighdy built sauropod closely related andsimilar in size to Diplodocus. The femur of A. altus is exceptional forits length and slenderness.

    Using the complete vertebra of A. altus as a guide, the height ofthe big A. fragillimus vertebrae was at least 2.4 m, and could havebeen more than 2.6m (8' 6"!). A. fragillimus was 2.2 to 2.5 timeslarger than A. altus and Diplodocus. The implications of suchdimensions are difficult to comprehend. Extrapolating the femur/vertebra ratio of A. altus to A. fragillimus results in a femur lengthof around 3.8 m. Hip height should have been about 9m (30'). Ifthe total length/vertebra height ratio was similar to that ofDiplodocus. then A. fragillimus was some 50m (approaching 170')long. Body mass should have been ten times higher than that of thesmaller diplodocids, about 125 tonnes. By working with the figuresthe size estimates can be forced down abour 10 to 20 percent, butthe fact is that the size of the vertebra could suppOrt even higherestimates.

    "Super-sauropods" could have brought down all bur the biggesttrees. The surprising slenderness of the giant Ampbicoelias vertebraand the Argmtinosaurus tibia suggests that even they were not closeto the maximum size possible! Indeed, it is hardly likely that wehave found the biggest sauropods, so species as heavy as 200 tonneblue whales cannot be ruled out. The skeletons alone ofsuper-sauropods weighed about 20 to 30 or more tonnes (morethan the biggest entire land mammals), so they were the mostpreservable and discoverable fossils ever. The scarcity of theirremains therefore suggests that their populations were extremelylow. This is contrary to the hypothesis that sauropods were energy-efficient reptiles with higher population densities than giantmammals. Besides, if 100 to 200 tonne sauropods had reptilianmetabolisms, then their hearts would have done only abour asmuch work as a big elephant's heart. It is difficult to see how suchweak hearts could have oxygenated the fifty tonnes of skeletalmuscles needed to carry and operate 125 tonnes of flesh and bones,or pumped blood at very high pressures all the way up the longnecks. Enormous heart tissues burning as much oxygen as an entireelephant were called for, raising total metabolisms to whale-likelevels. Consider that growing from a few to over a hundredthousand kilograms in a few decades meant that the biggestsauropods pur on up to ten kilograms a day, a feat exceeded by onlyenergetic whales calves fed great quantities of high nutrition-densitymilk.

    To be a giant on land probably requires an animal to be anaerobically capable athlete. This may be why terrestrial low-energyreptiles have never exceeded a tonne in mass, and endothermicmammals and dinosaurs have. The reason sauropods grew fivetimes bigger than land mammals cannot be attributed to theircontinuous growth-termination of growth and life almostcoincide in bull African elephants. Differing modes of reproductionare probably responsible. Over an adult lifespan of forty years, thenumber of eggs laid by a sauropod was fifty to five hundred timeshigher than the number of calves dropped by an elephant. Therehave to be a lot of adult elephants to raise the few highly dependentyoung that they produce. Only a few super-sauropods, each eatingas much as an entire elephant herd, could sustain a viablepopulation simply by breeding like rabbits. Even if all the adultswere lost, just a few juveniles needed to grow up and start thingsover again.

    AJuly attempt by a team from the Denver Museum of NaturalHistory, led by Kenneth Carpenter, to relocate the giantAmpbicoelias quarry with ground penetrating radar was notsuccessful, but it remains possible that more super-sauropod fossilswill be found at Garden Park.

    IhsaI' Fenu length (m) LastTI\IIk.=a TotaJlength (m) Tcmesml

    Amphicoelias 3.14.0' , 22-2.6 4{}{i()' 100.150fraa;IHmus

    AlyentinosauTUS & 23-2.4' 1.5 :n.35' 80-100AnlardosaurusBrachiosaurus 2.1·2.4 0.8 ~25 :»-50(=Uftrasauros)Swersaurus 3&45' 40-50

    Seismosaurus 30-34' 25-30AoatosaUFUS 1.7-1.8 1.35 22-23 17-20

    Gamarasaurus 1.5-1.8 0.7·1.0 15-18 14-23MamenchisauFUS 1.35 . 0.98 20.5 14

    AmphicoeNas altus 1.77 1075 12Barosaurus 1.5 1.1 2B 12Diplodocus 1.54 1.05 24.8 11OmeisauFUS 1.3 0.57 17.6 6.5

    "Estimated

  • •. "THE DINOSAUR REPORT FALL 1994 A.

    A Visual Guide toSuper-Sauropodsby Gregory S. Paul

    Indricotherium

    Argentinosaurus

    Ultrasauros1'1( ;"'+f1Y7~ hi

    Amphicoeliasfragillimus

    ==--===-- 4 m---~-- ----------------------------------------------