Friday, April 5, 2019
Diet and Digestion as Factors for Large Size in Sauropods
Diet and Digestion as Factors for Large sizing in SauropodsDinosaurs have captured the imagination since the  truly first fossils were discovered. The mystical creatures in many ancient cultures  may be attri neverthelessed to fossils weathering out of the earth. Nearly every culture had  nearly form of dragon in their mythology, a uniformity best explained by fossilized dinosaurs. Some scientists  conceptualize the fossils of ceratopsians  atomic number 18 the  reference of the mythological griffin.  bust of the  spell with dinosaurs is their size. With few animals reaching such(prenominal)  capacious proportions, the creatures that produced single bones as tall as a human   mustinessiness have seemed other worldly to ancient peoples. They still draw fascination today their features so different from anything  lifespan. As to a greater extent and more fossils  ar  prepare, explanations and extrapolations of the features, size, and behavior become more comprehensive. The un cognize e   lement of dinosaur life allows for wild projections and suppositions, and the drama of discovery and scientific discourse and hypothesis disproval  bear upon to draw the interest of people. As more is discovered, more questions are raised. One field of questions pertain to   sauropod dinosaur dinosaur size. Sauropods are distinct in their absolutely massive bodies. Far  braggyr than anything known by  livelihood humans, they are fascinating to consider. Trying to imagine their size is in  almost ways  bid  laborious to imagine the vastness of space. Without actually standing next to a mounted skeleton, there is no living reference to fix the size of sauropods in the imagination. Some estimates place the largest sauropods at 80,000kg, or 170,000 lbs. Conservative estimates  bring up weights of 40,000-50,000 kg. Sauropod height is in some specimens 60 feet, about 3  times the height of a two-story house. Some sauropods were a third of a football field in duration Such sizes are nearly    unfathomable, raising the question of how they got to be so massive and how their bodies  mattered. Modern herbivores are often  untold bigger than carnivores, because in general, their energy exp overthrowiture is less and the biomass of their  victuals source is  high. The huge size of sauropods is  emphatically tied to their  aliment, and it is  in all prob faculty that their   diet source and digestive mechanisms were a  study  chemical element in their  growing to such massive sizes.The food available to sauropods was  non very nutritive. Before the evolution and diversification of angiosperms  one hundred twenty-five million years ago,  saprophagous animals had to rely mainly on gymnosperms and ferns. The best measure of  whole shebang  nutritionary value is in the concentration of nitrogen and nonstructural carbohydrates like sugars. Protein levels largely correspond to the nitrogen levels in leaves. Because non-angiosperm  positions dont have the xylem transport network tha   t evolved, there is less nutrient flow, and therefore less nitrogen concentration. Ferns, cycads, and conifers, some of the main plant types available to the sauropod dinosaurs, were low in nutrition, and gymnosperms had secondary chemical defenses that made them  rough to eat and harder to digest (Midgley, 2005). A study by Zvereva and Kozlov (2006)  institute that the nitrogen concentration in gymnosperms drops in environments with elevated temperature and CO2 concentration, compared to the  new environment. Sugar levels tended to stay the same. The toughness of leaves increases when CO2 levels are higher(prenominal). The carbon-nitrogen ratio is  signifi brush offtly increased by elevated CO2. It is well-known that the environmental conditions during the age of dinosaurs was both higher in temperature and higher in CO2 concentration. This would have produced plant material signifi toilettly less nutritious than the plant  count today. The lesser food  prize supports the evolution    of large sauropod dinosaurs, as the Jarman-Bell Principle states that larger species can feed on diets of lesser quality food,  attestd by the relatively large size of modern font grazing mammals (Clauss, et. al., 2009). The physiology of the dinosaurs shows  march of partitioning. Some sauropods, such as diplodocids, were low browsers, who were un apparent to life their heads higher than a couple meters (Stevens and Parrish, 1999). This separated their food source from the mid to high browsers, limiting their available food source in some ways, yet also preventing competition amongst sauropod species. The lower plant quality requires that herbivorous animals  deliver more material in order to meet their nutritional and energy needs.Given the quality of available plants, extended digestion would have been  necessitate. Of several methods to  answer plant material, sauropods digestion was extremely  unrest heavy. They had very  unretentive oral processing. Their teeth were either br   oad and leaf like, with serrations in the crown, or later more  specialise and peg like. In most sauropods, teeth were present in the front part of the mouth but  non the back. The  eubstance plan in the early Mesozoic seems to have been to maximize food  use through teeth adapted for cropping, stripping, and pulling plant material but with  lesser oral processing.  in that location is some evidence in the tooth replacement rate and wear that teeth farther back in the mouths of sauropods were not used in oral processing or food acquisition, but rather served the  design of cheeks, to keep the food in the mouth (Schwarz, et. al., 2015). Prosauropods had some check development, but in the sauropod line, cheeks were  befuddled early (Chure, et. al., 2010). There have been no teeth found in association with sauropods that had a large horizontal surface, indicative of use for chewing (Christiansen, 1999). The heads of sauropods were extremely small compared to their body size. Any dental    batteries or cheeks would have increased the head mass and likely made it prohibitively heavy,  particularly at the end of a long neck. Christiansen also noted that the muzzle  width in sauropod dinosaurs is  proportionally  replete(p)r than the width in herbivorous animals. This enables sauropods to intake more food per bite,  change magnitude their intake rate. With relatively poor food nutrient quality, sauropods would need to consume a great amount of food to meet their daily needs. The length of the neck adds length to the digestive envelope, another way to maximize food consumption. The  overlook of oral processing, wide muzzle, and long neck work in conjunction to increase the consumption capability or sauropods, increasing their ability to meet their dietary needs and shortening their feeding time.With such high consumption and such little processing, digestion would be almost entirely focused in the gut. The particle size would be very large, making digesta harder to break    down, and the longer it takes to digest. There has been some debate about the presence of gastroliths and a  stomachic mill in sauropod dinosaurs. Gastric mills certainly would help break down the plant material into  small particle sizes and speed the digestion process. Unfortunately, the fossil record seems to rule out the presence of a gastric mill for most sauropod dinosaurs. Wings and Sander (2007) tested the hypothesis of sauropod gastroliths by testing gastrolith usage in farm ostriches. They found that rose quartz, which has the same properties as the white  nervure quartz found in association with sauropods, last the longest. Using granite cubes, they found that the general shape of the  rock n roll stays the same. They also found that the gastric milling process quickly roughens the face of the stone. None of the stones that he used in his test retained a shine, unlike those suspected to be sauropod gastroliths. He also found that gastroliths are about 1% of the total bod   y mass.  agree to them, the largest mass of gastroliths found in association with a sauropod is 15kg,  very much lower than 1% of the estimated sauropod body mass. Gastroliths likely scale with body mass simply because organ size roughly scales with body mass. The stomachs of sauropods could have been proportionally smaller than expected, although it is unlikely with the amount of food that they are estimated to have consumed. Using projections from living herbivores, it is estimated that the large sauropods would consume several hundred kilograms worth of food (Englemann, et. al., 2004). This estimate accounts for the proportional decrease in required food consumption for successively larger and larger organisms. In another study focusing on the  dispersion of gastrolith stones amongst sauropod remains, it was found that gastroliths were found with about 4% of sauropod fossils. There are some species of sauropod that have strong evidence of gastric mills, but they are not widesprea   d enough (Wings, 2014). With little to no processing of plant material, and with no evidence of any other break down processes, sauropods must have heavily relied on fermentation to digest their food. The relatively low density of nutrients in the gymnosperms at the time already required longer fermentation times. In order to achieve longer fermentation time, the gut size needs to increase, or the amount of food eaten needs to decrease. As the food particles must have been large, the time taken for fermentation would need to be even longer. With no internal  cottony tissues  preserve in the fossil record, it is hard to determine where fermentation took place in the gut. Fermentation demands a large digestive tract. Larger intestines allow for longer digesta retention, pulling more nutrients from the food. The large torsos of sauropods could certainly  change course and extensive gut. Estimates based on living animals gives a digesta retention time  among 8 and 16 days. Galapagos tur   tles, which do not chew their food, retain theirs for 11 days (Franz, et. al., 2009). Sauropod dinosaurs could have evolved to be so large because those individuals with larger guts had better survival chances than those who had smaller guts. As stated previously, the amount of food needed in relation to the body mass would likely have been much lower than other, smaller herbivorous organisms. As sauropods heads were evolved to take in the maximum amount of food, the intake of food would likely not have consumed most of the sauropods time. This time advantage would have been especially needed in semiarid environments, like the Jurassic area preserved in the Morrison formation. Although there is lush  phytology in the Morrison fossil record, it is not widespread instead found in clusters. The Morrison Formation suggests that the environment during the time of sauropods was savannah-like. The dense vegetation found was likely due to seasonal worker rains, or centered around areas of w   ater, such as lakes or streams. The vegetation likely moved, growing in different areas depending on rainfall (Englemann, et. al., 2004). Large herbivores would need to follow the seasonal changes, and  immigrate in search of new food sources. The size of the sauropods and the advantages of that size would have been a  study edge in a semi-arid environment. Such sizes would have also made locomotion more efficient.  semipermanent strides afforded by the overall large body size decreases the amount of energy per unit of distance. This, too, increases the ability of sauropods to migrate in search of food. Sauropod reliance on fermentation was so great that their methane production has been  cogitate to the warm climate of the Mesozoic era (Wilkinson, et. al., 2012). Assuming a more reptilian metabolism, one  penning puts the  global biomass of large sauropods at 200,000 kg/km2. Their total estimated annual methane emission is about 520 million tons. For comparison, modern day ruminant   s produce about 50-100 million tons of atmospheric methane, and the total modern day global emissions are about 500-600 million tons a year. The author notes that their estimate could have been overstated by a factor of two, but also understated by the same amount, depending on assumed metabolous function and density of sauropods. Sauropods large sizes and reliance on fermentation to digest food was a major influence on their environment, and created a positive feedback loop, where the temperature increase would push nutritive values of plants lower. Massive sauropod bodies were well adapted to their environment and digestion.Of course, such large sizes bring their own challenges. The most debated question is whether or not dinosaurs were endothermic or ectothermic, and whether endothermy was even possible in mega dinosaurs like the sauropods. At body  lower classes estimated between 20,000 kg and 80,000 kg, overheating would have been a huge problem in large sauropod dinosaurs. Whi   le it is unknown if they had special soft tissue adaptations in order to combat their size, it is speculated that their long necks and  track may have helped them keep cool by increasing surface area without adding too much internal volume (Eagle, et. al., 2011). The accelerated  maturement capable in endothermic animals is a major factor in favor of sauropod endothermy. Sauropods grew several size magnitudes, from hatchlings estimated to be about 10kg, to the hulking adult dinosaurs, in only a few decades (Sander  Clauss, 2008). But does the size of adult sauropods rule out endothermic metabolisms? According to Eagle and others (2011), endothermy was not impossible in large sauropods. In lower temperatures, more 13C-18O clumps form, which are preserved in the fossil record. The analysis of these clumps is not dependent on knowing the oxygen isotope composition of the  meet water. These clumps can be observed in the tooth bioapatite of dinosaurs. The accuracy of this method is 1C, w   ith precision within 1-2C. The accuracy of this method comes from tests of modern taxa, in which the isotopic temperature agrees with the expected temperature of the organisms. Eagle found the average body temperature of Brachiosaurus to be 38.2C  1C and the temperature of Camarasaurus to be 35.7C 1.3C including a sample from a different fossil site. These temperatures are within the range of modern mammals, and lower than the body temperatures of many birds, which can be greater than 40C. Eagle does note that the temperature reflects the temperature of tooth formation, which may differ from the main body temperature. dead body temperature is a product of metabolism, size, environmental temperature, and any special adaptations for the regulation of heat. The temperatures given for these sauropods is close to the temperatures estimated by  preferably research done by Gillooly (2006). Such temperatures in such large animals suggests that they were either ectothermic, had low  primary(   a) metabolic rates, or had some special methods of heat dissipation. Some researchers suggest that large sauropods were fermentative endotherms (Mackie, 2002). Because sauropods needed a lot of energy to reach their adult size in such little time, it is likely that they were endotherms who underwent some sort of change at maturation that prevented overheating as an adult. Metabolic changes through development is not unusual, so it is very possible that it also occurred in sauropods. They could have shifted from a higher metabolic endothermy during their  rapid growth phase, to a lesser metabolic homeothermy supported by the fermentation heat output from their  in full formed guts. Other evidence in support of endothermic sauropods are growth lines, or the lack there of, in sauropod bones (Kohler, et. al., 2012). Lines of arrested growth are normally associated with ectotherms, which have periods of rapid growth interspersed with periods of slow growth. These lines are found in mamma   ls as well, and in the majority of dinosaurs. The pattern the lines leave are not found in sauropod bones. This suggests unbroken, stead growth rates, highly unlikely in ectotherms. The evidence suggests endothermy in sauropods, even in large ones. Because endothermy requires more energy to maintain, sauropods would have had to consume a massive amount of food, unless they had a low basal metabolism. Hippopotami, while mammals and clearly not the size of sauropods, have  particularly long retention times because they have low food intake and enormous gut capacity. Their required energy for maintenance is remarkably low. This strategy is common in non-ruminant foregut fermentators and some small hindgut fermentators (Clauss, et. al., 2009). The metabolic process of sauropods is linked to their energetic needs and dietary restrictions. In some research, one of the byproducts of fermentation, heat, supports the endothermic theory, and endothermy in sauropods as juveniles helps explain    how they managed to reach their massive adult size. The herbivorous, fermentative nature of sauropods is not an obstacle in understanding their ability to function at such large sizes.Sauropods likely evolved to be large because of the abundance of plant materials, especially after many herbivores died out during both the Permian extinction, but also the Triassic-Jurassic extinction. Their adaptations allowed them to widely diversify and fill the  fresh opened ecological  recesss. Some have speculated that their large body size was driven by predation, as larger bodies, especially the size of sauropods, were a natural defense (Sander, et. al., 2011). Given the evidence, it is more likely that  opposition to predation was a lucky side effect of size, not the driving factor. It seems more likely that the  imaginativeness opportunities of plants drove the initial adaptation, especially with the diverse nature of sauropods and their apparent partitioning, than protection.The size of sau   ropods is inextricably linked to their diet and digestive methods. Much of the discussion of sauropod feeding is based on conjecture is based on living animals, that are obviously very different from sauropods. As there is are no known records of internal tissues, it is hard to know anything about how sauropods functioned internally beyond comparing them to existing behaviors and traits in todays animals.  counterbalance though todays herbivores are different than sauropods, patterns of herbivory are similar in very different taxa. The circumstantial evidence offered by analysis of modern organisms still enables scientists to attempt to fit sauropods within the known herbivorous patterns, with allowances for the unknowns.  maybe this analysis is just another element of the imaginative aspect of dinosaur life. Until more evidence is found, I believe that the evolution of the massive sauropods was in large part due to their diet and digestion. The nutrition offered by gymnosperms dema   nded higher levels of processing. The minimal oral digestion evidenced by small heads and non-chewing teeth led to greater digesta retention times in the gut. A greater gut size would have sped fermentation, compensating for the large particle size of the plant material and its low nutrient density. There is some evidence for resource partition amongst sauropods, both between different species and within the same species. Different tooth structure and browsing levels are some adaptations driven by available resources. The other benefits of the large body size of sauropods, in my opinion, do not seem likely to be as important in size evolution as the plants and their digestion. When the sauropods appeared, they filled an ecological niche left by previous extinctions, and quickly diversified, creating a hugely successful group of organisms, both in overall diversity, but also in longevity.Works CitedChristiansen, Per, 1999, On the Head Size of Sauropodomorph Dinosaurs Implications for    Ecology and Physiology. historic Biology, v. 13, iss. 4, p. 269-297.Chure, D., Britt, B. B., Whitlock, J. A., Wilson, J. A., 2010, First complete sauropod dinosaur skull from the Cretaceous of the Americas and the evolution of sauropod dentition. Naturwissenschaften, v. 97, iss. 4, p. 379-391.Eagle, R. A., Ttken, T., Martin, T. S., Tripati., A. K., Fricke, H. C., Connely, M., Cifelli, R. L., Eiler, J. M., 2011, Dinosaur Body Temperatures Determined from isotopic (13C- 18O) Ordering in  fogey Biominerals. Science, v. 333, n. 6041, p. 443-445.Englemann, G. F., Chure, D. J., Fiorillo, A. R., 2004, The implications of a dry climate for the paleoecology of the fauna of the Upper Jurassic Morrison Formation. Sedimentary Geology, v. 167, iss. 3-4, p. 397-308.Franz, R., Hummel, J., Kienzle, E., Klle, P., Gunga, H., Clauss, M., 2009, Allometry of visceral organs in living amniotes and its implications for sauropod dinosaurs. Proceedings of the Royal  ships company B, v. 276, iss. 1662, p. 1   731-1736.Gillooly, J. F., Allen, A. P., Charnov, E. L., 2006, Dinosaur Fossils Predict Body Temperatures. PLoS Biology, v. 4, iss. 8, p. 1467.Kohler, M., Marn-Moratalla, N., Jordana, X., Aanes, R., 2012, Seasonal bone growth and physiology in endotherms shed light on dinosaur physiology. Nature, v. 487, iss. 7407, p. 358-361.Mackie, Roderick I., 2002, Mutualistic Fermentative Digestion in the Gastrointestinal Tract Diversity and Evolution. Integrative and Comparative Biology, v. 42, n. 2, p. 319-326.Midgley, J. J., 2005, Why Dont Leaf-Eating Animals Prevent the Formation of  plant? Relative vs Absolute Dietary Requirements. The New Phytologist, v. 168, n. 2, p. 271- 273.Sander, P. Martin  Martin Clauss, 2008, Sauropod Gigantism. Science, v. 322, n. 5899, p. 200- 201.Sander, P. M., Christian, A., Clauss, M., Fechner, R., Gee, C. T., Griebeler, E., Gunga, H., Hummel, J., Mallison, H., Perry, S. F., Preuschoft, H., Rauhut, O. W. M., Remes, K., Ttken, T., Wings, O., Witzel, U., 2011, Bi   ology of the sauropod dinosaurs the evolution of gigantism.  biologic Reviews, v. 86, p. 117-155.Schwarz, D., Kosch, J. C. D., Fritsch G., Hildebrandt, 2015, Dentition and Tooth Replacement of Dicraeosaurus hansemanni (Dinosauria, Sauropoda, Diplodocoidea) from the Tendaguru Formation of Tanzania. Journal of Vertebrate Paleontology, v. 36.Stevens, Kent A.  J. Michael Parrish, 1999, Neck Posture and Feeding Habits of Two Jurassic Sauropod Dinosaurs. Science, v. 284, n. 5415, p. 798-800.Wilkinson, D. M., Nisbet, E. G., Ruxton, G. D., 2012, Could methane produced by sauropod dinosaurs have helped drive Mesozoic climate warmth?. Current Biology, v. 22, iss. 9, p. R292-R293.Wings, O., 2015, The  rarity of gastroliths in sauropod dinosaurs  a case study in the Late Jurassic Morrison Formation, western USA. Fossil Record, v. 18, iss. 1, p. 1-16.Wings, Oliver  P. Martin Sander, 2007, No gastric mill in sauropod dinosaurs new evidence from analysis of gastrolith mass and function in ostriche   s. Proceedings of the Royal Society B, v. 274, iss. 1610, p. 635-640.Zvereva, E. L.  M. V. Kozlov, 2006, Consequences of simultaneous elevation of carbon dioxide and temperature for plant-herbivore interactions a metaanalysis. Global  throw Biology, v. 12, iss. 1, p. 27-41.  
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