Dinosaurs Without Bones

by Anthony J. Martin

  p. 228 “For example, hippopotamuses do this by rapidly flicking their tails back and forth as they excrete, flinging poo with wild abandon.” Kranz, K.R. 1982. A note on the structure of tail hairs from a pygmy hippopotamus (Choeropsis liberiensis). Zoo Biology, 1: 237-241.

  p. 228 “Their digestion is also more complete than that of most mammals … birds excrete uric acid out of a single orifice, the cloaca.” Elbroch and Marks (2001).

  p. 228 “Nonetheless, raptors and other carnivorous birds are famous for their white-liquid sprays… .” Elbroch and Marks (2001).

  p. 228 “In a paper published in 2003, two researchers, Victor Meyer-Rochow and Jozsef Gal, became intrigued with the radial patterns of feces created by two penguin species… .” Meyer-Rochow, V.B., and Gal, J. 2003. Pressures produced when penguins pooh—calculations on avian defaecation. Polar Biology, 27: 56-58.

  p. 229 “In our species, vomiting mainly evicts toxic substances … but it can also be a reaction to severe physical or psychological trauma… .” Pelchat, M.L., and Rozin, P. 1982. The special role of nausea in the acquisition of food dislikes by humans. Appetite, 3: 341-351.

  p. 229 “Probably the most well known of such deposits are cough pellets, also known as gastric pellets.” Myhrvold, N.P. 2012. A call to search for fossilized gastric pellets. Historical Biology, 24: 505-517.

  p. 230 “Tracks associated with these cough pellets normally show the coughers with their feet together (side by side) and a pellet in front of the tracks.” Martin (2013).

  p. 231 “Crocodilians have extremely thorough digestion… .” Fisher, D.C., 1981. Crocodilian scatology, microvertebrate concentrations, and enamelless teeth. Paleobiology, 7: 262-275.

  p. 231 “So even though male ratites and some other birds, such as ducks and geese, have penises … these lack an enclosed urethra.” Brennan, P.L.R., and Prum, R.O. 2012. The erection mechanism of the ratite penis. Journal of Zoology, 286: 140-144.

  p. 232 “For humans, the standard descriptive method for feces is the Bristol scale, used in clinical situations.” Riegler, G., and Esposito, I. 2001. Bristol scale stool form: a still valid help in medical practice and clinical research. Techniques in Coloproctology, 5: 163-164.

  p. 232 “In between these two extremes, most of the weight of a stool (about 75%) comes from water, and of the ~25% solid stuff, almost a third of this can be from dead bacteria… .” Stephen, A.M., and Cummings, J.H. 1979. The microbial contribution to human faecal mass. Journal of Medical Microbiology, 13: 45-56.

  p. 232 “This tyranny started with seed plants, which developed in the Devonian Period (about 400 million years ago), then escalated into outright slavery… .” Gensel, P.G., and Edwards, D. 2003. Plants Invade the Land: Evolutionary and Environmental Persepctives. Columbia University Press, New York: 304 p.

  p. 233 “Most broadly, any former food item associated with the digestive tract of a dinosaur (or any other fossil animal, for that matter) is called a bromalite.” These definitions have been made by so many researchers and used in so many different ways, I hereby throw up (my hands, that is) and give up attempts to untangle the literature on who defined what and when. Instead, read this: Chin, K. 2012. What did dinosaurs eat: coprolites and other direct evidence of dinosaur diets. In Brett-Surman, M.K., Holtz, T.R., Jr., and Farlow, J.O. (editors), The Complete Dinosaur (2nd Edition). Indiana University Press, Bloomington, Indiana: 589-601.

  p. 234 “Coelophysis was a slender, greyhound-sized theropod and is one of the most abundantly represented dinosaurs in the fossil record… .” Holtz (2012).

  p. 234 “The ‘cannibalism’ hypothesis was originally interpreted and promoted by noted paleontologist Ned Colbert, who had excavated and studied Coelophysis since the late 1940s.” Nesbitt, S.J., Turner, A.H., Erickson, G.M., and Norell, M.A. 2006. Prey choice and cannibalistic behaviour in the theropod Coelophysis. Biology Letters, 2: 611-614.

  p. 234 “In 2002, paleontologist Robert Gay took a closer look at these Coelophysis specimens and found that the ‘juvenile Coelophysis’ bones were misidentified.” Gay, Robert J. 2002. The myth of cannibalism in Coelophysis bauri. Journal of Vertebrate Paleontology 22(3): 57A.

  p. 234 “Another study done by Sterling Nesbitt and others in 2006 confirmed that it was Hesperosuchus in the belly of the beast, not another Coelophysis.” Nesbitt et al. (2006).

  p. 234 “In yet another study in 2010, Gay pointed out that supposed ‘stomach contents’ in one Coelophysis made up a larger volume than its original stomach, so this was not an enterolite, either.” Gay, R. 2010. Evidence related to the cannibalism hypothesis in Coelophysis bauri from Ghost Ranch, New Mexico. In Notes on Early Mesozoic Theropods. Lulu Press: 9-24.

  p. 234 “Interestingly, this deposit, because of its position and jumble of bones, was interpreted as a regurgitalite.” Rinehart, L.F., Lucas, S.G., Heckert, A.B., Spielmann, J.A., and Celesky, M.D. 2009. The paleobiology of Coelophysis bauri (Cope) from the Upper Triassic (Apachean) Whitaker quarry, New Mexico, with detailed analysis of a single quarry block. New Mexico Museum of Natural History & Science, Bulletin 45: 260 p.

  p. 235 “For instance, in 2011, Jingmai O’Connor and two other paleontologists reported a specimen of Microraptor … with bird bones between its ribs.” O’Connor, J., Zhou, Z., and Xu, X. 2011. Additional specimen of Microraptor provides unique evidence of dinosaurs preying on birds. Proceedings of the National Academy of Sciences, 108: 19662-19665.

  p. 235 “A couple of Early Cretaceous theropods from China, Sinosauropteryx and Sinocalliopteryx, also have intriguing enterolites.” (1) Dong, Z., and Chen, P. 2000. A tiny fossil lizard in the stomach content of the feathered dinosaur Sinosauropteryx from northeastern China. Vert PalAsiatica, 38 (supplement): 10. (2) Xing, L., Bell, P.R., Persons, W.S., Ji, S., Miyashita, T., Burns, M.E., Ji, Q., and Currie, P.J. 2012. Abdominal contents from two large Early Cretaceous compsognathids (Dinosauria: Theropoda) demonstrate feeding on Confuciusornithids and Dromaeosaurids. PLoS One, 7: e44012. doi:10.1371/journal. pone.0044012.

  p. 235 “Another Sinosauropteryx contained jaws from three different small mammals… .” Hurum, J.H., Luo, Z.-X., and Kielan-Jaworowska, Z. 2006. Were mammals originally venomous? Acta Palaeontologica Polonica, 51: 1-11.

  p. 236 “Fortunately for Mesozoic mammals, at least one struck a blow against its dinosaurian oppressors… .” Hu, Y.M., Meng, J., and Wang, Y.Q. 2005. Large Mesozoic mammals fed on young dinosaurs. Nature, 433: 149-152.

  p. 236 “Based on other stomach contents, another feathered theropod, the Early Cretaceous Sinocalliopteryx of China, also ate birds and nonavian dinosaurs.” Xing et al. (2012).

  p. 236 “Ably representing North American theropods and their meals, David Varricchio—introduced in a previous chapter—discovered a Late Cretaceous tyrannosaurid (Daspletosaurus) in Montana with a few anomalous bones in its abdominal region.” Varricchio, D.J. 2001. Gut contents from a Cretaceous tyrannosaurid: implications for theropod dinosaur digestive tracts. Journal of Paleontology, 75: 401-406.

  p. 237 “The first specimen of this dinosaur was discovered in a clay pit mine in 1983; paleontologists Alan Charig and Angela Milner named it several years later, in 1986.” Charig, A.J., and Milner, A.C. 1986. Baryonyx, a remarkable new theropod dinosaur. Nature, 324: 359-361.

  p. 237 “In a more detailed study of this specimen published in 1997, Charig and Milner concluded that all of these traits made Baryonyx well suited for … eating fish.” Charig, A.J., and Milner, A.C. 1997. Baryonyx walkeri, a fish-eating dinosaur from the Wealden of Surrey. Bulletin of the Natural History Museum of London, Geology Series 53: 11-70.

  p. 238 “(As learned previously, paleontologists were further encouraged to adopt this formerly strange idea of fish-eating theropods when they found thousands of theropod swim tracks in Early Jurassic rocks of Utah.)” Milner et al. (2006).

  p. 238 “This same specimen of Baryonyx included lots of other ichnological extras, such as the remains of a juvenile ornithopod (identified as Iguanodon), gastrol
iths, and its own broken bones.” Charig and Milner (1997).

  p. 238 “Oddly enough, they mention ‘gastroliths’ in their 1986 paper, but only ‘an apparent gastrolith’ in the 1997 one.” Charig and Milner (1986, 1997).

  p. 238 “Only a few such trace fossils are known in dinosaurs, but one of the best is also in the most complete dinosaur from an entire continent, Minmi paravertebrata.” Molnar, R.E., and Clifford, H.T. 2001. An ankylosaurian cololite from the Lower Cretaceous of Queensland, Australia. In Carpenter, K. (editor), The Armored Dinosaurs. Indiana University Press, Bloomington, Indiana: 399-412.

  p. 239 “It turns out this specimen of Minmi was deposited in a Cretaceous sea, and no river was anywhere near its body when it was laid to rest in a shallow marine grave.” Molnar and Clifford (2001).

  p. 240 “One regurgitalite inferred from Late Triassic rocks contains pterosaur remains, but is thought to have come from a large fish rather than a land-dwelling animal.” Dalla Vecchia, F.M., Muscio, G., and Wild, R. 1989. Pterosaur remains in a gastric pellet from Upper Triassic (Norian) of Rio Seazza Valley (Udine, Italy). Gortania-Atti Museo Friul. St. Nat., 10: 121-132.

  p. 240 “I also mentioned one example interpreted in 2009, which was apparently fossilized beside the mouth of a Coelophysis.” Rinehart et al. (2009).

  p. 241 “Only one instance of fossilized dinosaur vomit, containing a mix of dinosaur and turtle bones in Early Cretaceous rocks of Mongolia, is inferred to have come from an actual dinosaur.” I mentioned this interpreted regurgitalite in my textbook (Martin, 2006), but unfortunately have not been able to hunt down the original reference, which I recall was in a Russian journal.

  p. 241 “Another possible dinosaurian up-chuck is embodied in a mass of four birds from the Early Cretaceous of Spain.” Sanz, J.L., Chiappe, L.M., Fernandez-Jalvo, Y., Ortega, F., Sanches-Chillon, B., Poyato-Ariza, F.J., and Perez-Moreno, B.P. 2001. An Early Cretaceous pellet. Nature, 409: 998–999.

  p. 242 “These deposits, found in Late Jurassic rocks of the U.K., consist of concentrated collections of belemnite shells.” Walker, S.E., and Brett, C.E. 2002. Post-Paleozoic patterns in marine predation: was there a Mesozoic and Cenozoic marine predatory revolution? In Kowalewski, M., and Kelley, P.H. (editors), The Fossil Record of Predation. Paleontological Society Special Publication, 8: 119-193.

  p. 243 “This urolite was mentioned in a poster presentation at a paleontology meeting in 2002 and understandably garnered much media attention… .” McCarville, K., and Bishop, G.A. 2002. To pee or not to pee: evidence for liquid urination in sauropod dinosaurs. Journal of Vertebrate Paleontology, 22 (Supplement to 3): 85.

  p. 244 “Known as the Purgatoire site (after the Purgatoire River, which runs through the area), it is one of the most spectacular dinosaur tracksites in the western U.S…” Lockley (1991).

  p. 245 “Given the controversy over the Morrison urolite, I was much relieved to later find out that paleontologists Marcelo Fernandes, Luciana Fernandes, and Paulo Souto had also interpreted dinosaur urolites… .” Fernandes, M.A., Fernandes, L.B., and Souto, P.B. 2004. Occurrence of urolites related to dinosaurs in the Lower Cretaceous of the Botucatu Formation, ParanaÅL Basin, Sao Paulo State, Brazil. Revista Brasileira de Paleontologia, 7: 263-268.

  p. 246 “Instead, they took two liters of water and poured it from 80 cm (2.6 ft) above and onto a loose sand surface with a slope having the same angle (about 30Åã) as the fossil ones.” Fernandes et al. (2004).

  p. 248 “Cincinnati’s a great city with a lot going for it, but it has the wrong age rocks (Ordovician Period, 450 million years old) and wrong rocks… .” Davis, R.A., and Meyer, D.L. 2009. A Sea Without Fish: Life in the Ordovician Sea of the Cincinnati Region. Indiana University Press, Bloomington, Indiana: 368 p.

  p. 249 “This means carnivorous dinosaur scat had a better chance of preserving than that coming from insectivores or herbivores… .” Chin, K. 2002. Analyses of coprolites produced by carnivorous vertebrates. Paleontological Society Papers, 8: 43-50.

  p. 249 “Second, anaerobic bacteria in the feces could have assisted in preserving it, in which their metabolic processes caused chemical reactions that made more minerals precipitate, and do so rapidly (geologically speaking).” Hollocher, T.C., Chin, K., Hollocher, K.T., and Kruge, M.A. 2001. Bacterial residues in coprolite of herbivorous dinosaurs: role of bacteria in mineralization of feces. Palaios, 16: 547-565.

  p. 249 “And third, rapid burial, such as from a nearby river flood, would have prevented fresh droppings from getting eaten, poked, prodded, sniffed, trampled, washed away, or otherwise damaged.” Thulborn, R.A., 1991. Morphology, preservation and palaeobiological significance of dinosaur coprolites. Palaeogeography, Palaeoclimatology, Palaeoecology, 83: 341-366.

  p. 250 “Of dinosaur coprolites identified thus far, the best understood ones are attributed to the Late Cretaceous hadrosaur Maiasaura of Montana.” Chin, K. 2007. The paleobiological implications of herbivorous dinosaur coprolites from the Upper Cretaceous Two Medicine Formation of Montana: why eat wood? Palaios, 22: 554-566.

  p. 250 “Other large coprolites are credited to Late Cretaceous tyrannosaurids, such as Tyrannosaurus rex; one of these is more than twice the length of a 12-inch sub sandwich.” (1) Chin, K., Tokaryk, T.T., Erickson, G.M., and Calk, L.C. 1998. A king-sized theropod coprolite. Nature, 393: 680-682. (2) Chin, K., Eberth, D.A., Schweitzer, M.H., Rando, T.A., Sloboda, W.J., and Horner, J.R. 2003. Remarkable preservation of undigested muscle tissue within a Late Cretaceous Tyrannosaurus coprolite from Alberta, Canada. Palaios, 18: 286-294.

  p. 251 “For example, Early Cretaceous coprolites from Belgium have bone fragments in them.” Poinar, G., Jr., and Boucot, A.J. 2006. Evidence of intestinal parasites of dinosaurs. Parasitology, 133: 245-250.

  p. 251 “This seemingly un-dinosaur-like behavior, which is extremely common in modern birds, was proposed for the bizarre theropod Mononykus. …” Senter (2005).

  p. 251 “Some beetles roll balls of this nutritious stuff as take-out, which they push into burrows, lay eggs on them, and seal off the burrow.” Bertone, M., Green, J., Washburn, S., Poore, M., Sorenson, C., and Watson, D.W. 2005. Seasonal activity and species composition of dung beetles (Coleoptera: Scarabaeidae and Geotrupidae) inhabiting cattle pastures in North Carolina (USA). Annals of the Entomological Society of America, 98: 309-321.

  p. 252 “As is often the case in ichnology, body parts had little to do with this discovery, which Chin and Gill documented in 1996.” Chin, K., and Gill, B.D. 1996. Dinosaurs, dung beetles, and conifers: participants in a Cretaceous food web. Palaios, 11: 280-285.

  p. 252 “Some of the burrows were open, but in others the insects had actively filled them, having packed a mixture of sediment and dung behind them and leaving distinctly visual ‘plugs.’” Chin and Gill (1996).

  p. 253 “Dung flies are relatively small and normally just lay their eggs on feces… .” Wiegmann, B., and Yeates, D.K. 2013. The Evolutionary Biology of Flies. Columbia University Press, New York: 512 p.

  p. 253 “Dung beetles today employ three different strategies in handling feces: tunneling, dwelling, or rolling.” Martin (2013).

  p. 254 “In a paper published in 2001, geochemist Thomas Hollocher, Karen Chin, and two other colleagues detected both abundant body fossils and chemical signatures of anaerobic bacteria in the coprolites.” Hollocher, T.C., Chin, K., Hollocher, K.T., and Kruge, M.A. 2001. Bacterial residues in coprolite of herbivorous dinosaurs: role of bacteria in mineralization of feces. Palaios, 16: 547-565.

  p. 254 “Once these researchers examined thin sections of the coprolites under microscopes, they realized that the calcite in the coprolites was probably precipitated in two stages… .” Hollocher et al. (2001).

  p. 254 “The fossilized wood lacked lignin, a connective tissue that holds wood fibers together.” Chin (2007).

  p. 257 “We did not know it at the time, but Karen Chin had already finished studying these snails and surmised how they got into feces in the first place.”
Chin, K., Hartman, J.H., and Roth, B. 2009. Opportunistic exploitation of dinosaur dung: fossil snails in coprolites from the Upper Cretaceous Two Medicine Formation of Montana. Lethaia, 42: 185-198.

  p. 258 “Speaking of ecosystems, individual dinosaurs harbored their own diverse microflora and microfauna … called a microbiome.” Because people tend to be anthropocentric (go figure), most literature about microbiomes focuses on them in humans. But here’s one in which the authors show how these microorganisms can influence non-human animal behavior: Ezenwa, V.O., Gerardo, N.M., Inouye, D.W., Medina, M., and Xavier, J.B. 2012. Animal behavior and the microbiome. Science, 338: 198-199.

  p. 258 “Not all of this flora and fauna are necessarily harmful to the host, though; for example, some gut bacteria help to produce vitamins K and B12… .” Hill, M.J. 1997. Intestinal flora and endogenous vitamin synthesis. European Journal of Cancer Prevention, 6: S43-45.

  p. 258 “Parasites fall into two broad categories: endoparasites (living inside), such as tapeworms, and ectoparasites (living outside), such as ticks.” Gunn, A., and Pitt, S.J. 2012. Parasitology: An Integrated Approach. John Wiley & Sons, New York: 442 p.

  p. 259 “In 2006, two paleontologists, George Poinar and Art Boucot, documented three types of endoparasites in a dinosaur coprolite… .” Poinar and Boucot (2006).

  p. 259 “The coprolites, first identified in 1903, ranged from 2 to 5 cm (1–2 in) wide and 11 to 13 cm (4.3–5 in) long, or human-sized.” Poinar and Boucot (2006).

  p. 260 “The first known coprolite attributed to a tyrannosaurid came out of fluvial (river) deposits in the Frenchman Formation of Saskatchewan.” Chin et al. (1998).

  p. 261 “Furthermore, toothmarks in Triceratops and other dinosaur bones show that T. rex punctured bone.” Erickson et al. (1996).

  p. 261 “Despite the impressive size of this T. rex coprolite, it was surpassed by one from the Late Cretaceous Dinosaur Park Formation of Alberta, reported in 2003.” Chin et al. (2003).

  p. 263 “Reported in 2005 by paleontologist Kurt Hollocher and three colleagues, these coprolites, like the tyrannosaurid ones, had bone fragments and were cemented with apatite.” Hollocher, K.T., Alcober, O.A., Colombi, C.E., and Hollocher, T.C. 2005. Carnivore coprolites from the Upper Triassic Ischigualasto Formation, Argentina: chemistry, mineralogy, and evidence for rapid initial mineralization. Palaios, 20: 51-63.