Solving the mystery of Megatherium diet.

Megatherium americanum, MACN.

Around 10,000 years ago, Argentina was home of numerous species of giant Xenarthrans, giant ground sloths (relative to tree sloth) and glyptodontids (relative to tiny extant armadillo). Sloths, characteristic of the mammal fauna of the Pleistocene of South America, show a great diversity with more than 80 genera, grouped in four families: Megatheriidae, Megalonychidae, Nothrotheriidae and Mylodontidae.

For more than a century different hypotheses on the dietary preferences of giant ground sloths have been proposed. In 1860, Owen gave an extensive explanations about their possible diet and behavior. He based his conclusions on the morphology of the skull, combined with peculiarities of the rest of the skeleton, but always by analogy with living tree sloth. He wrote: “Guided by the general rule that animals having the same kind of dentition have the same kind of food, I conclude that the Megatherium must have subsisted, like the Sloths, on the foliage of tree…”. In 1926, Angel Cabrera discussed the diet of Megatherium, rejecting some theories on myrmecophagy or insectivory, and agreed with Owen’s statements about a folivorous diet.

Megatherium americanum lower right tooth series. Scale bar: 5 cm (From M.S. Bargo and S.F. Vizcaíno, 2008)

The dietary preferences of extinct mammals can usually be evaluated through their tooth morphology, but the application of stable isotopes on fossil bones has yielded very important information to solve debates about the diet of extinct large mammal groups, by comparing the carbon and nitrogen isotopic composition of their bone collagen with those of coeval herbivorous and carnivorous taxa. Another isotopic approach is to mesure the difference between the carbon isotopic abundances of the collagen and the carbonate fractions of skeletal tissues. An animal with a herbivorous diet, exhibits significantly larger differences than a carnivore. The values measured on bone collagen from Megatherium, clearly fall in the same range as the large herbivores such as the equid Hippidion, the notoungulate Toxodon and the liptoptern Macrauchenia, for which there is no doubt about their herbivorous diet. Therefore, the hypotheses of insectivory or carnivory for these extinct mammals are not supported by the isotopic data.

 

References:

Hervé Bocherens et al. Isotopic insight on paleodiet of extinct Pleistocene megafaunal Xenarthrans from Argentina, Gondwana Research (2017). DOI: 10.1016/j.gr.2017.04.003

Bargo, M.S., Vizcaíno, S.F., 2008. Paleobiology of Pleistocene ground sloths (Xenarthra, Tardigrada): biomechanics, morphogeometry and ecomorphology applied to the masticatory apparatus. Ameghiniana 45: 175-196

Liaodactylus primus and the ecological evolution of Pterodactyloidea.

Skull of the newfound species Liaodactylus primus (Credit: Chang-Fu Zhou)

Skull of the newfound species Liaodactylus primus (Credit: Chang-Fu Zhou)

Pterosaurs are an extinct monophyletic clade of ornithodiran archosauromorph reptiles from the Late Triassic to Late Cretaceous. The group achieved high levels of morphologic and taxonomic diversity during the Mesozoic, with more than 150 species recognized so far. During their 149 million year history, the evolution of pterosaurs resulted in a variety of eco-morphological adaptations, as evidenced by differences in skull shape, dentition, neck length, tail length and wing span. Pterosaurs have traditionally been divided into two major groups, “rhamphorhynchoids” and “pterodactyloids”. Rhamphorhynchoids are characterized by a long tail, and short neck and metacarpus. Pterodactyloids have a much larger body size range, an elongated neck and metacarpus, and a relatively short tail. Darwinopterus from the early Late Jurassic of China appear to be a transitionary stage that partially fills the morphological gap between rhamphorhynchoids and pterodactyloids.

Pterodactyloidea, the most species-diverse group of pterosaurs, ruled the sky from Late Jurassic to the end of Cretaceous. Liaodactylus primus, a new specimen, discovered in northeast China’s Liaoning province, documents the only pre-Tithonian (145–152 Ma) pterodactyloid known with a complete skull, shedding new light on the origin of the Ctenochasmatidae, a group of exclusive filter feeders, and the timing of the critical transition from fish-catching to filter-feeding, a major ecological shift in the early history of the pterodactyloid clade. The holotype specimen is a nearly complete skull (133 mm long) and mandibles, with the first two cervical vertebrae preserved in articulation with the skull. The elongation of the rostrum, almost half the length of the skull, is accompanied by a significant increase in the number of marginal teeth, giving a total of 152 teeth in both sides of the upper and lower jaws. The teeth are closely spaced to form a ‘comb dentition’, a filter-feeding specialization.

Pterodaustro guinazui cast (Museo Argentino de Ciencias Naturales)

Pterodaustro guinazui cast (Museo Argentino de Ciencias Naturales)

Liaodactylus is the oldest known ctenochasmatid, predating the previously Tithonian (152 Ma) record (Gnathosaurus and Ctenochasma from Germany) by at least 8–10 Myr . The Ctenochasmatidae, represents a long-ranged clade (160–100 Ma), and the only pterodactyloid clade that crossed the Jurassic-Cretaceous transition. The group includes the Early Cretaceous Pterodaustro from Argentina. Popularly called the ‘flamingo pterosaur’, Pterodaustro represents the most remarkable filter-feeding pterosaur known from the fossil record, with a huge number (more than 1000) of densely spaced ‘teeth’ (elastic bristles) in its lower jaws, for filtering small crustaceans, microscopic plankton or algae from open water along lake shores.

Pterosaurs display an extraordinary eco-morphological disparity in feeding adaptations, expressed in skull, jaws and dentition. The Late Triassic Eopterosauria, the basalmost pterosaur clade, were mainly insectivorous. Jurassic insectivores include the Dimorphodontia, Campylognathoididae and Darwinoptera, whereas the Anurognathidae were the only Jurassic insectivores that survived the Jurassic–Cretaceous transition, but became extinct in the Early Cretaceous. The rise of the ctenochasmatid clade was the first major ecological shift in pterosaur evolution from insectivorous-piscivorous to filter-feeding. During Cretaceous time,  the Eupterodactyloidea, a group of advanced pterodactyloids, engaged in a variety of feeding adaptations, including filter-feeding, fish-eating, carnivory and scavenging, herbivory including frugivory, durophagy and omnivory. The Early Cretaceous tapejarids may have been herbivorous, while the pteranodontids, with large skull but tapering and toothless jaws were suitable for seizing fish in open-water environments. Finally, the Late Cretaceous azhdarchids have been hypothesized as foragers feeding on small animals and carrion in diverse terrestrial environments.

Time-calibrated cladogram showing stratigraphic range, eco-morphological diversity of pterosaur clades. (Adapted from Zhou et al., 2017)

Time-calibrated cladogram showing stratigraphic range, eco-morphological diversity of pterosaur clades. (Adapted from Zhou et al., 2017)

References:

Chang-Fu Zhou, Ke-Qin Gao, Hongyu Yi, Jinzhuang Xue, Quanguo Li, Richard C. Fox, Earliest filter-feeding pterosaur from the Jurassic of China and ecological evolution of Pterodactyloidea, 

Andres, B., Clark, J., & Xu, X. (2014). The earliest pterodactyloid and the origin of the group. Current Biology, 24(9), 1011-1016.

WITTON, M. P., 2010 Pteranodon and beyond: the history of giant pterosaurs from 1870 onwards. In: Moody, R.T.J., Buffetaut, E., Naish, D., Martill, D.M. (Eds.), Dinosaurs and Other Extinct Saurians: A Historical Perspective. Geological Society, London, Special Publications 343, 287–311.

 

Climate model simulations at the end of the Cretaceous.

gg_60212w_crater

Artist’s reconstruction of Chicxulub crater 66 million years ago.

About thirty years ago, the discovery of anomalously high abundance of iridium and other platinum group elements in the Cretaceous/Palaeogene (K-Pg) boundary led to the hypothesis that a 10 km asteroid collided with the Earth and caused one of the most devastating events in the history of life. The impact created the 180-kilometre wide Chicxulub crater causing widespread tsunamis along the coastal zones of the surrounding oceans and released an estimated energy equivalent of 100 teratons of TNT and produced high concentrations of dust, soot, and sulfate aerosols in the atmosphere.

To model the climatic effects of the impact, a team of scientist from the Potsdam Institute for Climate Impact Research (PIK), use literature information from geophysical impact modeling indicating that for a 2.9 km thick target region consisting of 30% evaporites and 70% water-saturated carbonates and a dunite projectile with 50% porosity, a velocity of 20 km/s and a diameter between 15 and 20 km, a sulfur mass of 100 Gt is produced. This is about 10,000 times the amount of sulfur released during the 1991 Pinatubo eruption. Additionally, for a sulfur mass of 100 Gt, about 1400 Gt of carbon dioxide are injected into the atmosphere, corresponding to an increase of the atmospheric CO2 concentration by 180 ppm. There could be additional CO2 emissions from ocean outgassing and perturbations of the terrestrial biosphere, adding a total of 360 ppm and 540 ppm of CO2. The main result is a severe and persistent global cooling in the decades after the impact. Global annual mean temperatures over land dropped to -32C in the coldest year and continental temperatures in the tropics reaching a mere -22C. This model is supported by a migration of cool, boreal dinoflagellate species into the subtropic Tethyan realm directly across the K–Pg boundary interval and the ingression of boreal benthic foraminifera into the deeper parts of the Tethys Ocean, interpreted to reflect millennial timescale changes in the ocean circulation after the impact (Vellekoop, 2014).

A time-lapse animation showing severe cooling due to sulfate aerosols from the Chicxulub asteroid impact 66 million years ago (Credit: PKI)

A time-lapse animation showing severe cooling due to sulfate aerosols from the Chicxulub asteroid impact 66 million years ago (Credit: PKI)

References:

Brugger J., G. Feulner, and S. Petri (2016), Baby, it’s cold outside: Climate model simulations of the effects of the asteroid impact at the end of the Cretaceous, Geophys. Res. Lett., 43,  doi:10.1002/2016GL072241.

Alvarez, L. W., W. Alvarez, F. Asaro, and H. V. Michel (1980), Extraterrestrial Cause for the Cretaceous-Tertiary Extinction, Science, 208 (4448), 1095{1108, doi: 10.1126/science.208.4448.1095.

Galeotti, S., H. Brinkhuis, and M. Huber (2004), Records of post Cretaceous-Tertiary boundary millennial-scale cooling from the western Tethys: A smoking gun for the impact-winter hypothesis?, Geology, 32, 529, doi:10.1130/G20439.1

Johan Vellekoop, Appy Sluijs, Jan Smit, Stefan Schouten, Johan W. H. Weijers, Jaap S. Sinningh Damsté, and Henk Brinkhuis, Rapid short-term cooling following the Chicxulub impact at the Cretaceous–Paleogene boundary, PNAS (2014) doi: 10.1073/pnas.1319253111

Sea level regulated tetrapod diversity dynamics through the Jurassic/Cretaceous interval.

The Jurassic/Cretaceous Boundary

Map of the Jurassic/Cretaceous Boundary by C. Scotese.

The Jurassic/Cretaceous (J/K) boundary, 145 Myr ago, remains as the less understood of major Mesozoic stratigraphic boundaries. Sedimentological, palynological and geochemical studies, indicate a climatic shift from predominantly arid to semi arid conditions in the latest Jurassic to more amicable humid conditions in the earliest Cretaceous. The continued fragmentation of Pangaea across the Late Jurassic and Early Cretaceous led to large-scale tectonic processes, on both regional and global scale, accompanied by some of the largest volcanic episodes in the history of the Earth; eustatic oscillations of the sea level; potentially heightened levels of anoxia, oceanic stagnation, and sulphur toxicity; along with two purported oceanic anoxic events in the Valanginian and Hauterivian. There’s also evidence of three large bolide impacts in the latest Jurassic, one of which might have been bigger than the end-Cretaceous Chicxulub impact.

Painting of a late Jurassic Scene on one of the large island in the Lower Saxony basin in northern Germany (From Wikimedia Commons)

Painting of a late Jurassic Scene on one of the large island in the Lower Saxony basin in northern Germany (From Wikimedia Commons)

The J/K interval represents a period of elevated extinction, and involves the persistent loss of diverse lineages, and the origins of many major groups that survived until the present day (e.g. birds). The magnitude of this drop in diversity ranges from around 33% for ornithischians to 75–80% loss for theropods and pterosaurs. Mammals suffered an overall loss of  diversity of 69%. Crocodyliforms suffered a major  decline across the Jurassic/Cretaceous boundary in both the marine and terrestrial realms; while non-marine turtles declined by 33% of diversity through the J/K boundary. In contrast, lepidosauromorphs greatly increased in diversity (48%) across the J/K boundary, reflecting the diversification of major extant squamate clades, including Lacertoidea, Scincoidea and Iguania.

In the marine realm, sauropterygians and ichthyosaurs show evidence for a notable decline in diversity across the J/K boundary, which continued into the Hauterivian for both groups.

Stratigraphic ranges of major Jurassic–Cretaceous theropod (A), sauropod (B), and ornithischian (C) dinosaur clades through the Middle Jurassic to Early Cretaceous (From Tennant et al,. 2016)

Stratigraphic ranges of major Jurassic–Cretaceous theropod (A), sauropod (B), and ornithischian (C) dinosaur clades through the Middle Jurassic to Early Cretaceous (From Tennant et al,. 2016)

Eustatic sea level is the principal mechanism controlling the Jurassic–Cretaceous diversity of tetrapods. Rising sea levels leads to greater division of landmasses through creation of marine barriers, modifying the spatial distribution of near-shore habitats and affecting the species–area relationship, which can lead to elevated extinctions. This fragmentation can also be a potential driver of biological and reproductive isolation and allopatric speciation, the combination of which we would expect to see manifest in the diversity signal. Additionally, the diversity of fully marine taxa was more probably affected by the opening and closure of marine dispersal corridors, whereas that of terrestrial and coastal taxa was more probably dependent on the availability of habitable ecosystems, including the extent of continental shelf area (Tennant, et al., 2016).

References:

Tennant J,P., Mannion P. D., Upchurch P., Sea level regulated tetrapod diversity dynamics through the Jurassic/Cretaceous interval, Nature Communications, ISSN: 2041-1723 DOI: 10.1038/ncomms12737

Tennant, J. P., Mannion, P. D., Upchurch, P., Sutton, M. D. and Price, G. D. (2016), Biotic and environmental dynamics through the Late Jurassic–Early Cretaceous transition: evidence for protracted faunal and ecological turnover. Biol Rev. doi:10.1111/brv.12255 

Butler, R. J., Benson, R. B. J., Carrano, M. T., Mannion, P. D. & Upchurch, P. Sea level, dinosaur diversity and sampling biases: investigating the ‘common cause’ hypothesis in the terrestrial realm. Proc. R. Soc. B 278, 1165–1170 (2011).

Climate Change and the Evolution of Mammals.

Wyoming_Bighorn_Basin

Bighorn Basin, Wyoming (Image: University of New Hampshire, College of Engineering and Physical Sciences).

Rapid global climate change can lead to rapid evolutionary responses. The Paleocene-Eocene Thermal Maximum (PETM; 55.8 million years ago), was a short-lived (~ 200,000 years) global warming event attributed to a rapid rise in the concentration of greenhouse gases in the atmosphere. It was suggested that this warming was initiated by the melting of methane hydrates on the seafloor and permafrost at high latitudes. This event was accompanied by other large-scale changes in the climate system, for example, the patterns of atmospheric circulation, vapor transport, precipitation, intermediate and deep-sea circulation, a rise in global sea level and ocean acidification.

The PETM onset is also marked by the largest deep-sea mass extinction among calcareous benthic foraminifera (including calcareous agglutinated taxa) in the last 93 million years. Similarly, planktonic foraminifera communities at low and high latitudes show reductions in diversity, while larger foraminifera are the most common constituents of late Paleocene–early Eocene carbonate platforms.

Phenacodus

Phenacodus by Heinrich Harder (1858-1935) . From Wikimedia Commons.

During the PETM, around 5 billion tons of CO2 was released into the atmosphere per year, and temperatures increased by 5 – 8°C. The rise in temperature coincided with a dramatic decrease in the body size of marine and terrestrial organisms. Dwarfing of mammalian taxa across the Palaeocene-Eocene Thermal Maximum (PETM) was first described in the Bighorn Basin, Wyoming. The basin has a remarkably fossil-rich sedimentary record of late Palaeocene to early Eocene age.  The interval of the Paleocene–Eocene Thermal Maximum is represented by a unique mammalian fauna composed by smaller, but morphologically similar species to those found later in the Eocene. Diminutive species include the early equid Sifrhippus sandrae, the phenacodontids Ectocion parvus and Copecion davisi. Two main hypotheses have been proposed to explain the observation of smaller body sizes during the global warming event. The first hypothesis is that mammal population decreased the average body-size in response to the environmental conditions that existed during the PETM global warming event. The second hypothesis is that the observed decrease in the average body-size was the result of extrinsic forces, such as the range extension of small species into the Bighorn Basin, displacing larger species (Burger, 2012). 

Comparison of the effects of anthropogenic emissions (total of 5000 Pg C over 500 years) and PETM carbon release (3000 Pg C over 6 kyr) on the surface ocean saturation state of calcite. From Zeebe, 2013

Comparison of the effects of anthropogenic emissions (total of 5000 Pg C over 500 years) and PETM carbon release (3000 Pg C over 6 kyr) on the surface ocean saturation state of calcite. From Zeebe, 2013

New findings revealed that the remarkable decrease in mean body size across the warming event, occurred through anagenetic change and immigration. However, species selection also was strong across the PETM but, intriguingly, favoured larger-bodied species, implying some unknown mechanism(s) by which warming events affect macroevolution (Rankin et al., 2015). 

Climate change is the major threat to biodiversity. The combination of global warming and the release of large amounts of carbon to the ocean-atmosphere system during the PETM has encouraged analogies to be drawn with modern anthropogenic climate change. Reduction in nutrients, food availability and water will probably have negative implications and are interrelated with climate change and shrinking organisms.  We need to understand how and why organisms are shrinking, and what it means for biodiversity and humanity.

References:

Rankin, B., Fox, J., Barron-Ortiz, C., Chew, A., Holroyd, P., Ludtke, J., Yang, X., Theodor, J. 2015. The extended Price equation quantifies species selection on mammalian body size across the Palaeocene/Eocene Thermal Maximum. Proceedings of the Royal Society B. doi: 10.1098/rspb.2015.1097

Barnosky, A. D. 2004 Biodiversity response to climate change in the middle Pleistocene: the Porcupine Cave fauna from Colorado. Berkeley, CA: University of California Press.

Burger, B.J., Northward range extension of a diminutive-sized mammal (Ectocion parvus) and the implication of body size change during the Paleoc…, Palaeogeogr. Palaeoclimatol. Palaeoecol. (2012), http://dx.doi.org/10.1016/j.palaeo.2012.09.008

Jablonski, D. 2008, Species selection: theory and data. Annu. Rev. Ecol. Evol. Syst. 39, 501–524.

Sheriden, J. A; Bickford, D. 2011, Shrinking body size as an ecological response to climate change. Nat. Clim.

Wright JD, Schaller MF (2013) Evidence for a rapid release of carbon at the Paleocene-Eocene thermal maximum. Proc Natl Acad Sci USA 110(40):15908–15913.

A Brief Introduction to Paleoecology.

Duria Antiquior famous watercolor by the geologist Henry de la Beche based on fossils found by Mary Anning. From Wikimedia Commons.

Duria Antiquior famous watercolor by the geologist Henry de la Beche based on fossils found by Mary Anning. From Wikimedia Commons.

Paleocology is a multidisciplinary science. It involves the reconstruction of past environments from geological and fossil  evidence. A more exhaustive definition was given by Valentí Rull in 2010: “the branch of ecology that studies the past of ecological systems and their trends in time using fossils and other proxies”. Paleoecology can be used to investigate (1) the rates of speciation and extinction, (2) biome shifts and ecosystem development and (3) adaptation, migration, and population change (Seppä, 2009).

Charles Lyell (1797–1875) and Roman Fedorovich Gekker (1900-1991)

Charles Lyell (1797–1875) and Roman Fedorovich Gekker (1900-1991)

The major philosophical concepts in paleoecology are uniformitarianism, analogy, and parsimony. The concept of uniformitarianism was created by James Hutton (1726–97) and  Charles Lyell (1797–1875). It can be summarized as ‘the present is the key to the past’ and is the basic principle of paleoecology. The concept of analogy involves the application of modern organismic features to ancient organisms, and of course parsimony is a central rule for any scientific inquiry. In 1933, the Russian paleontologist Roman Gekker published the first book dedicated to paleoecology: “Manual to Paleoecology”, based in his lectures about the Devonian Period. In this book he established the main objectives of Paleoecology. Later, in 1954, he wrote “Directions for Research in Paleoecology” and in 1957, he published “Introduction to Paleoecology”.

Scanning electron microscope image of different types of pollen grains. Image from Wikipedia.

Scanning electron microscope image of different types of pollen grains. Image from Wikipedia.

There are two major types of paleoecology: Quaternary paleoecology, concerned with the last 2.6 million years of Earth’s history, and Deep-time paleoecology, based on fossils from pre-Quaternary sediments over a wide range of timescales (Birks, 2013). In the last four decades, quantitative methods for reconstructing environmental variables have been developed from a range of biological proxies such as pollen, plant macrofossils, insects (chironomids, coleopterans), molluscs, ostracods, diatoms, chrysophycean cysts, testate amoebae, and cladocerans preserved in lake sediments and peat profiles, or dinoflagellate cysts, diatoms, pollen, foraminifera, coccolithophores, and radiolarians preserved in marine sediment records.

Lago Sarmiento in Southern Patagonia. Sediment cores recovered from lakes like this, help to reconstruct environmental changes. Photo credit: R. Dunbar.

Lago Sarmiento in Southern Patagonia. Sediment cores recovered from lakes like this, help to reconstruct environmental changes. Photo credit: R. Dunbar.

The dominant technique in Quaternary terrestrial paleoecology is the pollen analysis. Pollen analysis involves the quantitative examination of spores and pollen at successive horizons through a core, particularly in bog, marsh, lake or delta sediments (Armstrong, 2005). This method was created by Lennart von Post (1884–1950), a Swedish geologist and presented at the 16th Scandinavian meeting of natural scientists in Oslo. Since the 1980s, many fossil pollen data sets were developed specifically to reconstruct past climate change.

Reference:

Seddon, A. W. R., Mackay, A. W., Baker, A. G., Birks, H. J. B., Breman, E., Buck, C. E., Ellis, E. C., Froyd, C. A., Gill, J. L., Gillson, L., Johnson, E. A., Jones, V. J., Juggins, S., Macias-Fauria, M., Mills, K., Morris, J. L., Nogués-Bravo, D., Punyasena, S. W., Roland, T. P., Tanentzap, A. J., Willis, K. J., Aberhan, M., van Asperen, E. N., Austin, W. E. N., Battarbee, R. W., Bhagwat, S., Belanger, C. L., Bennett, K. D., Birks, H. H., Bronk Ramsey, C., Brooks, S. J., de Bruyn, M., Butler, P. G., Chambers, F. M., Clarke, S. J., Davies, A. L., Dearing, J. A., Ezard, T. H. G., Feurdean, A., Flower, R. J., Gell, P., Hausmann, S., Hogan, E. J., Hopkins, M. J., Jeffers, E. S., Korhola, A. A., Marchant, R., Kiefer, T., Lamentowicz, M., Larocque-Tobler, I., López-Merino, L., Liow, L. H., McGowan, S., Miller, J. H., Montoya, E., Morton, O., Nogué, S., Onoufriou, C., Boush, L. P., Rodriguez-Sanchez, F., Rose, N. L., Sayer, C. D., Shaw, H. E., Payne, R., Simpson, G., Sohar, K., Whitehouse, N. J., Williams, J. W., Witkowski, A. (2014), Looking forward through the past: identification of 50 priority research questions in palaeoecology. Journal of Ecology, 102: 256–267. doi: 10.1111/1365-2745.12195

Seppä, H. 2009. Palaeoecology. eLS DOI: 10.1002/9780470015902.a0003232

Walker, Mike J. C., and John J. Lowe. 2007. Quaternary science 2007: A 50-year retrospective.Journal of the Geological Society 164.6: 1073–1092. DOI: 10.1144/0016-76492006-195