Mammalian dwarfing during ancient greenhouse warming events.

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

The Paleocene-Eocene Thermal Maximum, known as PETM (approximately 55.8 million years ago), was a short-lived (~ 200,000 years) global warming event due 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 second largest hyperthermal of the early Eocene, known as ETM2, occurred about 2 million years after the PETM (approximately 53.7 Ma). Another smaller-amplitude hyperthermal, identified as “H2,” appears in the marine record about 100,000 years after ETM2 (approximately 53.6 Ma).

Sifrhippus sp. restoration in the Naturhistoriska Riksmuseet, Stockholm, Sweden (From Wikimedia Commons)

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. 

Fossils of early equids are common in lower Eocene deposits of the Bighorn Basin, making a comparison between the PETM and ETM2 hyperthermal events possible. Using tooth size as a proxy for body size, researchers found a statistically significant decrease in the body size of mammals’ during the PETM and ETM2. Teeth in adult mammals scale proportionally to body size. For instance, Sifrhippus demonstrated a decrease of at least 30% in body size during the first 130,000 years of the PETM, followed by a 76% rebound in body size during the recovery phase of the PETM. Arenahippus, an early horse the size of a small dog, decreased by about 14 percent in size during the ETM2. (D’Ambrosia et al., 2017)

Arenahippus jaw fragment (Image credit: University of New Hampshire)

Body size change during periods of climate change is commonly seen throughout historical and geological records. Studies of modern animal populations have also yielded similar body size results. Tropical trees, anurans and mammals have all demonstrated decreased size or growth rate during drought years. In the case of mammals, the observed decrease in the average body size could have been an evolutionary response to create a more efficient way to reduce body heat.

The combination of global warming and the release of large amounts of carbon to the ocean-atmosphere system during the PETM has encouraged analogies with the modern anthropogenic climate change, which has already led to significant shifts in the distribution, phenology and behaviour of organisms. Plus, the consequences of shrinkage are not yet fully understood. This underlines the urgency for immediate action on global carbon emission reductions.

 

 

References:

Abigail R. D’Ambrosia, William C. Clyde, Henry C. Fricke, Philip D. Gingerich, Hemmo A. Abels. Repetitive mammalian dwarfing during ancient greenhouse warming events. Science Advances, 2017; 3 (3): e1601430 DOI: 10.1126/sciadv.1601430

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

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

 

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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.

Response of marine ecosystems to the PETM.

Biotic change among foraminifera during and after the PETM. (From Speijer et al., 2012)

Biotic change during and after the PETM. (From Speijer et al., 2012)

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. During the PETM, around 5 billion tons of CO2 was released into the atmosphere per year, and temperatures increased by 5 – 9°C. 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 and a rise in global sea level.

Dinoflagellate cysts: Adnatosphaeridium robustum and Apectodinium augustum (From Sluijs and Brinkhuis, 2009)

Dinoflagellate cysts: Adnatosphaeridium robustum and Apectodinium augustum (From Sluijs and Brinkhuis, 2009)

The rapid warming at the beginning of the Eocene has been inferred from the widespread distribution of dinoflagellate cysts. One taxon in particularly, Apectodinium, spans the entire carbon isotope excursion (CIE) of the PETM. The distribution of Apectodinium is linked to high temperatures and increased food availability.

The most disruptive impact during the PETM was likely the exceptional ocean acidification and the rise of the calcite dissolution depth, affecting marine organisms with calcareous shells (Zachos et al., 2005). When CO2 dissolves in seawater, it produce carbonic acid. The carbonic acid dissociates in the water releasing hydrogen ions and bicarbonate. The formation of bicarbonate then removes carbonate ions from the water, making them less available for use by organisms.

Nannofossil abundance changes during the PETM. (From Kump, 2009.)

Nannofossil abundance changes during the PETM. (From Kump, 2009.)

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.

The response of most marine invertebrates (mollusks, echinoderms, brachiopods) to paleoclimatic change during the PETM is poorly documented.

Coccolithus bownii and Toweius pertusus

Coccolithus bownii and Toweius pertusus (Adapted from Bown and Pearson, 2009)

The PETM is also associated with dramatic changes among the calcareous plankton,characterized by the appearance of transient nanoplankton taxa of heavily calcified forms of Rhomboaster spp., Discoaster araneus, and D. anartios as well as Coccolithus bownii, a more delicate form.

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. The current rate of the anthropogenic carbon input  is probably greater than during the PETM, causing a more severe decline in ocean pH and saturation state. Also the biotic consequences of the PETM were fairly minor, while the current rate of species extinction is already 100–1000 times higher than would be considered natural. This underlines the urgency for immediate action on global carbon emission reductions.

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

 

References:

Maria Rose Petrizzo, The onset of the Paleocene–Eocene Thermal Maximum (PETM) at Sites 1209 and 1210 (Shatsky Rise, Pacific Ocean) as recorded by planktonic foraminifera, Marine Micropaleontology, Volume 63, Issues 3–4, 13 June 2007, Pages 187-200

Zeebe RE and Zachos JC. 2013 Long-term legacy ofmassive carbon input to the Earth system: Anthropocene versus Eocene. Phil Trans R Soc A 371: 20120006. http://dx.doi.org/10.1098/rsta.2012.0006.

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.

Kump, L.R., T.J. Bralower, and A. Ridgwell. 2009. Ocean acidification in deep time. Oceanography 22(4):94–107, http://dx.doi.org/10.5670/oceanog.2009.100

Raffi, I. and de Bernardi, B.: Response of calcareous nannofossils to the Paleocene-Eocene Thermal Maximum: observations on composition, preservation and calcification in sediments from ODP Site 1263 (Walvis Ridge – SW Atlantic), Mar. Micropaleontol., 69, 119–138, 2008.

Robert P. SPEIJER, Christian SCHEIBNER, Peter STASSEN & Abdel-Mohsen M. MORSI; Response of marine ecosystems to deep-time global warming: a synthesis of biotic patterns across the Paleocene-Eocene thermal maximum (PETM), Austrian Journal of Earth Sciences. Vienna. 2012. Volume 105/1.

Ocean acidification: Anthropocene versus Eocene.

Ammonia beccarii, Benthonic foraminifera. From Wikimedia Commons

Ammonia beccarii, Benthic foraminifera. From Wikimedia Commons

Last week a report from the International Programme on the State of the Ocean (IPSO) declared that ocean acidification has reached an unprecedented level in Earth’s history. Since the Industrial Revolution, the anthropogenic release of CO2 into the Earth’s atmosphere has increased a 40%.
Over the period from 1750 to 2000, the oceans have absorbed approximately one-third of the CO2 emitted by humans. The cost of this is the decrease in surface ocean pH, that cause dramatic effects on marine life.

When CO2 dissolves in seawater, it produce carbonic acid. The carbonic acid dissociates in the water releasing hydrogen ions and bicarbonate. The formation of bicarbonate then removes carbonate ions from the water, making them less available for use by organisms.

Other consequences of an increasingly acidic ocean include effects on metal speciation, reduced NH3/NH4+ ratios and alteration of underwater sound absorption.

Geological context for ocean acidification. (A) Candidate ocean acidification events. (B) Ocean surface pH calculated at 20 million year intervals. (C) Major changes in plankton assemblages. From Kump, 2009.

The geological record of ocean acidification may provide valuable insights for the future of Earth’s climate and how marine organisms could adapt to severe conditions.

The closest analog for today conditions is the Palaeocene–Eocene Thermal Maximum (PETM, approx. 56Ma), meaning greater similarities in continental configuration, ecosystem structure and function, and global carbon cycling.

The PETM was a short-lived (~ 200,000 years) global warming event when temperatures increased by 5-9°C. It was marked by the largest deep-sea mass extinction among calcareous benthic foraminifera in the last 93 million years. Similarly, planktonic foraminifer communities at low and high latitudes show reductions in diversity.

Nannofossil abundance changes during the PETM. From Kump, 2009.

The PETM is also associated with dramatic changes among the calcareous plankton,characterized by the appearance of transient nanoplankton taxa of heavily calcified forms of Rhomboaster spp., Discoaster araneus, and D. anartios as well as Coccolithus bownii, a more delicate form.

Because the PETM is considered the best analog to modern global warming, the changes in the assemblage of calcareous nanoplankton during this event could provide vital clues to the potential response of modern nanoplankton to ocean acidification.

Not only the magnitude but also the time scale of the carbon input is critical for its effect on ocean carbonate chemistry. The time scale of the anthropogenic carbon input is so short that the natural capacity of the surface reservoirs to absorb carbon is overwhelmed.

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

So, the anthropogenic carbon input rate is probably greater than during the PETM, causing a more severe decline in ocean pH and saturation state. Also the biotic consequences of the PETM were fairly minor, while the current rate of species extinction is already 100–1000 times higher than would be considered natural. This underlines the urgency for immediate action on global carbon emission reductions.

Trevor Manuel, a South African government minister and co-chair of the Global Ocean Commission stated that “Governments must respond as urgently as they do to national security threats – in the long run, the impacts are just as important”.

References:

Zeebe RE and Zachos JC. 2013 Long-term legacy ofmassive carbon input to the Earth system: Anthropocene versus Eocene. Phil Trans R Soc A 371: 20120006. http://dx.doi.org/10.1098/rsta.2012.0006.

Kump, L.R., T.J. Bralower, and A. Ridgwell. 2009. Ocean acidification in deep time. Oceanography 22(4):94–107, http://dx.doi.org/10.5670/oceanog.2009.100