Thermal performance of ecosystems: Modeling how physiological responses to temperature scale up in communities.

Understanding how ecosystems respond to their environmental temperature is a major challenge. Thermodynamic constraints on species' metabolic rates are expected to affect ecosystem characteristics, but species interactions and interspecific variation in physiological thermal response curves (TRC) may obscure ecosystem-level responses to temperature. As a result, macroecological patterns related to temperature are still poorly understood. We investigate how physiological TRC scale up to ecosystem-level thermal responses by modifying the Tangled Nature (TaNa) model, a stochastic network model of ecology and evolution. We include new parameterizations that make reproduction, death, and mutation temperature-dependent. We find that ecosystem survival probability depends on how the minimum fitness required for species survival varies with temperature. The thermal response of ecosystem survival probability is the only ecosystem property that is sensitive to interspecific variation in TRC. Species richness scales up directly from the TRC of mutation rate, and average species population sizes are inversely related to mutation rate, with Species Abundance Distributions (SADs) exhibiting more rare species in warmer temperatures. Interactions between species are also inversely related to mutation, with positive interactions occurring more frequently in colder temperatures. The abundance of surviving ecosystems is not sensitive to temperature. This work helps clarify the specific relationships between physiological responses to temperature and ecosystem-level repercussions when species are interacting and adapting to their thermal environments.

The Effect of Ocean Salinity on Climate and Its Implications for Earth's Habitability.

The influence of atmospheric composition on the climates of present-day and early Earth has been studied extensively, but the role of ocean composition has received less attention. We use the ROCKE-3D ocean-atmosphere general circulation model to investigate the response of Earth's present-day and Archean climate system to low versus high ocean salinity. We find that saltier oceans yield warmer climates in large part due to changes in ocean dynamics. Increasing ocean salinity from 20 to 50 g/kg results in a 71% reduction in sea ice cover in our present-day Earth scenario. This same salinity change also halves the pCO2 threshold at which Snowball glaciation occurs in our Archean scenarios. In combination with higher levels of greenhouse gases such as CO2 and CH4, a saltier ocean may allow for a warm Archean Earth with only seasonal ice at the poles despite receiving ∼20% less energy from the Sun.

Marine oxygen production and open water supported an active nitrogen cycle during the Marinoan Snowball Earth.

The Neoproterozoic Earth was punctuated by two low-latitude Snowball Earth glaciations. Models permit oceans with either total ice cover or substantial areas of open water. Total ice cover would make an anoxic ocean likely, and would be a formidable barrier to biologic survival. However, there are no direct data constraining either the redox state of the ocean or marine biological productivity during the glacials. Here we present iron-speciation, redox-sensitive trace element, and nitrogen isotope data from a Neoproterozoic (Marinoan) glacial episode. Iron-speciation indicates deeper waters were anoxic and Fe-rich, while trace element concentrations indicate surface waters were in contact with an oxygenated atmosphere. Furthermore, synglacial sedimentary nitrogen is isotopically heavier than the modern atmosphere, requiring a biologic cycle with nitrogen fixation, nitrification and denitrification. Our results indicate significant regions of open marine water and active biologic productivity throughout one of the harshest glaciations in Earth history.

Comment on "Long-term climate forcing by atmospheric oxygen concentrations".

Poulsen et al (Reports, 12 June 2015, p. 1238) argued that lower atmospheric oxygen levels during the Phanerozoic would have given a warmer climate. However, radiative and atmospheric structure changes under lower pressure both cause cooling, making their result unusual in that a hierarchy of models gives opposing results. Scrutiny of how radiative and cloud processes were represented, and a mechanistic explanation of the results, are required.

Habitability of waterworlds: runaway greenhouses, atmospheric expansion, and multiple climate states of pure water atmospheres.

There are four different stable climate states for pure water atmospheres, as might exist on so-called "waterworlds." I map these as a function of solar constant for planets ranging in size from Mars-sized to 10 Earth-mass. The states are as follows: globally ice covered (Ts ⪅ 245 K), cold and damp (270 ⪅ Ts ⪅ 290 K), hot and moist (350 ⪅ Ts ⪅ 550 K), and very hot and dry (Tsx2A86;900 K). No stable climate exists for 290 ⪅ T s ⪅ 350 K or 550 ⪅ Ts ⪅ 900 K. The union of hot moist and cold damp climates describes the liquid water habitable zone, the width and location of which depends on planet mass. At each solar constant, two or three different climate states are stable. This is a consequence of strong nonlinearities in both thermal emission and the net absorption of sunlight. Across the range of planet sizes, I account for the atmospheres expanding to high altitudes as they warm. The emitting and absorbing surfaces (optical depth of unity) move to high altitude, making their area larger than the planet surface, so more thermal radiation is emitted and more sunlight absorbed (the former dominates). The atmospheres of small planets expand more due to weaker gravity; the effective runaway greenhouse threshold is about 35 W m(-2) higher for Mars, 10 W m(-2) higher for Earth or Venus, but only a few W m(-2) higher for a 10 Earth-mass planet. There is an underlying (expansion-neglected) trend of increasing runaway greenhouse threshold with planetary size (40 W m(-2) higher for a 10 Earth-mass planet than for Mars). Summing these opposing trends means that Venus-sized (or slightly smaller) planets are most susceptible to a runaway greenhouse. The habitable zone for pure water atmospheres is very narrow, with an insolation range of 0.07 times the solar constant. A wider habitable zone requires background gas and greenhouse gas: N2 and CO2 on Earth, which are biologically controlled. Thus, habitability depends on inhabitance.

Tidal Venuses: triggering a climate catastrophe via tidal heating.

Traditionally, stellar radiation has been the only heat source considered capable of determining global climate on long timescales. Here, we show that terrestrial exoplanets orbiting low-mass stars may be tidally heated at high-enough levels to induce a runaway greenhouse for a long-enough duration for all the hydrogen to escape. Without hydrogen, the planet no longer has water and cannot support life. We call these planets "Tidal Venuses" and the phenomenon a "tidal greenhouse." Tidal effects also circularize the orbit, which decreases tidal heating. Hence, some planets may form with large eccentricity, with its accompanying large tidal heating, and lose their water, but eventually settle into nearly circular orbits (i.e., with negligible tidal heating) in the habitable zone (HZ). However, these planets are not habitable, as past tidal heating desiccated them, and hence should not be ranked highly for detailed follow-up observations aimed at detecting biosignatures. We simulated the evolution of hypothetical planetary systems in a quasi-continuous parameter distribution and found that we could constrain the history of the system by statistical arguments. Planets orbiting stars with masses<0.3 MSun may be in danger of desiccation via tidal heating. We have applied these concepts to Gl 667C c, a ∼4.5 MEarth planet orbiting a 0.3 MSun star at 0.12 AU. We found that it probably did not lose its water via tidal heating, as orbital stability is unlikely for the high eccentricities required for the tidal greenhouse. As the inner edge of the HZ is defined by the onset of a runaway or moist greenhouse powered by radiation, our results represent a fundamental revision to the HZ for noncircular orbits. In the appendices we review (a) the moist and runaway greenhouses, (b) hydrogen escape, (c) stellar mass-radius and mass-luminosity relations, (d) terrestrial planet mass-radius relations, and (e) linear tidal theories.

The runaway greenhouse: implications for future climate change, geoengineering and planetary atmospheres.

The ultimate climate emergency is a 'runaway greenhouse': a hot and water-vapour-rich atmosphere limits the emission of thermal radiation to space, causing runaway warming. Warming ceases only after the surface reaches approximately 1400 K and emits radiation in the near-infrared, where water is not a good greenhouse gas. This would evaporate the entire ocean and exterminate all planetary life. Venus experienced a runaway greenhouse in the past, and we expect that the Earth will in around 2 billion years as solar luminosity increases. But could we bring on such a catastrophe prematurely, by our current climate-altering activities? Here, we review what is known about the runaway greenhouse to answer this question, describing the various limits on outgoing radiation and how climate will evolve between these. The good news is that almost all lines of evidence lead us to believe that is unlikely to be possible, even in principle, to trigger full a runaway greenhouse by addition of non-condensible greenhouse gases such as carbon dioxide to the atmosphere. However, our understanding of the dynamics, thermodynamics, radiative transfer and cloud physics of hot and steamy atmospheres is weak. We cannot therefore completely rule out the possibility that human actions might cause a transition, if not to full runaway, then at least to a much warmer climate state than the present one. High climate sensitivity might provide a warning. If we, or more likely our remote descendants, are threatened with a runaway greenhouse, then geoengineering to reflect sunlight might be life's only hope. Injecting reflective aerosols into the stratosphere would be too short-lived, and even sunshades in space might require excessive maintenance. In the distant future, modifying Earth's orbit might provide a sustainable solution. The runaway greenhouse also remains relevant in planetary sciences and astrobiology: as extrasolar planets smaller and nearer to their stars are detected, some will be in a runaway greenhouse state.

Bistability of atmospheric oxygen and the Great Oxidation.

The history of the Earth has been characterized by a series of major transitions separated by long periods of relative stability. The largest chemical transition was the 'Great Oxidation', approximately 2.4 billion years ago, when atmospheric oxygen concentrations rose from less than 10(-5) of the present atmospheric level (PAL) to more than 0.01 PAL, and possibly to more than 0.1 PAL. This transition took place long after oxygenic photosynthesis is thought to have evolved, but the causes of this delay and of the Great Oxidation itself remain uncertain. Here we show that the origin of oxygenic photosynthesis gave rise to two simultaneously stable steady states for atmospheric oxygen. The existence of a low-oxygen (less than 10(-5) PAL) steady state explains how a reducing atmosphere persisted for at least 300 million years after the onset of oxygenic photosynthesis. The Great Oxidation can be understood as a switch to the high-oxygen (more than 5 x 10(-3) PAL) steady state. The bistability arises because ultraviolet shielding of the troposphere by ozone becomes effective once oxygen levels exceed 10(-5) PAL, causing a nonlinear increase in the lifetime of atmospheric oxygen. Our results indicate that the existence of oxygenic photosynthesis is not a sufficient condition for either an oxygen-rich atmosphere or the presence of an ozone layer, which has implications for detecting life on other planets using atmospheric analysis and for the evolution of multicellular life.