The timing of life history traits is central to lifetime fitness and nowhere is this more evident or well studied as in the phenology of flowering in governing plant reproductive success. Recent changes in the timing of environmental events attributable to climate change, such as the date of snowmelt at high altitudes, which initiates the growing season, have had important repercussions for some common perennial herbaceous wildflower species. The phenology of flowering at the Rocky Mountain Biological Laboratory (Colorado, USA) is strongly influenced by date of snowmelt, which makes this site ideal for examining phenological responses to climate change. Flower buds of Delphinium barbeyi, Erigeron speciosus, and Helianthella quinquenervis are sensitive to frost, and the earlier beginning of the growing season in recent years has exposed them to more frequent mid-June frost kills. From 1992 to 1998, on average 36.1% of Helianthella buds were frosted, but for 1999-2006 the mean is 73.9%; in only one year since 1998 have plants escaped all frost damage. For all three of these perennial species, there is a significant relationship between the date of snowmelt and the abundance of flowering that summer. Greater snowpack results in later snowmelt, later beginning of the growing season, and less frost mortality of buds. Microhabitat differences in snow accumulation, snowmelt patterns, and cold air drainage during frost events can be significant; an elevation difference of only 12 m between two plots resulted in a temperature difference of almost 28C in 2006 and a difference of 37% in frost damage to buds. The loss of flowers and therefore seeds can reduce recruitment in these plant populations, and affect pollinators, herbivores, and seed predators that previously relied on them. Other plant species in this environment are similarly susceptible to frost damage so the negative effects for recruitment and for consumers dependent on flowers and seeds could be widespread. These findings point out the paradox of increased frost damage in the face of global warming, provide important insights into the adaptive significance of phenology, and have general implications for flowering plants throughout the region and anywhere climate change is having similar impacts.

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Effects of climate change on phenology, frost damage, and floral abundance of montane wildflowers

CO2 currently accumulating in the atmosphere permeates into ocean surface layers, where it may impact on marine animals in addition to effects caused by global warming. At the same time, several countries are developing scenarios for the disposal of anthropogenic CO2 in the worlds' oceans, especially the deep sea. Elevated CO2 partial pressures (hypercapnia) will affect the physiology of water breathing animals, a phenomenon also considered in recent discussions of a role for CO2 in mass extinction events in earth history. Our current knowledge of CO2 effects ranges from effects of hypercapnia on acid-base regulation, calcification and growth to influences on respiration, energy turnover and mode of metabolism. The present paper attempts to evaluate critical processes and the thresholds beyond which these effects may become detrimental. CO2 elicits acidosis not only in the water, but also in tissues and body fluids. Despite compensatory accumulation of bicarbonate, acid-base parameters (pH, bicarbonate and CO2 levels) and ion levels reach new steady-state values, with specific, long-term effects on metabolic functions. Even though such processes may not be detrimental, they are expected to affect long-term growth and reproduction and may thus be harmful at population and species levels. Sensitivity is maximal in ommastrephid squid, which are characterized by a high metabolic rate and extremely pH-sensitive blood oxygen transport. Acute sensitivity is interpreted to be less in fish with intracellular blood pigments and higher capacities to compensate for CO2 induced acid-base disturbances than invertebrates. Virtually nothing is known about the degree to which deep-sea fishes are affected by short or long term hypercapnia. Sensitivity to CO2 is hypothesized to be related to the organizational level of an animal, its energy requirements and mode of life. Long-term effects expected at population and species levels are in line with recent considerations of a detrimental role of CO2 during mass extinctions in the earth's history. Future research is needed in this area to evaluate critical effects of the various CO2 disposal scenarios.

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Biological Impact of Elevated Ocean CO2 Concentrations: Lessons from Animal Physiology and Earth History

The deep sea, the largest ecosystem on Earth and one of the least studied, harbours high biodiversity and provides a wealth of resources. Although humans have used the oceans for millennia, technological developments now allow exploitation of fisheries resources, hydrocarbons and minerals below 2000 m depth. The remoteness of the deep seafloor has promoted the disposal of residues and litter. Ocean acidification and climate change now bring a new dimension of global effects. Thus the challenges facing the deep sea are large and accelerating, providing a new imperative for the science community, industry and national and international organizations to work together to develop successful exploitation management and conservation of the deep-sea ecosystem. This paper provides scientific expert judgement and a semi-quantitative analysis of past, present and future impacts of human-related activities on global deep-sea habitats within three categories: disposal, exploitation and climate change. The analysis is the result of a Census of Marine Life – SYNDEEP workshop (September 2008). A detailed review of known impacts and their effects is provided. The analysis shows how, in recent decades, the most significant anthropogenic activities that affect the deep sea have evolved from mainly disposal (past) to exploitation (present). We predict that from now and into the future, increases in atmospheric CO2 and facets and consequences of climate change will have the most impact on deep-sea habitats and their fauna. Synergies between different anthropogenic pressures and associated effects are discussed, indicating that most synergies are related to increased atmospheric CO2 and climate change effects. We identify deep-sea ecosystems we believe are at higher risk from human impacts in the near future: benthic communities on sedimentary upper slopes, cold-water corals, canyon benthic communities and seamount pelagic and benthic communities. We finalise this review with a short discussion on protection and management methods.

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Man and the last great wilderness: human impact on the deep sea.

Solubility of CO2 in water from −1.5 to 100 °C and from 0.1 to 100 MPa: evaluation of literature data and thermodynamic modelling

Currently rising CO 2 levels in atmosphere and marine surface waters as well as projected scenarios of CO 2 disposal in the ocean emphasize that CO 2 sensitivities need to be investigated in aquatic organisms, especially in animals which may well be the most sensitive. Moreover, to understand causes and effects, we need to identify the physiological processes that are sensitive to CO 2 beyond the current emphasis on calcification. Few animals may be acutely sensitive to moderate CO 2 increases, but subtle changes due to long-term exposure may already have started to be felt in a wide range of species. CO 2 effects identified in invertebrate fauna from habitats characterized by oscillating CO 2 levels include depressed metabolic rates and reduced ion exchange and protein synthesis rates. These result in shifts in metabolic equilibria and slowed growth. Long-term moderate hypercapnia has been observed to produce enhanced mortality with as yet unidentified cause and effect relationships. During future climate change, simultaneous shifts in temperature, CO 2 , and hypoxia levels will enhance sensitivity to environmental extremes relative to a change in just one of these variables. Some interactions between these variables result from joint effects on the same physiological mechanisms. Such interactions need to be considered in terms of future increases in atmospheric CO 2 and its uptake by the ocean as well as in terms of currently proposed mitigation scenarios. These include purposeful injection of CO 2 in the deep ocean or Fe fertilization of the surface ocean, which reduces subsurface O 2 levels. The resulting ecosystem shifts could develop progressively, rather than beyond specific thresholds, such that effects parallel CO 2 oscillations. It is unsure to what extent and how quickly species may adapt to permanently elevated CO 2 levels by microevolutionary compensatory processes.

Synergistic effects of temperature extremes, hypoxia, and increases in CO2 on marine animals : From earth history to global change : The ocean in a high-CO2 world

Solubility of CO2 and density of CO2 hydrate at 30 MPa

A field study of the effects of CO2 ocean disposal on mobile deep-sea animals

Experiments on the ocean sequestration of fossil fuel CO2 : pH measurements and hydrate formation

The rapidly rising levels of atmospheric and oceanic CO2 from the burning of fossil fuels has lead to well-established international concerns over dangerous anthropogenic interference with climate. Disposal of captured fossil fuel CO2 either underground, or in the deep ocean, has been suggested as one means of ameliorating this problem. While the basic thermodynamic properties of both CO2 and seawater are well known, the problem of interaction of the two fluids in motion to create a plume of high CO2/low pH seawater has been modeled, but not tested. We describe here a novel experiment designed to initiate study of this problem. We constructed a small flume, which was deployed on the sea floor at 4 km depth by a remotely operated vehicle, and filled with liquid CO2. Seawater flow was forced across the surface by means of a controllable thruster. Obtaining quantitative data on the plume created proved to be challenging. We observed and sensed the interface and boundary layers, the formation of a solid hydrate, and the low pH/high CO2 plume created, with both pH and conductivity sensors placed downstream. Local disequilibrium in the CO2 system components was observed due to the finite hydration reaction rate, so that the pH sensors closest to the source only detected a fraction of the CO2 emitted. The free CO2 molecules were detected through the decrease in conductivity observed, and the disequilibrium was confirmed through trapping a sample in a flow cell and observing an unusually rapid drop in pH to an equilibrium value.

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Deep ocean experiments with fossil fuel carbon dioxide: creation and sensing of a controlled plume at 4 km depth

In order to obtain useful information to assess the feasibility of CO2 confinement in the deep sea, a few experiments were conducted using a facility which can simulate the pressure and temperature condition of deep sea areas shallower than 3000 meters. The characteristics of CO2 clathrate, such as the effect of the mixing process of injected CO2 on the clathrate formation, and the stability of clathrate film formed at the CO2-water interface, were carefully examined. The unexpectedly high permeability of CO2 through clathrate film, which was determined from the observation of the shrinking process of a CO2 droplet and the rate of pressure decrease after CO2 injection, is an undesirable factor for the realization of CO2 confinement in the deep sea.

Izuo Aya

Papers

Solubility of CO2 and density of CO2 hydrate at 30 MPa

A field study of the effects of CO2 ocean disposal on mobile deep-sea animals

Experiments on the ocean sequestration of fossil fuel CO2 : pH measurements and hydrate formation

Deep ocean experiments with fossil fuel carbon dioxide: creation and sensing of a controlled plume at 4 km depth

Stability of Clathrate-Hydrate of Carbon Dioxide in Highly Pressurized water.