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Evolution of CAM and C 4 carbon-concentrating mechanisms

Mechanisms for concentrating carbon around the Rubisco enzyme, which drives the carbon-reducing steps in photosynthesis, are widespread in plants; in vascular plants they are known as crassulacean acid metabolism (CAM) and C 4 photosynthesis. CAM is common in desert succulents, tropical epiphytes, and aquatic plants and is characterized by nighttime fixation of CO2. The proximal selective factor driving the evolution of this CO2-concentrating pathway is low daytime CO2, which results from the unusual reverse stomatal behavior of terrestrial CAM species or from patterns of ambient CO2 availability for aquatic CAM species. In terrestrials the ultimate selective factor is water stress that has selected for increased water use efficiency. In aquatics the ultimate selective factor is diel fluctuations in CO2 availability for palustrine species and extreme oligotrophic conditions for lacustrine species. C4 photosynthesis is based on similar biochemistry but carboxylation steps are spatially separated in the leaf rather than temporally as in CAM. This biochemical pathway is most commonly associated with a specialized leaf anatomy known as Kranz anatomy; however, there are exceptions. The ultimate selective factor driving the evolution of this pathway is excessively high photorespiration that inhibits normal C3 photosynthesis under high light and high temperature in both terrestrial and aquatic habitats. CAM is an ancient pathway that likely has been present since the Paleozoic era in aquatic species from shallow-water palustrine habitats. While atmospheric CO2 levels have undoubtedly affected the evolution of terrestrial plant carbon-concentrating mechanisms, there is reason to believe that past atmospheric changes have not played as important a selective role in the aquatic milieu since palustrine habitats today are not generally carbon sinks, and the selective factors driving aquatic CAM are autogenic. Terrestrial CAM, in contrast, is of increasing selective value under extreme water deficits, and undoubtedly, high Mesozoic CO2 levels reduced the amount of landscape perceived by plants as water limited. Late Tertiary and Quaternary reductions in atmospheric CO2, coupled with increasing seasonality, were probably times of substantial species radiation and ecological expansion for CAM plants. C4 photosynthesis occurs in only about half as many families as CAM, and three-fourths of C4 species are either grasses or sedges. Molecular phylogenies indicate C4 is a more recent innovation than CAM and that it originated in the mid-Tertiary, 20–30 Ma, although some data support an earlier origin. While the timing of the origin of C4 remains controversial, the nearly explosive increase in C4 species is clearly documented in the late Miocene, 4–7 Ma. Increasing seasonality has been widely suggested as an important climatic stimulus for this C4 expansion. Alternatively, based on models of photosynthetic quantum yield at different temperatures and CO2 concentration, it has been hypothesized that the late Miocene C4 expansion resulted from declining atmospheric CO2 levels. This model is most appropriate for explaining the transition from C3 grasslands to C4 grasslands but by itself may not be sufficient to explain the more likely scenario of a late Miocene transition from C3 woodland/ savanna to C4 grasslands. A largely unexplored hypothesis is that climatic changes in late Miocene altered disturbance regimes, in particular the incidence of fires, which today are often associated with maintenance of C4 grasslands. Oceanic charcoal sediments that appear to represent Aeolian deposits from continental wildfires follow a strikingly similar pattern of explosive increase in late Miocene. Climate, CO2, and disturbance are not mutually exclusive explanations and probably all acted in concert to promote the expansion of C4 grasslands. More recently, late Quaternary changes in CO2 may have been responsible for driving major changes in the landscape distribution of C4 species. The theory is sound; however, many of the studies cited in support of this model are open to alternative interpretations, and none has eliminated climatic factors as important selective agents. CAM and C4 evolution required coupling of biochemical pathways with structural changes in photosynthetic tissues, succulence in CAM and Kranz in C4. This was apparently accomplished by piecemeal evolution beginning with mechanisms for recapturing respiratory CO2, although this need not have been so in aquatic CAM species. It has been proposed that the extreme rarity of both pathways in the same plant results from biochemical and structural incompatibilities (Sage 2002). Equally important is the fact that the selective environments are quite different, with CAM evolution thriving on stressful sites inhospitable to C3 species whereas C4 evolution has selected for rapid growth capable of outcompeting associated C3 plants.

International Journal of Plant Sciences↗

Determining CO 2 storage potential during miscible CO 2 enhanced oil recovery: Noble gas and stable isotope tracers

Rising atmospheric carbon dioxide (CO 2 ) concentrations are fueling anthropogenic climate change. Geologic sequestration of anthropogenic CO 2 in depleted oil reservoirs is one option for reducing CO 2 emissions to the atmosphere while enhancing oil recovery. In order to evaluate the feasibility of using enhanced oil recovery (EOR) sites in the United States for permanent CO 2 storage, an active multi-stage miscible CO 2 flooding project in the Permian Basin (North Ward Estes Field, near Wickett, Texas) was investigated. In addition, two major natural CO 2 reservoirs in the southeastern Paradox Basin (McElmo Dome and Doe Canyon) were also investigated as they provide CO 2 for EOR operations in the Permian Basin. Produced gas and water were collected from three different CO 2 flooding phases (with different start dates) within the North Ward Estes Field to evaluate possible CO 2 storage mechanisms and amounts of total CO 2 retention. McElmo Dome and Doe Canyon were sampled for produced gas to determine the noble gas and stable isotope signature of the original injected EOR gas and to confirm the source of this naturally-occurring CO 2 . As expected, the natural CO 2 produced from McElmo Dome and Doe Canyon is a mix of mantle and crustal sources. When comparing CO 2 injection and production rates for the CO 2 floods in the North Ward Estes Field, it appears that CO 2 retention in the reservoir decreased over the course of the three injections, retaining 39%, 49% and 61% of the injected CO 2 for the 2008, 2010, and 2013 projects, respectively, characteristic of maturing CO 2 miscible flood projects. Noble gas isotopic composition of the injected and produced gas for the flood projects suggest no active fractionation, while δ 13 C CO 2 values suggest no active CO 2 dissolution into formation water, or mineralization. CO 2 volumes capable of dissolving in residual formation fluids were also estimated along with the potential to store pure-phase supercritical CO 2 . Using a combination of dissolution trapping and residual trapping, both volumes of CO 2 currently retained in the 2008 and 2013 projects could be justified, suggesting no major leakage is occurring. These subsurface reservoirs, jointly considered, have the capacity to store up to 9 years of CO 2 emissions from an average US powerplant.

International Journal of Greenhouse Gas Control↗

Migration timing and tributary use of spawning flannelmouth sucker (Catostomus latipinnis)

Spawning phenology and associated migrations of fishes are often regulated by factors such as temperature and stream discharge, but flow regulation of mainstem rivers coupled with climate change might disrupt these cues and affect fitness. Flannelmouth sucker (Catostomus latipinnis) persisting in heavily modified river networks are known to spawn in tributaries that might provide better spawning habitat than neighboring mainstem rivers subject to habitat degradation (e.g., embedded sediments, altered thermal regimes, and disconnected floodplains). Passive integrative transponder (PIT) tag data and radio telemetry were used to quantify the timing and duration of flannelmouth sucker tributary spawning migrations in relation to environmental cues in McElmo Creek, a tributary to the San Juan River in the American Southwest. We also tested the extent of the tributary migration and assessed mainstem movements prior to and following tributary migrations. Additionally, multi-year datasets of PIT detections from other tributaries in the Colorado River basin were used to quantify interannual and cross-site variation in the timing of flannelmouth sucker spawning migrations in relation to environmental cues. The arrival and residence times of fish spawning in McElmo Creek varied among years with earlier migration and a three-week increase in residence time in relatively wet years compared to drier years. Classification tree analysis suggested a combination of discharge and temperature determined arrival timing. Of fish PIT tagged in the fall, 56% tagged within 10 km of McElmo Creek spawned in the tributary the following spring, as did 60% of radio-tagged fish, with a decline in its use corresponding to increased distance of tagging location. A broader analysis of four tributaries in the Colorado River basin, including McElmo Creek, found photoperiod and temperature of tributary and mainstem rivers were the most important variables in determining migration timing, but tributary and mainstem discharge also aided in classification success. The largest tributary, the Little Colorado River, had more residential fish or fish that stayed for longer periods (median = 30 days), while McElmo Creek fish stayed an average of just 10 days in 2022. Our results generally suggest that higher discharge, across years or across sites, results in extended use of tributaries by flannelmouth suckers. Conservation actions that limit water extraction and maintain natural flow regimes in tributaries, while maintaining open connection with mainstem rivers may benefit migratory species including flannelmouth suckers.

Journal of Fish Biology↗

Conservation implications of spatiotemporal variation in the terrestrial ecology of Western spadefoots

Conservation of species reliant on ephemeral resources can be especially challenging in the face of a changing climate. Western spadefoots ( Spea hammondii ) are small burrowing anurans that breed in ephemeral pools, but adults spend the majority of their lives underground in adjacent terrestrial habitat. Western spadefoots are of conservation concern throughout their range because of habitat loss, but little is known about the activity patterns and ecology of their terrestrial life stage. We conducted a radio-telemetry study of adult western spadefoots at 2 sites in southern California, USA, from December 2018 to November 2019 to characterize their survival, behavior, and movements from breeding through aestivation to refine conservation and management for the species. Western spadefoot survival varied seasonally, with risk of mortality higher in the active season than during aestivation. The probability of movement between successive observations was higher during the winter and spring and when atmospheric moisture was high and soil water content at 10-cm depth was low. The amount of rain between observations had the strongest effect on the probability of movement between observations; for every 20 mm of rainfall between observations, western spadefoots were 2.4 times more likely to move. When movements occurred, movement rates were highest when both relative humidity and soil water content at 10-cm depth were high. The conditions under which western spadefoots were likely active on the surface, likely to have moved, and moved at the highest rates are conditions that reduce the risk of desiccation of surface-active spadefoots. Western spadefoot home range areas varied between study sites and were mostly <1 ha, although 1 individual's home range area was >6 ha. Western spadefoots rapidly dispersed from the breeding pools, and asymptotic distances from the breeding pool were generally reached by June. The asymptotic distance from the breeding pool varied between sites, with the 95th percentile of the posterior predictive distribution reaching 486 m at 1 site and 187 m at the other. Western spadefoots did not select most habitat components disproportionately to their availability, but at Crystal Cove State Park, they avoided most evaluated vegetation types (graminoids, forbs, and shrubs). Spatial variation was evident in most evaluated western spadefoot behaviors; context-dependent behavior suggests that site-specific management is likely necessary for western spadefoots. Furthermore, comparison with an earlier study of western spadefoots at Crystal Cove State Park indicated substantial temporal variation in western spadefoot behavior. Therefore, basing management decisions on short-term studies might fail to meet conservation objectives. Better understanding the influences of spatial context and climatic variation on western spadefoot behavior will improve conservation efforts for this species.

California↗

Upper Rio Grande Basin water-resource status and trends: Focus area study review and synthesis

The Upper Rio Grande Basin (URGB) is a critical international water resource under pressure from a myriad of climatic, ecological, infrastructural, water-use, and legal constraints. The objective of this study is to provide a comprehensive assessment of the spatial distribution and temporal trends of selected water-budget components (snow processes, evapotranspiration (ET), streamflow processes, and groundwater storage) using integrated analyses, such as watershed modeling and water availability and use data in the URGB over the past three decades. A spatially distributed snow evolution modeling system simulated snowpack processes over 34 years (1984–2017). It highlighted snow water equivalent declines from -35 to -77 mm/decade with widespread variability across elevation zones and land cover types. Gridded actual ET data from the SSEBop model were developed and tested for the URGB and demonstrated that all land-cover types had significant decreasing trends (1986-2015) ranging from -14 to -80 mm/decade. Conductivity-mass-balance (CMB) hydrograph separation results found that baseflow forms a large component of total streamflow, ranging from 29 to 69% (49% average) of total streamflow at 17 URGB sites upstream of Albuquerque, NM. Three of 4 graphical hydrograph separation methods in the U.S. Geological Survey Groundwater Toolbox were found to be inappropriate for estimating baseflow in the URGB; the most promising method, baseflow index (BFI) Standard, was optimized using CMB data and tested at three URGB sites, with resulting overestimation of 0 to 47%. Simulated changes in groundwater storage were extracted from historical and recent groundwater-flow models of select alluvial basins (San Luis, Española, Middle Rio Grande, and Tularosa-Hueco). In general, decreases in groundwater storage were observed from 1903 to 2013 except for the San Luis alluvial basin (Colorado), where periods of recovery are observed. The PRMS hydrologic model was successfully calibrated for 9 near-native subbasins (Nash-Sutcliffe efficiency 0.47 to 0.85) and parameters translated to the remaining subbasins; compared to simulated near-native flows (with minimal influence of reservoirs or diversions), observed Rio Grande streamgage flows demonstrated reductions of 40% or more for New Mexico and Texas areas of the basin. Significant decreasing trends (1980-2015) in precipitation, snowmelt rate, streamflow, and baseflow were observed at many of the 12 streamgage basins studied, which suggests that the decreasing trends for actual ET may be related to overall decreasing water availability in the basin, with negative implications for agricultural production and groundwater abstraction. Water security concerns arise from our findings of higher fraction precipitation as rain, slower snowmelt rates leading to decreasing streamflow production, and an increasing fraction of baseflow, all of which will affect the timing and magnitude of water available for human needs in the basin.

Upper Rio Grande Basin↗

The geology of selected peat-forming environments in temperate and tropical latitudes

We studied peat in several geologic and climatic settings: (1) a glaciated terrain in cold-temperate Maine and Minnesota, U.S.A.; (2) an island in a temperate maritime climate in the Atlantic Ocean off the coast of Maine, U.S.A., where sea level is rising rapidly and changing the environment of peat accumulation; (3) swamps along the warm-temperate U.S. Atlantic and Gulf Coastal Plains, where sea level has changed often, thus creating sites for accumulation; and (4) in a tropical climate along the coast of Sarawak, Malaysia, and the delta of the Batang Hari River, Sumatra, Indonesia (Figs. 1 and 2). With the exception of the deposits on the Atlantic and Gulf Coastal Plains, most of the deposits described are domed bogs in which peat accumulation continued above the surface of the surrounding soil. The bogs of the U.S. Atlantic and Gulf Coastal Plains have almost level surfaces. All domed bogs are not entirely ombrotrophic (watered only from precipitation); multidomed bogs that rise from irregular or hilly surfaces may be crossed by streams that supply water to the bogs. The geologic processes or organic sedimentation, namely terrestrialization and paludification, are similar in all peat deposits considered here. Differences in geomorphology affecting the quantity and that quality of peat that has ash contents of less than 25%, which are desirable for commercial purposes, depend chiefly on: (1) high humidity, which is favorable to luxuriant growth of peat-forming vegetation; (2) a depositional setting that permits extensive accumulation relatively free from inorganic contamination from sea water and streams and from dust and volcanic ash; and (3) a stable regional water table that controls the rate of decomposition under aerobic conditions and protects the deposit against the ravages of fire. Differences in peat textures are due to the type of vegetation and to the degree of decomposition. The rate of decomposition is largely the result of the amount of oxidation and aerobic microbial activity. Stratigraphic distribution of various textures and amounts of inorganic components within a peat deposit is largely determined by the vertical positions occupied by peat-forming environments, such as pond, marsh, swamp and heath where vegetation accumulated, and the depth to zones of unoxygenated water. Peat also differs in the rate of accumulation. On the basis of carbon-14 dating, an estimated 8 m of peat in the tropical Batang Hari River deposit in Sumatra has been accumulating at the rate of about 1.5 m/1,000 yr, whereas peat in the cold-temperate deposit in Maine has been accumulating at the rate of 0.66 m/1,000 yr. Accumulation rates in domed deposits such as these are affected not only by factors controlling volume of biomass and aerobic decay but also by stream erosion and fires that remove peat. Such disconformities (see Fig. 2) within the deposit may be recognized by sudden vertical changes in degree of decomposition and/or the presence of charcoal. The trace-element content of peat deposits is affected by the environments of their settings. Samples of peat that have an ash content of less than 25% dry weight and that are from small, almost level swamp deposits along the Atlantic Coastal Plain of North Carolina were compared with similar samples from small domed bogs in Maine, a glaciated area. Samples from Nort Carolina, which are from deposits in thick fluvial and nearshore marine sediments far from the bedrock source, are generally higher in Ti, Cr and Pb. The Maine samples from deposits in glacial drift close to the bedrock source contain more Zn, Mn, P, Ca, Na and Fe. The kind and amount of trace elements within the deposits appear to relate largely to depositional setting, to kinds of bedrock source, and to the modes of transportation from source to peat swamp. Trace-element concentrations in the extensive Sumatra peat deposit, which represents a potentially commercial coal bed, are similar to those found in Appalachian coals except for As and Au, which are higher in the former. This similarity most likely implies that geochemical controls on mineral matter controlled the concentrations in both. Most peat deposits found in the world today are not precursors of economic coal beds because they are too local in extent, lack beds of commercial-quality peat greater than 6 m thick, and/or are too far from sea level to be rapidly by marine or marginal marine sediments before destruction by erosion and decomposition. However, the two domed deposits in Sarawak and Sumatra described above are parts of extensive, thick, low-sulfur, fuel-quality peat deposits, which have bases below the levels of rivers on coastal deltas. These are likely to be preserved in their lower part and to become precursors of tropical coal seams of potential economic importance. The lenticular and tabular peat deposits of the Atlantic and Gulf Coastal Plains, which have little chance of survival, have features resembling those in Tertiary peat deposits in alluvial settings of western North America. The quality, shape, and extent of modern and ancient peat deposits are controlled by the same factors - humidity, vegetation type, ground- and surface-water regimes, and physiographic and geologic setting. Thus, a study of modern peat deposits may help in studies of the areal distribution and the thickness and quality of ancient coal beds.

International Journal of Coal Geology↗