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At least 793 records · Page 44Linked to original sources
USGS2006

Integrating human impacts and ecological integrity into a risk-based protocol for conservation planning

Conservation planning aims to protect biodiversity by sustainng the natural physical, chemical, and biological processes within representative ecosystems. Often data to measure these components are inadequate or unavailable. The impact of human activities on ecosystem processes complicates integrity assessments and might alter ecosystem organization at multiple spatial scales. Freshwater conservation targets, such as populations and communities, are influenced by both intrinsic aquatic properties and the surrounding landscape, and locally collected data might not accurately reflect potential impacts. We suggest that changes in five major biotic drivers—energy sources, physical habitat, flow regime, water quality, and biotic interactions—might be used as surrogates to inform conservation planners of the ecological integrity of freshwater ecosystems. Threats to freshwater systems might be evaluated based on their impact to these drivers to provide an overview of potential risk to conservation targets. We developed a risk-based protocol, the Ecological Risk Index (ERI), to identify watersheds with least/most risk to conservation targets. Our protocol combines risk-based components, specifically the frequency and severity of human-induced stressors, with biotic drivers and mappable land- and water-use data to provide a summary of relative risk to watersheds. We illustrate application of our protocol with a case study of the upper Tennessee River basin, USA. Differences in risk patterns among the major drainages in the basin reflect dominant land uses, such as mining and agriculture. A principal components analysis showed that localized, moderately severe threats accounted for most of the threat composition differences among our watersheds. We also found that the relative importance of threats is sensitive to the spatial grain of the analysis. Our case study demonstrates that the ERI is useful for evaluating the frequency and severity of ecosystemwide risk, which can inform local and regional conservation planning.

Tennessee↗
USGS2024

Food-web dynamics of a floodplain mosaic overshadow the effects of engineered logjams for Pacific salmon and steelhead

Food webs vary in space and time. The structure and spatial arrangement of food webs are theorized to mediate temporal dynamics of energy flow, but empirical corroboration in intermediate-scale landscapes is scarce. River-floodplain landscapes encompass a mosaic of aquatic habitat patches and food webs, supporting a variety of aquatic consumers of conservation concern. How the structure and productivity of these patch-scale food webs change through time, and how floodplain restoration influences their dynamics, are unevaluated. We measured productivity and food-web dynamics across a mosaic of main-channel and side-channel habitats of the Methow River, WA, USA, during two study years (2009–2010; 2015–2016) and examined how food webs that sustained juvenile anadromous salmonids responded to habitat manipulation. By quantifying temporal variation in secondary production and organic matter flow across nontreated river-floodplain habitats and comparing that variation to a side channel treated with engineered logjams, we jointly confronted spatial food-web theory and assessed whether food-web dynamics in the treated side channel exceeded natural variation exhibited in nontreated habitats. We observed that organic matter flow through the more complex, main-channel food web was similar between study years, whereas organic matter flow through the simpler, side-channel food webs changed up to ~4-fold. In the side channel treated with engineered logjams, production of benthic invertebrates and juvenile salmonids increased between study years by 2× and 4×, respectively; however, these changes did not surpass the temporal variation observed in untreated habitats. For instance, juvenile salmonid production rose 17-fold in one untreated side-channel habitat, and natural aggregation of large wood in another coincided with a shift to community and food-web dominance by juvenile salmonids. Our findings suggest that interannual dynamism in material flux across floodplain habitat mosaics is interrelated with patchiness in food-web complexity and may overshadow the ecological responses to localized river restoration. Although this dynamism may inhibit detection of the ecological effects of river restoration, it may also act to stabilize aquatic ecosystems and buffer salmon and other species of conservation concern in the long term. As such, natural, landscape-level patchiness and dynamism in food webs should be integrated into conceptual foundations of process-based, river restoration.

Washington↗
USGS2019

Conservation of temporary wetlands

Temporary wetlands are characterized by frequent drying resulting in a unique, highly specialized assemblage of often rare or specialized plant and animal species. They are found on all continents and in a variety of landscape settings. Although accurate estimates of the abundance of temporary wetlands are available in only a few countries, global estimations identify a decline in number and quality. The key environmental factors driving the structure of ecological communities in temporary wetlands are the duration, timing, frequency and predictability of the aquatic and dry phases, which varies greatly with region and hydrogeomorphic setting. Temporary wetlands have been historically neglected, but improved social awareness of the functions and values of, and increases in scientific interest, suggest that this is changing. They play an ecological role in both global cycles (i.e., CO 2 emissions) and biodiversity (in proportion to their size, they contribute disproportionately to regional and global biodiversity). Moreover, they provide valuable ecosystem services including wildlife habitat, nutrient flux to adjacent ecosystems, flood control, water filtration, and cultural services. Effective conservation of temporary wetlands requires addressing threats (i.e., inconsistent and inadequate regulatory protections; climate change; changes in land use) and management challenges (i.e., management at both local and landscape scales; incomplete understanding of the ecosystem services provided by them; the need to enhance inventories). The most suitable approaches for conserving temporary wetlands include (1) regulations or other forms of protection; (2) sustainable management; (3) restoration and creation; and (4) collaborative conservation.

Book chapter↗
USGS2010

Regional estimates of ecological services derived from U.S. Department of Agriculture conservation programs in the Mississippi Alluvial Valley

The Mississippi Alluvial Valley (MAV) is the Nation?s largest floodplain and this once predominantly forested ecosystem provided significant habitat for a diverse flora and fauna, sequestered carbon in trees and soil, and stored floodwater, sediments, and nutrients within the floodplain. This landscape has been substantially altered by the conversion of nearly 75% of the riparian forests, predominantly to agricultural cropland, with significant loss and degradation of important ecosystem services. Large-scale efforts have been employed to restore the forest and wetland resources and the U.S. Department of Agriculture (USDA) Wetlands Reserve Program (WRP) and Conservation Reserve Program (CRP) represent some of the most extensive restoration programs in the MAV. The objective of the WRP is to restore and protect the functions and values of wetlands in agricultural landscapes with an emphasis on habitat for migratory birds and wetland-dependent wildlife, protection and improvement of water quality, flood attenuation, ground water recharge, protection of native flora and fauna, and educational and scientific scholarship. The degree to which these conservation practices can restore ecosystem functions and services is not well known. This project was initiated to quantify existing ecological services derived from USDA conservation practices in the MAV as part of the USDA Conservation Effects Assessment Project, Wetlands Component (CEAP-Wetlands). The U.S. Geological Survey (USGS), in collaboration with the USDA Natural Resources Conservation Service, the USDA Farm Service Agency, the U.S. Fish and Wildlife Service, and Ducks Unlimited, collected data on soils, vegetation, nitrogen cycling, migratory birds, and amphibians from 88 different sites between 2006 and 2008. Results from restored WRP sites were compared to baseline data from active agricultural cropland (AG) to evaluate changes in ecosystem services.

Arkansas, Kentucky, Louisiana, Mississippi, Missou↗
USGS2013

Influence of landscape characteristics on retention of expandable radiocollars on young ungulates

One tool used for wildlife management is the deployment of radiocollars to gain knowledge of animal populations. Understanding the influence of individual factors (e.g., species, collar characteristics) and landscape characteristics (e.g., forested cover, shrubs, and fencing) on retention of expandable radiocollars for ungulates is important for obtaining empirical data on factors influencing ecology of young-of-the-year ungulates. During 2001–2009, we captured and radiocollared 198 white-tailed deer (Odocoileus virginianus) fawns, 142 pronghorn (Antilocapra americana) fawns, and 73 mule deer (O. hemionus) fawns in South Dakota, Minnesota, and California, USA. We documented 72 (36.4%), 8 (5.6%), and 7 (9.6%) premature (2, SE = 0.1, n = 75) compared with areas where fawns shed collars (x = 3.24 km/km 2 , SE = 0.1, n = 56) prior to 270 days. Researchers of fawns should consider that radiocollars can be shed prematurely when estimating desired sample size to yield a suitable strength of inference about some natural process of interest.

California;Minnesota;South Dakota↗
USGS2018

Integrating continuous stocks and flows into state-and-transition simulation models of landscape change

State-and-transition simulation models (STSMs) provide a general framework for forecasting landscape dynamics, including projections of both vegetation and land-use/land-cover (LULC) change. The STSM method divides a landscape into spatially-referenced cells and then simulates the state of each cell forward in time, as a discrete-time stochastic process using a Monte Carlo approach, in response to any number of possible transitions. A current limitation of the STSM method, however, is that all of the state variables must be discrete. Here we present a new approach for extending a STSM, in order to account for continuous state variables, called a state-and-transition simulation model with stocks and flows (STSM-SF). The STSM-SF method allows for any number of continuous stocks to be defined for every spatial cell in the STSM, along with a suite of continuous flows specifying the rates at which stock levels change over time. The change in the level of each stock is then simulated forward in time, for each spatial cell, as a discrete-time stochastic process. The method differs from the traditional systems dynamics approach to stock-flow modelling in that the stocks and flows can be spatially-explicit, and the flows can be expressed as a function of the STSM states and transitions. We demonstrate the STSM-SF method by integrating a spatially-explicit carbon (C) budget model with a STSM of LULC change for the state of Hawai'i, USA. In this example, continuous stocks are pools of terrestrial C, while the flows are the possible fluxes of C between these pools. Importantly, several of these C fluxes are triggered by corresponding LULC transitions in the STSM. Model outputs include changes in the spatial and temporal distribution of C pools and fluxes across the landscape in response to projected future changes in LULC over the next 50 years. The new STSM-SF method allows both discrete and continuous state variables to be integrated into a STSM, including interactions between them. With the addition of stocks and flows, STSMs provide a conceptually simple yet powerful approach for characterizing uncertainties in projections of a wide range of questions regarding landscape change.

Hawai'i↗
USGS2009

Local versus landscape-scale effects of savanna trees on grasses

1. Savanna ecosystems – defined by the coexistence of trees and grasses – cover more than one‐fifth the world’s land surface and harbour most of the world’s rangelands, livestock and large mammal diversity. Savanna trees can have a variety of effects on grasses, with consequences for the wild and domestic herbivores that depend on them. 2. Studies of these effects have focused on two different spatial scales. At the scale of individual trees, many studies have shown net positive effects of trees on sub‐canopy grass nutrient concentrations and biomass. At the landscape scale, other studies have shown negative effects of high tree densities on grass productivity. These disparate results have led to different conclusions about the effects of trees on forage quality and ungulate nutrition in savannas. 3. We integrate these approaches by examining the effects of trees on grasses at both spatial scales and across a range of landscape‐scale tree densities. We quantified grass biomass, species composition and nutrient concentrations in these different contexts in an Acacia drepanolobium savanna in Laikipia, Kenya. 4. Individual trees had positive effects on grass biomass, most likely because trees enrich soil nitrogen. Grass leaf phosphorus in sub‐canopy areas, however, was depressed. The effects of individual trees could explain the effects of increasing landscape‐scale tree cover for the biomass of only two of the four dominant grass species. 5. The negative effects of trees on grass and soil phosphorus, combined with depressed grass productivity in areas of high tree cover, suggest that ungulate nutrition may be compromised in areas with many trees. 6. Synthesis . We conclude that few, isolated trees may have positive local effects on savanna grasses and forage, but in areas of high tree density the negative landscape‐scale effects of trees are likely to outweigh these positive effects. In savannas and other patchy landscapes, attempts to predict the consequences of changes in patch abundances for ecosystem services (e.g. rangeland productivity and carbon sequestration) will depend on our understanding of the extent to which local, patch‐scale dynamics do or do not predict landscape‐scale dynamics.

Journal of Ecology↗
USGS2022

Geomorphology shapes relationships between animal communities and ecosystem function in large rivers

Understanding how the Earth's surface (i.e. ‘nature's stage') influences connections between biodiversity and ecosystem function (BEF) is a central objective in ecology. Despite recent calls to examine these connections at multiple trophic levels and at more complex and realistic scales, little is known about how landscape structure shapes BEF relationships among animal communities in nature. We coupled high-resolution habitat mapping with extensive field sampling to quantify connections among the geophysical habitat templet, invertebrate assemblages and secondary production in two large North American riverscapes. Patterns of sediment size governed invertebrate assemblage structure, with particularly strong effects on composition, richness and taxonomic and functional diversity. These relationships propagated to drive positive relationships between biodiversity and secondary production that were modified by scale, context-dependencies and anthropogenic modification. Finally, leveraging spatially-explicit descriptions of geophysical and biological properties, we uncovered distinct and nested spatial scales of biodiversity and secondary production, and suggest that multiple geophysical processes simultaneously influence these patterns at different scales. Together, our findings advance our understanding of relationships between the physical templet and patterns of BEF, and help to predict how perturbations to the Earth's surface may propagate to influence biodiversity and energy flux through food webs.

Oikos↗
USGS2016

Managing climate change refugia for climate adaptation

Refugia have long been studied from paleontological and biogeographical perspectives to understand how populations persisted during past periods of unfavorable climate. Recently, researchers have applied the idea to contemporary landscapes to identify climate change refugia, here defined as areas relatively buffered from contemporary climate change over time that enable persistence of valued physical, ecological, and socio-cultural resources. We differentiate historical and contemporary views, and characterize physical and ecological processes that create and maintain climate change refugia. We then delineate how refugia can fit into existing decision support frameworks for climate adaptation and describe seven steps for managing them. Finally, we identify challenges and opportunities for operationalizing the concept of climate change refugia. Managing climate change refugia can be an important option for conservation in the face of ongoing climate change.

PLoS ONE↗
USGS2018

Patch and landscape responses of bird abundance to fragmentation in agroecosystems of east-central Argentina

Forest fragmentation in agroecosystems is linked to declines of avian species worldwide. Agriculture has greatly reduced native forest cover in east-central Argentina. Assessing the influence of fragmentation on forest bird populations is vital to inform reliable conservation and management strategies for the Espinal region of Argentina. We determined the relationships of vegetation structure within native forest patches, size and shape of these patches (patch scale), composition and spatial configuration (at landscape scale) to relative abundance of 17 forest bird species during austral fall-winter and spring-summer seasons. Birds were sampled from 2007 – 2009 in 45 forest patches within three landscape mosaics (30x30 km) representing a gradient of native forest fragmentation in east-central Argentina. We used an information-theoretic approach and approximated model inference to examine the effect of predictor environmental variables at two spatial scales on patterns of forest bird abundance. Density of trees within forest patches was the main predictor of bird abundance at the patch scale. Amount of forest and spatial configuration were also important. The abundance of several bird species was greater in patches with high density of trees and landscapes characterized by more forest cover and larger patches in close proximity. We found two main avian response patterns to forest fragmentation and patch characteristics. This information contributes reliable knowledge for the design of conservation measures of agricultural landscapes in the Espinal region of Argentina.

Avian Conservation and Ecology↗
USGS2019

Bighorn sheep habitat and model extrapolation across remote landscapes

Determining a species’ habitat use is an essential first step in any wildlife conservation action. We described habitat use, animal movements and probable lambing areas in a remote, restricted-access region of the Mojave Desert. Differences in habitat use between sexes was apparent, supporting the often-reported concept of risk-aversion by females. Animals exhibited low variability in distances travelled, although males travelled further and with more variability than females. All females demonstrated what we interpret as lambing behavior during the same 2 ½ month periods over the two years, strongly supporting our inference of lambing sites. Water appeared critical to animal long-range movements, with no animal moving beyond 8.5 km from known sources. Modeling habitat use across the landscape of concern is another necessary step for conservation of species, allowing managers to plan for and predict the outcomes of management actions. In ecology, models are often created within relatively small areas, then extrapolated across larger regions of concern. The ability to extrapolate ecological models may be especially useful across remote areas, where consistent access by wildlife managers may be highly restricted. These restrictions on access require that most data be collected remotely, necessitating the need for extrapolating models developed in other areas. We used data from GPS-collared desert bighorn sheep to describe and model habitat use across the Pintwater Range, located on the Nevada Test and Training Range of southern Nevada, a highly restricted military training ground. We tested the efficacy of habitat model extrapolation by comparing the performance of two models derived from adjacent but independent desert bighorn sheep populations. The predictive power of seasonal habitat models derived from adjacent mountain ranges was lower than those derived from the local population. However, the performance of the extrapolated model suggests it could still be a feasible alternative for estimating general habitat use.

Nevada↗
USGS2021

Native American fire management at an ancient wildland–urban interface in the Southwest United States

The intersection of expanding human development and wildland landscapes—the “wildland–urban interface” or WUI—is one of the most vexing contexts for fire management because it involves complex interacting systems of people and nature. Here, we document the dynamism and stability of an ancient WUI that was apparently sustainable for more than 500 y. We combine ethnography, archaeology, paleoecology, and ecological modeling to infer intensive wood and fire use by Native American ancestors of Jemez Pueblo and the consequences on fire size, fire–climate relationships, and fire intensity. Initial settlement of northern New Mexico by Jemez farmers increased fire activity within an already dynamic landscape that experienced frequent fires. Wood harvesting for domestic fuel and architectural uses and abundant, small, patchy fires created a landscape that burned often but only rarely burned extensively. Depopulation of the forested landscape due to Spanish colonial impacts resulted in a rebound of fuels accompanied by the return of widely spreading, frequent surface fires. The sequence of more than 500 y of perennial small fires and wood collecting followed by frequent “free-range” wildland surface fires made the landscape resistant to extreme fire behavior, even when climate was conducive and surface fires were large. The ancient Jemez WUI offers an alternative model for fire management in modern WUI in the western United States, and possibly other settings where local management of woody fuels through use (domestic wood collecting) coupled with small prescribed fires may make these communities both self-reliant and more resilient to wildfire hazards.

Arizona, New Mexico↗
USGS2019

Conservation challenges emerging from free-roaming horse management: a vexing social-ecological mismatch

Horses have been associated with human societies for millennia, and for many have come to symbolize wildness, power, resilience, and freedom. Although equids were extirpated from North America 10 000-12 000 years ago, descendants of domestic horses now roam freely in the USA and 17 other countries across six continents. In landscape-scale and experimental investigations, free-roaming horses (Equus caballus) have been shown to induce numerous alterations to native-ecosystem components and processes through influences on soil, water, plants, native ungulates, small mammals, birds, reptiles, and other aspects of biodiversity. However, we argue that the management of free-roaming horses both in the U.S. and globally has been complicated by “socio-ecological mismatches.” These three mismatches stem from discrepancies between: 1) steady-state (horse-management) goals vs. a stochastically variable environment; 2) different methods to evaluate evidence, resolve conflicting beliefs, and achieve objectives, in scientific vs. political processes; and 3) differing spatial and temporal scales between policy and management, and values of science vs. values of the general public. Such mismatches arise from an inability to reconcile conflicting processes and functions in a social-ecological system, often reflecting differences in the spatio-temporal scales at which diverse components operate. Here, we describe three types of mismatches, and illustrate how the ecological dynamics of aridlands generally fit poorly with existing approaches to horse management and policy. Such mismatches complicate cost-effective management of free-roaming horses and the ecosystems they inhabit, and reduce the palette of potential solutions.

Conservation Letters↗
USGS2020

Integrating multiple data sources and multi-scale land-cover data to model the distribution of a declining amphibian

Determining the spatial scale at which landscape features influence population persistence is an important task for conservation planning. One challenge is that sampling biases confound factors that influence species occurrence and survey effort. Recent developments in Point Process Models (PPMs) enable researchers to disentangle the sampling process from ecological drivers of species' distributions. Land-cover change is a driver of decline for the western spadefoot (Spea hammondii), which has been extirpated from much of its range in California. Assessing this species' status requires information on the current distribution of suitable habitat within its historical range, but little is known about the effect of the landscape surrounding breeding ponds on spadefoot occurrence. Critically, surveys for western spadefoots often occur along roads, potentially biasing data used to fit species distribution models. We created PPMs integrating historical presence/non-detection and presence-only data for western spadefoots and land-cover data at multiple spatial scales to model the distribution of this species while removing the influence of sampling bias. There was spatial sampling bias in presence-only data; records were more likely to be reported near roads and urban centers and PPMs that removed sampling bias outperformed models that ignored sampling bias. The occurrence of western spadefoots was positively related to the proportion of grassland within a 2000 m buffer. The remaining habitat for western spadefoots is largely found in the foothills surrounding California's Central Valley. Our study illustrates how PPMs can improve projections of habitat suitability and our understanding of the drivers of species' distributions.

California↗
USGS2022

Modeling functional flows in California rivers

Environmental flows are critical to the recovery and conservation of freshwater ecosystems worldwide. However, estimating desired ranges of environmental flows across large, diverse landscapes is challenging. To advance protections of environmental flows for streams in California, USA, we developed a statewide modeling approach focused on functional components of the natural flow regime. Functional flow components in California streams—fall pulse flows, wet season peak flows and base flows, the spring flow recession, and dry season baseflows—support essential physical and ecological processes in riverine ecosystems. These functional flow components can be represented by functional flow metrics (FFMs) and quantified by their magnitude, timing, frequency, duration, and rate-of-change from daily streamflow records. After quantifying FFMs at reference-quality streamflow gages in California, we used machine-learning methods to estimate their natural range of values for all stream reaches in the state based on physical watershed characteristics and climatic factors. We found that the models performed well in predicting FFMs in streams across a diversity of landscape and climate contexts, according to several model performance criteria. Using the predicted FFM values, we established initial estimates of ecological flows that are expected to support critical functions and are broadly protective of ecosystem health. Modeling functional flows statewide offers a pathway for increasing the pace and scale of environmental flow protections in California and beyond.

California↗
USGS2022

Functional wetland loss drives emerging risks to waterbird migration networks

Migratory waterbirds (i.e., shorebirds, wading birds, and waterfowl) rely on a diffuse continental network of wetland habitats to support annual life cycle needs. Emerging threats of climate and land-use change raise new concerns over the sustainability of these habitat networks as water scarcity triggers cascading ecological effects impacting wetland habitat availability. Here we use important waterbird regions in Oregon and California, United States, as a model system to examine patterns of landscape change impacting wetland habitat networks in western North America. Wetland hydrology and flooded agricultural habitats were monitored monthly from 1988 to 2020 using satellite imagery to quantify the timing and duration of inundation—a key delimiter of habitat niche values associated with waterbird use. Trends were binned by management practice and wetland hydroperiods (semi-permanent, seasonal, and temporary) to identify differences in their climate and land-use change sensitivity. Wetland results were assessed using 33 waterbird species to detect non-linear effects of network change across a diversity of life cycle and habitat needs. Pervasive loss of semi-permanent wetlands was an indicator of systemic functional decline. Shortened hydroperiods caused by excessive drying transitioned semi-permanent wetlands to seasonal and temporary hydrologies—a process that in part counterbalanced concurrent seasonal and temporary wetland losses. Expansion of seasonal and temporary wetlands associated with closed-basin lakes offset wetland declines on other public and private lands, including wildlife refuges. Diving ducks, black terns, and grebes exhibited the most significant risk of habitat decline due to semi-permanent wetland loss that overlapped important migration, breeding, molting, and wintering periods. Shorebirds and dabbling ducks were beneficiaries of stable agricultural practices and top-down processes of functional wetland declines that operated collectively to maintain habitat needs. Outcomes from this work provide a novel perspective of wetland ecosystem change affecting waterbirds and their migration networks. Understanding the complexity of these relationships will become increasingly important as water scarcity continues to restructure the timing and availability of wetland resources.

California, Nevada, Oregon↗
USGS2022

Broad-scale geographic and temporal assessment of northern long-eared bat (Myotis septentrionalis) maternity colony-landscape association

As the federally threatened northern long-eared bat Myotis septentrionalis continues to decline due to white-nose syndrome (WNS) impacts, the application of effective conservation measures is needed but often hindered by the lack of ecological data. To date, recommended management practices have been adopted in part from other federally listed sympatric species such as the endangered Indiana bat M. sodalis . During the maternity season, these measures have largely focused on conservation of known day-roost habitat, often with little consideration for foraging habitat, particularly riparian areas. We examined acoustic activity of northern long-eared bats relative to day-roost and capture data at coastal and interior sites in the District of Columbia, New York, Pennsylvania, Virginia, and West Virginia, USA, over the course of 6 summers (2015-2020), where maternity activity was still documented after the initial arrival and spread of WNS. Acoustic activity of northern long-eared bats relative to forest cover decreased at the acoustic site level (fine scale) but increased at the sampling region level (coarse scale). We observed a positive association of northern long-eared bat acoustic activity with riparian areas. Additionally, we observed higher levels of activity during pregnancy through early lactation period of the reproductive cycle prior to juvenile volancy. Our findings suggest the need for more explicit inclusion of forested riparian habitats in northern long-eared bat conservation planning. Acoustic sampling in spring and early summer rather than mid- to late summer and in forested riparian areas is the most effective strategy for identifying potential active northern long-eared bat maternity colonies on the local landscape.

Maryland, New York, Pennsylvania, Virginia, and We↗
USGS2003

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↗