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Vegetation response to early holocene warming as an analog for current and future changes: Special section

Temperatures in southwestern North America are projected to increase 3.5-4 ??C over the next 60-90 years. This will precipitate ecological shifts as the ranges of species change in response to new climates. During this shift, rapid-colonizing species should increase, whereas slow-colonizing species will at first decrease, but eventually become reestablished in their new range. This successional process has been estimated to require from 100 to over 300 years in small areas, under a stable climate, with a nearby seed source. How much longer will it require on a continental scale, under a changing climate, without a nearby seed source? I considered this question through an examination of the response of fossil plant assemblages from the Grand Canyon, Arizona, to the most recent rapid warming of similar magnitude that occurred at the start of the Holocene, 11,700 years ago. At that time, temperatures in southwestern North America increased about 4 ??C over less than a century. Grand Canyon plant species responded at different rates to this warming climate. Early-successional species rapidly increased, whereas late-successional species decreased. This shift persisted throughout the next 2700 years. I found two earlier, less-extreme species shifts following rapid warming events around 14,700 and 16,800 years ago. Late-successional species predominated only after 4000 years or more of relatively stable temperature. These results suggest the potential magnitude, duration, and nature of future ecological changes and have implications for conservation plans, especially those incorporating equilibrium assumptions or reconstituting past conditions. When these concepts are extended to include the most rapid early-successional colonizers, they imply that the recent increases in invasive exotics may be only the most noticeable part of a new resurgence of early-successional vegetation. Additionally, my results challenge the reliability of models of future vegetation and carbon balance that project conditions on the basis of assumptions of equilibrium within only a century. ?? 2009 Society for Conservation Biology.

Conservation Biology

Investigating the Environmental Effects of Agriculture Practices on Natural Resources: Scientific Contributions of the U.S. Geological Survey to Enhance the Management of Agricultural Landscapes

The U.S. Geological Survey (USGS) enhances and protects the quality of life in the United States by advancing scientific knowledge to facilitate effective management of hydrologic, biologic, and geologic resources. Results of selected USGS research and monitoring projects in agricultural landscapes are presented in this Fact Sheet. Significant environmental and social issues associated with agricultural production include changes in the hydrologic cycle; introduction of toxic chemicals, nutrients, and pathogens; reduction and alteration of wildlife habitats; and invasive species. Understanding environmental consequences of agricultural production is critical to minimize unintended environmental consequences. The preservation and enhancement of our natural resources can be achieved by measuring the success of improved management practices and by adjusting conservation policies as needed to ensure long-term protection.

Fact Sheet

Benthic habitats and offshore geological resources of Kaloko-Honokohau National Historical Park, Hawai‘i

Kaloko-Honokōhau National Historical Park (KAHO) is one of three National Park lands located along the western coast of the Island of Hawai‘i and the only one to include submerged lands and marine resources within its official boundaries. The park was established in 1978 and is 1,160 acres in size, including 596 acres of marine area. The submerged lands are currently managed by the State of Hawaii, Department of Land and Natural Resources, Division of Aquatic Resources (DLNR-DAR). Marine resources located within KAHO include coral reef and habitat for many marine animals, including the green sea turtle and a variety of fish and invertebrates. In addition, many archeological, cultural, and recreational resources are located within the marine realm of the park. Potential threats and stressors to the modern marine environment include ground-water and surface-water contamination, invasive plants and algae, fishing pressure and use of monofilament gill nets (which can ensnare marine life or become tangled on reefs and be left behind as fishing debris), and visitor use impacts, such as scuba diving and snorkeling. Illegal dumping, oil releases, boat groundings, and other physical damage to reef resources are potential threats from users of the nearby harbor. Special issues of concern for the park include establishing baseline conditions of the offshore resources before the development of adjacent coastal lands. Until this study, only a general knowledge of the distribution of benthic habitats and the characteristics of the offshore region of Kaloko-Honokōhau National Historical Park was available. In 2003, a collaborative project between the U.S. Geological Survey (USGS) Coastal and Marine Geology Program and the National Park Service (NPS) was initiated to develop detailed benthic-habitat classification maps for the marine lands within and adjacent to the park. The intent of this project is to provide baseline maps and a Geographic Information System (GIS) database and description of the biological and geological resources of these marine lands in order to facilitate the management, interpretation, and understanding of park resources. A benthic-habitat classification map was created for the park using existing color aerial photography, Scanning Hydrographic Operational Airborne Lidar Survey (SHOALS) bathymetric data, georeferenced underwater video, and still photography. Individual habitat polygons were classified using five basic attributes: (1) major structure or substrate, (2) dominant structure, (3) major biologic cover on the substrate, (4) percentage of major biological cover, and (5) geographic zone. Additional information regarding geology, morphology, and coral species were also noted. Marine resources located within KAHO include coral reef and habitat for many marine animals, including the green sea turtle and a variety of fish and invertebrates. In addition, many archeological, cultural, and recreational resources are located within the marine realm of the park. Potential threats and stressors to the modern marine environment include ground-water and surface-water contamination, invasive plants and algae, fishing pressure and use of monofilament gill nets (which can ensnare marine life or become tangled on reefs and be left behind as fishing debris), and visitor use impacts, such as scuba diving and snorkeling. Illegal dumping, oil releases, boat groundings, and other physical damage to reef resources are potential threats from users of the nearby harbor. Special issues of concern for the park include establishing baseline conditions of the offshore resources before the development of adjacent coastal lands. Until this study, only a general knowledge of the distribution of benthic habitats and the characteristics of the offshore region of Kaloko-Honokōhau National Historical Park was available. In 2003, a collaborative project between the U.S. Geological Survey (USGS) Coastal and Marine Geology Program and the National Park Service (NPS) was initiated to develop detailed benthic-habitat classification maps for the marine lands within and adjacent to the park. The intent of this project is to provide baseline maps and a Geographic Information System (GIS) database and description of the biological and geological resources of these marine lands in order to facilitate the management, interpretation, and understanding of park resources. A benthic-habitat classification map was created for the park using existing color aerial photography, Scanning Hydrographic Operational Airborne Lidar Survey (SHOALS) bathymetric data, georeferenced underwater video, and still photography. Individual habitat polygons were classified using five basic attributes: (1) major structure or substrate, (2) dominant structure, (3) major biologic cover on the substrate, (4) percentage of major biological cover, and (5) geographic zone. Additional information regarding geology, morphology, and coral species were also noted.

Hawai'i

Muted responses to chronic experimental nitrogen deposition on the Colorado Plateau

Anthropogenic nitrogen (N) deposition is significantly altering both community structure and ecosystem processes in terrestrial ecosystems across the globe. However, our understanding of the consequences of N deposition in dryland systems remains relatively poor, despite evidence that drylands may be particularly vulnerable to increasing N inputs. In this study, we investigated the influence of 7 years of multiple levels of simulated N deposition (0, 2, 5, and 8 kg N ha −1 year −1 ) on plant community structure and biological soil crust (biocrust) cover at three semi-arid grassland sites spanning a soil texture gradient. Biocrusts are a surface community of mosses, lichens, cyanobacteria, and/or algae, and have been shown to be sensitive to N inputs. We hypothesized that N additions would decrease plant diversity, increase abundance of the invasive annual grass Bromus tectorum, and decrease biocrust cover. Contrary to our expectations, we found that N additions did not affect plant diversity or B. tectorum abundance. In partial support of our hypotheses, N additions negatively affected biocrust cover in some years, perhaps driven in part by inter-annual differences in precipitation. Soil inorganic N concentrations showed rapid but ephemeral responses to N additions and plant foliar N concentrations showed no response, indicating that the magnitude of plant and biocrust responses to N fertilization may be buffered by endogenous N cycling. More work is needed to determine N critical load thresholds for plant community and biocrust dynamics in semi-arid systems and the factors that determine the fate of N inputs.

Arizona, Colorado, New Mexico, Utah

Introduced species as evolutionary traps

Invasive species can alter environments in such a way that normal behavioural decision-making rules of native species are no longer adaptive. The evolutionary trap concept provides a useful framework for predicting and managing the impact of harmful invasive species. We discuss how native species can respond to changes in their selective regime via evolution or learning. We also propose novel management strategies to promote the long-term co-existence of native and introduced species in cases where the eradication of the latter is either economically or biologically unrealistic.

Ecology Letters

Efficacy of oblique bubble screen deterrents on redirecting live eggs and larvae of grass carp (Ctenopharyngodon idella) at different developmental stages

Invasive carp have severely damaged aquatic ecosystems in the USA, particularly in the Mississippi River Basin. Behavioral deterrents have been developed in the last few decades to control population expansion into new ecosystems. However, none of these deterrents are capable of controlling early-life stage carp, which have limited or no mobility during their drifting stage in rivers. Capturing eggs and larvae in large numbers warrants new methods due to their distinct physical and biological properties as well as their lack of behavioral response. We tested a novel method to redirect downstream drifting eggs and larvae for their efficient removal in streams using an oblique bubble screen (OBS) deterrent. We investigated the effects of mean water velocity and airflow rate on redirection of eggs, pre-gas bladder inflation (GBI) larvae, near-GBI larvae, and dead larvae. Although similar OBS configurations had shown high efficacy redirecting plastic spheres (egg surrogates) in previous studies, they underperform redirecting live eggs and larvae. However, distinct patterns were identified for eggs, pre-GBI, near-GBI, and dead larvae. A detailed hydrodynamic analysis showed that eggs closely follow the larger scales of motions created by the OBS, and that larvae can actively respond to turbulence cues. This study yielded new insights into the movement of early-life stage grass carp in a turbulent flow with strong recirculation, and provided important data to improve the design of bubble screen dispersal barriers for invasive carp management and population control.

Water Resources Research

Status, ecology, and management of the invasive plant, Miconia calvescens DC (Melastomataceae) in the Hawaiian islands

Miconia calvescens (Melastomataceae), native to montane forests of the neotropics, has now invaded wet forests of both the Society and Hawaiian Islands. This tree, which grows up to 15 m tall, is potentially the most invasive and damaging weed of rainforests of Pacific islands. In moist conditions, it grows rapidly, tolerates shade, and produces abundant seed that is effectively dispersed by birds and accumulates in a large, persistent soil seed-bank. Introduced to the Hawaiian Islands in 1961, M. calvescens appears to threaten much of the biological diversity in native forests receiving 1800–2000 mm or more annual precipitation. Currently, M. calvescens is found on 4 Hawaiian islands— Hawaii, Maui, Oahu, and Kauai. Widespread awareness of this invader began in the early 1990s. Although biological control is being pursued, conventional control techniques (mechanical and chemical) to contain and eradicate it locally are underway.

Hawaii

An adaptive approach to invasive plant management on U.S. Fish and Wildlife Service-owned native prairies in the Prairie Pothole Region: decision support under uncertainity

Much of the native prairie managed by the U.S. Fish and Wildlife Service (Service) in the Prairie Pothole Region (PPR) is extensively invaded by the introduced cool-season grasses smooth brome (Bromus inermis) and Kentucky bluegrass (Poa pratensis). The central challenge to managers is selecting appropriate management actions in the face of biological and environmental uncertainties. We describe the technical components of a USGS management project, and explain how the components integrate and inform each other, how data feedback from individual cooperators serves to reduce uncertainty across the whole region, and how a successful adaptive management project is coordinated and maintained on a large scale. In partnership with the Service, the U.S. Geological Survey is developing an adaptive decision support framework to assist managers in selecting management actions under uncertainty and maximizing learning from management outcomes. The framework is built around practical constraints faced by refuge managers and includes identification of the management objective and strategies, analysis of uncertainty and construction of competing decision models, monitoring, and mechanisms for model feedback and decision selection. Nineteen Service field stations, spanning four states of the PPR, are participating in the project. They share a common management objective, available management strategies, and biological uncertainties. While the scope is broad, the project interfaces with individual land managers who provide refuge-specific information and receive updated decision guidance that incorporates understanding gained from the collective experience of all cooperators.

Prairie Pothole Region

The Fort Collins Science Center

With a focus on biological research, the U.S. Geological Survey Fort Collins Science Center (FORT) develops and disseminates science-based information and tools to support natural resource decision-making. This brochure succinctly describes the integrated science capabilities, products, and services that the FORT science community offers across the disciplines of aquatic systems, ecosystem dynamics, information science, invasive species science, policy analysis and social science assistance, and trust species and habitats.

Colorado

Recent scientific contributions by the U.S. Geological Survey in the San Francisco Bay and Sacramento–San Joaquin Delta Estuary

Introduction The San Francisco Bay and Sacramento–San Joaquin Delta Estuary (hereafter, Bay-Delta) is the largest estuary on the West Coast of the United States. The Bay-Delta covers more than 1,600 square miles and drains a watershed of more than 75,000 square miles, which is greater than 40 percent of California. The region surrounding the Bay-Delta is home to about 10 million people, and its habitats (fig. 1) support more than 800 plant and animal species. The waterways of the Bay-Delta are the central hub of California’s extensive freshwater delivery system, supplying water to more than 27 million Californians and 4 million acres of farmland in the Central Valley. The U.S. Geological Survey (USGS) is the primary science agency of the U.S. Department of the Interior, providing a broad range of Earth, water, biological, and mapping data and expertise to inform natural resource management across the country. This fact sheet focuses on research conducted by the USGS in the Bay-Delta region, mostly within the past 5 years. The fact sheet is organized across five major themes: water flow and water quality, fish and wildlife, wetland restoration, invasive species, and hazards. In each of these areas, the USGS works closely with Federal, State, and local agencies; academia; and non-governmental organizations to provide objective, evidence-based science. The data and knowledge gained from USGS research and monitoring are publicly available and are routinely used by agencies, including the U.S. Bureau of Reclamation, U.S. Army Corps of Engineers, U.S. Fish and Wildlife Service, California Department of Water Resources, California Department of Fish and Wildlife, California State Water Resources Control Board, Delta Stewardship Council, and many additional organizations and stakeholders to inform management of the Bay-Delta region and its resources.

California

Ecological tracers reveal resource convergence among prey fish species in a large lake ecosystem

1. We measured stable isotopes of carbon (δ 13 C) and nitrogen (δ 15 N) and fatty acid profiles in Lake Ontario alewife ( Alosa pseudoharengus ), rainbow smelt ( Osmerus mordax ), slimy sculpin ( Cottus cognatus ) and round goby ( Neogobius melanostomus ) collected from 1982 to 2008 to investigate how temporal variability in these ecological tracers can relate to ecosystem-level changes associated with the establishment of highly invasive dreissenid mussels. 2. Prey fish δ 15 N values remained relatively constant, with only slimy sculpin exhibiting a temporal increase in δ 15 N. In contrast, δ 13 C values for alewife, rainbow smelt and, especially, slimy sculpin became less negative over time and were consistent with the benthification of the Lake Ontario food web associated with dreissenids. 3. Principal components analysis revealed higher contributions of 14:0 and 16:1n-7 fatty acids and increasingly negative δ 13 C values in older samples in agreement with the greater historical importance of pelagic production for alewife, rainbow smelt and slimy sculpin. 4. Temporal declines in fatty acid unsaturation indices and Σn-3/Σn-6 ratios, and also increased 24:0/14:0 ratios for alewife, rainbow smelt and slimy sculpin, indicated the increasing importance of nearshore production pathways for more recently collected fish and resulted in values more similar to those for round goby. 5. These results indicate a temporal convergence of the food niche, whereas food partitioning has historically supported the coexistence of prey fish species in Lake Ontario. This convergence is consistent with changes in food-web processes associated with the invasion of dreissenid mussels.

Lake Ontario

Regeneration of native trees in the presence of invasive saltcedar in the Colorado River Delta, Mexico

Many riparian zones in the Sonoran Desert have been altered by elimination of the normal flood regime; such changes to the flow regime have contributed to the spread of saltcedar (Tamarix ramosissma Ledeb.), an exotic, salt-tolerant shrub. It has been proposed that reestablishment of a natural flow regime on these rivers might permit passive restoration of native trees, without the need for aggressive saltcedar clearing programs. We tested this proposition in the Colorado River delta in Mexico, which has received a series of large-volume water releases from U.S. dams over the past 20 years. We mapped the vegetation of the delta riparian corridor through ground and aerial surveys (1999–2002) and satellite imagery (1992–2002) and related vegetation changes to river flood flows and fire events. Although saltcedar is still the dominant plant in the delta, native cottonwood ( Populus fremontii S. Wats.) and willow (Salix gooddingii C. Ball) trees have regenerated multiple times because of frequent flood releases from U.S. dams since 1981. Tree populations are young and dynamic (ages 5–10 years). The primary cause of tree mortality between floods is fire. Biomass in the floodplain, as measured by the normalized difference vegetation index on satellite images, responds positively even to low-volume (but long-duration) flood events. Our results support the hypothesis that restoration of a pulse flood regime will regenerate native riparian vegetation despite the presence of a dominant invasive species, but fire management will be necessary to allow mature tree stands to develop.

Colorado River Delta

Fate and effects to the benthic community of a copper treatment to eradicate invasive mussels in a large western river, USA

Copper-based chemical treatments are commonly used to eradicate invasive mussels in small ponds and lakes, but their use in large rivers has been limited. In 2023, in response to a detection of invasive quagga mussels, a 10-km reach of the Snake River (Idaho, USA), was treated with an unprecedented 19,300 kg of chelated copper molluscicide to a target concentration of 1,000 µg/L for 10 days. We assessed the transport and fate of the copper and its exposure and effects on the nontarget benthic community downstream. Water samples were collected at seven locations throughout the treatment period, and sediment, periphyton, and benthic macroinvertebrates were collected pre- and posttreatment. Nearly half of the original mass of copper was removed from the water column via sedimentation, sorption to algae, or biological uptake within the 10-km treatment reach and the first 15 km downstream. Even so, dissolved copper concentrations exceeded the acute toxicity threshold at least as far as 28 km downstream for more than 2 weeks. Sediment copper increased by up to 8.3-fold, exceeding the consensus-based sediment quality threshold effect concentration at several sites. Effects on benthic macroinvertebrates varied by taxa. From 0–28 km downstream, invertebrate abundances decreased 52%–94%, with gastropods among the most affected. Of the unique taxa present at these sites pretreatment, 52%–64% were not found posttreatment but were replaced by other taxa, indicating a reorganization of the base of the food web. Additionally, from 0–15 km downstream, the percentage of individuals from tolerant taxa increased two to15-fold. Findings from this study can help watershed managers plan future invasive mussel responses while protecting culturally, economically, and ecologically important nontarget species in large rivers.

Idaho

Roads as conduits for exotic plant invasions in a semiarid landscape

Roads are believed to be a major contributing factor to the ongoing spread of exotic plants. We examined the effect of road improvement and environmental variables on exotic and native plant diversity in roadside verges and adjacent semiarid grassland, shrubland, and woodland communities of southern Utah (U.S.A.). We measured the cover of exotic and native species in roadside verges and both the richness and cover of exotic and native species in adjacent interior communities (50 m beyond the edge of the road cut) along 42 roads stratified by level of road improvement (paved, improved surface, graded, and four-wheel-drive track). In roadside verges along paved roads, the cover of Bromus tectorum was three times as great (27%) as in verges along four-wheel-drive tracks (9%). The cover of five common exotic forb species tended to be lower in verges along four-wheel-drive tracks than in verges along more improved roads. The richness and cover of exotic species were both more than 50% greater, and the richness of native species was 30% lower, at interior sites adjacent to paved roads than at those adjacent to four-wheel-drive tracks. In addition, environmental variables relating to dominant vegetation, disturbance, and topography were significantly correlated with exotic and native species richness and cover. Improved roads can act as conduits for the invasion of adjacent ecosystems by converting natural habitats to those highly vulnerable to invasion. However, variation in dominant vegetation, soil moisture, nutrient levels, soil depth, disturbance, and topography may render interior communities differentially susceptible to invasions originating from roadside verges. Plant communities that are both physically invasible (e.g., characterized by deep or fertile soils) and disturbed appear most vulnerable. Decision-makers considering whether to build, improve, and maintain roads should take into account the potential spread of exotic plants.

Conservation Biology

Climate change is advancing spring onset across the U.S. national park system

Many U.S. national parks are already at the extreme warm end of their historical temperature distributions. With rapidly warming conditions, park resource management will be enhanced by information on seasonality of climate that supports adjustments in the timing of activities such as treating invasive species, operating visitor facilities, and scheduling climate-related events (e.g., flower festivals and fall leaf-viewing). Seasonal changes in vegetation, such as pollen, seed, and fruit production, are important drivers of ecological processes in parks, and phenology has thus been identified as a key indicator for park monitoring. Phenology is also one of the most proximate biological responses to climate change. Here, we use estimates of start of spring based on climatically modeled dates of first leaf and first bloom derived from indicator plant species to evaluate the recent timing of spring onset (past 10–30 yr) in each U.S. natural resource park relative to its historical range of variability across the past 112 yr (1901–2012). Of the 276 high latitude to subtropical parks examined, spring is advancing in approximately three-quarters of parks (76%), and 53% of parks are experiencing “extreme” early springs that exceed 95% of historical conditions. Our results demonstrate how changes in climate seasonality are important for understanding ecological responses to climate change, and further how spatial variability in effects of climate change necessitates different approaches to management. We discuss how our results inform climate change adaptation challenges and opportunities facing parks, with implications for other protected areas, by exploring consequences for resource management and planning.

Ecosphere

Infection by the marine cestode Hepatoxylon trichiuri in returning Chinook salmon (Oncorhynchus tshawytscha) spawners in Patagonia: Implications for a novel fishery

Parasitological information may be useful for managing economically important fisheries by providing insights into population dynamics and health effects. The Chinook salmon ( Oncorhynchus tshawytscha ), an invasive species with high socio-ecological importance in South America, supports recreational and incipient artisanal fisheries. In Southern Chile, the recent approval of coastal fisheries for returning Chinook salmon in some regions highlights the importance of understanding salmon foraging, habitat use, and commercial value. However, such information is lacking. Parasites have been extensively used as cost-effective biological tags to identify fish stocks. We conducted the first parasitological examination of the marine tapeworm Hepatoxylon trichiuri larvae in returning adult Chinook salmon from the Cisnes River to assess its potential as a biological tag for returning Chinook salmon in Patagonia. Of 61 Chinook salmon from the Cisnes River, 52.46% were infected by H. trichiuri , with 1.65 ± 1.26 parasites per infected salmon (mean ± SD). Infection prevalence and intensity did not differ significantly between males and females. Infection intensity was higher in larger salmon but decreased with increasing salmon condition. Infection intensity was not significantly related to length, mass, or body condition. Infection prevalence and intensity of H. trichiuri varied spatially among Patagonian river-basin populations of Chinook salmon, thereby supporting the use of H. trichiuri as a biological tag. Our findings provide important parasitological metrics that may enhance future management practices by informing geographic variability in foraging locations of adult salmon and their commercial value for human consumption.

Cisnes River, Patagonia

Climate change adaptation for the US National Wildlife Refuge System

Since its establishment in 1903, the National Wildlife Refuge System (NWRS) has grown to 635 units and 37 Wetland Management Districts in the United States and its territories. These units provide the seasonal habitats necessary for migratory waterfowl and other species to complete their annual life cycles. Habitat conversion and fragmentation, invasive species, pollution, and competition for water have stressed refuges for decades, but the interaction of climate change with these stressors presents the most recent, pervasive, and complex conservation challenge to the NWRS. Geographic isolation and small unit size compound the challenges of climate change, but a combined emphasis on species that refuges were established to conserve and on maintaining biological integrity, diversity, and environmental health provides the NWRS with substantial latitude to respond. Individual symptoms of climate change can be addressed at the refuge level, but the strategic response requires system-wide planning. A dynamic vision of the NWRS in a changing climate, an explicit national strategic plan to implement that vision, and an assessment of representation, redundancy, size, and total number of units in relation to conservation targets are the first steps toward adaptation. This adaptation must begin immediately and be built on more closely integrated research and management. Rigorous projections of possible futures are required to facilitate adaptation to change. Furthermore, the effective conservation footprint of the NWRS must be increased through land acquisition, creative partnerships, and educational programs in order for the NWRS to meet its legal mandate to maintain the biological integrity, diversity, and environmental health of the system and the species and ecosystems that it supports.

Environmental Management

Ecological values of shallow-water habitats: Implications for the restoration of disturbed ecosystems

A presumed value of shallow-habitat enhanced pelagic productivity derives from the principle that in nutrient-rich aquatic systems phytoplankton growth rate is controlled by light availability, which varies inversely with habitat depth. We measured a set of biological indicators across the gradient of habitat depth within the Sacramento–San Joaquin River Delta (California) to test the hypothesis that plankton biomass, production, and pelagic energy flow also vary systematically with habitat depth. Results showed that phytoplankton biomass and production were only weakly related to phytoplankton growth rates whereas other processes (transport, consumption) were important controls. Distribution of the invasive clam Corbicula fluminea was patchy, and heavily colonized habitats all supported low phytoplankton biomass and production and functioned as food sinks. Surplus primary production in shallow, uncolonized habitats provided potential subsidies to neighboring recipient habitats. Zooplankton in deeper habitats, where grazing exceeded phytoplankton production, were likely supported by significant fluxes of phytoplankton biomass from connected donor habitats. Our results provide three important lessons for ecosystem science: (a) in the absence of process measurements, derived indices provide valuable information to improve our mechanistic understanding of ecosystem function and to benefit adaptive management strategies; (b) the benefits of some ecosystem functions are displaced by water movements, so the value of individual habitat types can only be revealed through a regional perspective that includes connectedness among habitats; and (c) invasive species can act as overriding controls of habitat function, adding to the uncertainty of management outcomes.

Ecosystems