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Environmental DNA (eDNA) analysis is increasingly used for biomonitoring and research of fish populations and communities by environmental resource managers and academic researchers. Although managers are much interested in expanding the use of eDNA as a survey technique, they are sceptical about both its utility (given that information is often limited to presence/absence of a species) and feasibility (given the need for proper laboratory facilities for sample processing). Nonetheless, under the right circumstances, eDNA analysis is cost-effective compared to many traditional aquatic survey methods and does not disturb habitat or harm the animals being surveyed. This article presents a case study in which eDNA analysis was successfully used to document the presence of a rare fish species in a waterway earmarked for restoration. The authors discuss the conditions that allowed this study to occur quickly and smoothly and speculate on how the goals of researchers and managers can be integrated for efficient and informative use of this tool.

We examined the extent to which supply-side, niche, and competition theories and concepts help explain a trematode community in which one species comprises 87% of the trematode individuals, and the remaining 15 species each have

Because natural ecosystems are complex, it is difficult to predict how their variability scales across space and levels of organization. The species-insurance hypothesis predicts that asynchronous dynamics among species should reduce variability when biomass is aggregated either from local species populations to local multispecies communities, or from metapopulations to metacommunities. Similarly, the spatial-insurance hypothesis predicts that asynchronous spatial dynamics among either local populations or local communities should stabilize metapopulation biomass and metacommunity biomass, respectively. In combination, both species and spatial insurance reduce variation in metacommunity biomass over time, yet these insurances are rarely considered together in natural systems. We partitioned the extent that species insurance and spatial insurance reduced the annual variation in macroalgal biomass in a southern California kelp forest. We quantified variability and synchrony at two levels of organization (population and community) and two spatial scales (local plots and region) and quantified the strength of species and spatial insurance by comparing observed variability and synchrony in aggregate biomass to null models of independent species or spatial dynamics based on cyclic-shift permutation. Spatial insurance was weak, presumably because large-scale oceanographic processes in the study region led to high spatial synchrony at both population- and community-level biomass. Species insurance was stronger due to asynchronous dynamics among the metapopulations of a few common species. In particular, a regional decline in the dominant understory kelp species Pterygophora californica was compensated for by the rise of three subdominant species. These compensatory dynamics were associated with positive values of the Pacific Decadal Oscillation, indicating that differential species tolerances to warmer temperature and nutrient-poor conditions may underlie species insurance in this system. Our results illustrate how species insurance can stabilize aggregate community properties in natural ecosystems where environmental conditions vary over broad spatial scales.

Local and global measurements of parasite prevalence and abundance are critical for understanding the dynamics that underlie the diversity, distribution, and evolution of infectious diseases. Here, we present a data set of gut helminths found in (1) raccoons throughout their range, based on primary literature from 1925-2017 and (2) raccoons in Santa Barbara County, California, USA surveyed from 2012 to 2015. The range-wide data set has 1,256 parasite entries from 217 literature sources across three continents and 32 states in the USA. This data set includes a list of all recorded raccoon gut helminths (n = 100) and their presence and prevalence in surveyed raccoon populations. The Santa Barbara data set includes gut helminth data from 182 raccoons from one Southern California County. In addition to the presence and abundance data for 13 parasite species, this data set includes measurements of 7,465 individual raccoon roundworms (Baylisascaris procyonis). For both range-wide and Santa Barbara data sets, we include information on parasite site of infection in host, sampling method, and sample size. We also provide geographic coordinates for infected raccoon populations (range-wide database) and individuals (Santa Barbara). In the associated metadata, we include sampling methods and summary figures for both the range-wide and Santa Barbara raccoon gut helminth records. There are no copyright or proprietary restrictions for research and/or teaching purposes. S. B. Weinstein and J. C. Van Wert contributed equally to this manuscript and are shared first authors.

Ecotones can increase free-living species richness, but little is known about how parasites respond to ecotones. Here, we use parasite communities in raccoons (Procyon lotor) to test the hypothesis that parasite communities can be divided into core and satellite species, each with fundamentally different responses to ecotones. We used published parasite surveys to classify parasites as common core or rare satellite species, and then surveyed raccoons in coastal California to examine how proximity to two aquatic ecotones altered parasite communities. Raccoons near ecotones had more satellite and fewer core parasite species. Specifically, the marine ecotone increased parasite diversity by adding satellite species to a persistent core community, whereas the freshwater ecotone shifted the community from core to satellite species without a net change in parasite richness. We hypothesize that increased parasite richness at the marine ecotone resulted from increased diet diversity, but that raccoons were sinks for some parasites. Increased exposure to rare parasites at ecotones has implications for wildlife health and provides insight into observed associations between ecotones and emerging disease.

Background: Every day, billions of people - especially those living in poverty - are exposed to infectious pathogens in the environment and are at risk of contracting 'environmentally mediated' infections: those with environmental reservoirs that affect disease persistence and control. The complex ecology of environmental pathogens creates a global health problem not easily solved with medical treatment alone. Methods: Here, we quantified the global disease burden caused by environmentally mediated infections and used a structural equation modeling approach to explore correlated factors at the global scale. Findings: We found that 80% of pathogen species known to infect humans are environmentally mediated, causing about 40% of today's burden of infectious disease (global loss of 130 million years of healthy life annually). More than 91% of environmentally mediated burden occurs in tropical countries, and the poorest countries carry the highest burdens across all latitudes. We found weak or absent effects of biodiversity or agricultural land use at the global scale. In contrast, the strongest proximate indicator of environmentally mediated infectious disease burden is rural poor livelihoods. Political stability and wealth are associated with improved sanitation, better health care, and lower proportions of rural poor people, indirectly resulting in lower burdens of environmentally mediated infections. Interpretation: The high and uneven burden of environmentally mediated infections highlights the need for innovative social and ecological interventions to complement biomedical advances in the pursuit of global health and sustainability goals. Funding Statement: SHS, IJJ, and GADL received support from the Stanford Institute for Innovation in Developing Economies Global Development and Poverty Initiative. SHS and GADL also received support from the Bill & Melinda Gates Foundation (OPP1114050), National Institutes of Health (NIH) grant #1R01TW010286, National Science Foundation Coupled Natural and Human Systems grant #1414102, and the National Institute for Mathematical and Biological Synthesis through the Working Group "Optimal Control of Neglected Tropical Diseases.” IJJ was also funded by National Science Foundation Graduate Research Fellowship #1656518. SHS, SRH, CLW, KDL, MBonds, and GADL were supported by a grant from the National Center for Ecological Analysis and Synthesis through the Science for Nature and People Partnership program. AJL was supported by the Davis Family E-IPER Fellowship at Stanford and the Stanford Interdisciplinary Graduate Fellowship from the Stanford Vice Provost for Graduate Education. NN was supported by the Stanford Bing Fellowship in Honor of Paul Ehrlich. CLW was supported by a Sloan Research Fellowship from the Alfred P. Sloan Foundation and by the Michigan Society of Fellows at the University of Michigan. AJP was supported by a Queensland Government Accelerate Postdoctoral Research Fellowship and the DARPA PREEMPT program Cooperative Agreement # D18AC00031. Declaration of Interests: The authors declare no competing interests.

WOS:000669987000010; Reef fishes are a treasured part of marine biodiversity, and also provide needed protein for many millions of people. Although most reef fishes might survive projected increases in ocean temperatures, corals are less tolerant. A few fish species strictly depend on corals for food and shelter, suggesting that coral extinctions could lead to some secondary fish extinctions. However, secondary extinctions could extend far beyond those few coral-dependent species. Furthermore, it is yet unknown how such fish declines might vary around the world. Current coral mass mortalities led us to ask how fish communities would respond to coral loss within and across oceans. We mapped 6964 coral-reef-fish species and 119 coral genera, and then regressed reef-fish species richness against coral generic richness at the 1 degrees scale (after controlling for biogeographic factors that drive species diversification). Consistent with small-scale studies, statistical extrapolations suggested that local fish richness across the globe would be around half its current value in a hypothetical world without coral, leading to more areas with low or intermediate fish species richness and fewer fish diversity hotspots.

"Food Web Complexity and Species Diversity" (Paine 1966) is the most-cited empirical article published in the American Naturalist. In short, Paine removed predatory sea stars (Pisaster ochraceus) from the rocky intertidal and watched the key prey species, mussels (Mytilus californianus), crowd out seven subordinate primary space-holding species. However, because these mussels are a foundational species, they provide three-dimensional habitat for over 300 associated species inhabiting the mussel beds; thus, removing sea stars significantly increases community-wide diversity. In any case, most ecologists cite Paine (1966) to support a statement that predators increase diversity by interfering with competition. Although detractors remained skeptical of top-down effects and keystone concepts, the paradigm that predation increases diversity spread. By 1991, "Food Web Complexity and Species Diversity" was considered a classic ecological paper, and after 50 years it continues to influence ecological theory and conservation biology.

Baylisascaris procyonis (raccoon roundworm) infection is common in raccoons and can cause devastating pathology in other animals, including humans. Limited information is available on the frequency of asymptomatic human infection. We tested 150 adults from California, USA, for B. procyonis antibodies; 11 were seropositive, suggesting that subclinical infection does occur.

Found throughout the tree of life and in every ecosystem, parasites are some of the most diverse, ecologically important animals on Earth-but in almost all cases, the least protected by wildlife or ecosystem conservation efforts. For decades, ecologists have been calling for research to understand parasites' important ecological role, and increasingly, to protect as many species from extinction as possible. However, most conservationists still work within priority systems for funding and effort that exclude or ignore parasites, or treat parasites as an obstacle to be overcome. Our working group identified 12 goals for the next decade that could advance parasite biodiversity conservation through an ambitious mix of research, advocacy, and management. Ecological Society of America; Georgetown Environment Initiative; Smithsonian National Museum of Natural HistorySmithsonian InstitutionSmithsonian National Museum of Natural History; Peter Buck Postdoctoral Fellowship from the Smithsonian National Museum of Natural History; National Science FoundationNational Science Foundation (NSF) [OCE-1829509]; Alfred P. Sloan Foundation Sloan Research FellowshipAlfred P. Sloan Foundation; University of Washington Innovation Award; University of Washington Royalty Research Fund awardUniversity of Washington The authors kindly thank the Ecological Society of America for supporting our workshop, as well as additional participants Kevin Burgio, Tad Dallas, and Roger Jovani; Laura Whitehouse, for her graphic design work on Fig. 1; Jonathan Wojcik for allowing the inclusion of his copyright Diplozoon illustration in Fig. 3; and dozens of collaborators and friends who have been part of the foundational work on parasite conservation, including Anna Phillips, Veronica Bueno, Carrie Cizauskas, Christopher Clements, Graeme Cumming, Eric Dougherty, Kevin Johnson, Wayne Getz, Nyeema Harris, Elizabeth Nichols, Sergey Mironov, Robert Poulin, and Heather Proctor. CJC gratefully acknowledges funding support from the Georgetown Environment Initiative, and research support from Anna Phillips and the Smithsonian National Museum of Natural History. KCB was supported by a Peter Buck Postdoctoral Fellowship from the Smithsonian National Museum of Natural History. CLW was supported by a grant from the National Science Foundation (OCE-1829509), an Alfred P. Sloan Foundation Sloan Research Fellowship, a University of Washington Innovation Award, and a University of Washington Royalty Research Fund award. Any use of trade, product, or firm names in this publication is for descriptive purposes only and does not imply endorsement by the U.S. Government. Public domain – authored by a U.S. government employee

To improve ability to detect white sharks without the need for tags, or visual census, we developed a species-specific environmental DNA (eDNA) assay that targets a 163 bp fragment of the white shark (Carcharodon carcharias) mitochondrial cytochrome B gene on a digital droplet PCR (ddPCR) platform. We used this marker to detect white shark DNA in 250 ml water samples taken from across two sites in Santa Barbara, California (USA) frequented by juvenile white sharks. We did not detect white shark DNA in samples from two neighboring sites where sharks are presumably absent, suggesting that eDNA can indicate nearby white sharks. This marker development, testing, and opportunistic application in a region with known distributions of white sharks indicates that environmental DNA could be developed further to monitor white sharks, thereby informing conservation planning and public safety. With the potential increase in white shark populations due to decades of protection, there is a need for fishery independent methods for assessing white shark distributions, and eDNA may provide be an ideal, non-intrusive tool for coastal assessments.

Two myxozoan species were observed in the kidney of topsmelt, Atherinops affinis , during a survey of parasites of estuarine fishes in the Carpinteria Salt Marsh Reserve, California. Fish collected on 3 dates in 2012 and 2013 were sectioned and examined histologically. Large extrasporogonic stages occurred in the renal interstitium of several fish from the first 2 collections (5/8, 11/20, respectively) and, in some fish, these replaced over 80% of the kidney. In addition, presporogonic and polysporogonic stages occurred in the lumen of the renal tubules, collecting ducts, and mesonephric ducts. The latter contained subspherical spores with up to 4 polar capsules, consistent with the genus Chloromyxum. For the third collection (15 May 2013, n = 30), we portioned kidneys for examination by histology, wet mount, and DNA extraction for small subunit ribosomal (SSU rDNA) gene sequencing. Histology showed the large extrasporogonic forms in the kidney interstitium of 3 fish and showed 2 other fish with subspherical myxospores in the lumen of the renal tubules with smooth valves and 2 spherical polar capsules consistent with the genus Sphaerospora. Chloromyxum-type myxospores were observed in the renal tubules of 1 fish by wet mount. Sequencing of the kidney tissue from this fish yielded a partial SSU rDNA sequence of 1,769 base pairs (bp). Phylogenetic reconstruction suggested this organism to be a novel species of Chloromyxum, most similar to Chloromyxum careni (84% similarity). In addition, subspherical myxospores with smooth valves and 2 spherical polar capsules consistent with the genus Sphaerospora were observed in wet mounts of 2 fish. Sequencing of the kidney tissue from 1 fish yielded a partial SSU rDNA sequence of 1,937 bp. Phylogenetic reconstruction suggests this organism to be a novel species of Sphaerospora most closely related to Sphaerospora epinepheli (93%). We conclude that these organisms represent novel species of the genera Chloromyxum and Sphaerospora based on host, location, and SSU rDNA sequence. We further conclude that the formation of large, histozoic extrasporogonic stages in the renal interstitium represents developmental stages of Chloromyxum species for the following reasons: (1) Large extrasporogonic stages were only observed in fish with Chloromyxum-type spores developing within the renal tubules, (2) a DNA sequence consistent with the Chloromyxum sp. was only detected in fish with the large extrasporogonic stages, and (3) several Sphaerospora species have extrasporogonic forms, but they are considerably smaller and are composed of far fewer cells.

A model for who eats and who is eaten There are many types of interactions between those that eat and those that are eaten. A multitude of theoretical equations describe these dynamics, from predator and prey to parasite and host. Lafferty et al. show that all forms of these relationships come down to fundamental consumer-resource interactions. Derived from the myriad complex interactions, a simple model can accommodate any such interaction, simplifying past models into a general theory of eat and be eaten. Science , this issue p. 854

Eliminating human parasitic disease often requires interrupting complex transmission pathways. Even when drugs to treat people are available, disease control can be difficult if the parasite can persist in nonhuman hosts. Here, we show that restoration of a natural predator of a parasite’s intermediate hosts may enhance drug-based schistosomiasis control. Our study site was the Senegal River Basin, where villagers suffered a massive outbreak and persistent epidemic after the 1986 completion of the Diama Dam. The dam blocked the annual migration of native river prawns (Macrobrachium vollenhoveni) that are voracious predators of the snail intermediate hosts for schistosomiasis. We tested schistosomiasis control by reintroduced river prawns in a before-after-control-impact field experiment that tracked parasitism in snails and people at two matched villages after prawns were stocked at one village’s river access point. The abundance of infected snails was 80% lower at that village, presumably because prawn predation reduced the abundance and average life span of latently infected snails. As expected from a reduction in infected snails, human schistosomiasis prevalence was 18 ± 5% lower and egg burden was 50 ± 8% lower at the prawn-stocking village compared with the control village. In a mathematical model of the system, stocking prawns, coupled with infrequent mass drug treatment, eliminates schistosomiasis from high-transmission sites. We conclude that restoring river prawns could be a novel contribution to controlling, or eliminating, schistosomiasis.

Human actions can affect wildlife and their nematode parasites. Species introductions and human-facilitated range expansions can create new host-parasite interactions. Novel hosts can introduce parasites and have the potential to both amplify and dilute nematode transmission. Furthermore, humans can alter existing nematode dynamics by changing host densities and the abiotic conditions that affect larval parasite survival. Human impacts on wildlife might impair parasites by reducing the abundance of their hosts; however, domestic animal production and complex life cycles can maintain transmission even when wildlife becomes rare. Although wildlife nematodes have many possible responses to human actions, understanding host and parasite natural history, and the mechanisms behind the changing disease dynamics might improve disease control in the few cases where nematode parasitism impacts wildlife.

Extrinsic environmental factors influence the distribution and population dynamics of many organisms, including insects that are of concern for human health and agriculture. This is particularly true for vector-borne infectious diseases, like malaria, which is a major source of morbidity and mortality in humans. Understanding the mechanistic links between environment and population processes for these diseases is key to predicting the consequences of climate change on transmission and for developing effective interventions. An important measure of the intensity of disease transmission is the reproductive number $R_0$. However, understanding the mechanisms linking $R_0$ and temperature, an environmental factor driving disease risk, can be challenging because the data available for parameterization are often poor. To address this we show how a Bayesian approach can help identify critical uncertainties in components of $R_0$ and how this uncertainty is propagated into the estimate of $R_0$. Most notably, we find that different parameters dominate the uncertainty at different temperature regimes: bite rate from 15-25$^\circ$ C; fecundity across all temperatures, but especially $\sim$25-32$^\circ$ C; mortality from 20-30$^\circ$ C; parasite development rate at $\sim$15-16$^\circ$C and again at $\sim$33-35$^\circ$C. Focusing empirical studies on these parameters and corresponding temperature ranges would be the most efficient way to improve estimates of $R_0$. While we focus on malaria, our methods apply to improving process-based models more generally, including epidemiological, physiological niche, and species distribution models., 27 pages, including 1 table and 3 figures

Foundation species define the ecosystems they live in, but ecologists have often characterized dominant plants as foundational without supporting evidence. Giant kelp has long been considered a marine foundation species due to its complex structure and high productivity; however, there is little quantitative evidence to evaluate this. Here, we apply structural equation modelling to a 15-year time series of reef community data to evaluate how giant kelp affects the reef community. Although species richness was positively associated with giant kelp biomass, most direct paths did not involve giant kelp. Instead, the foundational qualities of giant kelp were driven mostly by indirect effects attributed to its dominant physical structure and associated engineering influence on the ecosystem, rather than by its use as food by invertebrates and fishes. Giant kelp structure has indirect effects because it shades out understorey algae that compete with sessile invertebrates. When released from competition, sessile species in turn increase the diversity of mobile predators. Sea urchin grazing effects could have been misinterpreted as kelp effects, because sea urchins can overgraze giant kelp, understorey algae and sessile invertebrates alike. Our results confirm the high diversity and biomass associated with kelp forests, but highlight how species interactions and habitat attributes can be misconstrued as direct consequences of a foundation species like giant kelp.

Evolutionary transitions to parasitism are rare. In this study, we documented a potential step toward parasitism in the commensal clam Kurtiella pedroana (Bivalvia: Galeommatoidea). Galeommatoideans are known commensals of various invertebrates, including crustaceans. Emerita analoga (Decapoda: Hippidae) is an abundant intertidal mole crab inhabiting Pacific coast beaches in North and South America. Mole crabs collected from Monterey Bay, California, were measured and examined externally and internally for associated molluscs. Out of the 520 mole crabs, 37 large female individuals harbored 49 clams (prevalence of 7.11% and mean intensity of 1.3). Forty-one ectocommensal clams were attached by their byssal threads to the inside of the gill chambers or to the lateroventral surfaces. However, our key finding was 8 clams that lacked byssal threads and were living in the hemocoel of 6 crabs. These internal clams were smaller than the ectocommensals. Because these internal clams lacked access to their normal food, we hypothesize they might have fed on hemolymph as would a parasite. Clam larvae have no obvious exit from the hemocoel, implying that endoparasitism is a dead-end for K. pedroana. Regardless, facultative parasitism in a free-living or an ectocommensal is uncommon and suggests a pathway to parasitism.

Submitted manuscript version. Published version available at https://doi.org/10.1016/j.tree.2018.01.011. Species interactions can influence ecosystem functioning by enhancing or suppressing the activities of species that drive ecosystem processes, or by causing changes in biodiversity. However, one important class of species interactions – parasitism – has been little considered in biodiversity and ecosystem functioning (BD-EF) research. Parasites might increase or decrease ecosystem processes by reducing host abundance. Parasites could also increase trait diversity by suppressing dominant species or by increasing within-host trait diversity. These different mechanisms by which parasites might affect ecosystem function pose challenges in predicting their net effects. Nonetheless, given the ubiquity of parasites, we propose that parasite–host interactions should be incorporated into the BD-EF framework.

Most species aggregate in local patches. High host density in patches increases contact rate between hosts and parasites, increasing parasite transmission success. At the same time, for environmentally transmitted parasites, high host density can decrease infection risk to individual hosts, because infective stages are divided among all hosts in a patch, leading to safety in numbers. We tested these predictions using the California horn snail, Cerithideopsis californica (=Cerithidea californica), which is the first intermediate host for at least 19 digenean trematode species in California estuaries. Snails become infected by ingesting trematode eggs or through penetration by free-swimming miracidia that hatch from trematode eggs deposited with final-host (bird or mammal) feces. This complex life cycle decouples infective-stage production from transmission, raising the possibility of an inverse relationship between host density and infection risk at local scales. In a field survey, higher snail density was associated with increased trematode (infected snail) density, but decreased trematode prevalence, consistent with either safety in numbers, parasitic castration, or both. To determine the extent to which safety in numbers drove the negative snail-density-trematode-prevalence association, we manipulated uninfected snail density in 83 cages at eight sites within Carpinteria Salt Marsh (California, USA). At each site, we quantified snail density and used data on final-host (bird and raccoon) distributions to control for between-site variation in infective-stage supply. After three months, overall trematode infections per cage increased with snail biomass density. For egg-transmitted trematodes, per-snail infection risk decreased with snail biomass density in the cage and surrounding area, whereas per-snail infection risk did not decrease for miracidium-transmitted trematodes. Furthermore, both trematode recruitment and infection risk increased with infective-stage input, but this was significant only for miracidium-transmitted species. A model parameterized with our experimental results and snail densities from 524 field transects estimated that safety in numbers, when combined with patchy host density, halved per capita infection risk in this snail population. We conclude that, depending on transmission mode, host density can enhance parasite recruitment and reduce per capita infection risk.

To identify trematode diversity and life-cycles in the sub-Arctic Lake Takvatn, Norway, we characterised 120 trematode isolates from mollusc first intermediate hosts, metacercariae from second intermediate host fishes and invertebrates, and adults from fish and invertebrate definitive hosts, using molecular techniques. Phylogenies based on nuclear and/or mtDNA revealed high species richness (24 species or species-level genetic lineages) and uncovered trematode diversity (16 putative new species) from five families typical in lake ecosystems (Allocreadiidae, Diplostomidae, Plagiorchiidae, Schistosomatidae and Strigeidae). Sampling potential invertebrate hosts allowed matching of sequence data for different stages, thus achieving molecular elucidation of trematode life-cycles and exploration of host-parasite interactions. Phylogenetic analyses also helped identify three major mollusc intermediate hosts (Radix balthica, Pisidium casertanum and Sphaerium sp.) in the lake. Our findings increase the known trematode diversity at the sub-Arctic Lake Takvatn, showing that digenean diversity is high in this otherwise depauperate sub-Arctic freshwater ecosystem and indicating that sub-Arctic and Arctic ecosystems may be characterised by unique trematode assemblages.

Sapronotic disease agents have evolutionary and epidemiological properties unlike other infectious organisms. Their essential saprophagic existence prevents coevolution, and no host-parasite virulence trade-off can evolve. However, the host may evolve defenses. Models of pathogens show that sapronoses, lacking a threshold of transmission, cannot regulate host populations, although they can reduce host abundance and even extirpate their hosts. Immunocompromised hosts are relatively susceptible to sapronoses. Some particularly important sapronoses, such as cholera and anthrax, can sustain an epidemic in a host population. However, these microbes ultimately persist as saprophages. One-third of human infectious disease agents are sapronotic, including nearly all fungal diseases. Recognition that an infectious disease is sapronotic illuminates a need for effective environmental control strategies.

Mallards (Anas platyrhynchos) have been observed foraging on intertidal Pacific sand crabs (Hippidae, Emerita analoga) in the swash zone of sandy beaches around Coal Oil Point Reserve, California, and several other beaches on the west coast since at least November 2010. Unlike foraging shorebirds, Mallards do not avoid incoming swashes. Instead, the incoming swash lifts and deposits them down the beach. Shorebirds and diving ducks commonly feed on sand crabs, but sand crabs appear to be a novel behavior and food source for Mallards. Previous surveys of beaches did not report foraging Mallards on regional beaches, whereas foraging Mallards were common in contemporary (recent) surveys and anecdotal reports. Observations of this potentially new behavior were separated by as much as 1,300 km, indicating that this was not a local phenomenon. Mallards foraged singly, in pairs, and in flocks. An expansion of diet to sand crabs carries risks of exposure to surf, human disturbance, high salt intake, and t...

Managers are moving from a model of managing individual sectors, human activities, or ecosystem services to an ecosystem-based management (EBM) approach which attempts to balance the range of services provided by ecosystems. Applying EBM is often difficult due to inherent tradeoffs in managing for different services. This challenge particularly holds for estuarine systems, which have been heavily altered in most regions and are often subject to intense management interventions. Estuarine managers can often choose among a range of management tactics to enhance a particular service; although some management actions will result in strong tradeoffs, others may enhance multiple services simultaneously. Management of estuarine ecosystems could be improved by distinguishing between optimal management actions for enhancing multiple services and those that have severe tradeoffs. This requires a framework that evaluates tradeoff scenarios and identifies management actions likely to benefit multiple services. We created a management action-services matrix as a first step towards assessing tradeoffs and providing managers with a decision support tool. We found that management actions that restored or enhanced natural vegetation (e.g., salt marsh and mangroves) and some shellfish (particularly oysters and oyster reef habitat) benefited multiple services. In contrast, management actions such as desalination, salt pond creation, sand mining, and large container shipping had large net negative effects on several of the other services considered in the matrix. Our framework provides resource managers a simple way to inform EBM decisions and can also be used as a first step in more sophisticated approaches that model service delivery.

Most species interactions probably involve parasites. This review considers the extent to which marine ecologists should consider parasites to fully understand marine communities. Parasites are influential parts of food webs in estuaries, temperate reefs, and coral reefs, but their ecological importance is seldom recognized. Though difficult to observe, parasites can have substantial biomass, and they can be just as common as free-living consumers after controlling for body mass and trophic level. Parasites have direct impacts on the energetics of their hosts and some affect host behaviors, with ecosystem-level consequences. Although they cause disease, parasites are sensitive components of ecosystems. In particular, they suffer secondary extinctions due to biodiversity loss. Some parasites can also return to a system after habitat restoration. For these reasons, parasites can make good indicators of ecosystem integrity. Fishing can indirectly increase or decrease parasite populations and the effects of climate change on parasites are likely to be equally as complex.

Summary Parasites affect host behavior in several ways. They can alter activity, microhabitats or both. For trophically transmitted parasites (the focus of our study), decreased activity might impair the ability of hosts to respond to final-host predators, and increased activity and altered microhabitat choice might increase contact rates between hosts and final-host predators. In an analysis of trophically transmitted parasites, more parasite groups altered activity than altered microhabitat choice. Parasites that infected vertebrates were more likely to impair the host’s reaction to predators, whereas parasites that infected invertebrates were more likely to increase the host’s contact with predators. The site of infection might affect how parasites manipulate their hosts. For instance, parasites in the central nervous system seem particularly suited to manipulating host behavior. Manipulative parasites commonly occupy the body cavity, muscles and central nervous systems of their hosts. Acanthocephalans in the data set differed from other taxa in that they occurred exclusively in the body cavity of invertebrates. In addition, they were more likely to alter microhabitat choice than activity. Parasites in the body cavity (across parasite types) were more likely to be associated with increased host contact with predators. Parasites can manipulate the host through energetic drain, but most parasites use more sophisticated means. For instance, parasites target four physiological systems that shape behavior in both invertebrates and vertebrates: neural, endocrine, neuromodulatory and immunomodulatory. The interconnections between these systems make it difficult to isolate specific mechanisms of host behavioral manipulation.

Parasitologists have worked out many complex life cycles over the last ~150 yr, yet there have been few efforts to synthesize this information to facilitate comparisons among taxa. Most existing host-parasite databases focus on particular host taxa, do not distinguish final from intermediate hosts, and lack parasite life-history information. We summarized the known life cycles of trophically transmitted parasitic acanthocephalans, cestodes, and nematodes. For 973 parasite species, we gathered information from the literature on the hosts infected at each stage of the parasite life cycle (8,510 host-parasite species associations), what parasite stage is in each host, and whether parasites need to infect certain hosts to complete the life cycle. We also collected life-history data for these parasites at each life cycle stage, including 2,313 development time measurements and 7,660 body size measurements. The result is the most comprehensive data summary available for these parasite taxa. In addition to identifying gaps in our knowledge of parasite life cycles, these data can be used to test hypotheses about life cycle evolution, host specificity, parasite life-history strategies, and the roles of parasites in food webs.

Complex ecological networks appear robust to primary extinctions, possibly due to consumers' tendency to specialize on dependable (available and persistent) resources. However, modifications to the conditions under which the network has evolved might alter resource dependability. Here, we ask whether adaptation to historical conditions can increase community robustness, and whether such robustness can protect communities from collapse when conditions change. Using artificial life simulations, we first evolved digital consumer-resource networks that we subsequently subjected to rapid environmental change. We then investigated how empirical host–parasite networks would respond to historical, random and expected extinction sequences. In both the cases, networks were far more robust to historical conditions than new ones, suggesting that new environmental challenges, as expected under global change, might collapse otherwise robust natural ecosystems., Despite their complexity, ecological networks appear robust to species loss. Here, Strona and Lafferty use artificial life simulations and real-world data to show that such robustness applies to stable conditions, but can collapse when the environment changes.

Now-outdated estimates proposed that climate change should have increased the number of people at risk of malaria, yet malaria and several other infectious diseases have declined. Although some diseases have increased as the climate has warmed, evidence for widespread climate-driven disease expansion has not materialized, despite increased research attention. Biological responses to warming depend on the non-linear relationships between physiological performance and temperature, called the thermal response curve. This leads performance to rise and fall with temperature. Under climate change, host species and their associated parasites face extinction if they cannot either thermoregulate or adapt by shifting phenology or geographic range. Climate change might also affect disease transmission through increases or decreases in host susceptibility and infective stage (and vector) production, longevity, and pathology. Many other factors drive disease transmission, especially economics, and some change in time along with temperature, making it hard to distinguish whether temperature drives disease or just correlates with disease drivers. Although it is difficult to predict how climate change will affect infectious disease, an ecological approach can help meet the challenge.

Competition - colonization tradeoffs occur in many systems, and theory predicts that they can strongly promote species coexistence. However, there is little empirical evidence that observed competition - colonization tradeoffs are strong enough to maintain diversity in natural systems. This is due in part to a mismatch between theoretical assumptions and biological reality in some systems. We tested whether a competition - colonization tradeoff explains how a diverse trematode guild coexists in California horn snail populations, a system that meets the requisite criteria for the tradeoff to promote coexistence. A field experiment showed that subordinate trematode species tended to have higher colonization rates than dominant species. This tradeoff promoted coexistence in parameterized models but did not fully explain trematode diversity and abundance, suggesting a role of additional diversity maintenance mechanisms. Spatial heterogeneity is an alternative way to promote coexistence if it isolates competing species. We used scale transition theory to expand the competition - colonization tradeoff model to include spatial variation. The parameterized model showed that spatial variation in trematode prevalence did not isolate most species sufficiently to explain the overall high diversity, but could benefit some rare species. Together, the results suggest that several mechanisms combine to maintain diversity, even when a competition - colonization tradeoff occurs.

The California horn snail, Cerithideopsis californica, and the shore crabs, Pachygrapsus crassipes and Hemigrapsus oregonensis, compete for epibenthic microalgae, but the crabs also eat snails. Such intraguild predation is common in nature, despite models predicting instability. Using a series of manipulations and field surveys, we examined intraguild predation from several angles, including the effects of stage‐dependent predation along with direct consumptive and nonconsumptive predator effects on intraguild prey. In the laboratory, we found that crabs fed on macroalgae, snail eggs, and snails, and the size of consumed snails increased with predator crab size. In field experiments, snails grew less in the presence of crabs partially because snails behaved differently and were buried in the sediment (nonconsumptive effects). Consistent with these results, crab and snail abundances were negatively correlated in three field surveys conducted at three different spatial scales in estuaries of California, Baja California, and Baja California Sur: (1) among 61 sites spanning multiple habitat types in three estuaries, (2) among the habitats of 13 estuaries, and (3) among 34 tidal creek sites in one estuary. These results indicate that shore crabs are intraguild predators on California horn snails that affect snail populations via predation and by influencing snail behavior and performance.

Past theoretical models suggest fishing disease-impacted stocks can reduce parasite transmission, but this is a good management strategy only when the exploitation required to reduce transmission does not overfish the stock. We applied this concept to a red abalone fishery so impacted by an infectious disease (withering syndrome) that stock densities plummeted and managers closed the fishery. In addition to the non-selective fishing strategy considered by past disease-fishing models, we modelled targeting (culling) infected individuals, which is plausible in red abalone because modern diagnostic tools can determine infection without harming landed abalone and the diagnostic cost is minor relative to the catch value. The non-selective abalone fishing required to eradicate parasites exceeded thresholds for abalone sustainability, but targeting infected abalone allowed the fishery to generate yield and reduce parasite prevalence while maintaining stock densities at or above the densities attainable if the population was closed to fishing. The effect was strong enough that stock and yield increased even when the catch was one-third uninfected abalone. These results could apply to other fisheries as the diagnostic costs decline relative to catch value.

Linking marine epizootics to a specific aetiology is notoriously difficult. Recent diagnostic successes show that marine disease diagnosis requires both modern, cutting-edge technology (e.g. metagenomics, quantitative real-time PCR) and more classic methods (e.g. transect surveys, histopathology and cell culture). Here, we discuss how this combination of traditional and modern approaches is necessary for rapid and accurate identification of marine diseases, and emphasize how sole reliance on any one technology or technique may lead disease investigations astray. We present diagnostic approaches at different scales, from the macro (environment, community, population and organismal scales) to the micro (tissue, organ, cell and genomic scales). We use disease case studies from a broad range of taxa to illustrate diagnostic successes from combining traditional and modern diagnostic methods. Finally, we recognize the need for increased capacity of centralized databases, networks, data repositories and contingency plans for diagnosis and management of marine disease.

Most of the intertidal zone in estuaries of California, USA and Baja California, Mexico is covered with vascular vegetation. Shading by these vascular plants influences abiotic and biotic processes that shape benthic community assemblages. We present data on the effects of shading on the California horn snail, Cerithidea californica . This species is important because it is the most common benthic macrofaunal species in these systems and acts as an obligate intermediate host of several species of trematode parasites that infect several other species. Using observational and experimental studies, we found a negative effect of shade on the distribution and abundance of the California horn snail. We hypothesized that shading reduces the abundance of the epipelic diatoms that the snails feeds on, causing snails to leave shaded areas. We observed a negative relationship between vascular plant cover, sub-canopy light levels, and snail density in Mugu Lagoon. Then we experimentally manipulated light regimes, by clipping vegetation and adding shade structures, and found higher snail densities at higher light levels. In Goleta Slough, we isolated the effect of shade from vegetation by documenting a negative relationship between the shade created by two bridges and diatom and snail densities. We also found that snails moved the greatest distances over shaded channel banks compared to unshaded channel banks. Further, we documented the effect of water depth and channel bank orientation on shading in this system. An additional effect of shading is the reduction of temperature, providing an alternative explanation for some of our results. These results broaden our knowledge of how variation in the light environment influences the ecology of estuarine ecosystems.

Past models have suggested host–parasite coextinction could lead to linear, or concave down relationships between free-living species richness and parasite richness. I explored several models for the relationship between parasite richness and biodiversity loss. Life cycle complexity, low generality of parasites and sensitivity of hosts reduced the robustness of parasite species to the loss of free-living species diversity. Food-web complexity and the ordering of extinctions altered these relationships in unpredictable ways. Each disassembly of a food web resulted in a unique relationship between parasite richness and the richness of free-living species, because the extinction trajectory of parasites was sensitive to the order of extinctions of free-living species. However, the average of many disassemblies tended to approximate an analytical model. Parasites of specialist hosts and hosts higher on food chains were more likely to go extinct in food-web models. Furthermore, correlated extinctions between hosts and parasites (e.g. if parasites share a host with a specialist predator) led to steeper declines in parasite richness with biodiversity loss. In empirical food webs with random removals of free-living species, the relationship between free-living species richness and parasite richness was, on average, quasi-linear, suggesting biodiversity loss reduces parasite diversity more than previously thought.

We assessed how spatial and temporal heterogeneity and competition structure larval trematode communities in the pulmonate snail Lymnaea stagnalis . To postulate a dominance hierarchy, mark-release-recapture was used to monitor replacements of trematode species within snails over time. In addition, we sampled the trematode community in snails in different ponds in 3 consecutive years. A total of 7,623 snails (10,382 capture events) was sampled in 7 fishponds in the Jindřichův Hradec and Třeboň areas in South Bohemia (Czech Republic) from August 2006 to October 2008. Overall, 39% of snails were infected by a community of 14 trematode species; 7% of snails were infected with more than 1 trematode species (constituting 16 double- and 4 triple-species combinations). Results of the null-model analyses suggested that spatial heterogeneity in recruitment among ponds isolated trematode species from each other, whereas seasonal pulses in recruitment increased species interactions in some ponds. Competitive exclusion among trematodes led to a rarity of multiple infections compared to null-model expectations. Competitive relationships among trematode species were hypothesized as a dominance hierarchy based on direct evidence of replacement and invasion and on indirect evidence. Seven top dominant species with putatively similar competitive abilities (6 rediae and 1 sporocyst species) reduced the prevalence of the other trematode species developing in sporocysts only.

Nematomorph parasites manipulate crickets to enter streams where the parasites reproduce. These manipulated crickets become a substantial food subsidy for stream fishes. We used a field experiment to investigate how this subsidy affects the stream community and ecosystem function. When crickets were available, predatory fish ate fewer benthic invertebrates. The resulting release of the benthic invertebrate community from fish predation indirectly decreased the biomass of benthic algae and slightly increased leaf break-down rate. This is the first experimental demonstration that host manipulation by a parasite can reorganise a community and alter ecosystem function. Nematomorphs are common, and many other parasites have dramatic effects on host phenotypes, suggesting that similar effects of parasites on ecosystems might be widespread.

Parasite Niche Modeler (PaNic) is a free online software tool that suggests potential hosts for fish parasites. For a particular parasite species from the major helminth groups (Acanthocephala, Cestoda, Monogenea, Nematoda, Trematoda), PaNic takes data from known hosts (maximum body length, growth rate, life span, age at first maturity, trophic level, phylogeny, and biogeography) and hypothesizes similar fish species that might serve as hosts to that parasite. Users can give varying weights to host attributes and create custom models. In addition to suggesting plausible hosts (with varying degrees of confidence), the models indicate known host species that appear to be outliers in comparison to other known hosts. These unique features make PaNic an innovative tool for addressing both theoretical and applied questions in fish parasitology. PaNic can be accessed at http://purl.oclc.org/fishpest.

Organisms eating each other are only one of many types of well documented and important interactions among species. Other such types include habitat modification, predator interference and facilitation. However, ecological network research has been typically limited to either pure food webs or to networks of only a few (

Although many studies on the taxonomy of digenean trematodes of marine fishes have been completed in the Eastern Indo-Pacific (EIP) marine ecoregion, only a few have considered metacercarial stages. Here, the results are presented of a taxonomic survey of the digenean metacercariae of fishes from Palmyra Atoll, a remote and relatively pristine US National Wildlife Refuge located 1680 km SSW of Hawaii. Up to 425 individual fish were collected, comprising 42 fish species, from the sand flats bordering the lagoon of the atoll. Quantitative parasitological examinations of each fish were performed. Morphological descriptions of the encountered digenean metacercariae are provided, together with their prevalence, mean intensities, host and tissue-use. Up to 33,964 individuals were recovered representing 19 digenean metacercaria species from eight families. The species composition of digeneans in lagoon fishes at Palmyra Atoll is a subset of what has previously been reported for the EIP. Further, the large diversity and abundance of metacercariae reported in this study highlight the utility of including this group in future ecological research in the EIP marine ecoregion.

In September 2002, Hurricane Isidore devastated the Yucatán Peninsula, Mexico. To understand its effects on the parasites of aquatic organisms, we analyzed long-term monthly population data of the horn snail Cerithidea pliculosa and its trematode communities in Celestún, Yucatán, Mexico before and after the hurricane (February 2001 to December 2009). Five trematode species occurred in the snail population: Mesostephanus appendiculatoides, Euhaplorchis californiensis, two species of the genus Renicola and one Heterophyidae gen. sp. Because these parasites use snails as first intermediate hosts, fishes as second intermediate hosts and birds as final hosts, their presence in snails depends on food webs. No snails were present at the sampled sites for 6 months after the hurricane. After snails recolonised the site, no trematodes were found in snails until 14 months after the hurricane. It took several years for snail and trematode populations to recover. Our results suggest that the increase in the occurrence of hurricanes predicted due to climate change can impact upon parasites with complex life cycles. However, both the snail populations and their parasite communities eventually reached numbers of individuals and species similar to those before the hurricane. Thus, the trematode parasites of snails can be useful indicators of coastal lagoon ecosystem degradation and recovery.

The grass shrimp, Palaemonetes pugio, harbors a number of symbionts within its North American range. Here, we document the distribution and seasonality of 4 taxonomic groups that use P. pugio as a host in coastal Alabama. We conducted a regional survey of 4 symbionts of P. pugio over 3 seasons and compared assemblages across space and time. The most common parasite was the metacercarial stage of the microphallid trematode Microphallus turgidus, which remained consistently prevalent over the 3 seasons surveyed. We also monitored the prevalence of M. turgidus at 2 sites monthly. Prevalence fluctuated significantly among seasons at these sites, but spatial heterogeneity appears to have a stronger influence on regional parasite prevalence. Distributions of 3 of 4 symbionts overlapped in a single geographic area (Mon Louis Island, Alabama, U.S.A.); however, multispecies infections of individual hosts were normally distributed within host populations. Sites surrounding Mon Louis Island had substantiall...

Nematomorph parasites alter the behavior of their orthopteran hosts, driving them to water and creating a source of food for stream salmonids. We investigated whether nematomorphs could explain variation in terrestrial subsidies across several streams. In nine study streams, orthopterans comprise much of the stomach contents of trout (46 ± 31% on average). Total mass of ingested prey per trout biomass positively correlated with the mass of orthopterans ingested, suggesting that the orthopterans enhanced absolute mass of prey consumption by the trout population. The orthopterans ingested per trout biomass positively correlated with the abundance of nematomorphs in the stream, but not with the abundance of camel crickets (the dominant hosts) around the streams. Streams in conifer plantations had fewer nematomorphs than streams in natural deciduous forests. These results provide the first quantitative evidence that a manipulative parasite can explain variation in the allochthonous energy flow through and across ecosystems.

Parasites are ubiquitous in natural systems and ecosystem-level effects should be proportional to the amount of biomass or energy flow altered by the parasites. Here we quantified the extent to which a manipulative parasite altered the flow of energy through a forest-stream ecosystem. In a Japanese headwater stream, camel crickets and grasshoppers (Orthoptera) were 20 times more likely to enter a stream if infected by a nematomorph parasite (Gordionus spp.), corroborating evidence that nematomorphs manipulate their hosts to seek water where the parasites emerge as free-living adults. Endangered Japanese trout (Salvelinus leucomaenis japonicus) readily ate these infected orthopterans, which due to their abundance, accounted for 60% of the annual energy intake of the trout population. Trout grew fastest in the fall, when nematomorphs were driving energy-rich orthopterans into the stream. When infected orthopterans were available, trout did not eat benthic invertebrates in proportion to their abundance, leading to the potential for cascading, indirect effects through the forest-stream ecosystem. These results provide the first quantitative evidence that a manipulative parasite can dramatically alter the flow of energy through and across ecosystems., 寄生者（ハリガネムシ類）が駆動する渓畔生態系のエネルギー流の解明.京都大学プレスリリース.2011-04-13.