USGS ScienceSearch

Geology topics

Ryan F. Hechinger

Publications and source records attributed to Ryan F. Hechinger.

17 recordsLinked to original sources

Broadening the ecology of fear: Non-lethal effects arise from diverse responses to predation and parasitism

Research on the ‘ecology of fear’ posits that defensive prey responses to avoid predation can cause non-lethal effects across ecological scales. Parasites also elicit defensive responses in hosts with associated non-lethal effects, which raises the longstanding, yet unresolved question of how non-lethal effects of parasites compare with those of predators. We developed a framework for systematically answering this question for all types of predator–prey and host–parasite systems. Our framework reveals likely differences in non-lethal effects not only between predators and parasites, but also between different types of predators and parasites. Trait responses should be strongest towards predators, parasitoids and parasitic castrators, but more numerous and perhaps more frequent for parasites than for predators. In a case study of larval amphibians, whose trait responses to both predators and parasites have been relatively well studied, existing data indicate that individuals generally respond more strongly and proactively to short-term predation risks than to parasitism. Apart from studies using amphibians, there have been few direct comparisons of responses to predation and parasitism, and none have incorporated responses to micropredators, parasitoids or parasitic castrators, or examined their long-term consequences. Addressing these and other data gaps highlighted by our framework can advance the field towards understanding how non-lethal effects impact prey/host population dynamics and shape food webs that contain multiple predator and parasite species.

Proceedings of the Royal Society B

Parasitic nematodes of marine fishes from Palmyra Atoll, East Indo-Pacific, including a new species of Spinitectus (Nematoda, Cystidicolidae)

Here, we present the results of a taxonomic survey of the nematodes parasitizing fishes from the lagoon flats of Palmyra Atoll, Eastern Indo-Pacific. We performed quantitative parasitological surveys of 653 individual fish from each of the 44 species using the intertidal sand flats that border the atoll’s lagoon. We provide morphological descriptions, prevalence, and mean intensities of the recovered seven species of adult nematode ( Pulchrascaris chiloscyllii , Capillariidae gen. sp., Cucullanus bourdini, Cucullanus oceaniensis, Pseudascarophis sp., Spinitectus (Paraspinitectus) palmyraensis sp. nov., Philometra pellucida ) and three larval stages ( Pulchrascaris sp., Hysterothylacium sp., Cucullanus sp.). We recorded: Pulchrascaris chiloscyllii from Carcharhinus melanopterus ; Capillariidae gen. sp. from Chaetodon lunula, Lutjanus fulvus, and Ellochelon vaigiensis ; Cucullanus bourdin i from Arothron hispidus ; Cucullanus oceaniensis from Abudefduf sordidus ; Pseudascarophis sp. from Chaetodon auriga, Chaetodon lunula, and Mulloidichthys flavolineatus ; Spinitectus ( Paraspinitectus ) palmyraensis sp. nov. from Albula glossodonta ; Philometra pellucida from Arothron hispidus ; and three larval forms, Pulchrascaris sp. from Acanthurus triostegus, Acanthurus xanthopterus, Rhinecanthus aculeatus, Platybelone argalus, Carangoides ferdau, Carangoides orthogrammus, Caranx ignobilis, Caranx melampygus, Caranx papuensis, Chaetodon auriga, Chanos chanos, Amblygobius phalaena, Asterropteryx semipunctata, Valencienea sexguttata, Kyphosus cinerascens, Lutjanus fulvus, Lutjanus monostigma, Ellochelon vaigiensis, Mulloidichthys flavolineatus, Upeneus taeniopterus, Gymnothorax pictus, Abudefduf septemfasciatus, Abudefduf sordidus, and Stegastes nigricans ; Hysterothylacium sp. type MD from Acanthurus triostegus, Carangoides ferdau, Chaetodon lunula, Chanos chanos, Kyphosus cinerascens, Abudefduf sordidus, and Arothron hispidus ; and Cucullanus sp. from Caranx ignobilis . Spinitectus ( Paraspinitectus ) palmyraensis sp. nov. (Cystidicolidae) is described from the intestine of roundjaw bonefish Albula glossodonta . All the nematode species reported in this study represent new geographical records. We discuss how our survey findings compare to other areas of the Indo-Pacific, and the way the relatively numerical dominance of trophically transmitted larval stages likely reflect the intact food web of Palmyra Atoll, which includes a large biomass of large-bodied top predator sharks and ray-finned fishes.

ZooKeys

Parasitic copepods (Crustacea, Hexanauplia) on fishes from the lagoon flats of Palmyra Atoll, Central Pacific

We surveyed copepods parasitic on the fishes at Palmyra, a remote atoll in the Central Indo-Pacific faunal region. In total, we collected 849 individual fish, representing 44 species, from the intertidal lagoon flats at Palmyra and recovered 17 parasitic copepod species. The parasitic copepods were: Orbitacolax williamsi on Mulloidichthys flavolineatus ; Anuretes serratus on Acanthurus xanthopterus ; Caligus confusus on Carangoides ferdau , Carangoides orthogrammus , Caranx ignobilis , Caranx melampygus , and Caranx papuensis ; Caligus kapuhili on Chaetodon auriga and Chaetodon lunula ; Caligus laticaudus on Rhinecanthus aculeatus , Pseudobalistes flavimarginatus , M. flavolineatus , Upeneus taeniopterus , Chrysiptera glauca , and Epinephalus merra ; Caligus mutabilis on Lutjanus fulvus and Lutjanus monostigma ; Caligus randalli on C. ignobilis ; Caligus sp. on L. fulvus ; Caritus serratus on Chanos chanos ; Lepeophtheirus lewisi on A. xanthopterus ; Lepeophtheirus uluus on C. ignobilis ; Dissonus similis on Arothron hispidus ; Nemesis sp. on Carcharhinus melanopterus ; Hatschekia longiabdominalis on A. hispidus ; Hatschekia bicaudata on Chaetodon auriga and Chaetodon lunula ; Kroyeria longicauda on C. melanopterus and Lernanthropus sp. on Kyphosus cinerascens . All copepod species reported here have been previously reported from the Indo-Pacific but represent new geographical records for Palmyra, demonstrating large-scale parasite dispersion strategies.

ZooKeys

Sea‐level rise, habitat loss, and potential extirpation of a salt marsh specialist bird in urbanized landscapes

Sea‐level rise (SLR) impacts on intertidal habitat depend on coastal topology, accretion, and constraints from surrounding development. Such habitat changes might affect species like Belding's savannah sparrows ( Passerculus sandwichensis beldingi ; BSSP), which live in high‐elevation salt marsh in the Southern California Bight. To predict how BSSP habitat might change under various SLR scenarios, we first constructed a suitability model by matching bird observations with elevation. We then mapped current BSSP breeding and foraging habitat at six estuarine sites by applying the elevation‐suitability model to digital elevation models. To estimate changes in digital elevation models under different SLR scenarios, we used a site‐specific, one‐dimensional elevation model (wetland accretion rate model of ecosystem resilience). We then applied our elevation‐suitability model to the projected digital elevation models. The resulting maps suggest that suitable breeding and foraging habitat could decline as increased inundation converts middle‐ and high‐elevation suitable habitat to mudflat and subtidal zones. As a result, the highest SLR scenario predicted that no suitable breeding or foraging habitat would remain at any site by 2100 and 2110. Removing development constraints to facilitate landward migration of high salt marsh, or redistributing dredge spoils to replace submerged habitat, might create future high salt marsh habitat, thereby reducing extirpation risk for BSSP in southern California.

Ecology and Evolution

Monogenea of fishes from the lagoon flats of Palmyra Atoll in the Central Pacific

A survey of the monogeneans of fishes from the lagoon flats of Palmyra Atoll detected 16 species already reported from the Indo-West Pacific faunal region. A total of 653 individual fish from 44 species were collected from the sand flats bordering the lagoon of the atoll. Eighteen species of fish were infected with monogeneans. The monogenean species recovered were: Benedenia hawaiiensis on Acanthurus xanthopterus, Chaetodon auriga, Chaetodon lunula, Mulloidichthys flavolineatus, Pseudobalistes flavimarginatus and Rhinecanthus aculeatus; Ancyrocephalus ornatus on Arothron hispidus; Euryhaliotrema annulocirrus on Chaetodon auriga and Chaetodon lunula; Euryhaliotrema chrysotaeniae on Lutjanus fulvus; Euryhaliotrema grandis on Chaetodon auriga and Chaetodon lunula; Haliotrema acanthuri on Acanthurus triostegus; Haliotrema aurigae on Chaetodon auriga and Chaetodon lunula; Haliotrema dempsteri on Acanthurus xanthopterus; Haliotrema minutospirale on Mulloidichthys flavolineatus; Haliotrematoides patellacirrus on Lutjanus monostigma; Neohaliotrema bombini on Abudefduf septemfasciatus and Abudefduf sordidus; Acleotrema girellae and Acleotrema parastromatei on Kyphosus cinerascens; Cemocotylella elongata on Caranx ignobilis, Caranx melampygus and Caranx papuensis; Metamicrocotyla macracantha on Crenimugil crenilabris; and Pseudopterinotrema albulae on Albula glossodonta. All these monogenean–host combinations represent new geographical records. The monogenean species composition of the Palmyra Atoll is similar to that of the Hawaiian Islands. However, the number of species recovered was lower compared with other localities within the Indo-West Pacific, perhaps due to the geographical isolation of Palmyra Atoll.

Palmyra Atoll

The role of competition – colonization tradeoffs and spatial heterogeneity in promoting trematode coexistence

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.

Ecology

A lack of crowding? Body size does not decrease with density for two behavior-manipulating parasites

For trophically transmitted parasites that manipulate the phenotype of their hosts, whether the parasites do or do not experience resource competition depends on such factors as the size of the parasites relative to their hosts, the intensity of infection, the extent to which parasites share the cost of defending against the host’s immune system or manipulating their host, and the extent to which parasites share transmission goals. Despite theoretical expectations for situations in which either no, or positive, or negative density-dependence should be observed, most studies document only negative density-dependence for trophically transmitted parasites. However, this trend may be an artifact of most studies having focused on systems in which parasites are large relative to their hosts. Yet, systems are common where parasites are small relative to their hosts, and these trophically transmitted parasites may be less likely to experience resource limitation. We looked for signs of density-dependence in Euhaplorchis californiensis (EUHA) and Renicola buchanani (RENB), two manipulative trematode parasites infecting wild-caught California killifish (Fundulus parvipinnis). These parasites are small relative to killifish (suggesting resources are not limiting), and are associated with changes in killifish behavior that are dependent on parasite-intensity and that increase predation rates by the parasites’ shared final host (indicating the possibility for cost sharing). We did not observe negative density-dependence in either species, indicating that resources are not limiting. In fact, observed patterns indicate possible mild positive density-dependence for EUHA. Although experimental confirmation is required, our findings suggest that some behavior-manipulating parasites suffer no reduction in size, and may even benefit when "crowded" by conspecifics.

Integrative and Comparative Biology

Parasites affect food web structure primarily through increased diversity and complexity

Comparative research on food web structure has revealed generalities in trophic organization, produced simple models, and allowed assessment of robustness to species loss. These studies have mostly focused on free-living species. Recent research has suggested that inclusion of parasites alters structure. We assess whether such changes in network structure result from unique roles and traits of parasites or from changes to diversity and complexity. We analyzed seven highly resolved food webs that include metazoan parasite data. Our analyses show that adding parasites usually increases link density and connectance (simple measures of complexity), particularly when including concomitant links (links from predators to parasites of their prey). However, we clarify prior claims that parasites ‘‘dominate’’ food web links. Although parasites can be involved in a majority of links, in most cases classic predation links outnumber classic parasitism links. Regarding network structure, observed changes in degree distributions, 14 commonly studied metrics, and link probabilities are consistent with scale-dependent changes in structure associated with changes in diversity and complexity. Parasite and free-living species thus have similar effects on these aspects of structure. However, two changes point to unique roles of parasites. First, adding parasites and concomitant links strongly alters the frequency of most motifs of interactions among three taxa, reflecting parasites’ roles as resources for predators of their hosts, driven by trophic intimacy with their hosts. Second, compared to free-living consumers, many parasites’ feeding niches appear broader and less contiguous, which may reflect complex life cycles and small body sizes. This study provides new insights about generic versus unique impacts of parasites on food web structure, extends the generality of food web theory, gives a more rigorous framework for assessing the impact of any species on trophic organization, identifies limitations of current food web models, and provides direction for future structural and dynamical models.

PLoS Biology

Parasites as prey in aquatic food webs: implications for predator infection and parasite transmission

While the recent inclusion of parasites into food-web studies has highlighted the role of parasites as consumers, there is accumulating evidence that parasites can also serve as prey for predators. Here we investigated empirical patterns of predation on parasites and their relationships with parasite transmission in eight topological food webs representing marine and freshwater ecosystems. Within each food web, we examined links in the typical predator–prey sub web as well as the predator–parasite sub web, i.e. the quadrant of the food web indicating which predators eat parasites. Most predator– parasite links represented ‘concomitant predation’ (consumption and death of a parasite along with the prey/host; 58–72%), followed by ‘trophic transmission’ (predator feeds on infected prey and becomes infected; 8–32%) and predation on free-living parasite life-cycle stages (4–30%). Parasite life-cycle stages had, on average, between 4.2 and 14.2 predators. Among the food webs, as predator richness increased, the number of links exploited by trophically transmitted parasites increased at about the same rate as did the number of links where these stages serve as prey. On the whole, our analyses suggest that predation on parasites has important consequences for both predators and parasites, and food web structure. Because our analysis is solely based on topological webs, determining the strength of these interactions is a promising avenue for future research.

Oikos

A common scaling rule for abundance, energetics, and production of parasitic and free-living species

The metabolic theory of ecology uses the scaling of metabolism with body size and temperature to explain the causes and consequences of species abundance. However, the theory and its empirical tests have never simultaneously examined parasites alongside free-living species. This is unfortunate because parasites represent at least half of species diversity. We show that metabolic scaling theory could not account for the abundance of parasitic or free-living species in three estuarine food webs until accounting for trophic dynamics. Analyses then revealed that the abundance of all species uniformly scaled with body mass to the - 3/4 power. This result indicates "production equivalence," where biomass production within trophic levels is invariant of body size across all species and functional groups: invertebrate or vertebrate, ectothermic or endothermic, and free-living or parasitic.

Science

Homage to Linnaeus: How many parasites? How many hosts?

Estimates of the total number of species that inhabit the Earth have increased significantly since Linnaeus's initial catalog of 20,000 species. The best recent estimates suggest that there are ≈6 million species. More emphasis has been placed on counts of free-living species than on parasitic species. We rectify this by quantifying the numbers and proportion of parasitic species. We estimate that there are between 75,000 and 300,000 helminth species parasitizing the vertebrates. We have no credible way of estimating how many parasitic protozoa, fungi, bacteria, and viruses exist. We estimate that between 3% and 5% of parasitic helminths are threatened with extinction in the next 50 to 100 years. Because patterns of parasite diversity do not clearly map onto patterns of host diversity, we can make very little prediction about geographical patterns of threat to parasites. If the threats reflect those experienced by avian hosts, then we expect climate change to be a major threat to the relatively small proportion of parasite diversity that lives in the polar and temperate regions, whereas habitat destruction will be the major threat to tropical parasite diversity. Recent studies of food webs suggest that ≈75% of the links in food webs involve a parasitic species; these links are vital for regulation of host abundance and potentially for reducing the impact of toxic pollutants. This implies that parasite extinctions may have unforeseen costs that impact the health and abundance of a large number of free-living species.

Proceedings of the National Academy of Sciences of

Diversity increases biomass production for trematode parasites in snails

Increasing species diversity typically increases biomass in experimental assemblages. But there is uncertainty concerning the mechanisms of diversity effects and whether experimental findings are relevant to ecological process in nature. Hosts for parasites provide natural, discrete replicates of parasite assemblages. We considered how diversity affects standing-stock biomass for a highly interactive parasite guild: trematode parasitic castrators in snails. In 185 naturally occurring habitat replicates (individual hosts), diverse parasite assemblages had greater biomass than single-species assemblages, including those of their most productive species. Additionally, positive diversity effects strengthened as species segregated along a secondary niche axis (space). The most subordinate species—also the most productive when alone—altered the general positive effect, and was associated with negative diversity effects on biomass. These findings, on a previously unstudied consumer class, extend previous research to illustrate that functional diversity and species identity may generally both explain how diversity influences biomass production in natural assemblages of competing species.

Proceedings of the Royal Society B: Biological Sci

Ecosystem energetic implications of parasite and free-living biomass in three estuaries

Parasites can have strong impacts but are thought to contribute little biomass to ecosystems . We quantified the biomass of free-living and parasitic species in three estuaries on the Pacific coast of California and Baja California. Here we show that parasites have substantial biomass in these ecosystems. We found that parasite biomass exceeded that of top predators. The biomass of trematodes was particularly high, being comparable to that of the abundant birds, fishes, burrowing shrimps and polychaetes. Trophically transmitted parasites and parasitic castrators subsumed more biomass than did other parasitic functional groups. The extended phenotype biomass controlled by parasitic castrators sometimes exceeded that of their uninfected hosts. The annual production of free-swimming trematode transmission stages was greater than the combined biomass of all quantified parasites and was also greater than bird biomass. This biomass and productivity of parasites implies a profound role for infectious processes in these estuaries.

Nature

Endangered light-footed clapper rail affects parasite community structure in coastal wetlands

An extinction necessarily affects community members that have obligate relationships with the extinct species. Indirect or cascading effects can lead to even broader changes at the community or ecosystem level. However, it is not clear whether generalist parasites should be affected by the extinction of one of their hosts. We tested the prediction that loss of a host species could affect the structure of a generalist parasite community by investigating the role of endangered Light-footed Clapper Rails ( Rallus longirostris levipes ) in structuring trematode communities in four tidal wetlands in southern California, USA (Carpinteria Salt Marsh, Mugu Lagoon) and Mexico (Estero de Punta Banda, Bahia Falsa–San Quintín). We used larval trematode parasites in first intermediate host snails ( Cerithidea californica ) as windows into the adult trematodes that parasitize Clapper Rails. Within and among wetlands, we found positive associations between Clapper Rails and four trematode species, particularly in the vegetated marsh habitat where Clapper Rails typically occur. This suggests that further loss of Clapper Rails is likely to affect the abundance of several competitively dominant trematode species in wetlands with California horn snails, with possible indirect effects on the trematode community and changes in the impacts of these parasites on fishes and invertebrates.

Ecological Applications

Host diversity begets parasite diversity: Bird final hosts and trematodes in snail intermediate hosts

An unappreciated facet of biodiversity is that rich communities and high abundance may foster parasitism. For parasites that sequentially use different host species throughout complex life cycles, parasite diversity and abundance in ‘downstream’ hosts should logically increase with the diversity and abundance of ‘upstream’ hosts (which carry the preceding stages of parasites). Surprisingly, this logical assumption has little empirical support, especially regarding metazoan parasites. Few studies have attempted direct tests of this idea and most have lacked the appropriate scale of investigation. In two different studies, we used time-lapse videography to quantify birds at fine spatial scales, and then related bird communities to larval trematode communities in snail populations sampled at the same small spatial scales. Species richness, species heterogeneity and abundance of final host birds were positively correlated with species richness, species heterogeneity and abundance of trematodes in host snails. Such community-level interactions have rarely been demonstrated and have implications for community theory, epidemiological theory and ecosystem management.

Proceedings of the Royal Society B: Biological Sci

The introduced ribbed mussel (Geukensia demissa) in Estero de Punta Banda, Mexico: Interactions with the native cord grass, Spartina foliosa

Introduced populations of Guekensia demissa occur on the west coast of North America. They have been reported in San Francisco Bay, four southern California wetlands, and in Estero de Punta Banda (EPB), Baja California Norte, Mexico. We randomly sampled benthic invertebrates in four habitat types within EPB: marsh, channel, mudflat and pan. Geukensia demissa was the most abundant bivalve in the wetland at EPB. It was significantly associated with the native cordgrass, Spartina foliosa, and occurred at higher average densities in vegetated marsh sites (24/m2) and Spartina-dominated tidal channels (35/m2), compared to mudflat (0/m2), and pan (0/m 2) sites. We estimated that the total biomass of this invader was over four times that of the next most abundant bivalve, Tagelus spp., in EPB. We examined G. demissa for parasites and found that only a few native parasites colonized this introduced host at very low prevalences and intensities. We performed bird surveys to determine the habitat overlap and potential impact of this mussel on the EPB population of light-footed clapper rails (Rallus longirostrus levipes), an endangered species in the United States. The high abundance of G. demissa in EPB, its presence in clapper rail habitat, and its known effects on salt marsh habitat in it's native range, warrant further investigations of the impact of this invader in EPB and elsewhere. ?? Springer 2005.

Biological Invasions

Food webs including parasites, biomass, body sizes, and life stages for three California/Baja California estuaries

This data set presents food webs for three North American Pacific coast estuaries and a “Metaweb” composed of the species/stages compiled from all three estuaries. The webs have four noteworthy attributes: (1) parasites (infectious agents), (2) body-size information, (3) biomass information, and (4) ontogenetic stages of many animals with complex life cycles. The estuaries are Carpinteria Salt Marsh, California (CSM); Estero de Punta Banda, Baja California (EPB); and Bahía Falsa in Bahía San Quintín, Baja California (BSQ). Most data on species assemblages and parasitism were gathered via consistent sampling that acquired body size and biomass information for plants and animals larger than ∼1 mm, and for many infectious agents (mostly metazoan parasites, but also some microbes). We augmented this with information from additional published sources and by sampling unrepresented groups (e.g., plankton). We estimated free-living consumer–resource links primarily by extending a previously published version of the CSM web (which the current CSM web supplants) and determined most parasite consumer–resource links from direct observation. We recognize 21 possible link types including four general interactions: predators consuming prey, parasites consuming hosts, predators consuming parasites, and parasites consuming parasites. While generally resolved to the species level, we report stage-specific nodes for many animals with complex life cycles. We include additional biological information for each node, such as taxonomy, lifestyle (free-living, infectious, commensal, mutualist), mobility, and residency. The Metaweb includes 500 nodes, 314 species, and 11 270 links projected to be present given appropriate species' co-occurrences. Of these, 9247 links were present in one or more of the estuarine webs. The remaining 2023 links were not present in the estuaries but are included here because they may occur in other places or times. Initial analyses have examined and are examining the interrelationships among consumer strategy, body size, abundance, biomass, trophic level, life stages, and food-web structure and dynamics. Further use of these data may enable a more general exploration how infectious processes and parasites impact communities and ecosystems. Additionally, we present the data and metadata in a standardized format, attempting to provide a system-neutral template for future food-web assembly and publication.

Ecology