Trophic cascade induced by molluscivore predator alters pore-water biogeochemistry via competitive release of prey

Effects of predation may cascade down the food web. By alleviating interspecific competition among prey, predators may promote biodiversity, but the precise mechanisms of how predators alter competition have remained elusive. Here we report on a predator-exclosure experiment carried out in a tropical intertidal ecosystem, providing evidence for a three-level trophic cascade induced by predation by molluscivore Red Knots (Calidris canutus) that affects pore water biogeochemistry. In the exclosures the knots' favorite prey (Dosinia isocardia) became dominant and reduced the individual growth rate in an alternative prey (Loripes lucinalis). Dosinia, a suspension feeder, consumes suspended particulate organic matter (POM), whereas Loripes is a facultative mixotroph, partly living on metabolites produced by sulfur-oxidizing chemoautotrophic bacteria, but also consuming suspended POM. Reduced sulfide concentrations in the exclosures suggest that, without predation on Dosinia, stronger competition for suspended POM forces Loripes to rely on energy produced by endosymbiotic bacteria, thus leading to an enhanced uptake of sulfide from the surrounding pore water. As sulfide is toxic to most organisms, this competition-induced diet shift by Loripes may detoxify the environment, which in turn may facilitate other species. The inference that predators affect the toxicity of their environment via a multi-level trophic cascade is novel, but we believe it may be a general phenomenon in detritus-based ecosystems.     

Suppressing antagonistic bioengineering feedbacks doubles restoration success

In a seagrass restoration project, we explored the potential for enhancing the restoration process by excluding antagonistic engineering interactions (i.e., biomechanical warfare) between two ecosystem engineers: the bioturbating lugworm Arenicola marina and the sediment-stabilizing seagrass Zostera noltii Hornem. Applying a shell layer underneath half of our seagrass transplants successfully reduced adult lugworm density by over 80% and reduced lugworm-induced microtopography (a proxy for lugworm disturbance) at the wave-sheltered site. At the wave-exposed site adult lugworm densities and microtopography were already lower than at the sheltered site but were further reduced in the shell-treated units. Excluding lugworms and their bioengineering effects corresponded well with a strongly enhanced seagrass growth at the wave-sheltered site, which was absent at the exposed site. Enhanced seagrass growth in the present study was fully assigned to the removal of lugworms' negative engineering effects and not to any (indirect) evolving effects such as an altered biogeochemistry or sediment-stabilizing effects by the shell layer. The context-dependency implies that seagrass establishment at the exposed site is not constrained by negative ecosystem-engineering interactions only, but also by overriding physical stresses causing poor growth conditions. Present findings underline that, in addition to recent emphasis on considering positive (facilitating) interactions in ecological theory and practice, it is equally important to consider negative engineering interactions between ecosystem-engineering species. Removal of such negative interactions between ecosystem-engineering species can give a head start to the target species at the initial establishment phase, when positive engineering feedbacks by the target species on itself are still lacking. Though our study was carried out in a marine environment with variable levels of wave disturbance, similar principles may be expected to apply to other ecosystems that are inhabited by ecosystem engineers.     

Decomposing the mid-Holocene Tsuga decline in eastern North America

The mid-Holocene decline of Tsuga canadensis (hereafter Tsuga) populations across eastern North America is widely perceived as a synchronous event, driven by pests/pathogens, rapid climate change, or both. Pattern identification and causal attribution are hampered by low stratigraphic density of pollen-sampling and radiometric dates at most sites, and by absence of highly resolved, paired pollen and paleoclimate records from single sediment cores, where chronological order of climatic and vegetational changes can be assessed. We present an intensely sampled (contiguous 1-cm intervals) record of pollen and water table depth (inferred from testate amoebae) from a single core spanning the Tsuga decline at Irwin Smith Bog in Lower Michigan, with high-precision chronology. We also present an intensively sampled pollen record from Tower Lake in Upper Michigan. Both sites show high-magnitude fluctuations in Tsuga pollen percentages during the pre-decline maximum. The terminal decline is dated at both sites ca. 5000 cal yr BP, some 400 years later than estimates from other sites and data compilations. The terminal Tsuga decline was evidently heterochronous across its range. A transient decline ca. 5350 cal yr BP at both sites may correspond to the terminal decline at other sites in eastern North America. At Irwin Smith Bog, the terminal Tsuga decline preceded an abrupt and persistent decline in water table depths by approximately 200 years, suggesting the decline was not directly driven by abrupt climate change. The Tsuga decline may best be viewed as comprising at least three phases: a long-duration pre-decline maximum with high-magnitude and high-frequency fluctuations, followed by a terminal decline at individual sites, followed in turn by two millennia of persistently low Tsuga populations. These phases may not be causally linked, and may represent dynamics taking place at multiple temporal and spatial scales. Further progress toward understanding the phenomenon requires an expanded network of high-resolution pollen and paleoclimate chronologies.     

Plant-mediated and nonadditive effects of two global change drivers on an insect herbivore community

Warmer temperatures can alter the phenology and distribution of individual species. However, differences across species may blur community-level phenological responses to climate or cause biotic homogenization by consistently favoring certain taxa. Additionally, the response of insect communities to climate will be subject to plant-mediated effects, which may or may not overshadow the direct effect of rising temperatures on insects. Finally, recent evidence for the importance of interaction effects between global change drivers suggests that phenological responses of communities to climate may be altered by other drivers. We used a natural temperature gradient (generated by elevation and topology), combined with experimental nitrogen fertilization, to investigate the effects of elevated temperature and globally increasing anthropogenic nitrogen deposition on the structure and phenology of a seminatural grassland herbivore assemblage (lepidopteran insects). We found that both drivers, alone and in combination, severely altered how the relative abundance and composition of species changed through time. Importantly, warmer temperatures were associated with biotic homogenization, such that herbivore assemblages in the warmest plots had more similar species composition than those in intermediate or cool plots. Changes in herbivore composition and abundance were largely mediated by changes in the plant community, with increased nonnative grass cover under high treatment levels being the strongest determinant of herbivore abundance. In addition to compositional changes, total herbivore biomass more than doubled under elevated nitrogen and increased more than fourfold with temperature, bearing important functional implications for herbivores as consumers and as a prey resource. The crucial role of nonnative plant dominance in mediating responses of herbivores to change, combined with the frequent nonadditive (positive and negative) effects of the two drivers, and the differential responses of species, highlight that understanding complex ecosystem responses will benefit from multifactor, multitrophic experiments at community scales or larger.     

Underlying causes of habitat-associated differences in size of age-0 walleye pollock (Theragra chalcogramma) in the Gulf of Alaska

Age-0 walleye pollock (Theragra chalcogramma) caught in September in the Gulf of Alaska display habitat-associated differences in standard length (SL). Age-0 fish collected in the region around Sutwik Island and 375?km farther downstream near the Shumagin Islands most likely originate from the Shelikof Strait spawning aggregation. However, age-0 fish resulting from the same spawning aggregation differ in mean size up to 20?mm between areas by September. We examined the otoliths of the larval and age-0 stages of walleye pollock from these two areas in 2000 and 2001 to determine whether growth rate, hatch date, and/or temperature influenced fish size. Circulation models were used to determine whether transport of larvae from an upstream spawning group into the study areas could have occurred. Mean in situ temperature during sampling periods was not defined as a significant factor in altering growth rates. Overlapping hatch date distributions of the larval and age-0 fish in the Shumagin Island area confirmed that the fish were from the Shelikof Strait spawning group. Comparison of hatch date distributions in the upstream Sutwik Island area revealed larger/older larvae from an upstream spawning group mixed with larvae from the Shelikof Strait spawning group. Our results suggest that the offset of 20?mm SL between the groups of age-0 pollock was the result of a combination of enhanced survivorship of early-hatched larvae in the Sutwik area and the introduction and retention of the progeny of another spawning group originating upstream of Shelikof Strait.     

Assessing citizen contributions to butterfly monitoring in two large cities

Citizen science may be especially effective in urban landscapes due to the large pool of potential volunteers. However, there have been few evaluations of the contributions of citizen scientists to knowledge of biological communities in and around cities. To assess the effectiveness of citizen scientists' monitoring of species in urban areas, we compared butterfly data collected over 10 years in Chicago, Illinois (U.S.A.), and New York City, New York (U.S.A.). The dates, locations, and methods of data collection in Chicago were standardized, whereas data from New York were collected at any location at any time. For each city, we evaluated whether the number of observers, observation days (days on which observations were reported), and sampling locations were associated with the reported proportion of the estimated regional pool of butterfly species. We also compared the number of volunteers, duration of volunteer involvement, and consistency of sampling efforts at individual locations within each city over time. From 2001 to 2010, there were 73 volunteers in Chicago and 89 in New York. During this period, volunteers observed 86% and 89% of the estimated number of butterfly species present in Chicago and New York, respectively. Volunteers in New York reported a greater proportion of the estimated pool of butterfly species per year. In addition, more species were observed per volunteer and observation day in New York, largely due to the unrestricted sampling season in New York. Chicago volunteers were active for more years and monitored individual locations more consistently over time than volunteers in New York. Differences in monitoring protocol--especially length of sampling season and selection protocol for monitoring locations--influenced the relationship between species accrual and sampling effort, which suggests these factors are important in volunteer-based species-monitoring programs.     

Normalisation models for accounting for fat content in stable isotope measurements in salmonid muscle tissue

Stable isotope analysis is increasingly used in ecological studies. Because lipid content influences δ¹³C, lipids should be removed from lipid-rich samples before δ¹³C analysis. To account for differences in δ¹³C arising from differences in lipid content, relationships between lipid content, C:N ratio and Δδ¹³C with lipid removal can be used to normalise lipid content to uniform levels. We investigate these relationships for salmonid muscle and evaluate the suitability of previously published normalisation equations for these fish. Salmonids with a wide range of condition (muscle lipid content = 3–35% of dry weight) were considered. There were no consistent relationships between lipid content or C:N ratio and Δδ¹⁵N. There were linear relationships between C:N ratio and lipid content (L = −16.53 + 6.27 × C:N); C:N ratio and Δδ¹³C (Δδ¹³C = −1.87 + 0.65 × C:N); and lipid content and Δδ¹³C (Δδ¹³C = 0.01 + 0.10 × L), which should be used on salmonid stable isotope studies.     

Chemical profile of Festuca arundinacea extract showing allelochemical activity

The allelopathic potential of fescue (Festuca arundinacea Shreb.) was investigated under field and laboratory conditions. In the field, incorporation of fescue aboveground biomass into the soil provided an effective weed control as it significantly reduced both the number and the dry weight of all weed species. The effect on weeds was species specific. The reduction of the most abundant species, Picris echioides, with respect to the number of individuals and their dry weight was 67 and 45%. Fescue aboveground biomass, harvested immediately before soil incorporation, was utilized in a bioassay-oriented fractionation, aimed at identifying the compounds in the toxic extracts of the plant. While phytotoxic activity was detected for the n-butanol fraction, no activity was found for the aqueous one. Germination of an indicator species, lettuce (Lactuca sativa L.), was reduced by 33% at the higher extract concentration of 20 mg/mL, owing to the increase of the number of abnormal seedlings. The LC/MS profile of the extract revealed the presence of two alkaloids, seven flavonol glycosides, and two flavonol aglyca. Quercetin 3-O-rutinoside, isorhamnetin 3-O-rutinoside, and kaempferol 3-O-rutinoside were reported in fescue for the first time.     

How the social parasitic bumblebee <Emphasis Type="Italic">Bombus bohemicus</Emphasis> sneaks into power of reproduction

Social parasitism is widespread in many groups of social living hymenopteran species and has also evolved in the genus Bombus. Cuckoo bumblebees (subgenus Psithyrus) are obligate brood parasites in nests of other bumblebee species. After nest usurpation and the killing of the host queen, the parasite female has to control worker reproduction in order to accomplish and maintain reproductive dominance and to ensure her reproductive success. The aim of our study was to examine whether the generalist parasitic bumblebee Bombus bohemicus monopolizes and prevents worker reproduction by physical or chemical means and to identify possible odor compounds involved therein. We performed bioassays with callow workers of the host Bombus terrestris and have shown that B. bohemicus females are able to suppress host worker ovarian development, when these host workers are under the direct influence of the parasite female. Furthermore, by chemical analyses, we have demonstrated that the parasite females adjust to the odor profiles of their host queens in order to maintain the level of fertility signaling inside the host colony although the host queen is absent. We also found that host workers change their odor profile after nest usurpation by the parasite female and consequently, we suggest that the host and parasite are caught up in a chemical arms race.