Keystone Effects Research Articles

Phloem-feeding insects create parasitoid-free space for caterpillars

Seemingly small ecological changes can have large, ramifying effects that defy expectations. Such are keystone effects in ecosystems. Phloem-feeding insect herbivores can act as keystone species by altering community structure and species interactions via plant-mediated or ant-mediated mechanisms. Plant responses triggered by phloem feeders can disrupt tri-trophic interactions induced by leaf-chewing herbivores, while ants that tend phloem feeders can deter or prey on other arthropods. Here, we investigate how phloem-feeding herbivores change caterpillar-parasitoid interactions on Quercus alba (white oak) trees in natural forests. We factorially manipulated the presence of phloem-feeding insects as well as ant access on Q.alba branches over multiple years and sites and measured parasitism rates of co-occurring caterpillars. While 19.3% of caterpillars were parasitized when phloem feeders were removed, the presence of phloem feeders completely suppressed parasitism of caterpillars (0%). This stark pattern was consistent across the diverse community of phloem feeders and caterpillars. Our manipulation of ant access had no effect on parasitism of caterpillars, implicating a plant-mediated mechanism. We further assessed the mechanistic hypothesis that phloem feeders suppress plant emission of caterpillar-induced volatile compounds, which could disrupt host-location behavior by parasitoids of caterpillars. Phloem feeders indeed reduced concentrations of four volatile compounds, consistent with the putative plant volatile-mediated mechanism. Given the important role of parasitoids in controlling herbivore populations, this keystone effect of phloem feeders offers novel insight into community dynamics in forests and potentially other terrestrial ecosystems.

Effects of Hemiparasites in Grassland Restorations Are Not Universal

Root hemiparasites infiltrate the vascular tissue of host roots to acquire water and nutrients, which often reduces host growth. Hemiparasites are postulated to be keystone species in grassland communities if they suppress dominant species and increase plant community biodiversity, and ecosystem engineers if they increase nutrient accessibility for surrounding species. We examined keystone effects by evaluating species richness and evenness in 1 m2 plots in a recent prairie restoration where Castilleja sessiliflora was naturally present or absent, and in a longer-established prairie restoration with or without Pedicularis canadensis. We examined ecosystem engineer effects by determining nitrate and phosphate concentrations under, 25 cm from, and 50 cm from hemiparasites, and in the center of hemiparasite-free plots. On the C. sessiliflora site, plots with the hemiparasites had higher species richness due to more forbs and higher floristic quality, consistent with the keystone species hypothesis. Soil phosphate levels were also greater in plots with C. sessiliflora present, consistent with the hypothesis of ecosystem engineering by this hemiparasite. In contrast, plots with/without P. canadensis showed no associations of any community metrics with the hemiparasite, and no correspondence between the presence of hemiparasites and soil nutrients. Although hemiparasites can increase grassland community heterogeneity, the effect is not universal, and the direction and strength of effects likely depends on local conditions.

Breeding Density and Collision Mortality of Loggerhead Shrike (Lanius ludovicianus) in the Altamont Pass Wind Resource Area

Loggerhead shrike (Lanius ludovicianus) has declined across most or all of its geographic range. The species’ raptorial behavior requires maintenance of large territories, which means populations of breeding shrikes require large areas of habitat and are therefore sensitive to habitat loss and habitat fragmentation. We estimated breeding densities of loggerhead shrikes in the Altamont Pass Wind Resource Area (APWRA), California, where annual shrike mortality caused by wind turbine collisions was high until just before our study began in 2016. Based on surveys across an annual average 50 randomized sampling plots in 2016–2019, we estimated an average 129 breeding pairs/year across the 167.6-km2 APWRA. Relative to the size of the study area, density in the APWRA was relatively high compared to densities reported from other study sites across North America. It was higher than predicted by application of the Partners in Flight estimator, which was based on Breeding Bird Surveys along roads. We also found that loggerhead shrikes in the APWRA were limited by the availability of nest substrate and by California ground squirrels (Otospermophilus beecheyi) and their burrow complexes, which have keystone effects on vegetation and wildlife in the APWRA. To most effectively conserve loggerhead shrikes in the APWRA, wind turbine mortality should be minimized, ground squirrels conserved instead of eradicated as pests, and appropriate trees and shrubs cultivated where they are needed.

Coming undone: hemiparasite presence and effects in a prairie grassland diminish over time

Root hemiparasites acquire resources from neighboring plants' vascular systems and can limit host growth, depress community productivity, and exert keystone effects. The strength of these effects is posited to be greater where hosts are nutrient-stressed but studies of annual hemiparasites show effects to be short-lived and variable. We conducted a 10-year experiment testing whether fertilizer addition alters the impact of the clonal, perennial hemiparasite Pedicularis canadensis on a prairie community and examine whether short-term trends reflect longer-term effects on community dynamics. Hemiparasite removal in 1-m2 plots increased productivity over the first three field seasons, but later the difference between removal and non-removal plots diminished as P. canadensis disappeared from 24 of the 48 non-removal plots. Effects of hemiparasite removal were context independent relative to fertilizer and shade treatments, but fertilizer initially increased, and then subsequently suppressed P. canadensis biomass. In non-removal plots, hemiparasite biomass was negatively associated with total community dry mass, which was greater in fertilized plots. Initially, fertilizer promoted graminoids, but after seven more field seasons, non-legume forbs responded most strongly. Measures of biodiversity tended to increase with hemiparasite cover. Demographic data collected at two different times for P. canadensis show high survivorship of established plants, high seed input, with seedling survival greater in taller vegetation. Unlike annual hemiparasite populations, well-established P. canadensis buffer populations against large demographic swings. At the scale of a few square meters, this keystone species produces significant heterogeneity in a prairie, but its presence at that scale is transient over approximately one decade.

Plant community responses to bison reintroduction on the Northern Great Plains, United States: a test of the keystone species concept

Keystone species restoration, or the restoration of species whose effect on an ecosystem is much greater than their abundance would suggest, is a central justification for many wildlife reintroduction projects globally. Following restoration, plains bison (Bison bison L.) have been identified as a keystone species in the tallgrass prairie ecoregion, but we know of no research to document similar effects in the mixed‐grass prairie where restoration efforts are ongoing. This study addresses whether Northern Great Plains (NGP) mixed‐grass prairie plant communities exhibit traits consistent with four central keystone effects documented for bison in the tallgrass prairie. We collected species composition, diversity, abundance, bare ground cover, and plant height data in three treatments: where livestock (Bos taurus L.) continuously grazed, livestock were removed for 10 years, and bison have been introduced and resident for 10 years. We observed mixed support for bison acting as keystone species in this system. Supporting the keystone role of bison, we observed higher species richness and compositional heterogeneity (β‐diversity) in the bison treatment than either the livestock retention or livestock removal treatments. However, we observed comparable forb, bare ground, and plant height heterogeneity between bison‐restored sites and sites where livestock were retained, contradicting reported keystone effects in other systems. Our results suggest that after 10 years of being restored, bison partially fulfill their role as a keystone species in the mixed‐grass prairie, and we encourage continued long‐term data collection to evaluate their influence in the NGP.

Predation on competing mussel species: Patterns of prey consumption and its potential role in species coexistence

Theoretical and empirical studies have demonstrated that predators can modulate coexistence between prey species and can modify broad patterns of community structure. Although the keystone predator effect (i.e., local coexistence and species diversity mediated by predator consumption) is commonly reported, negative effects of predators on species coexistence are also common. The balance depends critically on the asymmetry of competition, predator preferences for different prey, and the effective mortality rates exerted by predator populations on prey per capita growth rates. Here we studied the role of predation on two competing mussels, the dominant competitor Perumytilus purpuratus that dominates in abundance at most shores, and the subordinate Semimytilus algosus, which is common, but persistently abundant only at a smaller set of shores. We examined the potential for keystone effects by quantifying and evaluating the patterns of mussel consumption rates of sea star (Heliaster helianthus) and crab (Acanthocyclus gayi) predators in laboratory experiments, and under natural field conditions. Field sites with contrasting mussel abundance and recruitment (i.e., different local prey population growth rates) were chosen. Laboratory experiments showed that crabs would first consume Semimytilus over Perumytilus in mixed mussel clumps, while the larger sea star predators made no distinction between mussel species. Regardless of the order in which mussel species were first ingested, consumption rates of the two mussel species through the experiment were not different for either predator species. A similar temporal pattern was partly observable in the field through specific differences in the order of mussel consumption; however, experimental mussels of both species where consumed after the first two months of exposure to predators at both sites. Thus, in this system predation could not foster coexistence between mussel species and it may instead favor the occurrence of local extinction of the subordinate competitor, especially at sites where the smaller and more selective crab predators are relatively more abundant and recruitment of the subordinate species is low. Existing patterns of locally abundant Semimytilus beds are probably related to high site-specific recruitment rates of this species and should be further explored.

The pre-launch characterization of SIMBIO-SYS/VIHI imaging spectrometer for the BepiColombo mission to Mercury. II. Spectral calibrations.

The Visible and near Infrared Hyperspectral Imager (VIHI) is the VIS-IR spectrometer with imaging capabilities aboard the ESA BepiColombo mission to Mercury. In this second paper, we report the instrument spectral characterization derived by the calibration campaign carried out before spacecraft integration. Complementary measurements concerning radiometric and linearity responses, as well as geometric performances, are described in Paper I [G. Filacchione et al., Rev. Sci. Instrum. 88, 094502 (2017)]. We have verified the VIHI spectral range, spectral dispersion, spectral response function, and spectral uniformity along the whole slit. Instrumental defects and optical aberrations due to smiling and keystone effects have been evaluated, and they are lower than the design requirement (<1/3 pixel). The instrumental response is uniform along the whole slit, while spectral dispersion is well represented by a second order curve, rather than to be constant along the spectral dimension.

Functional relationships reveal keystone effects of the gopher tortoise on vertebrate diversity in a longleaf pine savanna

Keystone species are important drivers of diversity patterns in many ecosystems. Their effects on ecological processes are fundamental to understanding community dynamics, making them attractive conservation targets for ecosystem management. However, many studies assume keystone effects are constant. By developing functional relationships of species’ effects and assessing how they vary with context, we can design more efficient conservation strategies to maintain keystone impacts. The threatened gopher tortoise (Gopherus polyphemus) is presumed to be a keystone species promoting biodiversity in endangered longleaf pine ecosystems of the Southeastern Coastal Plain, USA. Although many commensals use tortoise burrows, their putative keystone influence on emergent diversity patterns lacks critical evaluation. We quantified the functional relationship between tortoise burrow density and non-volant vertebrate diversity in a longleaf pine savanna, located in central Florida. Tortoise burrow density had a positive effect on vertebrate diversity and evenness but did not affect species richness. This relationship was robust across fire disturbance regimes and was the primary factor explaining diversity at the local scale. Our results demonstrate keystone effects of the gopher tortoise through an ecosystem engineering mechanism. Continued gopher tortoise population declines will have large, negative impacts on vertebrate diversity in this biodiversity hotspot. Therefore, maintaining gopher tortoise populations is critical to effectively conserve dependent species and the function of endangered longleaf pine ecosystems. We show that developing a functional understanding of keystone relationships (not a binomial categorization) can lead to important insights into community processes.

Parasites that change predator or prey behaviour can have keystone effects on community composition

Parasites play pivotal roles in structuring communities, often via indirect interactions with non-host species. These effects can be density-mediated (through mortality) or trait-mediated (behavioural, physiological and developmental), and may be crucial to population interactions, including biological invasions. For instance, parasitism can alter intraguild predation (IGP) between native and invasive crustaceans, reversing invasion outcomes. Here, we use mathematical models to examine how parasite-induced trait changes influence the population dynamics of hosts that interact via IGP. We show that trait-mediated indirect interactions impart keystone effects, promoting or inhibiting host coexistence. Parasites can thus have strong ecological impacts, even if they have negligible virulence, underscoring the need to consider trait-mediated effects when predicting effects of parasites on community structure in general and biological invasions in particular.

Spatial variation in keystone effects: small mammal diversity associated with black‐tailed prairie dog colonies

Species with extensive geographic ranges may interact with different species assemblages at distant locations, with the result that the nature of the interactions may vary spatially. Black‐tailed prairie dogs Cynomys ludovicianus occur from Canada to Mexico in grasslands of the western Great Plains of North America. Black‐tailed prairie dogs alter vegetation and dig extensive burrow systems that alter grassland habitats for plants and other animal species. These alterations of habitat justify the descriptor “ecological engineer,” and the resulting changes in species composition have earned them status as a keystone species. We examined the impact of black‐tailed prairie dogs on small mammal assemblages by trapping at on‐ and off‐colony locations at eight study areas across the species' geographic range. We posed 2 nested hypotheses: 1) prairie dogs function as a keystone species for other rodent species; and 2) the keystone role varies spatially. Assuming that it does, we asked what are the sources of the variation?Black‐tailed prairie dogs consistently functioned as a keystone species in that there were strong statistically significant differences in community composition on versus off prairie dog colonies across the species range in prairie grassland. Small mammal species composition varied along both latitudinal and longitudinal gradients, and species richness varied from 4 to 11. Assemblages closer together were more similar; such correlations approximately doubled when including only on‐ or off‐colony grids. Black‐tailed prairie dogs had a significant effect on associated rodent assemblages that varied regionally, dependent upon the composition of the local rodent species pool. Over the range of the black‐tailed prairie dog, on‐colony rodent richness and evenness were less variable, and species composition was more consistent than off‐colony assemblages.

Keystone effects of an alien top-predator stem extinctions of native mammals

Alien predators can have catastrophic effects on ecosystems and are thought to be much more harmful to biodiversity than their native counterparts. However, trophic cascade theory and the mesopredator release hypothesis predict that the removal of top predators will result in the reorganization of trophic webs and loss of biodiversity. Using field data collected throughout arid Australia, we provide evidence that removal of an alien top-predator, the dingo, has cascading effects through lower trophic levels. Dingo removal was linked to increased activity of herbivores and an invasive mesopredator, the red fox (Vulpes vulpes), and to the loss of grass cover and native species of small mammals. Using species distribution data, we predict that reintroducing or maintaining dingo populations would produce a net benefit for the conservation of threatened native mammals across greater than 2.42 x 10(6) km(2) of Australia. Our study provides evidence that an alien top predator can assume a keystone role and be beneficial for biodiversity conservation, and also that mammalian carnivores more generally can generate strong trophic cascades in terrestrial ecosystems.

Scaling up keystone effects from simple to complex ecological networks

AbstractPredicting the consequences of species loss requires extending our traditional understanding of simpler dynamic systems of few interacting species to the more complex ecological networks found in natural ecosystems. Especially important is the scaling up of our limited understanding of how and under what conditions loss of ‘keystone’ species causes large declines of many other species. Here we explore how these keystone effects vary among simulations progressively scaled up from simple to more complex systems. Simpler simulations of four to seven interacting species suggest that species up to four links away can strongly alter keystone effects and make the consequences of keystone loss potentially indeterminate in more realistically complex communities. Instead of indeterminacy, we find that more complex networks of up to 32 species generally buffer distant influences such that variation in keystone effects is well predicted by surprisingly local ‘top‐down’, ‘bottom‐up’, and ‘horizontal‘ constraints acting within two links of the keystone subsystem. These results demonstrate that: (1) strong suppression of the competitive dominant by the keystone may only weakly affect subordinate competitors; (2) the community context of the target species determines whether strong keystone effects are realized; (3) simple, measurable, and local attributes of complex communities may explain much of the empirically observed variation in keystone effects; and (4) increasing network complexityper sedoes not inherently make the prediction of strong keystone effects more complicated.

Keystone effects of the endangered Stephens' kangaroo rat ( Dipodomys stephensi)

Kangaroo rats ( Dipodomys) have been argued to exert keystone effects because they interact strongly with other species, and their removal results in major changes in community structure. We evaluated the nature and magnitude of such impacts by the endangered Stephens' kangaroo rat ( Dipodomys stephensi), employing an exclosure experiment to determine how the removal of this species impacted relative cover by vegetation, plant species diversity, abundance of a dominant annual plant genus ( Erodium), and predation of seeds from artificial trays. Fifteen months of D. stephensi removal resulted in significant increases in herb cover and Erodium abundance, and significant declines in bare ground and in seed predation. These results support the hypothesis that D. stephensi has major impacts on plant community structure and seems to function as a keystone species. It remains to be seen how the loss of this species from much of its range will influence local community composition and dynamics.

Gravelly flood deposits of Irvine Creek, Ontario, Canada

ABSTRACTIn May 1974 a powerful flood flushed the Grand River basin, Ontario. The effects on the bedload were drastic in a narrow (30 m) and deep (40 m) rock walled and floored gorge near Elora, Ontario. Along Irvine Creek, the tributary occupying the gorge, the gravel cover was reworked in several types of bars, predominantly transverse and point bars. The bars formed very rapidly in response to essentially steady, non‐uniform flow that developed during a brief period of high flood.Superimposed on major bars are several minor sedimentary features such as coarse transverse ribs, chute channels and bars, longitudinal ribs, imbrication clusters, backsets with well developed imbrication, that were formed under very high stream discharge. Structures like imbrication clusters, transverse ribs and small riffle bars require a ‘live bed’ situation to form, and they develop when stones come to a stop either because they cluster during transport, or because keystone effects occur along shallow channels. In Irvine Creek, very few sedimentary features were formed during waning and low flood stages: only some shadow deposits and a few Ostler lenses. The few fines that were available were lost downstream or filled in lower parts of gravel beds.This study confirms that in streams that experience strong seasonal fluctuations in discharge, bedforms that develop during high floods have a high probability of preservation. In gravelly deposits, foreset structures and plane beds are most commonly preserved, although they may be difficult to recognize in old deposits, which may appear massive, particularly if the gravel has been infilled with finer pebbles and sand. In the case of Irvine Creek, all deposits are organized, and lateral and vertical variations in textures, particularly imbrication and packing, are very useful in the recognition of sedimentary structures.