Growing evidence suggests that organisms with narrow niche requirements are particularly disadvantaged in small habitat patches, typical of fragmented landscapes. However, the mechanisms behind this relationship remain unclear. Dietary specialists may be particularly constrained by the availability of their food resources as habitat area shrinks. For herbivorous insects, host plants may be filtered out of small habitat fragments by neutral sampling processes and deterministic plant community shifts due to altered microclimates, edge effects and browsing by ungulates. We examined the relationship between forest fragment area and the abundance of dietary‐specialist and dietary‐generalist larval Lepidoptera (caterpillars) and their host plants in the northeastern USA. We surveyed caterpillars and their host plants over 3 years in equal‐sized plots within 32 forest fragments varying in area between 3 and 1014 ha. We tested whether the abundances and species richness of dietary specialists increased more than those of dietary generalists with increasing fragment area and, if so, whether the difference could be explained by reduced host plant availability or increased browsing by white‐tailed deer ( Odocoileus virginianus ). The overall abundance of dietary specialists was positively related to fragment area; the relationship was substantially weaker for dietary generalists. There was notable variation among species within diet breadth groups, however. There was no effect of fragment area on the diversity of dietary‐specialist or dietary‐generalist caterpillars. Deer activity was not related to the abundances of either dietary‐generalist or dietary‐specialist caterpillars. Plant community composition was strongly associated with fragment area. Larger fragments were more likely to include host plants for both dietary‐specialist and dietary‐generalist caterpillars. Deer activity was correlated with decreased host plant availability for both groups, with a slightly stronger impact on host plants of dietary specialists. Although dietary specialists were more likely to lack host plants in fragments, the relationship between fragment area and host availability did not depend on caterpillar diet breadth. This study provides further evidence that decreasing patch area disproportionately impacts specialist consumers. Because this relationship was derived from equal‐sized plots, it is robust to some criticisms levelled at fragmentation research. The mechanisms for specialist consumer declines, however, remain elusive.
How consumer diversity determines consumption efficiency is a central issue in ecology. In the context of predation and biological control, this relationship concerns predator diversity and predation efficiency. Reduced predation efficiency can result from different predator taxa eating each other in addition to their common prey (interference due to intraguild predation). By contrast, multiple predator taxa with overlapping but complementary feeding niches can generate increased predation efficiency on their common prey (enemy complementarity). When viewed strictly from an ecological perspective, intraguild predation and enemy complementarity are opposing forces. However, from an evolutionary ecology perspective, predators facing strong intraguild predation may evolve traits that reduce their predation risk, possibly leading to niche complementarity between enemies; thus, selection from intraguild predation may lead to enemy complementarity rather than opposing it. As specialized predators that live in or on their hosts, parasitoids are subjected to intraguild predation from generalist predators that consume the parasitoids' hosts. The degree to which parasitoid-predator interactions are ruled by interference versus enemy complementarity has been debated. Here, we address this issue with field experiments in a forest community consisting of multiple species of trees, herbivorous caterpillars, parasitoids, ants, and birds. Our experiments and analyses found no interference effects, but revealed clear evidence for complementarity between parasitoids and birds (not ants). Parasitism rates by hymenopterans and dipterans were negatively associated with bird predation risk, and the variation in the strength of this negative association suggests that this enemy complementarity was due to parasitoid avoidance of intraguild predation. We further argue that avoidance of intraguild predation by parasitoids and other arthropod predators may explain enigmatic patterns in vertebrate-arthropod-plant food webs in a variety of terrestrial ecosystems.
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.
Supplementary Figure from Impaired Proteolysis of Noncanonical RAS Proteins Drives Clonal Hematopoietic Transformation
As plant species expand their upper limits of distribution under current warming, some retain both traditional climate space and biotic environment while others encounter novel conditions. The latter is the case for Rhododendron campanulatum, a woody shrub that grows both above and below treeline at our study site in the Eastern Himalayas where a very conspicuous, stable treeline was defined by a nearly contiguous canopy of tall Abies spectabilis trees, many of which are over a century old. Prior work showed that treeline had remained static in this region while R. campanulatum expanded its elevational range limit. We tested local adaptation of R. campanulatum by performing reciprocal transplants between the species' current elevational range limit (4023 m above sea level [asl]) and just above treeline (3876 m asl). Contrary to expectation, the coldest temperatures of late winter and early mid-spring were experienced by plants at the lower elevation: R. campanulatum at species' limit (upper site) were covered by snow for a longer period (40 more days) and escaped the coldest temperatures suffered by conspecifics at treeline (lower site). The harsher spring conditions at treeline likely explain why leaves were smaller at treeline (15.3 cm(2)) than at species limit (21.3 cm(2)). Contrary to results from equivalent studies in other regions, survival was reduced more by downslope than by upslope movement, again potentially due to extreme cold temperatures observed at treeline in spring. Upslope transplantation had no effect on mortality, but mortality of species limit saplings transplanted downslope was three times higher than that of residents at both sites. A general expectation is that locals should survive better than foreign transplants, but survival of locals and immigrants at our species limit site was identical. However, those species limit saplings that survived the transplant to treeline grew faster than both locals at treeline and the transplants at species limit. Overall, we found asymmetric adaptation: Compared with treeline saplings, those at species limit (147 m above treeline) were more tolerant of extremes in the growing season but less tolerant of extremes in winter and early mid-spring, displaying local adaptation in a more complex manner than simply home advantage, and complicating predictions about impacts of future regional climate change.
Climate change reinvigorates two debates: first, about relative roles of taxonomic, genetic and phenetic criteria in prioritizing entities for conservation and, second, on the role of hybrids in biodiversity assessments, as climatechange-induced range shifts create sympatry between formerly allopatric species/subspecies, thereby generating novel hybrids. We argue that rapid climate change necessitates a shift in the ethical balance sheet for conservation prioritization. Here we use the extensive ecotypic variation in a climate-sensitive butterfly, Euphydryas editha, to illustrate a case in which conservation biologists choosing extant populations as sources for reintroductions to extinct sites, translocations to novel sites, or genetic rescue of inbred populations should first select populations with appropriate phenotypes for climate adaptation and preferences for host and habitat and only then, within that group, rely on subspecies identity or genomic similarity between source and target populations. Preferences evolve rapidly, and we show that they differed sufficiently between an endangered subspecies and potential sources genomically close to it that restoration efforts would be doomed. Conversely, introductions from genomically more distant, ecologically similar populations would succeed and hybridize. Evolutionary biologists have argued that hybridizations, whether caused by climate change or genetic rescue projects, may be essential for retaining genetic diversity and adaptive potential at species/genus level. Conservation practitioners, in contrast, tend to oppose genetic change, arguing to preserve "purity" of existing groups. Our example of the role of climate change in conservation of a species with rapid evolution of critical traits and a mosaic pattern of local adaptation supports the evolutionary biologists' argument.
Despite growing evidence that "connectedness" of humans with nature creates multiple benefits for both humans and nature, these benefits are not fully considered by health and conservation policymakers. Studies are scattered across scientific disciplines including health, education, psychology and biology, making it difficult to get a complete overview. Here, we conduct a systematic review, focused on recent meta-analyses that investigate impacts of psychological and/or physical connection with nature on human health and well-being and on attitudes and actions that promote nature conservation. By "psychological connection" we mean the extent to which people see themselves as part of nature and by "physical connection" we mean contact with natural areas. We identified 16 relevant meta-analyses covering 832 independent studies. We found consistent conclusions across geographically diverse experimental studies that physical connection with nature improved human cognition, social skills, physical and mental health, and psychological connection to nature. Experiments also showed that psychological connection with nature had significant positive impact on pro-environmental behaviors and values. Correlational studies supported experimental results and, in addition, found psychological connection with nature positively correlated with mental and physical health. Studies are biased toward adults rather than children and away from southern regions (Africa, Oceania and South America). Overall, our review suggests a critical role for psychological and physical human-nature connections in developing a sustainable future. Although experimental studies are rare, conducting cross-cultural experimental research is needed if governmental and non-governmental stakeholders, researchers and citizens are to develop appropriate actions toward achieving United Nations Sustainable Development Goals.
Studies in birds and trees show climatic stresses distributed across species' ranges, not only at range limits. Here, new analyses from the butterfly Euphydryas editha reveal mechanisms generating these stresses: geographic mosaics of natural selection, acting on tradeoffs between climate adaptation and fitness traits, cause some range-central populations to evolve to limits of climatic tolerance, while others remain resilient. In one ecotype, selection for predator avoidance drives evolution to limits of thermal tolerance. In a second ecotype, the endangered Bay Checkerspot, selection on fecundity drives evolution to the climate-sensitive limit of ability to complete development within the lifespans of ephemeral hosts, causing routinely high mortality from insect-host phenological asynchrony. The tradeoff between maternal fecundity and offspring mortality generated similar values of fitness on different dates, partly explaining why fecundity varied by more than an order of magnitude. Evolutionary response to the tradeoff rendered climatic variability the main driver of Bay Checkerspot dynamics, and increases in this variability, associated with climate change, were a key factor behind permanent extinction of a protected metapopulation. Finally, we discuss implications for conservation planning of our finding that adaptive evolution can reduce population-level resilience to climate change and generate geographic mosaics of climatic stress. This article is part of the theme issue 'Species' ranges in the face of changing environments (Part II)'.
Evolutionary change impacts the rate at which insect pests, pollinators, or disease vectors expand or contract their geographic ranges. Although evolutionary changes, and their ecological feedbacks, strongly affect these risks and associated ecological and economic consequences, they are often underappreciated in management efforts. Greater rigor and scope in study design, coupled with innovative technologies and approaches, facilitates our understanding of the causes and consequences of eco-evolutionary dynamics in insect range shifts. Future efforts need to ensure that forecasts allow for demographic and evolutionary change and that management strategies will maximize (or minimize) the adaptive potential of range-shifting insects, with benefits for biodiversity and ecosystem services.
Internationally agreed sustainability goals are being missed. Here, we conduct global meta-analyses to assess how the extent to which humans see themselves as part of nature-known as human-nature connectedness (HNC)-can be used as a leverage point to reach sustainability. A meta-analysis of 147 correlational studies shows that individuals with high HNC had more pronature behaviours and were significantly healthier than those with low HNC. A meta-analysis of 59 experimental studies shows significant increases in HNC after manipulations involving contact with nature and mindfulness practices. Surprisingly, this same meta-analysis finds no significant effect of environmental education on HNC. Thus, HNC is positively linked to mind-sets that value sustainability and behaviours that enhance it. Further, we argue that HNC can be enhanced by targeted practices, and we identify those most likely to succeed. Our results suggest that enhancing HNC, via promotion of targeted practices, can improve sustainability and should be integrated into conservation policy.
Insects have been key players in the assessments of biodiversity impacts of anthropogenically driven environmental change, including the evolutionary and ecological impacts of climate change. Populations of Edith’s Checkerspot Butterfly (Euphydryas editha) adapt rapidly to diverse environmental conditions, with numerous high-impact studies documenting these dynamics over several decades. However, studies of the underlying genetic bases of these responses have been hampered by missing genomic resources, limiting the ability to connect genomic responses to environmental change. Using a combination of Oxford Nanopore long reads, haplotype merging, HiC scaffolding followed by Illumina polishing, we generated a highly contiguous and complete assembly (contigs n = 142, N50 = 21.2 Mb, total length = 607.8 Mb; BUSCOs n = 5,286, single copy complete = 97.8%, duplicated = 0.9%, fragmented = 0.3%, missing = 1.0%). A total of 98% of the assembled genome was placed into 31 chromosomes, which displayed large-scale synteny with other well-characterized lepidopteran genomes. The E. editha genome, annotation, and functional descriptions now fill a missing gap for one of the leading field-based ecological model systems in North America.
Scientists often need to know whether pairs of entities tend to occur together or independently. Standard approaches to this issue use co-occurrence indices such as Jaccard, Sørensen-Dice, and Simpson. We show that these indices are sensitive to the prevalences of the entities they describe and that this invalidates their interpretability. We propose an index, α, that is insensitive to prevalences. Published datasets reanalyzed with both α and Jaccard's index (J) yield profoundly different biological inferences. For example, a published analysis using J contradicted predictions of the island biogeography theory finding that community stability increased with increasing physical isolation. Reanalysis of the same dataset with the estimator [Formula: see text] reversed that result and supported theoretical predictions. We found similarly marked effects in reanalyses of antibiotic cross-resistance and human disease biomarkers. Our index α is not merely an improvement; its use changes data interpretation in fundamental ways.
We review evidence for predators as agents of selection for the evolution of dietary specialization by insect herbivores. Since being proposed in the 1980s, this “enemy-free space” hypothesis has garnered much attention, but relatively little testing. Most tests have used caterpillars to address the community-level prediction that generalist predators preferentially attack dietary generalist herbivore species relative to their specialist counterparts. Both qualitative and quantitative aspects of our review found mostly supporting evidence for this prediction. We also reviewed literature on various mechanisms proposed for the superior anti-predator defenses of specialist herbivores, including sequestration of plant allelochemicals, specific camouflage, increased behavioral efficiency, and greater ability to suppress herbivore-induced plant volatiles. Considerable evidence supports allelochemical sequestration as a primary mechanism for the anti-predator advantage of specialists. Specific camouflage and increased feeding efficiency seem likely mechanisms based on evidence from limited testing. In contrast, the evidence indicates that specialists are no better than generalists at suppressing herbivore-induced volatiles. The mechanistic part of our review prompted us to propose a refinement of the enemy-free space hypothesis: the greater the dependence of anti-predator defenses on taxonomically specific host plant traits, the stronger the selection for the evolution of dietary specialization in herbivores. This refinement makes new predictions intended to spur mechanistic and evolutionary tests of the enemy-free space hypothesis.
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Abstract Remote sensing imagery can provide critical information on the magnitude and extent of damage caused by forest pests and pathogens. However, monitoring short‐term changes in deciduous forest condition caused by defoliating insects is challenging and requires approaches that directly account for seasonal vegetation dynamics. We implemented a previously published harmonic modeling approach for forest condition monitoring in Google Earth Engine and systematically assessed the relative ability of condition change products generated using various model parameterizations for predicting pest abundances and defoliation during the 2016–2018 gypsy moth (Lymantria dispar) outbreak in southern New England. Our comparisons revealed that most models made reasonable predictions of changes in canopy condition and egg and larval abundances of L. dispar, indicating a strong correlation between our harmonic‐based estimates of condition change and defoliator activity. The greatest differences in predictive ability were in the spectral domain, with assessments based on Tasseled Cap Greenness, Simple Ratio, and the Enhanced Vegetation Index ranking among the top models, and the commonly used Normalized Difference Vegetation Index consistently exhibiting poorer performance. We also observed notable differences in the magnitude of scores for different baseline periods. Additionally, we found that Landsat‐based condition scores better explained larval abundance than egg mass counts, which have historically been used as a proxy for later‐season larval abundance, indicating that our remote sensing approach may be more accurate and cost‐effective for generating consistent retrospective assessments of L. dispar population abundance in addition to estimates of canopy damage. These findings provide important linkages between spectral changes detected using a harmonic modeling approach and biophysical aspects of defoliator activity, with potential to extend monitoring and prediction to regional or even continental scales.
As species' poleward range limits expand under climate change, generalists are expected to be better colonists than specialists, extending their ranges faster. This effect of specialization on range shifts has been shown, but so has the reverse cause–effect: in a global meta‐analysis of butterfly diets, it was range expansions themselves that caused increases in population‐level diet breadth. What could drive this unexpected process? We provide a novel behavioral mechanism by showing that, in a butterfly with extensive ecotypic variation, Edith's checkerspot, diet breadths increased after colonization events as diversification of individual host preferences pulled novel hosts into population diets. Subsequently, populations that persisted reverted toward monophagy. We draw together three lines of evidence from long‐term studies of 15 independently evolving populations. First, direct observations showed a significant increase in specialization across decades: in recent censuses, eight populations used fewer host genera than in the 1980s while none used more. Second, behavioral preference‐testing experiments showed that extinctions and recolonizations at two sites were followed, at first by diversification of heritable preference ranks and increases in diet breadth, and subsequently by homogenization of preferences and contractions of diet breadth. Third, we found a significant negative association in the 1980s between population‐level diet breadth and genetic diversity. Populations with fewer mtDNA haplotypes had broader diets, extending to 3–4 host genera, while those with higher haplotype diversity were more specialized. We infer that diet breadth had increased in younger, recently colonized populations. Preference diversification after colonization events, whether caused by (cryptic) host shifts or by release of cryptic genetic variation after population bottlenecks, provides a mechanism for known effects of range shifts on diet specialization. Our results explain how colonizations at expanding range margins have increased population‐level diet breadths, and predict that increasing specialization should accompany population persistence as current range edges become range interiors.
This review was solicited as an autobiography. The "problems" in my title have two meanings. First, they were professional difficulties caused by my decision to study oviposition preferences of butterflies that were not susceptible to traditional preference-testing designs. Until I provided video, my claim that the butterflies duplicate natural post-alighting host-assessment behavior when placed on hosts by hand was not credible, and the preference-testing technique that I had developed elicited skepticism, anger, and derision. The second meaning of "problems" is scientific. Insect preference comes with complex dimensionality that interacts with host acceptability. Part Two of this review describes how my group's work in this area has revealed unexpected axes of variation in plant-insect interactions-axes capable of frustrating attempts to derive unequivocal conclusions from apparently sensible experimental designs. The possibility that these complexities are lurking should be kept in mind as preference and performance experiments are devised.
This review was solicited as an autobiography. The "problems" in my title have two meanings. First, they were professional difficulties caused by my decision to study oviposition preferences of butterflies that were not susceptible to traditional preference-testing designs. Until I provided video, my claim that the butterflies duplicate natural post-alighting host-assessment behavior when placed on hosts by hand was not credible, and the preference-testing technique that I had developed elicited skepticism, anger, and derision. The second meaning of "problems" is scientific. Insect preference comes with complex dimensionality that interacts with host acceptability. Part Two of this review describes how my group's work in this area has revealed unexpected axes of variation in plant-insect interactions-axes capable of frustrating attempts to derive unequivocal conclusions from apparently sensible experimental designs. The possibility that these complexities are lurking should be kept in mind as preference and performance experiments are devised.
Dynamics of herbivorous insect diet breadth are important in generation of novel pests, biological control of weeds and as indicators of global change impacts. But what forces and events drive these dynamics? Here we present evidence for a novel scenario: that specialization increases in persistent populations, but that, at the species level, this trend is countered by effects of colonizations. Colonizations cause host shifts, which are followed by non-adaptive evolutionary expansions of diet breadth, adding transitory hosts during adaptation to the principal novel host. We base this thesis on long-term study of 15 independently-evolving populations of Edith’s Checkerspot butterfly, eight of which used fewer host genera in recent censuses than in the 1980’s, while none used more - a significant increase in specializaton. At the same time, two extintion/recolonization events were followed by temporary expansions of diet breadth. Behavioural experiments showed that these expansions were driven by within-population diversification of individual oviposition preferences. These results may explain an old puzzle: a significant negative association between population-level diet breadth and mtDNA diversity. Populations with fewer mtDNA haplotypes had broader diets, suggesting that diet breadth increases in younger, recently-colonized populations. A recent global meta-analysis of butterfly diets, using biogeographic data, explains latitudinal patterns of diet breadth by showing that poleward range expansions have caused reduced specialization. This implies broad applicability of our results, which provide a plausible mechanism for the latitudinal trends: colonizations at expanding range margins would increase population-level diet breadths, while population persistence in range interiors would facilitate increasing specialization.
Abstract Alpine treelines are expected to shift upward due to recent climate change. However, interpretation of changes in montane systems has been problematic because effects of climate change are frequently confounded with those of land use changes. The eastern Himalaya, particularly Langtang National Park, Central Nepal, has been relatively undisturbed for centuries and thus presents an opportunity for studying climate change impacts on alpine treeline uncontaminated by potential confounding factors. We studied two dominant species, Abies spectabilis (AS) and Rhododendron campanulatum (RC), above and below the treeline on two mountains. We constructed 13 transects, each spanning up to 400 m in elevation, in which we recorded height and state (dead or alive) of all trees, as well as slope, aspect, canopy density, and measures of anthropogenic and animal disturbance. All size classes of RC plants had lower mortality above treeline than below it, and young RC plants (<2 m tall) were at higher density above treeline than below. AS shows little evidence of a position change from the historic treeline, with a sudden extreme drop in density above treeline compared to below. Recruitment, as measured by size–class distribution, was greater above treeline than below for both species but AS is confined to ~25 m above treeline whereas RC is luxuriantly growing up to 200 m above treeline. Synthesis. Evidence suggests that the elevational limits of RC have shifted upward both because (a) young plants above treeline benefited from facilitation of recruitment by surrounding vegetation, allowing upward expansion of recruitment, and (b) temperature amelioration to mature plants increased adult survival. We predict that the current pure stand of RC growing above treeline will be colonized by AS that will, in turn, outshade and eventually relegate RC to be a minor component of the community, as is the current situation below the treeline.
Eric Aaron合作论文数Department of Computer Science
Wesleyan University2