Chuan-Kai Ho

Invertebrates, ecosystem services and climate change

The sustainability of ecosystem services depends on a firm understanding of both how organisms provide these services to humans and how these organisms will be altered with a changing climate. Unquestionably a dominant feature of most ecosystems, invertebrates affect many ecosystem services and are also highly responsive to climate change. However, there is still a basic lack of understanding of the direct and indirect paths by which invertebrates influence ecosystem services, as well as how climate change will affect those ecosystem services by altering invertebrate populations. This indicates a lack of communication and collaboration among scientists researching ecosystem services and climate change effects on invertebrates, and land managers and researchers from other disciplines, which becomes obvious when systematically reviewing the literature relevant to invertebrates, ecosystem services, and climate change. To address this issue, we review how invertebrates respond to climate change. We then review how invertebrates both positively and negatively influence ecosystem services. Lastly, we provide some critical future directions for research needs, and suggest ways in which managers, scientists and other researchers may collaborate to tackle the complex issue of sustaining invertebrate‐mediated services under a changing climate.

Is Diet Quality an Overlooked Mechanism for Bergmann’s Rule?

Bergmann’s rule (body size increases with latitude) has long interested biologists; however, its mechanism remains unclear. An overlooked mechanism (latitudinal variation in plant quality) might help explain Bergmann’s rule. We studied three herbivores. In the field, the planthopper Prokelisia and the sea hare Aplysia, but not the long‐horned grasshopper Orchelimum, were larger at high latitudes, following Bergmann’s rule. In the laboratory, all three species grew larger or faster on high‐latitude plants. High‐latitude diets increased Prokelisia size and Aplysia growth rates by 8% and 72%, respectively, enough to explain the increase in field body size toward high latitudes. Therefore, latitudinal variation in herbivore body size could be influenced by latitudinal variation in plant quality, which may directly or indirectly also affect body size in detritivores, parasitoids, and predators. Studies of Bergmann’s rule should consider the influence of biotic factors on body size in addition to abiotic factors such as temperature and precipitation.

Environmental gradients and herbivore feeding preferences in coastal salt marshes

Current theories of plant-herbivore interactions suggest that plants may differ in palatability to herbivores as a function of abiotic stress; however, studies of these theories have produced mixed results. We compared the palatability of eight common salt marsh plants that occur across elevational and salinity stress gradients to six common leaf-chewing herbivores to determine patterns of plant palatability. The palatability of every plant species varied across gradients of abiotic stress in at least one comparison, and over half of the comparisons indicated significant differences in palatability. The direction of the preferences, however, was dependent on the plant and herbivore species studied, suggesting that different types of stress affect plants in different ways, that different plant species respond differently to stress, and that different herbivore species measure plant quality in different ways. Overall, 51% of the variation in the strength of the feeding preferences could be explained by a knowledge of the strength of the stress gradient and the type of gradient, plant and herbivore studied. This suggests that the prospects are good for a more complex, conditional theory of plant stress and herbivore feeding preferences that is based on a mechanistic understanding of plant physiology and the factors underlying herbivore feeding preferences.

Effects of an Omnivorous Katydid, Salinity, and Nutrients on a Planthopper-Spartina Food Web

Top–down and bottom–up effects interact to structure communities, especially in salt marshes, which contain strong gradients in bottom–up drivers such as salinity and nutrients. How omnivorous consumers respond to variation in prey availability and plant quality is poorly understood. We used a mesocosm experiment to examine how salinity, nutrients, an omnivore (the katydid Orchelimum fidicinium) and an herbivore (the planthopper Prokelisia spp.) interacted to structure a simplified salt marsh food web based on the marsh grass Spartina alterniflora. Bottom–up effects were strong, with both salinity and nutrients decreasing leaf C/N and increasing Prokelisia abundance. Top–down effects on plants were also strong, with both the herbivore and the omnivore affecting S. alterniflora traits and growth, especially when nutrients or salt were added. In contrast, top–down control by Orchelimum of Prokelisia was independent of bottom–up conditions. Orchelimum grew best on a diet containing both Spartina and Prokelisia, and in contrast to a sympatric omnivorous crab, did not shift to an animal-based diet when prey were present, suggesting that it is constrained to consume a mixed diet. These results suggest that the trophic effects of omnivores depend on omnivore behavior, dietary constraints, and ability to suppress lower trophic levels, and that omnivorous katydids may play a previously unrecognized role in salt marsh food webs.

Preference and performance in plant-herbivore interactions across latitude--a study in U.S. Atlantic salt marshes.

High-latitude plants are often more palatable to herbivores than low-latitude conspecifics. Does increased plant palatability lead to better herbivore performance? Our field and laboratory work investigated (A) whether high-latitude plants have traits indicating that they should be higher-quality foods for herbivores; (B) whether geographic differences in plant quality are more important than local adaptation of herbivores. We studied 3 plant species and 6 invertebrate herbivores in U.S. Atlantic Coast. Past studies had shown high-latitude individuals of these plants are more palatable than low-latitude conspecifics. We documented plant traits and herbivore performance (body size) in the field across latitude. We collected individuals from different latitudes for factorial (plant region x herbivore region) laboratory experiments, examining how herbivore performance was affected by plant region, herbivore region, and their interaction (i.e., local adaptation). Field surveys suggested high-latitude plants were likely of higher quality to herbivores. Leaf nitrogen content in all plant species increased toward high latitudes, consistent with lower leaf C/N and higher leaf chlorophyll content at high latitudes. Furthermore, leaf toughness decreased toward higher latitudes in 1 species. The body size of 4 herbivore species increased with latitude, consistent with high-latitude leaves being of higher quality, while 2 grasshopper species showed the opposite pattern, likely due to life-history constraints. In the laboratory, high-latitude plants supported better performance in 4 herbivore species (marginal in the 5th). The geographic region where herbivores were collected affected herbivore performance in all 6 species; however, the pattern was mixed, indicating a lack of local adaptation by herbivores to plants from their own geographic region. Our results suggest that more-palatable plants at high latitudes support better herbivore growth. Given that geographic origin of either plants or herbivores can affect herbivore performance, the nature of plant-herbivore interactions is likely to change if climate change “reshuffles” plant and herbivore populations across latitude.

The Interactive Effects of Pulsed Grazing Disturbance and Patch Size Vary among Wetland Arthropod Guilds

Pulse disturbances and habitat patch size can determine community composition independently or in concert, and may be particularly influential on small spatial scales for organisms with low mobility. In a field experiment, we investigated whether the effects of a pulsed disturbance that simulated a grazing event varied with habitat patch size. We focused on the short-term responses of multiple co-occurring emergent salt marsh arthropods with differing levels of mobility and dispersal potential. As part of a marsh restoration project, two types of emergent marsh structures were created: small circular mounds (0.5 m diameter) separated by several meters of aquatic habitat, and larger, elongated terraces (>50 m long). Study plots (0.25 m2) were established on both structures; in a subset of plots, we simulated a pulsed grazing disturbance event by clipping the aboveground tissue of emergent plants, primarily Spartina alterniflora. At the end of the two-month recovery period, Ischnodemus (Hemiptera: Blissidae) density was over 50% lower in disturbed treatments within both large (terrace) and small (mound) patches. Predatory spider treatment responses were similar to Ischnodemus responses, suggesting a trophic relationship between those two arthropod groups. Alternatively, spiders may have been directly affected by the loss of shelter in the disturbed plots. Prokelisia (Homoptera: Delphacidae), which are generally more mobile than Ischnodemus, were not affected by disturbance treatment or by patch size, suggesting the potential for rapid recolonization following disturbance. Larval stem borers decreased by an order of magnitude in disturbed plots, but only in the large patches. In general, the disturbance effects of vegetation removal on arthropod density and community composition were stronger than patch size effects, and there were few interactions between pulsed disturbance and patch size. Rather, emergent marsh arthropod responses to disturbance and habitat area treatments were linked to the dispersal potential and mobility of each individual taxon.

Warming impact on herbivore population composition affects top-down control by predators

Understanding warming impact on herbivores facilitates predicting plant/crop dynamics in natural/agricultural systems. However, it remains unclear how warming will affect herbivore population size and population composition, consequently altering herbivore colonization in a tri-trophic system (plant-herbivore-predator or crop-pest-biocontrol agent). We studied a soybean-aphid-lady beetle system, by conducting (1) a laboratory warming experiment to examine warming impact (+2 °C or +4 °C) on the aphid population size and composition (alate proportion), and (2) a field colonization experiment to examine whether the warming-induced effect subsequently interacts with predators (lady beetles) in affecting aphid colonization. The results showed that warming affected the initial aphid population composition (reduced alate proportion) but not population size; this warming-induced effect strengthened the top-down control by lady beetles and slowing aphid colonization. In other words, biocontrol on crop pests by predators could improve under 2–4 °C warming. Furthermore, aphid colonization was affected by an interaction between the alate proportion and predator (lady beetle) presence. This study suggests that warming affects herbivore population composition and likely mediates top-down control on herbivore colonization by predators. This mechanism may be crucial but underappreciated in climate change ecology because population composition (wing form, sex ratio, age/body size structure) shifts in many species under environmental change.