Induced responses to competition and herbivory: natural selection on multi-trait phenotypic plasticity

Herbivory and competition are two of the most common biotic stressors for plants. When occurring simultaneously, responses to one interaction can constrain the induction of responses to the other interaction due to resource limitation and other interactive effects. Thus, to maximize fitness when interacting with competitors and herbivores, plants are likely to express particular combinations of plastic responses. This study reports the interactive effects of herbivory and competition on responses induced in Tithonia tubaeformis plants and describes how natural selection acts on particular plastic responses and on their different combinations. Competition induced a stem elongation response, expressed through an increase in height and mean internode length, together with a decrease in basal diameter. Interestingly, realized resistance increased in both competition and herbivory treatments, suggesting a plastic response in both constitutive and induced resistance traits. Particular combinations of plastic responses defined three plant phenotypes: vigorous, elongated, and resistant plants. The ecological context in which plants grew modified the traits and the particular combinations of plastic responses that were favored by selection. Vigorous plants were favored by selection in all environments, except when they were damaged by herbivores in the absence of neighbors. The combination of responses defining an elongated plant phenotype was favored by selection in crowded conditions. Resistance was negatively selected in the absence of competition and herbivory but favored in the presence of both interactions. In addition, contextual analyses detected that population structure in heterogeneous environments can also influence the outcomes of selection. These findings suggest that natural selection can act on particular combinations of plastic responses, which may allow plants to adjust their phenotypes to those that promote greater fitness under particular ecological conditions.     

Minimizing opportunity costs to aquatic connectivity restoration while controlling an invasive species

Controlling invasive species is critical for conservation but can have unintended consequences for native species and divert resources away from other efforts. This dilemma occurs on a grand scale in the North American Great Lakes, where dams and culverts block tributary access to habitat of desirable fish species and are a lynchpin of long‐standing efforts to limit ecological damage inflicted by the invasive, parasitic sea lamprey (Petromyzon marinus). Habitat restoration and sea‐lamprey control create conflicting goals for managing aging infrastructure. We used optimization to minimize opportunity costs of habitat gains for 37 desirable migratory fishes that arose from restricting sea lamprey access (0–25% increase) when selecting barriers for removal under a limited budget (US$1–105 million). Imposing limits on sea lamprey habitat reduced gains in tributary access for desirable species by 15–50% relative to an unconstrained scenario. Additional investment to offset the effect of limiting sea‐lamprey access resulted in high opportunity costs for 30 of 37 species (e.g., an additional US$20–80 million for lake sturgeon [Acipenser fulvescens]) and often required ≥5% increase in sea‐lamprey access to identify barrier‐removal solutions adhering to the budget and limiting access. Narrowly distributed species exhibited the highest opportunity costs but benefited more at less cost when small increases in sea‐lamprey access were allowed. Our results illustrate the value of optimization in limiting opportunity costs when balancing invasion control against restoration benefits for diverse desirable species. Such trade‐off analyses are essential to the restoration of connectivity within fragmented rivers without unleashing invaders.     

Hemiparasites generate environmental heterogeneity and enhance species coexistence in salt marshes.

Tidal inundation and salinity are considered to be controlling factors in salt marsh species distributions. Parasitic plants may also influence community organization as parasite-host interactions may play a functional role in stress amelioration due to physiological mechanisms for salinity tolerance and resource acquisition. Endangered root hemiparasites (Cordylanthus maritimus ssp. palustris and Cordylanthus mollis ssp. mollis) occupy unique habitat within fragmented northern California tidal wetlands. My objective was to examine the effects of these root hemiparasites on soil salinity, aeration, and community composition. I compared experimentally established bare patches, shaded and unshaded, and parasite removal patches to controls with hemiparasites across intertidal elevation gradients. Plant community composition, soil salinity, and redox potential were measured as response variables. In this field removal experiment, I demonstrated that parasite-host associations can enhance the amelioration of physical stress conditions in the salt marsh exceeding the passive role of shading by vegetation. Consumer-driven reduction of physical stress resulted in increased plant species richness, and the effect was most pronounced with elevated salinity and hypoxia stress. Although previous studies have demonstrated that removal of dominant plant biomass by herbivores can increase physical stress in salt marshes, this is one of the first examples of a positive indirect effect of a consumer on community diversity through physical stress relief. Greater understanding of biological interactions coupled with abiotic factors may improve rare plant conservation and salt marsh restoration success.     

Do plant density, nutrient availability, and herbivore grazing interact to affect phlorotannin plasticity in the brown seaweed Ascophyllum nodosum

Plants have different strategies to cope with herbivory, including induction of chemical defences and compensatory growth. The most favourable strategy for an individual plant may depend on the density at which the plants are growing and on the availability of nutrients, but this has not been tested previously for marine plant–herbivore interactions. We investigated the separate and interactive effects of plant density, nutrient availability, and herbivore grazing on the phlorotannin (polyphenolic) production in the brown seaweed Ascophyllum nodosum. Seaweed plants grown at low or high densities were exposed either to nutrient enrichment, herbivorous littorinid gastropods (Littorina obtusata), or a combination of nutrients and herbivores in an outdoor mesocosm experiment for 2 weeks. Seaweeds grown at a low density tended to have higher tissue nitrogen content compared to plants grown at a high density when exposed to elevated nutrient levels, indicating that there was a density dependent competition for nitrogen. Herbivore grazing induced a higher phlorotannin content in plants grown under ambient, but not enriched, nutrient levels, indicting either that phlorotannin plasticity is more costly when nutrients are abundant or that plants responded to herbivory by compensatory growth. However, there were no significant interactive or main effects of plant density on the seaweed phlorotannin content. The results indicate that plants in both high and low densities induce chemical defence, and that eutrophication may have indirect effects on marine plant–herbivore interactions through alterations of plant chemical defence allocation.     

Ecological differences and coexistence in a guild of microparasites: Bartonella in wild rodents

The study of ecological differences among coexisting microparasites has been largely neglected, but it addresses important and unusual issues because there is no clear distinction in such cases between conventional (resource) and apparent competition. Here patterns in the population dynamics are examined for four species of Bartonella (bacterial parasites) coexisting in two wild rodent hosts, bank voles (Clethrionomys glareolus) and wood mice (Apodemus sylvaticus). Using generalized linear modeling and mixed effects models, we examine, for these four species, seasonal patterns and dependencies on host density (both direct and delayed) and, having accounted for these, any differences in prevalence between the two hosts. Whereas previous studies had failed to uncover species differences, here all four were different. Two, B. doshiae and B. taylorii, were more prevalent in wood mice, and one, B. birtlesii, was more prevalent in bank voles. B. birtlesii, B. grahamii, and B. taylorii peaked in prevalence in the fall, whereas B. doshiae peaked in spring. For B. birtlesii in bank voles, density dependence was direct, but for B. taylorii in wood mice density dependence was delayed. B. birtlesii prevalence in wood mice was related to bank vole density. The implications of these differences for species coexistence are discussed.     

入侵种喜旱莲子草对土壤养分的表型可塑性反应

对外来入侵种喜旱莲子草（Alternanthera philoxeroides（Mart．）Griseb．）在不同土壤氮养分处理中的表型可塑性反应进行了研究。结果表明：土壤中氮养分水平对喜旱莲子草克隆生长有显著影响，如分枝强度、苇节长度随氮含量升高均有不同程度的增加；氮养分最高的处理中根系生物量投资最小，而茎、叶生物量投资最大，对照处理中则表现山根冠比最大。研究结果表明，在不同土壤养分条件下，喜旱莲子草不同表型结构有可能同时对环境选择作出反应，在种群生存与维持、生长、繁殖等功能方面实现种群个体各器官生物量投资的优化配置来适应多样化的环境，以增强其入侵能力，     

Trade-offs, temporal variation, and species coexistence in communities with intraguild predation

Intraguild predation/parasitism (IGP: competing species preying on or parasitizing each other) is widespread in nature, but the mechanisms by which intraguild prey and predators coexist remain elusive. Theory predicts that a trade-off between resource competition and IGP should allow local niche partitioning, but such trade-offs are expressed only at intermediate resource productivity and cannot explain observations of stable coexistence at high productivity. Coexistence must therefore involve additional mechanisms beside the trade-off, but very little is known about the operation of such mechanisms in nature. Here I present the first experimental test of multiple coexistence mechanisms in a natural community exhibiting IGP. The results suggest that, when resource productivity constrains the competition-IGP trade-off, a temporal refuge for the intraguild prey can not only promote coexistence, but also change species abundances to a pattern qualitatively different from that expected based on the trade-off or a refuge alone. This is the first empirical study to demonstrate a mechanism for why communities with IGP do not lose species diversity in highly productive environments. These results have implications for diversity maintenance in multi-trophic communities, and the use of multiple natural enemies in biological control.     

Morphological variation and phenotypic plasticity of buoyancy in the macroalga Turbinaria ornata across a barrier reef

Many aspects of morphology of benthic algae (length, surface area-to-volume ratio, and blade undulation) are plastic traits that vary in response to physical factors (such as light or water flow environment). This study examines whether frond buoyancy is a plastic trait, and whether differences in morphology including buoyancy affect the potential persistence of macroalgae in habitats characterized by different water flow regimes. Fronds of the tropical alga Turbinaria ornata in protected backreef environments in Moorea, French Polynesia possess pneumatocysts (gas-filled floats) and experience positive buoyant forces, whereas fronds in wave-exposed forereef sites either lack pneumatocysts entirely or have very small, rudimentary pneumatocysts and experience negative buoyant forces. Forereef fronds transplanted to the backreef developed pneumatocysts and experienced increased buoyant force indicating that buoyancy is a phenotypically plastic trait in T. ornata. In comparing the potential for dislodgement by drag, drag was greater on forereef fronds at low flow speeds as these fronds were stiffer and did not bend over at low flow speeds and therefore were less streamlined in the flow than backreef algae, which bent easily. The environmental stress factor (ESF) (a measure of the likelihood of detachment for a frond in its habitat) was higher for forereef than backreef fronds at all flow speeds. When examined with respect to the flow velocities likely in their respective habitats however, the chance of detachment for backreef and forereef was similar. Neither backreef nor forereef fronds were predicted to break under normal, non-storm conditions, but both were predicted to break in storms. Strong forereef morphologies are well suited to habitats characterized by rapid flow, whereas the weaker, buoyant, tall backreef fronds are well suited to habitats where crowding and shading is common but hydrodynamic forces are low.     

Regional coexistence and local dominance in Chaoborus: species sorting along a predation gradient

Variation in the intensity of predation across the well-known environmental gradient of freshwater habitats from small, ephemeral ponds to large, permanent lakes is a key factor in the development and maintenance of aquatic community structure. Here, we present data on the distribution and abundance of four species of Chaoborus (Diptera: Chaoboridae) across this environmental gradient. Chaoborus show a distinct pattern of species sorting when aquatic systems are divided into fish and fishless environments, and this pattern is consistent with species traits known to influence their vulnerability to fish predation (i.e., pigmentation, diel vertical migration [DVM] behavior, and body size). To test whether fish are the drivers of this pattern, we created a gradient in fish density by stocking bluegill sunfish (Lepomis macrochirus) into 15 experimental ponds in southwestern Michigan, USA, and then allowed Chaoborus species to colonize. There was clear evidence of species sorting along the predation gradient; Chaoborus americanus was most abundant in the fishless ponds, C. flavicans was neutral in response to fish, and C. punctipennis and C. albatus were most abundant at high fish biomass, a response consistent with their field pattern. Furthermore, prey preference experiments confirm that size selective predation and differences in Chaoborus species traits contribute to the pattern of Chaoborus abundance and distribution.     

Soil vs. canopy seed storage and plant species coexistence in species-rich Australian shrublands

The fire-prone shrublands of southwestern Australia are renowned for their high plant species diversity and prominence of canopy seed storage (serotiny). We compared species richness, abundance, and life history attributes for soil and canopy seed banks in relation to extant vegetation among four sites with different substrate conditions and high species turnover (50-80%) to identify whether this unusual community-level organization of seed storage might contribute to maintenance of high species richness. Soil seed bank (SSB) densities were low to moderate (233-1435 seeds/m2) compared with densities for other Mediterranean-type vegetation and were lowest for sites with highest canopy seed bank (CSB) species richness and lowest nutrient availability, but not richness or abundance of resprouters. Annuals were infrequent in the lowest nutrient sites, but there was no evidence that small SSB size was due to low seed inputs or a trade-off between seed production/storage and seed size in response to low nutrient availability. Sorensen's similarity between SSB and extant vegetation was 26-43% but increased to 54-57% when the CSB was included, representing levels higher than reported for most other ecosystems. Resprouting species were well represented in both the SSB and CSB, and there was no evidence for lower seed production in resprouters than in non-sprouters overall. The SSB and CSB held no species in common and were characterized by markedly different seed dispersal attributes, with winged or small seeds in the CSB and seeds dispersed by ants, birds, and wind (though none with wings) in the SSB. There was no evidence of spatial differentiation in the distribution of seeds of SSB species between vegetated and open microsites that might facilitate species coexistence, but most woody non-sprouters showed aggregation at scales of 1-2 m, implying limited seed dispersal. High similarity between overall seed bank (SSB + CSB) and extant species composition, high number of resprouting species, and seed dispersal processes before (SSB) and after fire (CSB) leading to differential spatial aggregation of post-fire recruits from the two seed bank types may buffer species composition against rapid change and provide a mechanism for maintaining species coexistence at the local scale.     

No reproductive character displacement in male advertisement signals of Hyla japonica in relation to the sympatric H. suweonensis

Reproductive interaction between closely related taxa may leave a distinctive signature in which populations of interacting taxa are more dissimilar in sympatry than in allopatry. An ideal condition for such a pattern of reproductive character displacement (RCD) may occur when a population has limited gene flow and experiences strong selection pressure, exerted by an interacting taxon in areas of sympatry. In Korea, there are two closely related treefrog species: Hyla japonica, which is distributed widely throughout the country, and Hyla suweonensis, which occurs sympatrically in a narrow strip of western coastal plains. H. suweonensis is only found within the distribution of H. japonica. These two species have a similar single-note call structure. Here, we tested the possibility of RCD in H. japonica by examining geographic variation in advertisement calls. Although means of temporal and spectral characters were significantly different between the two species, sympatric populations of H. japonica and H. suweonensis overlapped in distributions of most characters. Furthermore, allopatric and sympatric H. japonica populations did not differ in all call characters. Weak genetic differentiation between sympatric and allopatric populations of H. japonica implied either substantial gene flow or recent genetic isolation. Possible explanations for no RCD in male advertisement calls of H. japonica include a difference in fine temporal characteristics between the two species, migration between sympatric and allopatric localities in H. japonica, RCD in female preferences in H. japonica, and weak selection pressure by H. suweonensis.     

Divergent effects of climate change on the egg-laying opportunity of species in cold and warm regions

Climate warming can substantially impact embryonic development and juvenile growth in oviparous species. Estimating the overall impacts of climate warming on oviparous reproduction is difficult because egg-laying events happen throughout the reproductive season. Successful egg laying requires the completion of embryonic development as well as hatching timing conducive to offspring survival and energy accumulation. We propose a new metric—egg-laying opportunity (EO)—to estimate the annual hours during which a clutch of freshly laid eggs yields surviving offspring that store sufficient energy for overwintering. We estimated the EO within the distribution of a model species, Sceloporus undulatus, under recent climate condition and a climate-warming scenario by combining microclimate data, developmental functions, and biophysical models. We predicted that EO will decline as the climate warms at 74.8% of 11,407 sites. Decreasing hatching success and offspring energy accounted for more lost EO hours (72.6% and 72.9%) than the occurrence of offspring heat stress (59.9%). Nesting deeper (at a depth of 12 cm) may be a more effective behavioral adjustment for retaining EO than using shadier (50% shade) nests because the former fully mitigated the decline of EO under the considered warming scenario at more sites (66.1%) than the latter (28.3%). We advocate for the use of EO in predicting the impacts of climate warming on oviparous animals because it encapsulates the integrative impacts of climate warming on all stages of reproductive life history.     

Modeling outcomes of approaches to sustained human and snow leopard coexistence

The snow leopard (Uncia uncia) is in danger of extinction. Killing to protect livestock is among the primary causes of its decline. Efforts to mitigate this threat have focused on balancing the need to conserve the snow leopard with the needs of local people in snow leopard habitat, many of whom rely on raising livestock for their livelihoods. Conservation of the snow leopard has the characteristics of a public good, and outside funding is required to support conservation efforts. There are 5 commonly discussed approaches to resolving this issue: (1) direct payments for conservation, (2) investments in protection from predation, (3) damage compensation payments, (4) investments in better livestock husbandry, and (5) leases of pastureland for wild prey. After a review of these 5 conservation strategies, an economic–ecologic model, which includes the interactions between the snow leopard, its wild prey, and livestock, is used to evaluate the 2 most promising conservation strategies. The model reveals that investments in protection from predation and leases of pastureland for wild prey are effective but only in delaying the eventual extinction of the snow leopard. To preserve the snow leopard, these approaches must be applied more aggressively and new ones explored.     

Abundance and survival rates of green turtles in an urban environment: coexistence of humans and an endangered species

Longitudinal capture-mark-recapture data were used to estimate abundance and survival rates for green turtles (Chelonia mydas) in San Diego Bay, California, USA. These turtles were closely associated with warm effluent from a power plant during winter months. The life stage distribution of green turtles in the bay ranged from post-pelagic juveniles to adults (44.0–110.4 cm straight carapace length). During 99 capture sessions between December 2, 1990, and March 25, 2009, 96 individual green turtles were caught. To estimate abundance and survival rates, robust-design mark-recapture models were fitted to capture-recapture histories using software MARK. The estimated annual survival rate was 0.861 (SE = 0.147, 95% CI = 0.356–0.986), whereas annual abundance ranged from 16 (SE = 6.3, 95% CI = 4–29) to 61 (SE = 13.2, 95% CI = 36–88). This study provides the first survival rate and abundance estimates for a green turtle foraging population in the highly industrialized San Diego Bay.     

Ecological and physiological controls of species composition in green macroalgal blooms

Green macroalgal blooms have substantially altered marine community structure and function, specifically by smothering seagrasses and other primary producers that are critical to commercial fisheries and by creating anoxic conditions in enclosed embayments. Bottom-up factors are viewed as the primary drivers of these blooms, but increasing attention has been paid to biotic controls of species composition. In Washington State, USA, blooms are often dominated by Ulva spp. intertidally and Ulvaria obscura subtidally. Factors that could cause this spatial difference were examined, including competition, grazer preferences, salinity, photoacclimation, nutrient requirements, and responses to nutrient enrichment. Ulva specimens grew faster than Ulvaria in intertidal chambers but not significantly faster in subtidal chambers. Ulva was better able to acclimate to a high-light environment and was more tolerant of low salinity than Ulvaria. Ulvaria had higher tissue N content, chlorophyll, chlorophyll b: chlorophyll a, and protein content than Ulva. These differences suggest that nitrogen availability could affect species composition. A suite of five grazers preferred Ulva to Ulvaria in choice experiments. Thus, bottom-up factors allow Ulva to dominate the intertidal zone while resistance to grazers appears to allow Ulvaria to dominate the subtidal zone. While ulvoid algae are in the same functional-form group, they are not functionally redundant.     

不同底质和透明度下马来眼子菜的表型可塑性研究

马来眼子菜(Potamogeton malaianusMiq.)是目前太湖沉水植物优势种之一。文章比较分析了3种不同底质和透明度情况下马来眼子菜的生长特征,探索其在太湖不断扩展的原因及其适应性。分别选取粘土质粉砂、粉砂和下蜀黄土底质上生长的马来眼子菜进行观察试验。结果表明,这3种底质在粒度组成、营养盐和分布特征上具有显著差异。在太湖粘土质粉砂和粉砂底质上马来眼子菜的生物量比下蜀黄土底质上的马来眼子菜高,其生物量随着底质营养盐的增加而增加。马来眼子菜与觅光相关的形态指标,如高度、节间距、节数、叶数、叶长及叶面积均随着底质营养盐的增加而显著增加。马来眼子菜的冠层高度与水体的透明度呈显著负相关,表明其对剧烈变化的湖泊光环境具有很强的适应能力。马来眼子菜在不同底质上的形态可塑性是其优先占据湖泊资源成为优势种的重要原因。     

Seed arrival and ecological filters interact to assemble high-diversity plant communities

Two prominent mechanisms proposed to structure biodiversity are niche-based ecological filtering and chance arrival of propagules from the species pool. Seed arrival is hypothesized to play a particularly strong role in high-diversity plant communities with large potential species pools and many rare species, but few studies have explored how seed arrival and local ecological filters interactively assemble species-rich communities in space and time. We experimentally manipulated seed arrival and multiple ecological filters in high-diversity, herbaceous-dominated groundcover communities in longleaf pine savannas, which contain the highest small-scale species richness in North America (up to > 40 species/m2). We tested three hypotheses: (1) local communities constitute relatively open-membership assemblages, in which increased seed arrival from the species pool strongly increases species richness; (2) ecological filters imposed by local fire intensity and soil moisture influence recruitment and richness of immigrating species; and (3) ecological filters increase similarity in the composition of immigrating species. In a two-year factorial field experiment, we manipulated local fire intensity by increasing pre-fire fuel loads, soil moisture using rain shelters and irrigation, and seed arrival by adding seeds from the local species pool. Seed arrival increased species richness regardless of fire intensity and soil moisture but interacted with both ecological filters to influence community assembly. High-intensity fire decreased richness of resident species, suggesting an important abiotic filter. In contrast, high-intensity fire increased recruitment and richness of immigrating species, presumably by decreasing effects of other ecological filters (competition and resource limitation) in postfire environments. Drought decreased recruitment and richness of immigrating species, whereas wet soil conditions increased recruitment but decreased or had little effect on richness. Moreover, some ecological filters (wet soil conditions and, to a lesser extent, high-intensity fire) increased similarity in the composition of immigrating species, illustrating conditions that influence deterministic community assembly in species-rich communities. Our experiment provides insights into how dispersal-assembly mechanisms may interact with niche-assembly mechanisms in space (spatial variation in disturbance) and time (temporal variation in resource availability) to structure high-diversity communities and can help guide conservation of threatened longleaf pine ecosystems in the face of habitat fragmentation and environmental change.     

Contest behaviour in the speckled wood butterfly (Pararge aegeria): seasonal phenotypic plasticity and the functional significance of flight performance

Although contemporary animal contest theory emphasises the importance of physical asymmetries in resolving disputes, such asymmetries do not obviously settle fights in all groups. Territorial male butterflies, for example, compete via elaborate non-contact aerial interactions in which success is determined by relative persistence. Prior research suggests that the resolution of these contests is not clearly related to physical variables such as body size or energy reserves. However, given that the contests involve elaborate aerial manoeuvres, one long-standing suggestion is that asymmetries in flight performance, and thus flight morphology, may be important. We addressed this hypothesis via a manipulative investigation into the biophysical correlates of contest success in the speckled wood butterfly, Pararge aegeria. This species possesses the ability for significant adaptive phenotypic plasticity in relevant flight morphological parameters. We took advantage of this plasticity to rear 90 individuals of markedly varying flight morphologies, which we then pitted against each other in a semi-controlled experimental fashion. Multiple logistic and lognormal analyses provided little evidence for the relevance of morphological parameters, including relative flight musculature, wing loading and wing aspect ratio (wing length relative to area), to the outcome and/or duration of experimental contests. Instead, we found a positive effect of age upon contest success. Given that ability for high acceleration is strongly linked to variation in these morphological parameters, our findings suggest that flight performance is not a strong determinant of resource-holding potential in this notably territorial butterfly.     

Patterns in soil fertility and root herbivory interact to influence fine-root dynamics

Fine-scale soil nutrient enrichment typically stimulates root growth, but it may also increase root herbivory, resulting in trade-offs for plant species and potentially influencing carbon cycling patterns. We used root ingrowth cores to investigate the effects of microsite fertility and root herbivory on root biomass in an aggrading upland forest in the coastal plain of South Carolina, USA. Treatments were randomly assigned to cores from a factorial combination of fertilizer and insecticide. Soil, soil fauna, and roots were removed from the cores at the end of the experiment (8-9 mo), and roots were separated at harvest into three diameter classes. Each diameter class responded differently to fertilizer and insecticide treatments. The finest roots (< 1.0 mm diameter), which comprised well over half of all root biomass, were the only ones to respond significantly to both treatments, increasing when fertilizer and when insecticide were added (each P < 0.0001), with maximum biomass found where the treatments were combined (interaction term significant, P < 0.001). These results suggest that root-feeding insects have a strong influence on root standing crop with stronger herbivore impacts on finer roots and within more fertile microsites. Thus, increased vulnerability to root herbivory is a potentially significant cost of root foraging in nutrient-rich patches.