Costs of inducible defence along a resource gradient

In addition to having constitutive defence traits, many organisms also respond to predation by phenotypic plasticity. In order for plasticity to be adaptive, induced defences should incur a benefit to the organism in, for example, decreased risk of predation. However, the production of defence traits may include costs in fitness components such as growth, time to reproduction, or fecundity. To test the hypothesis that the expression of phenotypic plasticity incurs costs, we performed a common garden experiment with a freshwater snail, Radix balthica, a species known to change morphology in the presence of molluscivorous fish. We measured a number of predator-induced morphological and behavioural defence traits in snails that we reared in the presence or absence of chemical cues from fish. Further, we quantified the costs of plasticity in fitness characters related to fecundity and growth. Since plastic responses may be inhibited under limited resource conditions, we reared snails in different densities and thereby levels of competition. Snails exposed to predator cues grew rounder and thicker shells, traits confirmed to be adaptive in environments with fish. Defence traits were consistently expressed independent of density, suggesting strong selection from predatory molluscivorous fish. However, the expression of defence traits resulted in reduced growth rate and fecundity, particularly with limited resources. Our results suggest full defence in predator related traits regardless of resource availability, and costs of defence consequently paid in traits related to fitness.     

Invertebrate food webs along a stream resource gradient

1. The flow of energy through food webs with similar species can vary with both space and time. The river continuum concept (RCC) provides a useful framework for predicting variability in the biota and food availability along streams. We estimated the flow of organic matter (g m^?2 year^?1) through food webs, arrayed along a stream, that had different resource inputs. Four sites were sampled along the Little Tennessee River, North Carolina, U.S.A.: two fifth order sites, one sixth and one seventh order site. The dominant resource is leaf detritus in the upstream reach (the upstream fifth order site), algae in the mid‐reaches (the downstream fifth and sixth order sites), and suspended material downstream (seventh order site). 2. Eleven genera, contributing from 50 to 66% of the total macroinvertebrate secondary production of each site, were studied. We estimated organic matter flow from resource to consumer by combining previously measured rates of invertebrate secondary production with gut content analyses and assimilation efficiencies. 3. Organic matter flow through food webs increased in a downstream direction, while the structure of the food webs remained constant. The total food consumed by the taxa analysed increased from 34 g m^?2 year^?1 at the upstream site to 730 g m^?2 year^?1 at the most downstream site. We estimate that the organic matter consumed by the entire macroinvertebrate community ranged from 66 to 1164 g m^?2 year^?1. These results indicate that there is variation in the magnitude of organic matter flow through the food webs along this river continuum. 4. The dominant food resource consumed also changed along the gradient. Leaf detritus consumption decreased from 58% of the total consumption upstream to 6% downstream, whereas consumption of amorphous detritus increased from 18 to 64%. The proportion of animal material consumed also increased from 3 to 27%. The total consumption of autochthonous resources (diatoms and filamentous algae) increased along the continuum (from 6.41 to 34.05 g m^?2 year^?1). We conclude that these results are related to variation in resource availability, dietary shifts and invertebrate secondary production. These results link resource availability to energy flow, a relationship originally suggested by the RCC.     

Competition intensity along a productivity gradient in a low-diversity grassland

Competition among plants often shifts from roots to shoots as productivity increases and species composition changes. We examined competition in an old field with low diversity to test whether this shift occurred along a productivity gradient without species turnover. Forty plots received one of four nitrogen treatments (0, 5, or 15 g added N m-2 yr-1 or 400 g m-2 yr-1 of sawdust added to immobilize N) annually for 5 yr. All N levels were dominated by the perennial grasses Agropyron cristatum and Bromus inermis. Transplants of Agropyron were grown with all neighbors, roots of neighbors, or no neighbors present to measure total, root, and shoot competition. Transplant growth was 22%-165% higher in subplots without neighbors present, which indicates that competition occurred. Competition from neighbor roots was primarily responsible for suppression of transplant growth over the entire productivity gradient. In contrast to previous field experiments that found either an increase in total competition intensity or a shift from root to shoot competition with increasing productivity, we found neither. Increases in total competition intensity or shifts from root to shoot competition found along other gradients may be caused by changes in species composition and not by increased resources or neighbor biomass. These results suggest that different competitive mechanisms may operate in low-diversity vegetation than in more diverse natural vegetation.     

Ant foraging strategies vary along a natural resource gradient

Food selection by foragers is sensitive to the availability of resources, which may vary along geographical gradients. Hence, selectivity of food types by foragers is expected to track these resource gradients. Here we addressed this hypothesis and asked if foraging decisions of seed-eating ants differ along a geographic gradient of habitat productivity. The study was carried out for two years at five sites along a natural climatic gradient, ranging from arid to Mediterranean, where plant productivity varies six-fold across a short geographic distance of 250 km. We found that in ant colonies of the genus Messor, collective foraging decisions differed along the gradient. Specifically, at the high-productivity sites, a stronger association was found between plant seed availability and selectivity, suggesting that colonies respond more accurately to within-patch variation in food amounts. In contrast, colonies in low-productivity sites foraged in patches with higher concentration of seeds, suggesting that they respond more accurately to among-patch variation in food amounts. Moreover, at the high-productivity sites, colonies were more discriminating in their choice of food and preferred bigger seeds, while in the low-productivity sites, where smaller seeds were relatively more abundant, food collection depended mostly on seed availability. An experiment with artificial seed patches performed along the same climatic gradient, revealed no difference in food selectivity across sites when food type and availability were similar, and a general preference for bigger over medium-sized seeds. Overall, our findings suggest that resource availability is an important factor explaining food choice along a climatic gradient and imply that in low-productivity regions small-seeded species incur high predation pressure, whereas in high-productivity regions, large-seeded species suffer higher predation. This could have important consequences for plant species composition, particularly at the face of climate change, which could dramatically alter the foraging decisions of granivores.     

Competition and coexistence of rhizomatous perennial plants along a nutrient gradient

I studied competition and coexistence of three tall clonal perennial plant species, Calamagrostis epigejos (L.) Roth, Solidago canadensis L., and Tanacetum vulgare L. along a gradient of soil productivity over five years. A replacement series field experiment was conducted with high, moderate and low fertility levels in 1m×1m plots. There were significant effects of soil type on ramet density (P<0.001), mean height (P<0.01), and total biomass (P<0.01). Ramet density, mean height, and total biomass increased with increasing soil fertility. There were also significant effects of mixture on ramet density (P<0.01), but not on mean height and total biomass for all species. Significant neighbor effects on ramet density and total biomass (P<0.01) were found for Solidago, showing that it is important whether Tanacetum or Calamagrostis is its neighbor within mixtures. During the five years there was only one case of competitive exclusion: Calamagrostis excluded Solidago on the most fertile substrate in the fifth growing season. In most cases species coexisted over the five years. Each of the three species was able to dominate in at least one combination of substrate type and mixture. The experiment showed that asymmetric competition for light on substrates of high fertility, symmetric competition for nutrients on nutrient-poor soil and positive interactions especially on substrates of intermediate fertility played a role. A founder effect was evident in aggregated mixtures of Calamagrostis and Solidago on the nutrient-rich substrate. A conceptual model of the relative importance of root competition for soil nutrients, shoot competition for light, and positive interactions along the fertility gradient is presented. The model emphasizes that positive interactions play an important role over a broad range of the productivity scale with a peak at intermediate levels of fertility. On the substrate of high productivity shoot competition for light is more important than positive interactions and root competition for soil nutrients as well. The competitive superiority of Calamagrostis on the most productive substrate was evident only in the long run. Rare events like extreme summer drought or selective herbivore pressure caused a switch in dominance in mixtures with Solidago, respectively Tanacetum. The guerrilla growth strategy of Calamagrostis and interference competition through a dense cover of aboveground biomass and litter could further cause competitive exclusion.     

Competition between Australian native and introduced grasses along a nutrient gradient

Abstract Seven grass species were grown in monocultures and in multispecies mixtures along a gradient of total nutrient levels that ranged from 1/64 to 16× the normal level of nutrient solution. The seven grasses represented three ecological groups: (i) three perennial species native to Australia (Themeda triandra, Poa labillardieri and Danthonia carphoides); (ii) two introduced annuals (Vulpia bromoides and Hordeum leporinum); and (iii) two introduced perennials (Lolium perenne and Dactylis glomerata). We hypothesized that the native grasses would prove less competitive when grown at increased nutrient levels than those introduced from Europe. Results supported the hypothesis. The native species were unable to compete in mixtures even at the lowest nutrient level, where T. triandra was the most productive species in monoculture. Lolium perenne and Dactylis glomerata dominated mixtures at intermediate nutrient levels. The responses of the annual introduced grasses differed in that Vulpia bromoides showed an optimum at intermediate nutrient levels in both monoculture and in mixtures, whereas Hordeum leporinum dominated at the highest nutrient levels in mixture but was suppressed by V. bromoides, L. perenne and D. glomerata at intermediate levels. The results are discussed in terms of predicting species responses in mixtures from their performance in monocultures as well as in terms of previous observations on the sequential changes in botanical composition of south‐eastern Australian grasslands after 150 years of continuous grazing by sheep.     

Competition between Salicornia europaea and Atriplex prostrata (Chenopodiaceae) along an experimental salinity gradient

Salicornia europaea L. is a halophyte that often occupies the lowestand most saline (>3.5% total salt) areas of salt marshes. Atriplexprostrata Boucher is less salt tolerant than S. europaea and oftengrows in a less saline (<2.0% total salts) zone adjacent to S. europaea. The purpose of this experiment was to determine thecompetitive outcome when these two species are grown at differentsalinities to ascertain the extent salinity and competition affect plantzonation. Plants were grown in a de Wit replacement series at 85, 170,and 340 mM NaCl in half-strength Hoagland's no. 2 nutrient solution fortwo months. There was a significant effect of salt concentration,competition, and their interaction on biomass production of S. europaea plants. However, only salt concentration significantly affectedbiomass production of A. prostrata plants. Results of thisexperiment confirmed the results of other studies that demonstrated thatthe more salt tolerant species were less competitive at lower salinities. Atriplex prostrata was the better competitor at 85 mM NaCl, whereasS. europaea was the better competitor at 340 mM NaClbecause growth of A. prostrata was inhibited. At 170 mMNaCl, A. prostrata biomass production decreased more than S. europaea biomass in mixed culture.     

Porcupine disturbances and vegetation pattern along a resource gradient in a desert

Summary In the Negev Desert, Israel, the Indian crested porcupine, Hystrix indica, digs similar sized, discrete, elongated pits (257±21.3 cm³; n=144) while foraging for below-ground plant storage organs. In these digs, soil moisture content is higher than in the surrounding soil matrix. The digs disturb population and community structure due to porcupine consumption or damage of 18 species of plants, and repopulation by 55 plant species. Over the past 14 years we have studied dig dynamics on a rocky hillslope with three distinct habitats as regards soil moisture content. Midslope soil moisture is the highest, decreasing towards upper and lower slope. We have counted a total of 6,609 digs in the area: 2141 on the upper, 3211 in the middle and 1257 on the lower part of the slope. The number of digs at midslope is significantly higher than on the other parts of the slope (ANOVA; P<0.0001). There is a significant (P<0.05) correlation between the mean number of porcupine digs and the cumulative rainfall amount for the 2 years prior to dig formation. To study plant repopulation in digs, all plants in 144 digs along the slope and from equal sized plots in the undistarbed soil matrix were collected. In all, 288 samples with 20 584 plants were collected, 2042 from the matrix and 18,542 from digs. Of the 55 species, we focused on the abundance patterns of Filago desertorum, Picris cyanocarpa and Bromus rubens, which made up 69.5% of all the individuals in the digs and 68.3% in the matrix. Our results showed that all three species increased in abundance in the digs as compared to the matrix. F. desertorum density increased by a factor of 2.9, P. cyanocarpa by 9.5 and B. rubens by 12.0. There were species-specific responses in abundance to the location of the digs along the moisture gradient. The only species whose abundance responded to the moisture gradient was F. desertorum. P. cyanocarpa demonstrated peak abundance in the location with the poorest moisture regime, while B. rubens showed peak abundance at the intermediate part of the moisture gradient. We suggest a scheme for integrating the increase in density and the species-specific responses to the digs along a water gradient based on R.H. Whittaker's view of individual species abundances along an environmental gradient.     

The gradostat: A model of competition along a nutrient gradient

The general mathematical theory of the gradostat is presented for two competitors. The gradostat provides a mechanism for studying competition along a nutrient gradient. In the two vessel case, the results are complete and the conditions are testable. In then-vessel case, the relevant conditions are stated in terms of the stability modulii of certain matrices and are testable for any specific case.     

Species richness in a model with resource gradient

In order to study the dependence of the species richness on heterogeneity of the habitat, we introduce an extended model of annual plants which combines the features of the island model and of gradient heterogeneity resources. First, we consider a native population of plants living on a square lattice of linear size L. After equilibration of this native population, seeds of several different species j = 2, ... , k of annual plants invade the system; they compete among themselves and the native ones. The system is exposed to a one-dimensional water gradient, and each species is characterised by a tolerance to a surplus of water, τ(j). We study the influences of the properties of the gradient of the resource (GR) on the species richness (SR) present in the system. We have shown that the relation between GR and SR is not straightforward and that several cases could be distinguished: For a large class of control parameters, SR increases linearly with GR. However, when the values of the control parameters are such as to create wide inhabitable regions, the relation between SR and GR ceases to have a monotonic character. We have also demonstrated that the average species richness as a function of the resource availability has a hump shape. Our results can be simply explained within our model and are in agreement with several previous field and theoretical works.     

Differences in hyporheic-zone microbial community structure along a heavy-metal contamination gradient

The hyporheic zone of a river is nonphotic, has steep chemical and redox gradients, and has a heterotrophic food web based on the consumption of organic carbon entrained from downwelling surface water or from upwelling groundwater. The microbial communities in the hyporheic zone are an important component of these heterotrophic food webs and perform essential functions in lotic ecosystems. Using a suite of methods (denaturing gradient gel electrophoresis, 16S rRNA phylogeny, phospholipid fatty acid analysis, direct microscopic enumeration, and quantitative PCR), we compared the microbial communities inhabiting the hyporheic zone of six different river sites that encompass a wide range of sediment metal loads resulting from large base-metal mining activity in the region. There was no correlation between sediment metal content and the total hyporheic microbial biomass present within each site. However, microbial community structure showed a significant linear relationship with the sediment metal loads. The abundances of four phylogenetic groups (groups I, II, III, and IV) most closely related to alpha-, beta-, and gamma-proteobacteria and the cyanobacteria, respectively, were determined. The sediment metal content gradient was positively correlated with group III abundance and negatively correlated with group II abundance. No correlation was apparent with regard to group I or IV abundance. This is the first documentation of a relationship between fluvially deposited heavy-metal contamination and hyporheic microbial community structure. The information presented here may be useful in predicting long-term effects of heavy-metal contamination in streams and provides a basis for further studies of metal effects on hyporheic microbial communities.     

Fine-scale spatial organization of tallgrass prairie vegetation along a topographic gradient

Fine-scale spatial patterns of native tallgrass prairie vegetation were studied on Konza Prairie, Kansas, USA. Three sites, upland, slope, and lowland, were sampled in an ungrazed watershed. Presence of vascular plant species was recorded in two 25.6 m long transects of contiguous 5×5 cm micro-quadrats on each topographic position. Spatial patterns of species and functional groups were analyzed by information theory models ofJuhász-Nagy. Within-community variability of coexistence was expressed by the diversity and spatial dependence of local species combinations. Considerable diversity in the local coexistence of species was found on each site. Upland and hillside communities were richer and more diverse in species combinations than lowland. Spatial scale effected coexistence relationships. The maxima of information theory estimates varied between 15 and 30 cm. There was no trend in the variation of characteristic scales along the topographical gradient. Above 10 m, all sites tended to be homogeneous. The analysis of spatial associations revealed that variability in the local coexistence of species was strongly constrained in all topographic positions. Overall spatial association of species was the lowest on lowland. The characteristic scales of maximum association were between 1.2 m and 3 m at all sites. The maxima of information theory estimates for the functional group-based data appeared at smaller plot sizes than for the species based analyses. Only weak spatial associations were detected among the functional groups indicating that individuals of functional groups coexist well at small scales, and form combinations close to random expectations. The length of transects did not effect the relative associations. Strong positive correlations were found between the number of components (species or functional groups) and the maxima, of information theory models suggesting that richness is a good predictor of within-community coexistence relations. However, there was no relationship between richness and the characteristic scales of community patterns.     

Investment in defense and cost of predator-induced defense along a resource gradient

An organism’s investment in different traits to reduce predation is determined by the fitness benefit of the defense relative to the fitness costs associated with the allocation of time and resources to the defense. Inherent tradeoffs in time and resource allocation should result in differential investment in defense along a resource gradient, but competing models predict different patterns of investment. There are currently insufficient empirical data on changes in investment in defensive traits or their costs along resource gradients to differentiate between the competing allocation models. In this study, I exposed tadpoles to caged predators along a resource gradient in order to estimate investment in defense and costs of defense by assessing predator-induced plasticity. Induced defenses included increased tail depth, reduced feeding, and reduced swimming activity; costs associated with these defenses were reduced developmental rate, reduced growth, and reduced survival. At low resource availability, these costs predominately resulted in reduced survival, while at high resource availability the costs yielded a reduced developmental rate. Defensive traits responded strongly to predation risk, but did not respond to resource availability (with the exception of feeding activity), whereas traits construed as costs of defenses showed the opposite pattern. Therefore, defensive traits were highly sensitive to predation risk, while traits construed as costs of defense were highly sensitive to resource allocation tradeoffs. This difference in sensitivity between the two groups of traits may explain why the correlation between the expression of defensive traits and the expression of the associated defense costs was weak. Furthermore, my results indicate that genetic linkages and mechanistic integration of multiple defensive traits and their associated costs may constrain time and resource allocation in ways that are not addressed in existing models. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Seasonal dynamics of shallow-hyporheic-zone microbial community structure along a heavy-metal contamination gradient

Heavy metals contaminate numerous freshwater streams and rivers worldwide. Previous work by this group demonstrated a relationship between the structure of hyporheic microbial communities and the fluvial deposition of heavy metals along a contamination gradient during the fall season. Seasonal variation has been documented in microbial communities in numerous terrestrial and aquatic environments, including the hyporheic zone. The current study was designed to assess whether relationships between hyporheic microbial community structure and heavy-metal contamination vary seasonally by monitoring community structure along a heavy-metal contamination gradient for more than a year. No relationship between total bacterial abundance and heavy metals was observed (R(2) = 0.02, P = 0.83). However, denaturing gradient gel electrophoresis pattern analysis indicated a strong and consistent linear relationship between the difference in microbial community composition (populations present) and the difference in the heavy metal content of hyporheic sediments throughout the year (R(2) = 0.58, P < 0.001). Correlations between heavy-metal contamination and the abundance of four specific phylogenetic groups (most closely related to the alpha, beta, and gamma-proteobacteria and cyanobacteria) were apparent only during the fall and early winter, when the majority of organic matter is deposited into regional streams. These seasonal data suggest that the abundance of susceptible populations responds to heavy metals primarily during seasons when the potential for growth is highest.     

Concordance among stream assemblages and spatial autocorrelation along a fragmented gradient

Patterns of spatial autocorrelation of biota and distributional similarity (concordance) between assemblages of different organism groups have important implications in both theoretical ecology and biodiversity conservation. Here we report environmental gradients and spatial distribution patterns of taxonomic composition among stream fish, benthic macroinvertebrate, and diatom assemblages along a fragmented stream in south‐western France. We quantified spatial patterns of lotic assemblage structure along this stream, and we tested for concordance in distribution patterns among the three taxonomic groups. Our results showed that both environmental characteristics and stream assemblages were spatially autocorrelated. For stream fish and diatom assemblages, these patterns reflected assemblage changes along the longitudinal stream gradient, whereas environmental variables and benthic macroinvertebrates exhibited a more patchy spatial pattern. Cross‐taxa concordance was significant between stream fish and diatoms, and between stream fish and benthic macroinvertebrates. The assemblage concordance between stream fish and diatoms could be attributed to similar responses along the longitudinal gradient, whereas those between stream fish and benthic macroinvertebrates may result from biotic interactions. Based on potential dispersal capacities of taxa, our results validated the hypotheses that weakly dispersing taxa exhibit greater concordance than highly dispersing ones and that dispersal capacities affect how taxonomic groups respond to their local environment. Both diatoms and highly dispersing stream fish were affected by stream fragmentation (i.e. the number of dams between sites), while the effect of fragmentation was not significant for invertebrates that fly well in their adult stage, thus emphasizing the importance of the way of dispersal. These results suggest that addressing the effects of dispersal capacity on stream assemblage patterns is crucial to identifying mechanisms behind patterns and to better understanding the determinants of stream biodiversity.     

Temperature and trophic structure are driving microbial productivity along a biogeographical gradient

Temperature is known to influence ecosystem processes through its direct effect on biological rates such as respiration and nutrient cycling. These changes can then indirectly affect ecologically processes by altering trophic dynamics, the persistence of a species in a given environment, and, consequently, its distribution. However, it is not known if this direct effect of temperature on biological rates is singularly the most important factor for the functioning of ecosystems, or if trophic structure and the adaptation of a species to the local environment also play an essential role. Understanding the relative importance of these factors is crucial for predicting the impact that climate change will have on species and ecosystems. To achieve a more complete understanding of the impact of changing temperatures, it is necessary to integrate perspectives from biogeography, such as the influences of species distribution and local adaptation, with ecosystem and community ecology. By using the microbial community inhabiting the water‐filled leaves of Sarracenia purpurea, we tested the importance of temperature, trophic structure, and local adaptation on ecosystem functioning. We accomplished this by collecting communities along a natural temperature gradient and maintaining these communities in a common garden, factorial experiment. To test for the importance of local adaptation and temperature, the origin of each community was crossed with the temperature from each site. Additionally, to test the importance of top‐down trophic regulation for ecosystem functioning, the presence of the mosquito larvae top predator was manipulated. We found that temperature has a greater effect on ecosystem functioning than origin, and that top‐down trophic regulation increased with temperature. Our results emphasize the synergistic effects of temperature and biotic interactions when predicting the consequences of global warming on ecosystem functioning.     

Evolution of resource use along a gradient of stress leads to increased facilitation

The stress‐gradient hypothesis (SGH) posits that the relative importance of facilitative interactions versus negative interactions increases as levels of abiotic stress increase. Originally formulated in empirical studies of plant populations, in recent years the SGH has been found to describe how interactions change in response to stress in a wide range of species including algae, mussels and moths. However, there has been little theory attempting to predict patterns from first principles in relation to different types of interactions. Here, we use mathematical models of microbial populations to investigate whether patterns consistent with the SGH arise when species interact through resource use and allelopathy. Evolution alters the degree to which competition for resource use versus facilitation (cross‐feeding) occurs. Our results are consistent with the SGH; species interactions evolve to be more facilitative as average stress intensifies. This occurs because at greater stress the species evolve to become specialists on either of the two resources thereby decreasing overlap in resource use and increasing facilitation through cross‐feeding. In addition, the production of toxic allelopathic compounds decreases as stress intensifies due to density‐dependent effects. Our results suggest that the SGH could arise through fundamental interactions that are common to many organisms and therefore that the SGH could be a more widespread phenomenon than previously recognised.