Precopulatory but not postcopulatory male reproductive traits diverge in response to mating system manipulation in Drosophila melanogaster

Competition between males creates potential for pre‐ and postcopulatory sexual selection and conflict. Theory predicts that males facing risk of sperm competition should evolve traits to secure their reproductive success. If those traits are costly to females, the evolution of such traits may also increase conflict between the sexes. Conversely, under the absence of sperm competition, one expectation is for selection on male competitive traits to relax thereby also relaxing sexual conflict. Experimental evolution studies are a powerful tool to test this expectation. Studies in multiple insect species have yielded mixed and partially conflicting results. In this study, we evaluated male competitive traits and male effects on female costs of mating in Drosophila melanogaster after replicate lines evolved for more than 50 generations either under enforced monogamy or sustained polygamy, thus manipulating the extent of intrasexual competition between males. We found that in a setting where males competed directly with a rival male for access to a female and fertilization of her ova polygamous males had superior reproductive success compared to monogamous males. When comparing reproductive success solely in double mating standard sperm competition assays, however, we found no difference in male sperm defense competitiveness between the different selection regimes. Instead, we found monogamous males to be inferior in precopulatory competition, which indicates that in our system, enforced monogamy relaxed selection on traits important in precopulatory rather than postcopulatory competition. We discuss our findings in the context of findings from previous experimental evolution studies in Drosophila ssp. and other invertebrate species.     

The effect of nitrogen availability and water conditions on competition between a facultative CAM plant and an invasive grass

Abstract Plants with crassulacean acid metabolism (CAM) are increasing their abundance in drylands worldwide. The drivers and mechanisms underlying the increased dominance of CAM plants and CAM expression (i.e., nocturnal carboxylation) in facultative CAM plants, however, remain poorly understood. We investigated how nutrient and water availability affected competition between Mesembryanthemum crystallinum (a model facultative CAM species) and the invasive C₃ grass Bromus mollis that co‐occur in California's coastal grasslands. Specifically we investigated the extent to which water stress, nutrients, and competition affect nocturnal carboxylation in M. crystallinum. High nutrient and low water conditions favored M. crystallinum over B. mollis, in contrast to high water conditions. While low water conditions induced nocturnal carboxylation in 9‐week‐old individuals of M. crystallinum, in these low water treatments, a 66% reduction in nutrient applied over the entire experiment did not further enhance nocturnal carboxylation. In high water conditions M. crystallinum both alone and in association with B. mollis did not perform nocturnal carboxylation, regardless of the nutrient levels. Thus, nocturnal carboxylation in M. crystallinum was restricted by strong competition with B. mollis in high water conditions. This study provides empirical evidence of the competitive advantage of facultative CAM plants over grasses in drought conditions and of the restricted ability of M. crystallinum to use their photosynthetic plasticity (i.e., ability to switch to CAM behavior) to compete with grasses in well‐watered conditions. We suggest that a high drought tolerance could explain the increased dominance of facultative CAM plants in a future environment with increased drought and nitrogen deposition, while the potential of facultative CAM plants such as M. crystallinum to expand to wet environments is expected to be limited.     

Effects of biotic interactions on modeled species' distribution can be masked by environmental gradients

A fundamental goal of ecology is to understand the determinants of species' distributions (i.e., the set of locations where a species is present). Competition among species (i.e., interactions among species that harms each of the species involved) is common in nature and it would be tremendously useful to quantify its effects on species' distributions. An approach to studying the large‐scale effects of competition or other biotic interactions is to fit species' distributions models (SDMs) and assess the effect of competitors on the distribution and abundance of the species of interest. It is often difficult to validate the accuracy of this approach with available data. Here, we simulate virtual species that experience competition. In these simulated datasets, we can unambiguously identify the effects that competition has on a species' distribution. We then fit SDMs to the simulated datasets and test whether we can use the outputs of the SDMs to infer the true effect of competition in each simulated dataset. In our simulations, the abiotic environment influenced the effects of competition. Thus, our SDMs often inferred that the abiotic environment was a strong predictor of species abundance, even when the species' distribution was strongly affected by competition. The severity of this problem depended on whether the competitor excluded the focal species from highly suitable sites or marginally suitable sites. Our results highlight how correlations between biotic interactions and the abiotic environment make it difficult to infer the effects of competition using SDMs.     

Interactions among salt marsh plants vary geographically but not latitudinally along the California coast

The strength of species interactions often varies geographically and locally with environmental conditions. Competitive interactions are predicted to be stronger in benign environments while facilitation is expected to be stronger in harsh ones. We tested these ideas with an aboveground neighbor removal experiment at six salt marshes along the California coast. We determined the effect of removals of either the dominant species, Salicornia pacifica, or the subordinate species on plant cover, aboveground biomass and community composition, as well as soil salinity and moisture. We found that S. pacifica consistently competed with the subordinate species and that the strength of competition varied among sites. In contrast with other studies showing that dominant species facilitate subordinates by moderating physical stress, here the subordinate species facilitated S. pacifica shortly after removal treatments were imposed, but the effect disappeared over time. Contrary to expectations based on patterns observed in east coast salt marshes, we did not see patterns in species interactions in relation to latitude, climate, or soil edaphic characteristics. Our results suggest that variation in interactions among salt marsh plants may be influenced by local‐scale site differences such as nutrients more than broad latitudinal gradients.     

Spatial heterogeneity and scale‐dependent habitat selection for two sympatric raptors in mixed‐grass prairie

Sympatric predators are predicted to partition resources, especially under conditions of food limitation. Spatial heterogeneity that influences prey availability might play an important role in the scales at which potential competitors select habitat. We assessed potential mechanisms for coexistence by examining the role of heterogeneity in resource partitioning between sympatric raptors overwintering in the southern Great Plains. We conducted surveys for wintering Red‐tailed hawk (Buteo jamaicensis) and Northern Harrier (Circus cyanea) at two state wildlife management areas in Oklahoma, USA. We used information from repeated distance sampling to project use locations in a GIS. We applied resource selection functions to model habitat selection at three scales and analyzed for niche partitioning using the outlying mean index. Habitat selection of the two predators was mediated by spatial heterogeneity. The two predators demonstrated significant fine‐scale discrimination in habitat selection in homogeneous landscapes, but were more sympatric in heterogeneous landscapes. Red‐tailed hawk used a variety of cover types in heterogeneous landscapes but specialized on riparian forest in homogeneous landscapes. Northern Harrier specialized on upland grasslands in homogeneous landscapes but selected more cover types in heterogeneous landscapes. Our study supports the growing body of evidence that landscapes can affect animal behaviors. In the system we studied, larger patches of primary land cover types were associated with greater allopatry in habitat selection between two potentially competing predators. Heterogeneity within the scale of raptor home ranges was associated with greater sympatry in use and less specialization in land cover types selected.     

Estimating tree and stand sapwood area in spatially heterogeneous southeastern Australian forests

Aims Natural and anthropogenic changes in forests can have important influences on transpiration and water production. Understanding the effects of increasing disturbances, due for example to climate change and forest harvesting, requires detailed information on how forest density and structural attributes relate to transpiration. Mean annual transpiration of eucalypt forest communities is often strongly correlated with total cross-sectional sapwood area. Our aim was to test an efficient method for estimating sapwood area at 1.3 m height (SA 1.3) in a large number of trees to understand the spatial heterogeneity of tree and stand sapwood area within and between forest communities, and develop allometric relationships that predict SA 1.3 with forest inventory data. We also apply tree competition models to determine the degree to which the relationship between SA 1.3 and tree basal area at 1.3 m height (BA 1.3) is influenced by competition.Methods We visited 25 recently harvested southeastern Australian forest sites consisting of 1379 trees and 5 Eucalyptus species to evaluate a new efficient data collection method for estimating SA 1.3 with tree taper and stump dimensions data using mixed effects models. The locations of 784 stumps within one 5-ha site were accurately mapped using an unmanned aerial vehicle (UAV), and four distance-dependent tree competition models were applied across the site to explain within-stand variation in the ratio of SA 1.3 to BA 1.3. Data from 24 additional sites, consisting of ten 15 m radial plots per site, were used to analyse within-site variation in R Ha (the ratio of stand sapwood area SA Ha to stand basal area BA Ha). The radial plots were merged within each site to evaluate between-site variations in R Ha across the landscape. For predicting SA Ha with forest inventory data, we computed the relationship between SA Ha and a new index of total stem perimeter per hectare, defined as ? B A H a N T, where N T is tree stocking density.Important findings Our 1379 measured stems represent the most comprehensive measure of sapwood area, surpassing the 757 measured stems in native eucalypt forests published in literature. The species-specific R Ha varied considerably across sites and therefore extrapolating SA Ha with spatially distributed BA Ha maps and a generalized R Ha would introduce local uncertainty. We found that the species-specific stem perimeter index was more effective at capturing variability in SA Ha across the landscape using forest composition, structure and density data (R 2 : 0.72–0.77). The strong correlation between tree SA 1.3 and BA 1.3 improved slightly using tree competition models (R 2 increased from 0.86 to 0.88). Relating SA Ha to routinely measured forest inventory attributes within permanent plots and Light Detection and Ranging (LiDAR) data may provide opportunities to map forest water use in time and space across large areas disturbed by wildfire and logging.     

Soil heterogeneity affects ramet placement of Hydrocotyle vulgaris

Aims Soil heterogeneity is common in natural habitats. It may trigger foraging responses (placing more ramets and/or roots in nutrient-rich patches than in nutrient-poor patches) and further affect the growth of plants. However, the impact of soil heterogeneity on competitive interactions has been little tested.Methods We conducted a greenhouse experiment to investigate the effects of soil heterogeneity on intraspecific competition with a stoloniferous herb Hydrocotyle vulgaris. We grew one (without competition) or nine ramets (with competition) of H. vulgaris under a homogeneous environment and two heterogeneous environments differing in patch size (large or small patches). In the heterogeneous treatment, the soil consisted of the same number of nutrient-rich and nutrient-poor patches arranged in a chessboard manner, and in the homogeneous treatment, the soil was an even mixture of the same amount of the nutrient-rich and the nutrient-poor soil.Important findings Irrespective of intraspecific competition, H. vulgaris showed foraging responses to soil heterogeneity in the large patch treatment, e.g. it produced significantly more biomass, ramets, aboveground mass and root mass in the nutrient-rich patches than in the nutrient-poor patches. In the small patch treatment, foraging responses were observed when intraspecific competition was present, but responses were not observed when there was no competition. However, we find a significant effect of soil heterogeneity on neither overall growth nor competitive intensity of H. vulgaris. Our results suggest that foraging responses to soil heterogeneity may not necessarily be adaptive and intraspecific competition may not be influenced by soil heterogeneity.     

Testing experimentally the effect of soil resource mobility on plant competition

Aims The volume of soil beyond a plant's roots from which that plant is able to acquire a particular nutrient depends upon the mobility of that nutrient in the soil. For this reason it has been hypothesized that the strength of competitive interactions between plants vary with soil nutrient mobility. We aimed to provide an experimental test of this hypothesis.Methods We devised two experimental systems to investigate specifically the effect of nutrient transport rates upon intraspecific competition. In the first, the exchange of rhizosphere water and dissolved nutrients between two connected pots, each containing one plant, was manipulated by alternately raising and lowering the pots. In the second experiment, the roots systems of two competing plants were separated by partitions of differing porosity, thereby varying the plants' access to water and nutrients in the other plant′s rhizosphere. In this second experiment, we also applied varying amounts of nutrients to test whether higher nutrient input would reduce competition when competition for light is avoided, and applied different water levels to affect nutrient concentrations without changing nutrient supply.Important findings In both experiments, lower mobility reduced competitive effects on plant biomass and on relative growth rate (RGR), as hypothesized. In the second experiment, however, competition was more intense under high nutrient input, suggesting that low nutrient supply rates reduced the strength of the superior competitor. Competitive effects on RGR were only evident under the low water level, suggesting that under lower nutrient concentrations, competitive effects might be less pronounced. Taken together, our results provide the first direct experimental evidence that a reduction in nutrient mobility can reduce the intensity of competition between plants.     

Sperm displacement behavior of the cuttlefish Sepia esculenta (Cephalopoda: Sepiidae)

Sperm displacement behavior of cuttlefish (Sepia esculenta) was observed in a tank. Before ejaculation, male cuttlefish used their arms III to scrape out sperm masses attached to the buccal membranes of females. The removed sperm mass debris was directly visible and countable. Active sperm were present within the removed sperm debris, implying that the aim of this behavior is to remove competing male sperm. However, many sperm masses remained on the female buccal membrane even after the removal behavior, showing that sperm removal in S. esculenta is incomplete. The duration of sperm removal (an indicator of male investment in that process) was unaffected by the body sizes of mated pair, the duration of spermatangia placement at the current mating (for the hypothesis that the sperm removal serves to creat attachment space of spermatophores), or the estimated amount of sperm masses deposited from previous matings. Moreover, male S. esculenta performed sperm removal regardless of whether the last male to mate with the partner was himself, suggesting males remove not only the sperm of rivals but also their own. Although the number of removed sperm masses increased with the time spent on removal of sperm, male cuttlefish may shorten the duration of sperm removal to avoid the risk of mating interruption. We conclude that this time restriction would likely influence the degree of partial sperm removal in S. esculenta. A digital video image relating to the article is available at .This revised version was published online in April 2005 with corrections in the abstract.     

Belowground Interactions in a Vine (Vitis vinifera L.)-tall Fescue (Festuca arundinacea Shreb.) Intercropping System: Water Relations and Growth

In the Mediterranean area, the introduction of cover crops in vineyards is hampered by the risk of severe competition for water. Belowground interactions are still not very clear in this perennial-herbaceous association. This work was aimed at characterizing the development of the root systems of associated crops and the soil water dynamics. It also investigated whether water competition could be the cause of vine vigour and yield reductions. Experiments were conducted in a 4-year-old association (vine – tall fescue) and in a weed controlled vineyard. Water transfers in the soil were estimated on the basis of the soil water potential and soil hydrodynamic properties. The vine root system was concentrated in the soil under the row, whereas the intercrop highly colonized a soil compartment under the inter-row to a depth of approximately 1 m. Despite this spatial complementarity in root distribution, intercropping reduced the amount of soil water available for the vine crop. The low soil water content reduced soil water conductivity thereby limiting water transfers, despite a significant gradient in the soil water potential. This conductivity did not differ significantly between treatments but the intercrop enhanced the infiltration of winter rainwater, probably by limiting surface runoff. There was temporal complementarity in this association since the period of intense water uptake by the intercrop occurred earlier than noted for the vine under bare soil conditions. Nevertheless, the competition for water was limited by better refilling of the soil water profile during winter in the intercropped treatment. The intercrop clearly interacted with the vine and decreased its vegetative vigor. Since predawn leaf water potential and stomatal conductance did not differ among treatments, mechanism(s) other than competition for water (e.g. nutrient competition, allelopathy) may be responsible.