Semi-arid Plant Communities Research Articles

Does a history of population co-occurrence predict plant performance, community productivity, or invasion resistance?

A history of species co-occurrence in plant communities is hypothesized to lead to greater niche differentiation, more efficient resource partitioning, and more productive, resistant communities as a result of evolution in response to biotic interactions. A similar question can be asked of co-occurring populations: do individual species or community responses differ when communities are founded with plants sharing a history of population co-occurrence (sympatric) or originating from different locations (allopatric)? Using shrub, grass, and forb species from six locations in the western Great Basin, USA, we compared establishment, productivity, reproduction, phenology, and resistance to invaders for experimental communities with either sympatric or allopatric population associations. Each community type was planted with six taxa in outdoor mesocosms, measured over three growing seasons, and invaded with the annual grass Bromus tectorum in the final season. For most populations, the allopatric or sympatric status of neighbors was not important. However, in some cases it was beneficial for some species from some locations to be planted with allopatric neighbors, while others benefited from sympatric neighbors, and some of these responses had large effects. For instance, the Elymus population that benefited the most from allopatry grew 50% larger with allopatric neighbors than in single origin mesocosms. This response affected invasion resistance, as B. tectorum biomass was strongly affected by productivity and phenology of Elymus spp., as well as Poa secunda. Our results demonstrate that while community composition can affect plant performance in semi-arid plant communities, assembling communities from sympatric populations is not sufficient to ensure high productivity and invasion resistance. Instead, we observed an idiosyncratic interaction between sampling effects and evolutionary history, with the potential for seed source of individual populations to have community-level effects. This article is protected by copyright. All rights reserved.

Plant diversity patterns along an elevation gradient: the relative impact of environmental and spatial variation on plant diversity and assembly in arid and semi-arid regions

The spatial distribution pattern of species diversity and its driving variables are essential to understanding how biodiversity will respond to the threat of future climatic instability. This study examines how the alpha and beta diversity indices vary along the elevation gradient and which factors are more responsible for arid and semi-arid plant diversity and community assembly. There were 121 species from 90 genera and 42 families found at elevations ranging from 1,200 to 2,600 m within the Ningxia Helan Mountain National Nature Reserve in China. These were primarily shrubs and herbs with characteristics adapted to extreme temperatures. The diversity of both the shrub layer and the herb layer was highest at mid-elevation, underscoring the importance of protecting mid-elevation ecosystems. Climatic distance and topographic distance were more influential than spatial distance in driving the species composition change of shrubs and herbs. In the herb layer, climatic, topographic, and spatial factors contributed more to community assembly than shrub communities. In this case, improving the environment in the community or introducing herbs could help the restoration of these places. These findings provide insight into biogeographic patterns, biodiversity growth mechanisms, and community formation processes. Moreover, it can improve projections of climate change within and across ecosystems and provide the scientific basis for the use and management of plant resources in arid and semi-arid regions.

Facilitation enhances ecosystem function with non‐random species gains

Facilitation, an ecological interaction assembling plant communities worldwide, has been shown to modulate both species richness and ecosystem functions. Such a biodiversity–ecosystem functioning (BEF) relationship can be decomposed into different components not only related to species losses and gains but also to the identity of the species and the context in which they live. Using an extension of the classical BEF approach named CAFE (community assembly and the functioning of ecosystems), we quantified the contribution of these components to the BEF relationship in a Spanish semiarid plant community shaped by facilitation. We used species richness as a measure for biodiversity and plant cover as a proxy of multiple ecosystem functions including plant productivity, soil protection, soil fertility and microbial productivity. Nurse plants doubled the number of species that live beneath them relative to open ground, but caused a five‐fold increase in plant cover. The disproportionate increase of plant cover was a consequence of the identity of the species enhanced by nurse plants, which were more productive than the average. We discuss these results in terms of sampling effects (i.e. the higher probability of richer communities to harbour hyperproductive species) and complementary effects (i.e. richer communities enhancing productivity through resource partitioning, abiotic facilitation or biotic feedbacks). The enhancement of ecosystem functions that plant facilitation produces by incorporating species with high functional values to the community may reverberate among other trophic levels and propagate beyond the local scale where the ecological interaction is produced.

Extensive pollen-mediated gene flow across intensively managed landscapes in an insect-pollinated shrub native to semiarid habitats.

Our knowledge of the impact of landscape fragmentation on gene flow patterns is mainly drawn from tropical and temperate ecosystems, where landscape features, such as the distance of a tree to the forest edge, drive connectivity and mating patterns. Yet, the structure of arid and semiarid plant communities - with open canopies and a scattered distribution of trees - differs greatly from those that are well-characterized in the literature. As a result, we ignore whether the documented consequences of landscape fragmentation on plant mating and gene flow patterns also hold for native plant communities in arid and semiarid regions. We investigated the relative contribution of plant traits, pollinator activity, and individual neighbourhood in explaining variation in mating and gene flow patterns of an insect-pollinated semiarid arborescent shrub, Ziziphus lotus, at three sites embedded in highly altered agriculture landscapes. We used 14 SSRs, seed paternity analyses, and individual mixed effect mating models (MEMMi) to estimate the individual mating variables and the pollen dispersal kernel at each site. Individual spatial location, flower density, and floral visitation rate explained most of the variation of mating variables. Unexpectedly, individual correlated paternity was very low and shrubs surrounded by the most degraded matrix exhibited an increased fraction of pollen immigration and a high effective number of pollen donors per mother shrub. Overall, our results reveal that an active pollinator assemblage ensures highly efficient mating, and maintains pollen-mediated gene flow and notable connectivity levels, even in highly altered landscapes, potentially halting genetic isolation within and between distant sites.

Water regulation and soil retention services in semiarid ecosystems of southeastern Iran, 2018–2020

Water regulation and soil retention are regulating services that play an important role in the sustainability of arid ecosystems. A study was carried out to assess the potential of semiarid plant communities in providing water regulation and soil retention ecosystem services. Concrete plots were made around plants and bare surface (control) to measure sediment and runoff after each rainfall event from 2018 to 2020. The potential of plant communities in providing water regulation and soil retention services was estimated using density and the mean area of patches. The shrub–grasses and shrub–annuals communities had the highest and lowest potential for providing water regulation and soil retention services, showing the plant communities with priority for conservation and restoration, respectively. Land degradation was the most important factor affecting these services in the region. Soil retention and water regulation were, respectively, reduced 50% and 50%, 62% and 70%, and 75% and 90% in areas with moderate, high and severe degradation compared to areas with low degradation. Rainfall events with depth higher than 15 mm significantly reduced these services in all plant communities. However, proper management moderated the effects of heavy rains in the shrub–grasses plant community. Therefore, ecosystems must be managed in the way that assures the greatest density of cushion semishrubs to have the highest potential in providing water regulation and soil retention services.

Effects of Elevated Temperature and CO2 Concentration on Seedling Growth of Ventenata dubia (Leers) Coss. and Bromus tectorum L.

The impacts of climate change are expected to alter the abundance and distribution of invasive annual grasses, such as Bromus tectorum L. (cheatgrass) and Ventenata dubia (Leers) Coss. (ventenata). High temperature extremes will be more frequent and for longer periods, and increased atmospheric CO2 is expected to double even with the most conservative estimates. Climate change draws concern for the potential success of winter annual grasses in arid and semi-arid plant communities. Information on B. tectorum’s growth response to climate change in laboratory and field experiments are available for monocultures; however, more knowledge is needed on the response when growing with other invasive grasses, such as V. dubia. We examined differences in seedling growth for V. dubia and B. tectorum growing alone and with each other under current (4 °C/23 °C at 400 ppm CO2) and elevated (10.6 °C/29.6 °C at 800 ppm CO2) climate conditions. There was one trial per climate scenario with 10 replications per competition type (inter-, intra-specific competition for each species). Bromus tectorum was larger than V. dubia across climate and competition treatments, but contrary to previous studies, both species were smaller in the elevated climate treatment. Ventenata dubia allocated more growth to its roots than B. tectorum across both climate treatments, indicating V. dubia may have a competitive advantage for soil resources now and in the future.

Consistent spatial patterns across several plant communities within a region indicate that the same processes may be acting on Andean deserts and semideserts

AbstractQuestionsSpatial pattern analysis may be used to derive hypotheses on the processes that shape plant communities. Do different mature semidesert communities exhibit the same major spatial patterns, suggesting that similar processes shape them?LocationNine localities in the Bolivian Andes.MethodsWe mapped all woody individuals at each locality. We performed uni‐, bi‐, and multivariate spatial analyses of nine Andean, mature semiarid plant communities to test for similarity in spatial patterns.ResultsWe found that the nine communities exhibited, in most of the main spatial pattern attributes, close similarities (community aggregation, importance of plant interactions, life form differences, etc.). Some peculiarities were likely due to differences in life form or site history.ConclusionsAt least for Andean semiarid communities, the complete analysis of a representative plot conveys reliable information on spatial relationships and, hence, on potential underlying mechanisms. This shows that the use of 1–2 representative plots can be used to extrapolate to similar communities within a given region. Our results can be used as a plausible hypothesis to be tested in other zones of the planet.

Facilitation and herbivory during restoration of California coastal sage scrub

Using nurse plants to facilitate native plant recruitment in degraded habitats is a common restoration practice across various arid and semiarid environments. Living trees or shrubs are typically considered nurse plants, whereas dead shrubs left in the landscape from prolonged drought are understudied prospective facilitators for native plant recruitment. The interaction between nurse plants and biotic pressures, such as herbivory, on native recruitment is also not well understood in semiarid plant communities. We investigated the effects of facilitation and herbivory on native seedling germination, growth, and survival in the restoration of degraded coastal sage scrub (CSS) habitat. Native shrub seedlings (Artemisia californica and Salvia mellifera) were planted, and native annual species (Amsinckia intermedia, Deinandra fasciculata, Phacelia distans, and Pseudognaphalium californicum) were sown in three Shrub Type treatments (live shrub, dead shrub, and exposed areas), with a nested Cage treatment (no cage and cage) in each Shrub Type treatment. Annual species grew equally well in all Shrub Type treatments; shrub seedlings grew largest in exposed areas. While there was little evidence of facilitation for all species tested, there were strong positive effects of caging on growth and establishment of all species. Caging palatable native species or planting species with anti‐herbivory traits around target plants may be more strategic approaches compared to using nurse plants in restoring degraded CSS after extended drought.

Plant-plant interactions influence phylogenetic diversity at multiple spatial scales in a semi-arid mountain rangeland.

Molecular phylogenies are increasingly used to understand how biotic interactions and environment shape phylogenetic community structure (PCS). However, we do not understand the effects of plant-plant interactions and environment on PCS and phylogenetic diversity across spatial scales, particularly in rangelands. Here, we ask: (1) do plant-plant interactions and environment affect PCS and phylogenetic diversity differently across the three spatial scales of the patch, the community, and the habitat? (2) What are the impacts of dominant cushion-nurse plants on the phylogenetic structure of plant communities? We assessed the PCS of semi-arid plant communities along an elevation gradient at the patch, community and habitat scales. Then, we assessed co-occurrence patterns along two sample slopes. Our results indicated important roles for biotic interactions and environmental filtering in determining phylogenetic diversity, with biotic interactions, in particular, having a stronger tendency to increase phylogenetic diversity. This is most likely due to the asymmetrical effects of nurse plants across the three spatial scales on our two different slopes. The impact of biotic interactions caused non-random phylogenetic patterns in more severe environments. In conclusion, biotic interactions influence phylogenetic diversity by altering PCS across aspects and along elevation gradients.

Species-specific interference exerted by the shrub Cistus clusii Dunal in a semi-arid Mediterranean gypsum plant community

BackgroundThe gypsovag shrub Cistus clusii is locally dominant in semi-arid gypsum plant communities of North-Eastern Spain. This species commonly grows in species-poor patches even though it has nurse potential, suggesting interference on neighbouring species. Other Cistus species exert a chemically mediated interference on plant communities, suggesting that it might be a common phenomenon in this genus. This study aimed investigating whether C. clusii exerts chemically mediated interference on neighbouring species in gypsum plant communities. We tested in a greenhouse whether aqueous extracts from C. clusii leaves (L), roots (R) and a mixture of both (RL) affected germination, seedling survival, and growth of nine native species of gypsum communities, including C. clusii itself. We further assessed in the field richness and abundance of plants under the canopy of C. clusii compared to Gypsophila struthium (shrub with a similar architecture having a nurse role) and in open patches. Finally, we specifically assessed in the field the influence of C. clusii on the presence of the species tested in the greenhouse experiment.ResultsAqueous extracts from C. clusii (R and RL) negatively affected either germination or survival in four of nine species. In the field, richness and abundance of plants were lower under the canopy of C. clusii than under G. struthium, but higher than in open patches. Specifically, five of nine species were less frequent than expected under the canopy of C. clusii.ConclusionsCistus clusii shows species-specific interference with neighbouring species in the community, which may be at least partially attributable to its phytotoxic activity. To our knowledge, this is the first report of species-specific interference by C. clusii.

Size-by-environment interactions: a neglected dimension of species' responses to environmental variation.

In both plant and animal systems, size can determine whether an individual survives and grows under different environmental conditions. However, it is unclear whether and when size-dependent responses to exogenous environmental fluctuations affect population dynamics. Size-by-environment interactions create pathways for environmental fluctuations to influence population dynamics by allowing for negative covariation between sizes within vital rates (e.g. small and large individuals have negatively covarying survival rates) and/or size-dependent variability in a vital rate (e.g. survival of large individuals varies less than small individuals through time). Whether these phenomena affect population dynamics depends on how they are mediated by elasticities (they must affect the sizes and vital rates that matter) and their projected impacts will depend on model functional form (the impact of reduced variance depends on the relationship between the environment and vital rate). We demonstrate these ideas with an analysis of fifteen species from five semiarid plant communities. We find that size-by-environment interactions are common but do not impact long-term population dynamics. Size-by-environment interactions may yet be important for other species. Our approach can be applied to species in other ecosystems to determine if and how size-by-environment interactions allow them to cope with, or exploit, fluctuating environments.

Evidence for chemical interference effect of an allelopathic plant on neighboring plant species: A field study.

Many studies have reported the phytotoxicity of allelopathic compounds under controlled conditions. However, more field studies are required to provide realistic evidences for the significance of allelopathic interference in natural communities. We conducted a 2-years field experiment in a semiarid plant community (NE Spain). Specifically, we planted juvenile individuals and sowed seeds of Salsola vermiculata L., Lygeum spartum L. and Artemisia herba-alba Asso. (three co-dominant species in the community) beneath adult individuals of the allelopathic shrub A. herba-alba, and assessed the growth, vitality, seed germination and seedling survival of those target species with and without the presence of chemical interference by the incorporation of activated carbon (AC) to the soil. In addition, juveniles and seeds of the same three target species were planted and sown beneath the canopy of adults of S. vermiculata (a shrub similar to A. herba-alba, but non-allelopathic) and in open bare soil to evaluate whether the allelopathic activity of A. herba-alba modulates the net outcome of its interactions with neighboring plants under contrasting abiotic stress conditions. We found that vitality of A. herba-alba juveniles was enhanced beneath A. herba-alba individuals when AC was present. Furthermore, we found that the interaction outcome in A. herba-alba microsite was neutral, whereas a positive outcome was found for S. vermiculata microsite, suggesting that allelopathy may limit the potential facilitative effects of the enhanced microclimatic conditions in A. herba-alba microsite. Yet, L. spartum juveniles were facilitated in A. herba-alba microsite. The interaction outcome in A. herba-alba microsite was positive under conditions of very high abiotic stress, indicating that facilitative interactions predominated over the interference of allelopathic plants under those conditions. These results highlight that laboratory studies can overestimate the significance of allelopathy in nature, and consequently, results obtained under controlled conditions should be interpreted carefully.

Plant community changes after land abandonment control CO2 balance in a dry environment

Human activities can dramatically alter natural plant communities which, after disturbance cessation, undergo secondary succession. In arid environments plant succession is quite slow, and its link to the carbon (C) cycle is not well known. We assessed changes in C balance on a semiarid plant community along a chronosequence spanning ca. 100 years after land abandonment in an arid environment in SE Spain to examine temporal changes in C following human disturbance. We selected 5 individuals of the dominant plant species along five plant community stages differing in the time since land abandonment occurred, and we used a closed-chamber infrared gas analyzer method to estimate the contribution of whole plants and bare soil to community C exchange. We estimated CO2 fluxes for each plant community stage and calculated temporal differences along the chronosequence. Plant community composition and plant cover changed throughout the chronosequence. Carbon balance was related to changes in plant photosynthesis and plant and soil respiration along the chronosequence. Overall, community C exchange shifted from source to sink as plant colonization progressed. It took 65 years for the system to recover the equivalent C sink capacity of the undisturbed site. Recovery of arid plant communities after land abandonment may enhance long-term C sequestration and significantly contribute to C balance at the global level.

Phytotoxic effects of volatile and water soluble chemicals of Artemisia herba-alba

We investigated the phytotoxic effects of volatile and water soluble chemicals produced by the shrub Artemisia herba-alba Asso. We conducted a number of germination and early seedling growth bioassays on species that co-exist with A. herba-alba in natural semiarid plant communities (Salsola vermiculata L. Lygeum spartum L. Pinus halepensis Mill. and A. herba-alba itself). In addition, we assessed the phytotoxic effects of a mixture of phenols that were identified in the aqueous extract of A. herba-alba on the germination of such species. We found that volatile chemicals inhibited the germination of P. halepensis seeds, promoted the growth of P. halepensis seedlings and reduced the root biomass of S. vermiculata seedlings, while water soluble chemicals promoted the growth of L. spartum and P. halepensis seedlings. On the other hand, both volatile and water soluble compounds inhibited the germination of A. herba-alba seeds, highlighting its strong autotoxic nature. We did not find a phytotoxic effect of phenols identified (catechol, protocatechuic and vanillic acids) on species that co-exists with A. herba-alba, although they had a moderate autotoxic effect. Our results suggest that both the volatile and water soluble chemicals might be involved in chemical interactions among plants in semiarid environments.

Rangeland monitoring using remote sensing: comparison of cover estimates from field measurements and image analysis

Rangeland monitoring is important for evaluating and assessing semi-arid plant communities. Remote sensing provides an effective tool for rapidly and accurately assessing rangeland vegetation and other surface attributes such as bare soil and rock. The purpose of this study was to evaluate the efficacy of remote sensing as a surrogate for field-based sampling techniques in detecting ground cover features (i.e., trees, shrubs, herbaceous cover, litter, surface), and comparing results with field-based measurements collected by the Utah Division of Wildlife Resources Range Trent Program. In the field, five 152 m long transects were used to sample plant, litter, rock, and bare-ground cover using the Daubenmire ocular estimate method. At the same location of each field plot, a 4-band (R,G,B,NIR), 25 cm pixel resolution, remotely sensed image was taken from a fixed-wing aircraft. Each image was spectrally classified producing 4 cover classes (tree, shrub, herbaceous, surface). No significant differences were detected between canopy cover collected remotely and in the field for tree (P = 0.652), shrub (P = 0.800), and herbaceous vegetation (P = 0.258). Surface cover was higher in field plots (P < 0.001), likely in response to the methods used to sample surface features by field crews. Accurately classifying vegetation and other features from remote sensed information can improve the efficiency of collecting vegetation and surface data. This information can also be used to improve data collection frequency for rangeland monitoring and to efficiently quantify ecological succession patterns.

Effects of the allelopathic plantArtemisia herba-albaAsso on the soil seed bank of a semi-arid plant community

Effects of the allelopathic plant<i>Artemisia herba-alba</i>Asso on the soil seed bank of a semi-arid plant community

Factors driving species assemblage in Mediterranean soil seed banks: from the large to the fine scale.

Many studies have analysed the mechanisms that determine plant coexistence in standing vegetation, but the determinants of soil seed bank species assemblies have rarely been studied. In gypsum soil communities, aerial vegetation and seed banks are tightly connected in space and time, but the mechanisms involved in their organization may differ. The aim of this study is to understand the relative importance of biotic and abiotic factors controlling soil seed bank composition and structure. Persistent and complete (i.e. persistent plus transient) soil seed banks were investigated at two spatial scales in a very species-rich semi-arid community dominated by annuals. A water addition treatment equivalent to 50 % annual increase in average precipitation (abiotic factor) was applied for two consecutive years, and the relationships of the soil seed bank to the biological soil crust (BSC), above-ground vegetation and the presence of Stipa tenacissima tussocks (biotic factors) were simultaneously evaluated. As expected, the standing vegetation was tightly related to seed abundance, species richness and composition in both seed banks. Remarkably, BSC cover was linked to a decrease in seed abundance and species richness in the persistent seed bank, and it even determined complete seed bank composition at the fine spatial scale. However, this effect disappeared at coarser scales, probably because of the high spatial heterogeneity induced by BSCs. In contrast to findings on standing vegetation, Stipa and the irrigation treatment for two consecutive years had no effect on soil seed banks. Soil seed bank assemblies in our semi-arid plant community were the result of above-ground vegetation dynamics and of the direct filtering processes on seed fate operated by the spatially heterogeneous BSCs. Cover of BSCs was negatively correlated with seed abundance and species richness, and affected seed species composition in the soil. Changes in species composition and enrichment when the BSC cover is low suggest that BSCs promote a fine scale niche differentiation in the soil seed bank and thereby potentially enhance species coexistence and high species diversity in these communities.

Plant-plant interactions as a mechanism structuring plant diversity in a Mediterranean semi-arid ecosystem.

Plant–plant interactions are among the fundamental processes that shape structure and functioning of arid and semi‐arid plant communities. Despite the large amount of studies that have assessed the relationship between plant–plant interactions (i.e., facilitation and competition) and diversity, often researchers forget a third kind of interaction, known as allelopathy. We examined the effect of plant–plant interactions of three dominant species: the perennial grass Lygeum spartum, the allelopathic dwarf shrub Artemisia herba‐alba, and the nurse shrub Salsola vermiculata, on plant diversity and species composition in a semi‐arid ecosystem in NE Spain. Specifically, we quantified the interaction outcome (IO) based on species co‐occurrence, we analyzed diversity by calculation of the individual species–area relationship (ISAR), and compositional changes by calculation of the Chao‐Jaccard similarity index. We found that S. vermiculata had more positive IO values than L. spartum, and A. herba‐alba had values between them. Lygeum spartum and A. herba‐alba acted as diversity repellers, whereas S. vermiculata acted as a diversity accumulator. As aridity increased, A. herba‐alba transitioned from diversity repeller to neutral and S. vermiculata transitioned from neutral to diversity accumulator, while L. spartum remained as diversity repeller. Artemisia herba‐alba had more perennial grass species in its local neighborhood than expected by the null model, suggesting some tolerance of this group to its “chemical neighbor”. Consequently, species that coexist with A. herba‐alba were very similar among different A. herba‐alba individuals. Our findings highlight the role of the nurse shrub S. vermiculata as ecosystem engineer, creating and maintaining patches of diversity, as well as the complex mechanism that an allelopathic plant may have on diversity and species assemblage. Further research is needed to determine the relative importance of allelopathy and competition in the overall interference of allelopathic plants.

Notice of release of Amethyst Germplasm hoary tansyaster: selected class of natural germplasm

The US Department of Agriculture (USDA), Natural Resources Conservation Service (NRCS), Aberdeen Plant Materials Center, Aberdeen, Idaho, announces the release of Amethyst Germplasm hoary tansyaster ( Machaeranthera canescens (Pursh) A. Gray [Asteraceae]), a selected class natural track germplasm identified by NRCS accession number 9076670 for conservation plantings in the Intermountain US. Amethyst Germplasm hoary tansyaster is a native forb collected near St Anthony Sand Dunes in Fremont County, Idaho. This forb flowers and produces seed during the first growing season and provides late summer and early fall blossoms for native pollinators. It is recommended for pollinator plantings, wildlife areas, and rangeland rehabilitation seedings in arid to semiarid plant communities where native forb diversity is desirable.

Effect of livestock grazing in the partitions of a semiarid plant–plant spatial signed network

In recent times, network theory has become a useful tool to study the structure of the interactions in ecological communities. However, typically, these approaches focus on a particular kind of interaction while neglecting other possible interactions present in the ecosystem. Here, we present an ecological network for plant communities that consider simultaneously positive and negative interactions, which were derived from the spatial association and segregation between plant species. We employed this network to study the structure and the association strategies in a semiarid plant community of Cabo de Gata-Níjar Natural Park, SE Spain, and how they changed in 4 sites that differed in stocking rate. Association strategies were obtained from the partitions of the network, built based on a relaxed structural balance criterion. We found that grazing simplified the structure of the plant community. With increasing stocking rate species with no significant associations became dominant and the number of partitions decreased in the plant community. Independently of stocking rate, many species presented an associative strategy in the plant community because they benefit from the association to certain ‘nurse’ plants. These ‘nurses’ together with species that developed a segregating strategy, intervened in most of the interactions in the community. Ecological networks that combine links with different signs provide a new insight to analyze the structure of natural communities and identify the species which play a central role in them.