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Although the role played by phylogeny in the assembly of plant communities remains as a priority to complete the theory of species coexistence, experimental evidence is lacking. It is still unclear to what extent phylogenetic diversity is a driver or a consequence of species assembly processes. We experimentally explored how phylogenetic diversity can drive the community level responses to drought conditions in annual plant communities. We manipulated the initial phylogenetic diversity of the assemblages and the water availability in a common garden experiment with two irrigation treatments: average natural rainfall and drought, formed with annual plant species of gypsum ecosystems of Central Spain.
We recorded plant survival and the numbers of flowering and fruiting what are marketing strategies pdf per species in each assemblage. GLMMs were performed for the proportion of surviving, flowering, fruiting what information would a phylogenetic tree give us per species and is long distance relationships worth it what information would a phylogenetic tree give us proportion of surviving species and plants per pot.
In water limited conditions, high phylogenetic diversity favored species coexistence over time with higher plant survival and how can i make my pdf file smaller flowering and fruiting plants per species and more species and plants surviving per pot. Our results agree with the existence of niche complementarity and the convergence of water economy strategies as major mechanisms for promoting species coexistence in plant assemblages in semiarid Mediterranean habitats.
Our findings point to high phylogenetic diversity among neighboring plants as a plausible feature underpinning the coexistence of species, because the success of each species in terms of surviving and producing offspring in drought conditions was greater when the initial phylogenetic diversity was higher. Our study is what information would a phylogenetic tree give us step forward to understand how phylogenetic relatedness is connected to the mechanisms determining the maintenance of biodiversity.
The current theoretical framework and evidence suggest that both stochastic 12 and deterministic mechanisms 34567 operate simultaneously on the assembly of plant communities 8910 Abiotic and biotic filters—mostly acting at the regional and the fine spatial scales, respectively—are important drivers of species assembly in drylands 12together with facilitation that has been described as an important coexistence mechanism in stressful environments i.
Plant trait-based community ecology is recognized as an invaluable tool to understand these processes because it provides morphological or physiological trait-based indices in order to identify the role played by each species at the community level in a niche complementarity context Thus, a species will become part what information would a phylogenetic tree give us a realized species assemblage only if it possesses suitable traits to pass through the filters imposed by restrictive environmental conditions and it reduces niche overlap with neighbor species In the last two decades, the toolbox of community ecologists has incorporated analyses of the phylogenetic patterns of plant communities to understand assembly processes 16 It is evident that historical and evolutionary mechanisms related to migration and speciation are critical for the formation of the regional species pool, but it is not clear how the phylogenetic diversity that describes the degree of relatedness among species can provide information about assembly processes that occur at the ecological time scale 5 A phylogeny should summarize the ecological requirements of coexisting species because it synthesizes the morphological, physiological, and phenological changes in each species throughout evolutionary time in a reduced geographical domain 1920 However, phylogenetic distance among species could indicate not only what information would a phylogenetic tree give us differences, but also competitive inequalities differences in species competitive abilities which should drive competitive exclusion 22 Indeed, the identification of niche differences should be even more feasible throughout the phylogenetic than the functional approach 1424because the latter would require the analysis of several traits most of which might be hard or impossible to measure 16 Thus, phylogenetic diversity could represent more reliably niche differences than functional diversity 2627282930but see Ref.
Many studies have aimed to detect assembly mechanisms based on the observed phylogenetic diversities under field conditions i. For instance, coexistence of phylogenetically close species is usually interpreted as a result of habitat filtering processes and can be indicative of habitat use as a conserved trait along phylogeny 16 However, these types of low phylogenetic diversity assemblages can also result from competition among species when the competitive ability under certain environmental conditions is associated with whole clades By contrast, high phylogenetic diversity responses could be associated with facilitation among species 3536but also with competition processes when competitive exclusion occurs between close relatives with patent niche overlap 1637 Furthermore, if niche convergence occurs among distantly related taxa, high phylogenetic diversity will also be observed in the resulting species assemblages under competitive scenarios Consequently, progress needs to be made in order to elucidate the causal relationships among phylogenetic diversity and assembly mechanisms by directly manipulating the what information would a phylogenetic tree give us diversity of whole assemblages i.
This has rarely been attempted with vascular plants to the best of our knowledge but see Refs. A wide consensus exists on the need for experimental approaches to specifically analyze the mechanisms involved in the assembly of plant communities 56. Ephemeral plant communities in the central Tagus valley, which naturally form high species density assemblages at fine spatial scales up to 38 species per 0.
These features allow the design and implementation of what information would a phylogenetic tree give us communities containing selected species under controlled conditions in common gardens Shifts of assembly mechanisms in a regional what information would a phylogenetic tree give us pool greatly depend on the harshness of the abiotic conditions 45especially dealing with resource availability 12 Since water availability is the main limiting resource in semi-arid Mediterranean ecosystems 47it strongly affects plant community dynamics 48particularly species richness and composition Furthermore, species-specific interactions i.
In the present study, we manipulated both the level of phylogenetic relatedness among coexisting what information would a phylogenetic tree give us i. We aimed to evaluate the effects of the phylogenetic diversity of assemblages on surrogates of community performance i. In the coexistence theory context 6community performance is the net sum of all the differences in fitness of the species that form an assemblage The fitness inequalities among species may cause some of them to disappear, and thus the decrease in the number of species per sampling unit registered throughout the experiment indicated the limitations imposed by the experimental treatments.
The two main hypotheses tested in this study are see our conceptual framework in Fig. By contrast, if phylogenetic relatedness predicts the competitive ability of species, in the manner that closely related species can compete more efficiently for the same resources 16then species will be more likely to coexist in low phylogenetic diversity scenarios. Previous studies have suggested that the competition among closely related species is symmetric, i.
Thus, in high phylogenetic diversity assemblages, a few species are expected to perform better than the rest, so the species richness will decline faster in these scenarios than in low phylogenetic diversity ones under severe drought treatments. By contrast, if the functional traits related to water economy are convergent among distantly related taxa, then we expect phylogenetically diverse assemblages to be more resistant to drought than those that are closely related.
Finally, if drought resistance would randomly occur along phylogeny, we expect that the response of species assemblages to water limitation would not show a clear pattern in different experimental scenarios. Conceptual model illustrating the hypotheses on the mechanisms involved in the assembly of the annual plant community related to phylogenetic diversity.
Conversely, if phylogenetic relatedness predicts the competitive ability of species, then coexistence will be more likely to occur in low phylogenetic diversity scenarios i. In contrast, if water economy traits in the species pool are convergent among distantly related taxa, phylogenetically diverse assemblages will be more resistant to drought than those formed by close relatives. The target plant community comprised annual plant communities on gypsum soils in the Tagus valley, central Spain, which has a semiarid Mediterranean climate with mean annual temperatures around The dominant vegetation comprises gypsophilous dwarf shrubs e.
The annual plant communities are what information would a phylogenetic tree give us from a rich regional floristic pool over species in the middle Tagus valley 43 of ephemeral, highly life-cycle synchronized plants October—early Junegenerating high species density assemblages at fine spatial scales up to 38 species per 0. We established 6 experimental scenarios, but we finally maintained 4 of them because two of the scenarios did not fulfill what information would a phylogenetic tree give us requirements to enter the experiment seed germination was not enough at each plotthus, we finally used 28 species to build the species assemblages see below.
We prepared a common garden experiment with experimental assemblages and more than seedlings. The experimental design consisted of manipulating the phylogenetic diversity of starting experimental assemblages together with water availability treatments. The plant emergence of species in these communities is highly synchronized, so we prepared different phylogenetic combinations at this early demographic stage for our experimental treatments.
MPD index The SES. MPD is a standardized phylogenetic index that contrasts the observed Mean Pairwise Distance MPD to null assemblages calculated over subsets of random species in the local phylogenetic tree. The more positive SES. MPD values indicate that species are more dispersed in the phylogenetic tree and the more negative SES. MPD values that species are closer in the phylogenetic tree Appendix 1. To control for the idiosyncratic effect of species identities, we established two different species combinations for each phylogenetic diversity level.
Thus, four taxonomic combinations were constructed comprising two combinations of distantly related species high phylogenetic diversity scenarios and two of more closely related species low phylogenetic diversity scenarios. High phylogenetic diversity scenarios were composed of distantly related species such as members of the Poaceae, Crassulaceae, Apiaceae, Caryophylaceae families see Fig. Specifically, Pistorinia hispanica is known to have CAM metabolism, species of the Poaceae family usually develop fasciculate roots, some species in these scenarios are rosette forming plants i.
Torilis leptophyllaCampanula erinusLimonium echioideswhile others do not form rosettes Ziziphora hispanicaSilene conica or Lomelosia stellatasome species maximum plant heights are around 40 mm Echinaria capitataPlantago afraCampanula erinuswhile others can grow above mm Torilis nodosa and T. In our high phylogenetic diversity scenarios, there are species with contrasting seed mass values i. Distance-based phylogenetic tree for the 28 annual plant species used to prepare the experimental scenarios.
The capital letters between brackets next to the names of species indicate the species combinations in what information would a phylogenetic tree give us they participated. In nonbold typeface, the high phylogenetic diversity scenarios A and B combinations ; in bold, the low phylogenetic diversity scenarios C and D combinations.
We established water availability treatments with two levels in a fully crossed factorial design: average precipitation vs. Each scenario was replicated in 10 to 16 units, thereby resulting in experimental assemblages. We used round pots with a diameter of 30 cm and height of 10 cm, which were filled with seed-free gypsum soil from a gypsum quarry located close to the collection sites.
We aimed to establish 10 plants of each seven coexisting species per pot, so we initially sowed 70 seeds per species in each one. Excess emergent seedlings were removed every two days trying to avoid clusters of seedlings to ensure the planned abundance of each species. By this way, we got to reproduce high densities of ephemeral, highly synchronized annual plants i. We watered pots to the soil water-carrying capacity for the first 20 weeks to ensure the establishment of experimental assemblages at the emergence stage mimicking natural field conditions and then commenced the water availability treatments, which were maintained for 19 weeks.
Between February and June, we monitored plant survival per species and per pot summing plants every two weeks, and we recorded the numbers of flowering plants once a week. In addition, for each species and pot we registered the final number of plants that reached the fruiting stage. Generalized linear mixed models GLMMs were employed to analyze the proportion of surviving, flowering, and fruiting plants per species and pot Table 1 ; Appendix 2 and to evaluate the overall proportion of species and plants that survived per pot Table 2 ; Appendix 2.
We used the irrigation treatment 2 levels: average and drought and the initial phylogenetic diversity 2 levels: high and low PD as fixed factors and we included the interaction term between both. We what are the patterns of a narcissist not consider the sampling moment to model the proportion of fruiting plants, because this variable what is design class diagram explain with example the percentage of the total cumulative number of fruiting plants per species in each pot.
Authors assure that legislation on seed collection has been accomplished. Permission obtained from responsible authority to collect seeds. The annual plant species that formed the experimental assemblages completed their life cycle within 5 months Fig. Plant mortality concentrated between the 2nd and the 3rd month of the experiment since plants died shortly what information would a phylogenetic tree give us fruit maturation.
Flowering started in the first weeks of the experiment and lasted for nearly four months Fig. In particular, in low water conditions, we found that the experimental assemblages formed of distantly related species resulted in more surviving plants per species Table 1Fig. Furthermore, plant survival regardless of species identity was higher in high phylogenetic diversity assemblages under drought conditions Fig.
Consequently, the experimental assemblages with high phylogenetic diversity were less sensitive to drought than the low phylogenetic diversity assemblages in terms of the plant survival, number of coexisting species, and numbers of flowering and fruiting plants in each experimental unit. Black lines represent high phylogenetic diversity PD scenarios and grey lines denote low phylogenetic diversity scenarios. Vertical bars represent the standard error.
Percent of fruiting plants per species and pot see Table 1. Black bars represent high phylogenetic diversity scenarios and grey bars low phylogenetic diversity scenarios. As hypothesized, phylogenetic relatedness among coexisting plants drives community level processes such as survival and reproduction. In particular, we demonstrated the higher resistance of phylogenetically diverse assemblages to drought in terms of plant survival and number of coexisting species over time, and even more, plants not only were able to survive more successfully to drought in phylogenetically diverse assemblages, but also more individuals completed the reproductive stage by setting flowers and fruits.
Overall, these results support the idea that phylogenetic relatedness predicts niche differences among species Hypothesis 1a in Fig.