Functional rarity and evenness are key facets of biodiversity to boost multifunctionality.

The functional traits of organisms within multispecies assemblages regulate biodiversity effects on ecosystem functioning. Yet how traits should assemble to boost multiple ecosystem functions simultaneously (multifunctionality) remains poorly explored. In a multibiome litter experiment covering most of the global variation in leaf trait spectra, we showed that three dimensions of functional diversity (dispersion, rarity, and evenness) explained up to 66% of variations in multifunctionality, although the dominant species and their traits remained an important predictor. While high dispersion impeded multifunctionality, increasing the evenness among functionally dissimilar species was a key dimension to promote higher multifunctionality and to reduce the abundance of plant pathogens. Because too-dissimilar species could have negative effects on ecosystems, our results highlight the need for not only diverse but also functionally even assemblages to promote multifunctionality. The effect of functionally rare species strongly shifted from positive to negative depending on their trait differences with the dominant species. Simultaneously managing the dispersion, evenness, and rarity in multispecies assemblages could be used to design assemblages aimed at maximizing multifunctionality independently of the biome, the identity of dominant species, or the range of trait values considered. Functional evenness and rarity offer promise to improve the management of terrestrial ecosystems and to limit plant disease risks.

Applying the Multi-Scale Integrated Analysis of Societal and Ecosystem Metabolism (MuSIASEM) to characterize the society-agriculture-forest system: the case of Huayopata, Cuzco (Peru).

The divergence between the disappearance of primary forests and the appearance of secondary forests indicates a set of circumstances that simultaneously converge in what we called the society-agriculture-forest complex. Such circumstances vary between places and over time and are associated with internal factors-factors originating within the reference system-and external factors-factors originating outside the borders of the reference system-restrict the use of standard strategies for any reality. We present a quantitative model that helps to understand the relationships of the society-agriculture-forest complex as a whole. This comprehensive understanding will allow a clearer discussion of the positive and negative consequences of prioritizing actions on any of the system components. Our model establishes a set of quantitative relationships among: (i) the requirements of food and timber products for society to maintain its structure and functionality, (ii) the level of openness of the society with respect to other societies for the exchange of resources, and (iii) the interface between the economic productive systems and the ecological productive systems. To test the model, the case of Huayopata (Peru) was studied. Findings suggest that the abandonment of agricultural production and, particularly, of tea favors the appearance of secondary forests. However, projects by the Congress of Peru to reactivate the production of tea without adequate technological support to improve the current processes that use firewood for boilers would put the forestry system at risk. In addition, a potential worker 'pull factor' could reconfigure the food system and impact on the local agricultural sector. Supplementary Information: The online version contains supplementary material available at 10.1007/s10668-022-02457-6.

The evolutionary assembly of forest communities along environmental gradients: recent diversification or sorting of pre-adapted clades?

Recent studies have demonstrated that ecological processes that shape community structure and dynamics change along environmental gradients. However, much less is known about how the emergence of the gradients themselves shape the evolution of species that underlie community assembly. In this study, we address how the creation of novel environments leads to community assembly via two nonmutually exclusive processes: immigration and ecological sorting of pre-adapted clades (ISPC), and recent adaptive diversification (RAD). We study these processes in the context of the elevational gradient created by the uplift of the Central Andes. We develop a novel approach and method based on the decomposition of species turnover into within- and among-clade components, where clades correspond to lineages that originated before mountain uplift. Effects of ISPC and RAD can be inferred from how components of turnover change with elevation. We test our approach using data from over 500 Andean forest plots. We found that species turnover between communities at different elevations is dominated by the replacement of clades that originated before the uplift of the Central Andes. Our results suggest that immigration and sorting of clades pre-adapted to montane habitats is the primary mechanism shaping tree communities across elevations.

Temperature sensitivity of soil respiration rates enhanced by microbial community response.

Soils store about four times as much carbon as plant biomass, and soil microbial respiration releases about 60 petagrams of carbon per year to the atmosphere as carbon dioxide. Short-term experiments have shown that soil microbial respiration increases exponentially with temperature. This information has been incorporated into soil carbon and Earth-system models, which suggest that warming-induced increases in carbon dioxide release from soils represent an important positive feedback loop that could influence twenty-first-century climate change. The magnitude of this feedback remains uncertain, however, not least because the response of soil microbial communities to changing temperatures has the potential to either decrease or increase warming-induced carbon losses substantially. Here we collect soils from different ecosystems along a climate gradient from the Arctic to the Amazon and investigate how microbial community-level responses control the temperature sensitivity of soil respiration. We find that the microbial community-level response more often enhances than reduces the mid- to long-term (90 days) temperature sensitivity of respiration. Furthermore, the strongest enhancing responses were observed in soils with high carbon-to-nitrogen ratios and in soils from cold climatic regions. After 90 days, microbial community responses increased the temperature sensitivity of respiration in high-latitude soils by a factor of 1.4 compared to the instantaneous temperature response. This suggests that the substantial carbon stores in Arctic and boreal soils could be more vulnerable to climate warming than currently predicted.

Microbial responses to warming enhance soil carbon loss following translocation across a tropical forest elevation gradient.

Tropical soils contain huge carbon stocks, which climate warming is projected to reduce by stimulating organic matter decomposition, creating a positive feedback that will promote further warming. Models predict that the loss of carbon from warming soils will be mediated by microbial physiology, but no empirical data are available on the response of soil carbon and microbial physiology to warming in tropical forests, which dominate the terrestrial carbon cycle. Here we show that warming caused a considerable loss of soil carbon that was enhanced by associated changes in microbial physiology. By translocating soils across a 3000 m elevation gradient in tropical forest, equivalent to a temperature change of ± 15 °C, we found that soil carbon declined over 5 years by 4% in response to each 1 °C increase in temperature. The total loss of carbon was related to its original quantity and lability, and was enhanced by changes in microbial physiology including increased microbial carbon-use-efficiency, shifts in community composition towards microbial taxa associated with warmer temperatures, and increased activity of hydrolytic enzymes. These findings suggest that microbial feedbacks will cause considerable loss of carbon from tropical forest soils in response to predicted climatic warming this century.

Adaptation of soil microbial growth to temperature: Using a tropical elevation gradient to predict future changes.

Terrestrial biogeochemical feedbacks to the climate are strongly modulated by the temperature response of soil microorganisms. Tropical forests, in particular, exert a major influence on global climate because they are the most productive terrestrial ecosystem. We used an elevation gradient across tropical forest in the Andes (a gradient of 20°C mean annual temperature, MAT), to test whether soil bacterial and fungal community growth responses are adapted to long-term temperature differences. We evaluated the temperature dependency of soil bacterial and fungal growth using the leucine- and acetate-incorporation methods, respectively, and determined indices for the temperature response of growth: Q10 (temperature sensitivity over a given 10oC range) and Tmin (the minimum temperature for growth). For both bacterial and fungal communities, increased MAT (decreased elevation) resulted in increases in Q10 and Tmin of growth. Across a MAT range from 6°C to 26°C, the Q10 and Tmin varied for bacterial growth (Q10-20  = 2.4 to 3.5; Tmin  = -8°C to -1.5°C) and fungal growth (Q10-20  = 2.6 to 3.6; Tmin  = -6°C to -1°C). Thus, bacteria and fungi did not differ significantly in their growth temperature responses with changes in MAT. Our findings indicate that across natural temperature gradients, each increase in MAT by 1°C results in increases in Tmin of microbial growth by approximately 0.3°C and Q10-20 by 0.05, consistent with long-term temperature adaptation of soil microbial communities. A 2°C warming would increase microbial activity across a MAT gradient of 6°C to 26°C by 28% to 15%, respectively, and temperature adaptation of microbial communities would further increase activity by 1.2% to 0.3%. The impact of warming on microbial activity, and the related impact on soil carbon cycling, is thus greater in regions with lower MAT. These results can be used to predict future changes in the temperature response of microbial activity over different levels of warming and over large temperature ranges, extending to tropical regions.

Microbes follow Humboldt: temperature drives plant and soil microbial diversity patterns from the Amazon to the Andes.

More than 200 years ago, Alexander von Humboldt reported that tropical plant species richness decreased with increasing elevation and decreasing temperature. Surprisingly, coordinated patterns in plant, bacterial, and fungal diversity on tropical mountains have not yet been observed, despite the central role of soil microorganisms in terrestrial biogeochemistry and ecology. We studied an Andean transect traversing 3.5 km in elevation to test whether the species diversity and composition of tropical forest plants, soil bacteria, and fungi follow similar biogeographical patterns with shared environmental drivers. We found coordinated changes with elevation in all three groups: species richness declined as elevation increased, and the compositional dissimilarity among communities increased with increased separation in elevation, although changes in plant diversity were larger than in bacteria and fungi. Temperature was the dominant driver of these diversity gradients, with weak influences of edaphic properties, including soil pH. The gradients in microbial diversity were strongly correlated with the activities of enzymes involved in organic matter cycling, and were accompanied by a transition in microbial traits towards slower-growing, oligotrophic taxa at higher elevations. We provide the first evidence of coordinated temperature-driven patterns in the diversity and distribution of three major biotic groups in tropical ecosystems: soil bacteria, fungi, and plants. These findings suggest that interrelated and fundamental patterns of plant and microbial communities with shared environmental drivers occur across landscape scales. These patterns are revealed where soil pH is relatively constant, and have implications for tropical forest communities under future climate change.

Solar radiation and functional traits explain the decline of forest primary productivity along a tropical elevation gradient.

One of the major challenges in ecology is to understand how ecosystems respond to changes in environmental conditions, and how taxonomic and functional diversity mediate these changes. In this study, we use a trait-spectra and individual-based model, to analyse variation in forest primary productivity along a 3.3 km elevation gradient in the Amazon-Andes. The model accurately predicted the magnitude and trends in forest productivity with elevation, with solar radiation and plant functional traits (leaf dry mass per area, leaf nitrogen and phosphorus concentration, and wood density) collectively accounting for productivity variation. Remarkably, explicit representation of temperature variation with elevation was not required to achieve accurate predictions of forest productivity, as trait variation driven by species turnover appears to capture the effect of temperature. Our semi-mechanistic model suggests that spatial variation in traits can potentially be used to estimate spatial variation in productivity at the landscape scale.

Variation in leaf wettability traits along a tropical montane elevation gradient.

Leaf wetting is often considered to have negative effects on plant function, such that wet environments may select for leaves with certain leaf surface, morphological, and architectural traits that reduce leaf wettability. However, there is growing recognition that leaf wetting can have positive effects. We measured variation in two traits, leaf drip tips and leaf water repellency, in a series of nine tropical forest communities occurring along a 3300-m elevation gradient in southern Peru. To extend this climatic gradient, we also assembled published leaf water repellency values from 17 additional sites. We then tested hypotheses for how these traits should vary as a function of climate. Contrary to expectations, we found that the proportion of species with drip tips did not increase with increasing precipitation. Instead, drip tips increased with increasing temperature. Moreover, leaf water repellency was very low in our sites and the global analysis indicated high repellency only in sites with low precipitation and temperatures. Our findings suggest that drip tips and repellency may not solely reflect the negative effects of wetting on plant function. Understanding the drivers of leaf wettability traits can provide insight into the effects of leaf wetting on plant, community, and ecosystem function.

Scaling leaf respiration with nitrogen and phosphorus in tropical forests across two continents.

Leaf dark respiration (Rdark ) represents an important component controlling the carbon balance in tropical forests. Here, we test how nitrogen (N) and phosphorus (P) affect Rdark and its relationship with photosynthesis using three widely separated tropical forests which differ in soil fertility. Rdark was measured on 431 rainforest canopy trees, from 182 species, in French Guiana, Peru and Australia. The variation in Rdark was examined in relation to leaf N and P content, leaf structure and maximum photosynthetic rates at ambient and saturating atmospheric CO2 concentration. We found that the site with the lowest fertility (French Guiana) exhibited greater rates of Rdark per unit leaf N, P and photosynthesis. The data from Australia, for which there were no phylogenetic overlaps with the samples from the South American sites, yielded the most distinct relationships of Rdark with the measured leaf traits. Our data indicate that no single universal scaling relationship accounts for variation in Rdark across this large biogeographical space. Variability between sites in the absolute rates of Rdark and the Rdark  : photosynthesis ratio were driven by variations in N- and P-use efficiency, which were related to both taxonomic and environmental variability.

Scale dependence of canopy trait distributions along a tropical forest elevation gradient.

Average responses of forest foliar traits to elevation are well understood, but far less is known about trait distributional responses to elevation at multiple ecological scales. This limits our understanding of the ecological scales at which trait variation occurs in response to environmental drivers and change. We analyzed and compared multiple canopy foliar trait distributions using field sampling and airborne imaging spectroscopy along an Andes-to-Amazon elevation gradient. Field-estimated traits were generated from three community-weighting methods, and remotely sensed estimates of traits were made at three scales defined by sampling grain size and ecological extent. Field and remote sensing approaches revealed increases in average leaf mass per unit area (LMA), water, nonstructural carbohydrates (NSCs) and polyphenols with increasing elevation. Foliar nutrients and photosynthetic pigments displayed little to no elevation trend. Sample weighting approaches had little impact on field-estimated trait responses to elevation. Plot representativeness of trait distributions at landscape scales decreased with increasing elevation. Remote sensing indicated elevation-dependent increases in trait variance and distributional skew. Multiscale invariance of LMA, leaf water and NSC mark these traits as candidates for tracking forest responses to changing climate. Trait-based ecological studies can be greatly enhanced with multiscale studies made possible by imaging spectroscopy.

The variation of productivity and its allocation along a tropical elevation gradient: a whole carbon budget perspective.

Why do forest productivity and biomass decline with elevation? To address this question, research to date generally has focused on correlative approaches describing changes in woody growth and biomass with elevation. We present a novel, mechanistic approach to this question by quantifying the autotrophic carbon budget in 16 forest plots along a 3300 m elevation transect in Peru. Low growth rates at high elevations appear primarily driven by low gross primary productivity (GPP), with little shift in either carbon use efficiency (CUE) or allocation of net primary productivity (NPP) between wood, fine roots and canopy. The lack of trend in CUE implies that the proportion of photosynthate allocated to autotrophic respiration is not sensitive to temperature. Rather than a gradual linear decline in productivity, there is some limited but nonconclusive evidence of a sharp transition in NPP between submontane and montane forests, which may be caused by cloud immersion effects within the cloud forest zone. Leaf-level photosynthetic parameters do not decline with elevation, implying that nutrient limitation does not restrict photosynthesis at high elevations. Our data demonstrate the potential of whole carbon budget perspectives to provide a deeper understanding of controls on ecosystem functioning and carbon cycling.

Leaf-level photosynthetic capacity in lowland Amazonian and high-elevation Andean tropical moist forests of Peru.

We examined whether variations in photosynthetic capacity are linked to variations in the environment and/or associated leaf traits for tropical moist forests (TMFs) in the Andes/western Amazon regions of Peru. We compared photosynthetic capacity (maximal rate of carboxylation of Rubisco (Vcmax ), and the maximum rate of electron transport (Jmax )), leaf mass, nitrogen (N) and phosphorus (P) per unit leaf area (Ma , Na and Pa , respectively), and chlorophyll from 210 species at 18 field sites along a 3300-m elevation gradient. Western blots were used to quantify the abundance of the CO2 -fixing enzyme Rubisco. Area- and N-based rates of photosynthetic capacity at 25°C were higher in upland than lowland TMFs, underpinned by greater investment of N in photosynthesis in high-elevation trees. Soil [P] and leaf Pa were key explanatory factors for models of area-based Vcmax and Jmax but did not account for variations in photosynthetic N-use efficiency. At any given Na and Pa , the fraction of N allocated to photosynthesis was higher in upland than lowland species. For a small subset of lowland TMF trees examined, a substantial fraction of Rubisco was inactive. These results highlight the importance of soil- and leaf-P in defining the photosynthetic capacity of TMFs, with variations in N allocation and Rubisco activation state further influencing photosynthetic rates and N-use efficiency of these critically important forests.

Predicting trait-environment relationships for venation networks along an Andes-Amazon elevation gradient.

Understanding functional trait-environment relationships (TERs) may improve predictions of community assembly. However, many empirical TERs have been weak or lacking conceptual foundation. TERs based on leaf venation networks may better link individuals and communities via hydraulic constraints. We report measurements of vein density, vein radius, and leaf thickness for more than 100 dominant species occurring in ten forest communities spanning a 3,300 m Andes-Amazon elevation gradient in Peru. We use these data to measure the strength of TERs at community scale and to determine whether observed TERs are similar to those predicted by physiological theory. We found strong support for TERs between all traits and temperature, as well weaker support for a predicted TER between maximum abundance-weighted leaf transpiration rate and maximum potential evapotranspiration. These results provide one approach for developing a more mechanistic trait-based community assembly theory.

Global variability in leaf respiration in relation to climate, plant functional types and leaf traits.

Leaf dark respiration (Rdark ) is an important yet poorly quantified component of the global carbon cycle. Given this, we analyzed a new global database of Rdark and associated leaf traits. Data for 899 species were compiled from 100 sites (from the Arctic to the tropics). Several woody and nonwoody plant functional types (PFTs) were represented. Mixed-effects models were used to disentangle sources of variation in Rdark . Area-based Rdark at the prevailing average daily growth temperature (T) of each site increased only twofold from the Arctic to the tropics, despite a 20°C increase in growing T (8-28°C). By contrast, Rdark at a standard T (25°C, Rdark (25) ) was threefold higher in the Arctic than in the tropics, and twofold higher at arid than at mesic sites. Species and PFTs at cold sites exhibited higher Rdark (25) at a given photosynthetic capacity (Vcmax (25) ) or leaf nitrogen concentration ([N]) than species at warmer sites. Rdark (25) values at any given Vcmax (25) or [N] were higher in herbs than in woody plants. The results highlight variation in Rdark among species and across global gradients in T and aridity. In addition to their ecological significance, the results provide a framework for improving representation of Rdark in terrestrial biosphere models (TBMs) and associated land-surface components of Earth system models (ESMs).

Climate Warming and Soil Carbon in Tropical Forests: Insights from an Elevation Gradient in the Peruvian Andes.

The temperature sensitivity of soil organic matter (SOM) decomposition in tropical forests will influence future climate. Studies of a 3.5-kilometer elevation gradient in the Peruvian Andes, including short-term translocation experiments and the examination of the long-term adaptation of biota to local thermal and edaphic conditions, have revealed several factors that may regulate this sensitivity. Collectively this work suggests that, in the absence of a moisture constraint, the temperature sensitivity of decomposition is regulated by the chemical composition of plant debris (litter) and both the physical and chemical composition of preexisting SOM: higher temperature sensitivities are found in litter or SOM that is more chemically complex and in SOM that is less occluded within aggregates. In addition, the temperature sensitivity of SOM in tropical montane forests may be larger than previously recognized because of the presence of "cold-adapted" and nitrogen-limited microbial decomposers and the possible future alterations in plant and microbial communities associated with warming. Studies along elevation transects, such as those reviewed here, can reveal factors that will regulate the temperature sensitivity of SOM. They can also complement and guide in situ soil-warming experiments, which will be needed to understand how this vulnerability to temperature may be mediated by altered plant productivity under future climatic change.

Serving IPV Survivors in Culturally Diverse Communities: Perspectives From Current Service Providers.

This qualitative study examines current IPV service providers' perspectives on service delivery methods that best reach and serve IPV survivors from culturally diverse communities. Semi-structured interviews were conducted with 11 service providers, and transcripts were analyzed for themes related to service providers' experiences. Five themes emerged from the data that suggest best practices for reaching and serving survivors from culturally diverse backgrounds, including understanding survivors' backgrounds, promoting trust and inclusivity, building community relationships, providing culturally responsive education on IPV, and supporting current and future staff with training. Study findings provide implications for the education and training of future service providers.

Woody plant taxonomic, functional, and phylogenetic diversity decrease along elevational gradients in Andean tropical montane forests: Environmental filtering and arrival of temperate taxa.

Mountains are paramount for exploring biodiversity patterns due to the mosaic of topographies and climates encompassed over short distances. Biodiversity research has traditionally focused on taxonomic diversity when investigating changes along elevational gradients, but other facets should be considered. For first time, we simultaneously assessed elevational trends in taxonomic, functional, and phylogenetic diversity of woody plants in Andean tropical montane forests and explored their underlying ecological and evolutionary causes. This investigation covered four transects (traversing ca. 2200 m a.s.l.) encompassing 114 plots of 0.1 ha across a broad latitudinal range (ca. 10°). Using Hill numbers to quantify abundance-based diversity among 37,869 individuals we observed a consistent decrease in taxonomic, functional, and phylogenetic diversity as elevation increased, although the decrease was less pronounced for higher Hill orders. The exception was a slight increase in phylogenetic diversity when dominant species were over-weighted. The decrease in taxonomic and functional diversity might be attributed to an environmental filtering process towards highlands, where the increasingly harsher conditions exclude species and functional strategies. Besides, the differences in steepness decrease between Hill orders suggest that rare species disproportionately contribute to functional diversity. For phylogenetic diversity the shifting elevational trend between Hill orders indicates a greater than previously considered influence in central Andean highlands of tropical lowlands originated species with strong niche conservatism relative to distantly related temperate lineages. This could be explained by a decreasing presence and abundance of temperate, extratropical taxa towards the central Andes relative to northern or southern Andes, where they are more prevalent.

Historical Assembly of Andean Tree Communities.

Patterns of species diversity have been associated with changes in climate across latitude and elevation. However, the ecological and evolutionary mechanisms underlying these relationships are still actively debated. Here, we present a complementary view of the well-known tropical niche conservatism (TNC) hypothesis, termed the multiple zones of origin (MZO) hypothesis, to explore mechanisms underlying latitudinal and elevational gradients of phylogenetic diversity in tree communities. The TNC hypothesis posits that most lineages originate in warmer, wetter, and less seasonal environments in the tropics and rarely colonize colder, drier, and more seasonal environments outside of the tropical lowlands, leading to higher phylogenetic diversity at lower latitudes and elevations. In contrast, the MZO hypothesis posits that lineages also originate in temperate environments and readily colonize similar environments in the tropical highlands, leading to lower phylogenetic diversity at lower latitudes and elevations. We tested these phylogenetic predictions using a combination of computer simulations and empirical analyses of tree communities in 245 forest plots located in six countries across the tropical and subtropical Andes. We estimated the phylogenetic diversity for each plot and regressed it against elevation and latitude. Our simulated and empirical results provide strong support for the MZO hypothesis. Phylogenetic diversity among co-occurring tree species increased with both latitude and elevation, suggesting an important influence on the historical dispersal of lineages with temperate origins into the tropical highlands. The mixing of different floras was likely favored by the formation of climatically suitable corridors for plant migration due to the Andean uplift. Accounting for the evolutionary history of plant communities helps to advance our knowledge of the drivers of tree community assembly along complex climatic gradients, and thus their likely responses to modern anthropogenic climate change.

Tropical forest lianas have greater non-structural carbohydrate concentrations in the stem xylem than trees.

Lianas (woody vines) are important components of tropical forests and are known to compete with host trees for resources, decrease tree growth and increase tree mortality. Given the observed increases in liana abundance in some forests and their impacts on forest function, an integrated understanding of carbon dynamics of lianas and liana-infested host trees is critical for improved prediction of tropical forest responses to climate change. Non-structural carbohydrates (NSC) are the main substrate for plant metabolism (e.g., growth, respiration), and have been implicated in enabling tree survival under environmental stress, but little is known of how they vary among life-forms or of how liana infestation impacts host tree NSC. We quantified stem total NSC (NSC) concentrations and its fractions (starch and soluble sugars) in trees without liana infestation, trees with more than 50% of the canopy covered by lianas, and the lianas infesting those trees. We hypothesized that i) liana infestation depletes NSC storage in host trees by reducing carbon assimilation due to competition for resources; ii) trees and lianas, which greatly differ in functional traits related to water transport and carbon uptake, would also have large differences in NSC storage, and that As water availability has a significant role in NSC dynamics of Amazonian tree species, we tested these hypotheses within a moist site in western Amazonia and a drier forest site in southern Amazonia. We did not find any difference in NSC, starch or soluble sugar concentrations between infested and non-infested trees, in either site. This result suggests that negative liana impact on trees may be mediated through mechanisms other than depletion of host tree NSC concentrations. We found lianas have higher stem NSC and starch than trees in both sites. The consistent differences in starch concentrations, a long term NSC reserve, between life forms across sites reflect differences in carbon gain and use of lianas and trees. Soluble sugar concentrations were higher in lianas than in trees in the moist site but indistinguishable between life forms in the dry site. The lack of difference in soluble sugars between trees and lianas in the dry site emphasize the importance of this NSC fraction for plant metabolism of plants occurring in water limited environments. Abstract in Portuguese and Spanish are available in the supplementary material.