Climate-land-use interactions shape tropical mountain biodiversity and ecosystem functions.

Agriculture and the exploitation of natural resources have transformed tropical mountain ecosystems across the world, and the consequences of these transformations for biodiversity and ecosystem functioning are largely unknown1-3. Conclusions that are derived from studies in non-mountainous areas are not suitable for predicting the effects of land-use changes on tropical mountains because the climatic environment rapidly changes with elevation, which may mitigate or amplify the effects of land use4,5. It is of key importance to understand how the interplay of climate and land use constrains biodiversity and ecosystem functions to determine the consequences of global change for mountain ecosystems. Here we show that the interacting effects of climate and land use reshape elevational trends in biodiversity and ecosystem functions on Africa's largest mountain, Mount Kilimanjaro (Tanzania). We find that increasing land-use intensity causes larger losses of plant and animal species richness in the arid lowlands than in humid submontane and montane zones. Increases in land-use intensity are associated with significant changes in the composition of plant, animal and microorganism communities; stronger modifications of plant and animal communities occur in arid and humid ecosystems, respectively. Temperature, precipitation and land use jointly modulate soil properties, nutrient turnover, greenhouse gas emissions, plant biomass and productivity, as well as animal interactions. Our data suggest that the response of ecosystem functions to land-use intensity depends strongly on climate; more-severe changes in ecosystem functioning occur in the arid lowlands and the cold montane zone. Interactions between climate and land use explained-on average-54% of the variation in species richness, species composition and ecosystem functions, whereas only 30% of variation was related to single drivers. Our study reveals that climate can modulate the effects of land use on biodiversity and ecosystem functioning, and points to a lowered resistance of ecosystems in climatically challenging environments to ongoing land-use changes in tropical mountainous regions.

Primary productivity and habitat protection predict elevational species richness and community biomass of large mammals on Mt. Kilimanjaro.

Despite their diversity and their large functional and cultural importance, the patterns and predictors of large mammal diversity along elevational gradients on tropical mountains remain poorly understood. Today, large mammals are threatened by human disturbances such as habitat destruction and hunting and may increasingly depend on the conservation of protected areas. Here, we use field data on the diversity of large mammals along a 3.6 km elevational gradient on Mt. Kilimanjaro to evaluate the importance of climate, net primary productivity and human impact for the distribution, species richness and community biomass of wild mammals. Mammal species richness was explored with camera traps on 66 study plots along an elevational gradient from 870 to 4,550 m a.s.l.. We applied path analysis and variance partitioning analysis to unravel the direct and indirect effects of temperature, precipitation, primary productivity, land use, land area, the protection of habitats and the occurrence of domestic mammals on the diversity of wild mammals. Both species richness and community biomass of wild mammals showed a unimodal distribution with elevation, peaking in the montane zone of Mt. Kilimanjaro. However, the peak shifted significantly to lower elevations when only protected habitats were considered. Wild mammal diversity increased with net primary productivity, protection of habitats and temperature. Our study underscores the importance of energy resources for the establishment of diversity gradients in large mammals. While temperature has been revealed as a direct predictor of diversity in most ectothermic taxa, in endothermic organisms temperature has stronger indirect effects, via a modulation of net primary productivity. Moreover, our study reveals how patterns of diversity on tropical mountains are influenced by human impact, pointing to the pivotal role of protected areas for the long-term conservation of mountain biodiversity.

Leaf traits mediate changes in invertebrate herbivory along broad environmental gradients on Mt. Kilimanjaro, Tanzania.

Temperature, primary productivity, plant functional traits, and herbivore abundances are considered key predictors of leaf herbivory but their direct and indirect contributions to community-level herbivory are not well understood along broad climatic gradients. Here, we determined elevational herbivory patterns and used a path analytical approach to disentangle the direct and indirect effects of climate, land use, net primary productivity (NPP), herbivore abundance, and plant functional traits on community-level invertebrate herbivory along the extensive elevational and land use gradients at Mt. Kilimanjaro, Tanzania. We recorded standing leaf herbivory caused by leaf chewers, leaf miners and leaf gallers on 55 study sites distributed in natural and anthropogenic habitats along a 3,060 m elevation gradient. We related the total community-level herbivory to climate (temperature and precipitation), NPP, plant functional traits (specific leaf area, leaf carbon-to-nitrogen [CN] ratio and leaf nitrogen-to-phosphorus [NP] ratio) and herbivore abundances. Leaf herbivory ranged from 5% to 11% along the elevation gradient. Total leaf herbivory showed unimodal pattern in natural habitats but a strongly contrasting bimodal pattern in anthropogenic habitats. We also detected some variation in the patterns of leaf herbivory along environmental gradients across feeding guilds with leaf chewers being responsible for a disproportionally large part of herbivory. Path analyses indicated that the variation in leaf herbivory was mainly driven by changes in leaf CN and NP ratios which were closely linked to changes in NPP in natural habitats. Similarly, patterns of leaf herbivory in anthropogenic habitats were best explained by variation in leaf CN ratios and a negative effect of land use. Our study elucidates the strong role of leaf nutrient stoichiometry and its linkages to climate and NPP for explaining the variation in leaf herbivory along broad climatic gradients. Furthermore, the study suggests that climatic changes and nutrient inputs in the course of land use change may alter leaf herbivory and consequently energy and nutrient fluxes in terrestrial habitats.

A dataset of occurrence of wild bees and their interaction with foraging plants along a livestock grazing gradient of northern Tanzania.

A dataset describing the occurrence of wild bees and their interaction with forage plants along livestock grazing gradient is critical in understanding bee-plant interaction networks and in developing conservation plans to ensure ecosystem services in human-modified landscapes. Despite this need, bee-plant datasets are scarce in Africa, and Tanzania is no exception. Therefore, in this article, we present a dataset of wild bee species richness, occurrence, and distribution collected across sites with different levels of livestock grazing intensity and forage resources thereby. The data presented in this paper supports a research article by Lasway et al., 2022 describing the effects of grazing intensity on East African bee assemblages. The paper presents primary data on bee species, collection method, date of collection, bee family, identifier, plant forage resource, forage plant life form, forage plant family, location (GPS coordinates), grazing intensity category, mean annual temperature (°C), and elevation (m asl). The data were collected intermittently between August 2018 and March 2020 from 24 study sites distributed along three levels of livestock grazing intensity with eight replicates for each: low, moderate, and high livestock grazing intensity. In each study site, two 50 × 50 m study plots were set from which bees and floral resources were sampled and quantified. The two plots were placed in a way to capture the overall structural heterogeneity of the respective habitat by placing the two plots in contrasting microhabitats where possible. For example, in moderately livestock-grazed habitats, plots were placed on sites with and without tree or shrub cover to ensure representativeness. This paper presents a dataset comprising 2,691 bee individuals from 183 species representing 55 genera of the five bee families: Halictidae (74), Apidae (63), Megachilidae (40), Andrenidae (5), and Colletidae (1). In addition, the dataset comprises 112 species of flowering plants that were identified as potential forage resources for bees. This paper supplements rare but critical data on bee pollinators in Northern Tanzania and advances our knowledge of the potential drivers of bee-pollinator whose populations diversity are declining globally. The dataset will also promote collaborations among researchers who would wish to combine and extend their data for further analysis to gain a broader understanding of the phenomenon on a larger spatial scale.

Plant traits mediate the effects of climate on phytophagous beetle diversity on Mt. Kilimanjaro.

Patterns of insect diversity along elevational gradients are well described in ecology. However, it remains little tested how variation in the quantity, quality, and diversity of food resources influence these patterns. Here we analyzed the direct and indirect effects of climate, food quantity (estimated by net primary productivity), quality (variation in the specific leaf area index, leaf nitrogen to phosphorus and leaf carbon to nitrogen ratio), and food diversity (diversity of leaf traits) on the species richness of phytophagous beetles along the broad elevation and land use gradients of Mt. Kilimanjaro, Tanzania. We sampled beetles at 65 study sites located in both natural and anthropogenic habitats, ranging from 866 to 4,550 m asl. We used path analysis to unravel the direct and indirect effects of predictor variables on species richness. In total, 3,154 phytophagous beetles representing 19 families and 304 morphospecies were collected. We found that the species richness of phytophagous beetles was bimodally distributed along the elevation gradient with peaks at the lowest (˜866 m asl) and upper mid-elevations (˜3,200 m asl) and sharply declined at higher elevations. Path analysis revealed temperature- and climate-driven changes in primary productivity and leaf trait diversity to be the best predictors of changes in the species richness of phytophagous beetles. Species richness increased with increases in mean annual temperature, primary productivity, and with increases in the diversity of leaf traits of local ecosystems. Our study demonstrates that, apart from temperature, the quantity and diversity of food resources play a major role in shaping diversity gradients of phytophagous insects. Drivers of global change, leading to a change of leaf traits and causing reductions in plant diversity and productivity, may consequently reduce the diversity of herbivore assemblages.

A synopsis of the Bee occurrence data of northern Tanzania.

BACKGROUND: Bees (Hymenoptera: Apoidea: Anthophila) are the most important group of pollinators with about 20,507 known species worldwide. Despite the critical role of bees in providing pollination services, studies aiming at understanding which species are present across disturbance gradients are scarce. Limited taxononomic information for the existing and unidentified bee species in Tanzania make their conservation haphazard. Here, we present a dataset of bee species records obtained from a survey in nothern Tanzania i.e. Kilimanjaro, Arusha and Manyara regions. Our findings serve as baseline data necessary for understanding the diversity and distribution of bees in the northern parts of the country, which is a critical step in devising robust conservation and monitoring strategies for their populations. NEW INFORMATION: In this paper, we present information on 45 bee species belonging to 20 genera and four families sampled using a combination of sweep-netting and pan trap methods. Most species (27, ~ 60%) belong to the family Halictidae followed by 16 species (35.5%) from the family Apidae. Megachilidae and Andrenidae were the least represented, each with only one species (2.2%). Additional species of Apidae and Megachilidae sampled during this survey are not yet published on Global Biodiversity Information Facility (GBIF), once they will be available on GBIF, they will be published in a subsequent paper. From a total of 953 occurrences, highest numbers were recorded in Kilimanjaro Region (n = 511), followed by Arusha (n = 410) and Manyara (n = 32), but this pattern reflects the sampling efforts of the research project rather than real bias in the distributions of bee species in northern Tanzania.

Species richness is more important for ecosystem functioning than species turnover along an elevational gradient.

Many experiments have shown that biodiversity enhances ecosystem functioning. However, we have little understanding of how environmental heterogeneity shapes the effect of diversity on ecosystem functioning and to what extent this diversity effect is mediated by variation in species richness or species turnover. This knowledge is crucial to scaling up the results of experiments from local to regional scales. Here we quantify the diversity effect and its components-that is, the contributions of variation in species richness and species turnover-for 22 ecosystem functions of microorganisms, plants and animals across 13 major ecosystem types on Mt Kilimanjaro, Tanzania. Environmental heterogeneity across ecosystem types on average increased the diversity effect from explaining 49% to 72% of the variation in ecosystem functions. In contrast to our expectation, the diversity effect was more strongly mediated by variation in species richness than by species turnover. Our findings reveal that environmental heterogeneity strengthens the relationship between biodiversity and ecosystem functioning and that species richness is a stronger driver of ecosystem functioning than species turnover. Based on a broad range of taxa and ecosystem functions in a non-experimental system, these results are in line with predictions from biodiversity experiments and emphasize that conserving biodiversity is essential for maintaining ecosystem functioning.

The Conservation of Native Honey Bees Is Crucial.

Recent studies have emphasized the role of the western honey bee, Apis mellifera, as a managed agricultural species worldwide, but also as a potential threat to endangered wild pollinators. This has resulted in the suggestion that honey bees should be regulated in natural areas to conserve wild pollinators. We argue that this perspective fails to appreciate the multifaceted nature of honey bees as native or introduced species with either managed or wild colonies. Wild populations of A. mellifera are currently imperiled, and natural areas are critical for the conservation of local subspecies and genotypes. We propose that a differentiation between managed and wild populations is required and encourage integrated conservation planning for all endangered wild bees, including A. mellifera.