Modified forests are vital for species communities and ecological functionality in a heterogeneous South African landscape

Land-use change is a major threat to forest ecosystems worldwide. Therefore, understanding the effects of human forest modification on biodiversity is an important task for conservation ecologists. The main objective of my dissertation was to evaluate how different intensities of forest modification contribute to the maintenance of species diversity and ecosystem functionality in a human-modified landscape. For this purpose, I based my studies in a heterogeneous landscape around two nature reserves, Vernon Crookes and Oribi Gorge, in KwaZulu-Natal, South Africa. I selected six most representative types of scarp forest modification – ranging from continuous forest and natural forest fragments in nature reserves to fragments within plantations and agricultural matrix, forested gardens, and secondary forest. In a total of 36 study sites, I assessed flower-visiting insects using insect traps and recorded local bird assemblages with point counts. Further, I observed flower visitation and seed removal on the native and widespread tree Celtis africana (Ulmacea) to analyse whether forest modification affects pollination and seed dispersal services. To assess how forest configuration affects the dispersal of animals, I carried out an in-depth study on the movement behaviour of bird assemblages within and among forest patches in the Vernon Crookes region. By means of direct observations and bird mist-netting, I followed up bird movements across nine forest fragments belonging to three different forest types. In all these three projects I give special attention to the responses of the different functional groups of a species community. The richness of flower-visiting insects, community composition and flower visitation on C. africana differed significantly among the different forest types and between two study seasons in 2009 and 2010. Both flower visitor richness and flower visitation rates were strongly enhanced in the human-modified forests. This could be explained by a high abundance of large-bodied pollinators in these sites. In particular, feral honey bees (Apis mellifera) played a major role in the pollination of C. africana trees located in forest fragments within plantations and agriculture, forested gardens and secondary forests. However, effective fruit set of C. africana was not enhanced by an increase of flower visitation, possibly due to the tree’s capability of wind pollination. This implies that even though forest modification can strongly alter insect assemblages, pollination services for trees with unspecialized flowers may remain resilient at a landscape scale. Bird species richness was not significantly different among forest types. However, I found a significant increase in bird abundance in modified forests. In particular, fragments within agriculture, forested gardens, and secondary forests attracted a large number of forest generalists, shrubland and open country species. The abundance of forest specialists however, was much lower in modified forests. Changes in the composition of bird functional groups were also confirmed by multivariate analysis, which clearly separated bird communities by forest type. I found the highest abundance of frugivores visiting C. africana in natural forest fragments, fragments within agriculture, forested gardens and secondary forests. That was also true for the estimated total fruit removal per C. africana tree, even though the differences among the forest types were not significant. In summary, I could show that overall bird abundance and seed removal services can be enhanced in modified forests. However, the results also underline the importance of protected natural forest for bird specialist species sensitive to human disturbance. I found a very high movement activity of the overall bird community among the nine forest fragments that was significantly structured by bird functional groups. Especially, frugivorous birds, forest specialists and large-bodied species showed the highest dispersal abilities across the landscape. These results might be facilitated by overall high fragment quality, providing food and shelter, as well as the close proximity among the forest fragments within the landscape. Yet, a fourth-corner analysis revealed that even though modified forests were rather attractive to frugivores, forest specialists as well as large-bodied species, there was still a high affinity of the latter functional groups to natural forest fragments, close canopy cover and large fragment size. Only a small proportion of the overall bird community was recorded to steadily persist in the forest fragments. In particular, patches in the agricultural landscape were frequently used by resident insectivores and forest generalists. Ultimately, these findings suggest that remnant forest fragments may represent valuable stepping-stones as well as permanent habitat for many forest birds and thus, will help to maintain regional bird assemblages in human-modified landscapes. Overall, my results strongly suggest that modified forests contribute to the maintenance of species diversity and ecosystem functionality in a human-modified landscape. With respect to a vast increase of human-modified forests worldwide, evidence of a high conservation potential of these habitats is encouraging news for conservation managers. In particular, modified forests that are located in close proximity to protected areas have high conservation priority as they may expand buffer zones around natural forests in human-modified landscapes. Generalizations, however, should be considered with caution. My findings strongly emphasize that human-modified forests do not completely compensate for the overall loss of natural habitat. High sensitivity of forest specialist species and overall changes in local community composition demonstrate that natural forests are essential to maintain species diversity at a larger scale. Additionally, high flexibility towards habitat changes of many species in the study region might be based on the patchy historic distribution of scarp forest that has strongly been determined by terrain and orographic conditions of the environment. Thus, it is possible that an evolutionary adaptation has lessened the vulnerability of the region’s fauna and flora to the present anthropogenic forest fragmentation. Ultimately, most of the forest types in the study region are characterized by high habitat quality, including for example resource availability, structural heterogeneity and close proximity to further forest patches, so that altogether, they contribute to the high species diversity. Consequently, the maintenance of structurally rich forest habitat is essential to maintain species diversity and ecological functionality in human-modified landscapes

Metatranscriptomics reveals contrasting effects of elevation on the activity of bacteria and bacterial viruses in soil

Soil microbial diversity affects ecosystem functioning and global biogeochemical cycles. Soil bacterial communities catalyse a diversity of biogeochemical reactions and have thus sparked considerable scientific interest. One driver of bacterial community dynamics in natural ecosystems has so far been largely neglected: the predator-prey interactions between bacterial viruses (bacteriophages) and bacteria. To generate ground level knowledge on environmental drivers of these particular predator-prey dynamics, we propose an activity-based ecological framework to simultaneous capture community dynamics of bacteria and bacteriophages in soils. An ecological framework and specifically the analyses of community dynamics across latitudinal and elevational gradients have been widely used in ecology to understand community-wide responses of innumerable taxa to environmental change, in particular to climate. Here, we tested the hypothesis that the activity of bacteria and bacteriophages codeclines across an elevational gradient. We used metatranscriptomics to investigate bacterial and bacteriophage activity patterns at five sites across 400 elevational metres in the Swiss Alps in 2015 and 2017. We found that metabolic activity (transcription levels) of bacteria declined significantly with increasing elevation, but activity of bacteriophages did not. We showed that bacteriophages are consistently active in soil along the entire gradient, making bacteriophage activity patterns divergent from that of their putative bacterial prey. Future efforts will be necessary to link the environment-activity relationship to predator-prey dynamics, and to understand the magnitude of viral contributions to carbon, nitrogen and phosphorus cycling when infection causes bacterial cell death, a process that may represent an overlooked component of soil biogeochemical cycles

Scatter-hoarding birds disperse seeds to sites unfavorable for plant regeneration

Scatter-hoarding birds provide effective long-distance seed dispersal for plants. Transporting seeds far promotes population spread, colonization of new areas, and connectivity between populations. However, whether seeds transported over long distances are deposited in habitats favorable to plant regeneration has rarely been investigated, mainly due to methodological constraints. To investigate dispersal patterns and distances of Swiss stone pine (Pinus cembra) seeds we utilized advances in tracking technology to track the movements of their sole disperser, the spotted nutcracker (Nucifraga caryocatactes). We found routine individual movements between single seed harvesting and seed caching site. Harvesting sites of individual birds overlapped, whereas seed caching sites were separated and located on average 5.3 km away from the harvesting site. Interestingly, most distant caching sites were located at low elevations and in spruce forest, where Swiss stone pine does not naturally occur. This suggests that nutcrackers disperse seeds over long distances but that a large portion of these seeds are cached outside the known pine habitat. Therefore, we conclude that the implications of such long-distance seed dispersal movements for plant populations should be carefully considered in combination with the effects of habitat quality on plant recruitment

Downsizing of animal communities triggers stronger functional than structural decay in seed-dispersal networks

Downsizing of animal communities due to defaunation is prevalent in many ecosystems. Yet, we know little about its consequences for ecosystem functions such as seed dispersal. Here, we use eight seed-dispersal networks sampled across the Andes and simulate how downsizing of avian frugivores impacts structural network robustness and seed dispersal. We use a trait-based modeling framework to quantify the consequences of downsizing—relative to random extinctions—for the number of interactions and secondary plant extinctions (as measures of structural robustness) and for long-distance seed dispersal (as a measure of ecosystem function). We find that downsizing leads to stronger functional than structural losses. For instance, 10% size-structured loss of bird species results in almost 40% decline of long-distance seed dispersal, but in less than 10% of structural loss. Our simulations reveal that measures of the structural robustness of ecological networks underestimate the consequences of animal extinction and downsizing for ecosystem functioning.Fil: Donoso, Isabel. Senckenberg Biodiversity and Climate Research Centre; AlemaniaFil: Sorensen, Marjorie C.. Senckenberg Biodiversity and Climate Research Centre; Alemania. University of Guelph; Canadá. Goethe Universitat Frankfurt; AlemaniaFil: Blendinger, Pedro Gerardo. Universidad Nacional de Tucumán. Instituto de Ecología Regional. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Tucumán. Instituto de Ecología Regional; ArgentinaFil: Kissling, W. Daniel. University of Amsterdam; Países BajosFil: Neuschulz, Eike Lena. Senckenberg Biodiversity and Climate Research Centre; AlemaniaFil: Mueller, Thomas. Senckenberg Biodiversity and Climate Research Centre; AlemaniaFil: Schleuning, Matthias. Senckenberg Biodiversity and Climate Research Centre; Alemani

Avian seed dispersal may be insufficient for plants to track future temperature change on tropical mountains

AIM: Climate change causes shifts in species ranges globally. Terrestrial plant species often lag behind temperature shifts, and it is unclear to what extent animal-dispersed plants can track climate change. Here, we estimate the ability of bird-dispersed plant species to track future temperature change on a tropical mountain. LOCATION: Tropical elevational gradient (500–3500 m.a.s.l.) in the Manú biosphere reserve, Peru. TIME PERIOD: From 1960–1990 to 2061–2080. TAXA: Fleshy-fruited plants and avian frugivores. METHODS: Using simulations based on the functional traits of avian frugivores and fruiting plants, we quantified the number of long-distance dispersal (LDD) events that woody plant species would require to track projected temperature shifts on a tropical mountain by the year 2070 under different greenhouse gas emission scenarios [representative concentration pathway (RCP) 2.6, 4.5 and 8.5]. We applied this approach to 343 bird-dispersed woody plant species. RESULTS: Our simulations revealed that bird-dispersed plants differed in their climate-tracking ability, with large-fruited and canopy plants exhibiting a higher climate-tracking ability. Our simulations also suggested that even under scenarios of strong and intermediate mitigation of greenhouse gas emissions (RCP 2.6 and 4.5), sufficient upslope dispersal would require several LDD events by 2070, which is unlikely for the majority of woody plant species. Furthermore, the ability of plant species to track future changes in temperature increased in simulations with a low degree of trait matching between plants and birds, suggesting that plants in generalized seed-dispersal systems might be more resilient to climate change. MAIN CONCLUSION: Our study illustrates how the functional traits of plants and animals can inform predictive models of species dispersal and range shifts under climate change and suggests that the biodiversity of tropical mountain ecosystems is highly vulnerable to future warming. The increasing availability of functional trait data for plants and animals globally will allow parameterization of similar models for many other seed-dispersal systems

Similar composition of functional roles in Andean seed-dispersal networks, despite high species and interaction turnover

The species composition of local communities varies in space, and its similarity generally decreases with increasing geographic distance between communities, a phenomenon known as distance decay of similarity. It is, however, not known how changes in local species composition affect ecological processes, that is, whether they lead to differences in the local composition of species' functional roles. We studied eight seed-dispersal networks along the South American Andes and compared them with regard to their species composition and their composition of functional roles. We tested (1) if changes in bird species composition lead to changes in the composition of bird functional roles, and (2) if the similarity in species composition and functional-role composition decreased with increasing geographic distance between the networks. We also used cluster analysis to (3) identify bird species with similar roles across all networks based on the similarity in the plants they consume, (i) considering only the species identity of the plants and (ii) considering the functional traits of the plants. Despite strong changes in species composition, the networks along the Andes showed similar composition of functional roles. (1) Changes in species composition generally did not lead to changes in the composition of functional roles. (2) Similarity in species composition, but not functional-role composition, decreased with increasing geographic distance between the networks. (3) The cluster analysis considering the functional traits of plants identified bird species with similar functional roles across all networks. The similarity in functional roles despite the high species turnover suggests that the ecological process of seed dispersal is organized similarly along the Andes, with similar functional roles fulfilled locally by different sets of species. The high species turnover, relative to functional turnover, also indicates that a large number of bird species are needed to maintain the seed-dispersal process along the Andes.Fil: Dehling, D. Matthias. University of Canterbury; Nueva ZelandaFil: Peralta, Guadalupe. University of Canterbury; Nueva ZelandaFil: Bender, Irene Maria Antoinetta. Universidad Nacional de Tucumán. Instituto de Ecología Regional. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Tucumán. Instituto de Ecología Regional; ArgentinaFil: Blendinger, Pedro Gerardo. Universidad Nacional de Tucumán. Instituto de Ecología Regional. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Tucumán. Instituto de Ecología Regional; ArgentinaFil: Böhning Gaese, Katrin. Goethe Universitat Frankfurt; AlemaniaFil: Muñoz, Marcia C.. Universidad de la Salle; ColombiaFil: Neuschulz, Eike Lena. Senckenberg Biodiversität Und Klima Forschungszentrum; AlemaniaFil: Quitián, Marta. Senckenberg Biodiversität Und Klima Forschungszentrum; AlemaniaFil: Saavedra, Francisco. Universidad Mayor de San Andrés; BoliviaFil: Santillán, Vinicio. Senckenberg Biodiversität Und Klima Forschungszentrum; AlemaniaFil: Schleuning, Matthias. Senckenberg Biodiversität Und Klima Forschungszentrum; AlemaniaFil: Stouffer, Daniel B.. University of Canterbury; Nueva Zeland

Avian seed dispersal may be insufficient for plants to track future temperature change on tropical mountains

[Aim] Climate change causes shifts in species ranges globally. Terrestrial plant species often lag behind temperature shifts, and it is unclear to what extent animal-dispersed plants can track climate change. Here, we estimate the ability of bird-dispersed plant species to track future temperature change on a tropical mountain.[Location] Tropical elevational gradient (500–3500 m.a.s.l.) in the Manú biosphere reserve, Peru. [Time period] From 1960–1990 to 2061–2080. [Taxa] Fleshy-fruited plants and avian frugivores. [Methods] Using simulations based on the functional traits of avian frugivores and fruiting plants, we quantified the number of long-distance dispersal (LDD) events that woody plant species would require to track projected temperature shifts on a tropical mountain by the year 2070 under different greenhouse gas emission scenarios [representative concentration pathway (RCP) 2.6, 4.5 and 8.5]. We applied this approach to 343 bird-dispersed woody plant species. [Results] Our simulations revealed that bird-dispersed plants differed in their climate-tracking ability, with large-fruited and canopy plants exhibiting a higher climate-tracking ability. Our simulations also suggested that even under scenarios of strong and intermediate mitigation of greenhouse gas emissions (RCP 2.6 and 4.5), sufficient upslope dispersal would require several LDD events by 2070, which is unlikely for the majority of woody plant species. Furthermore, the ability of plant species to track future changes in temperature increased in simulations with a low degree of trait matching between plants and birds, suggesting that plants in generalized seed-dispersal systems might be more resilient to climate change. [Main conclusion] Our study illustrates how the functional traits of plants and animals can inform predictive models of species dispersal and range shifts under climate change and suggests that the biodiversity of tropical mountain ecosystems is highly vulnerable to future warming. The increasing availability of functional trait data for plants and animals globally will allow parameterization of similar models for many other seed-dispersal systems.Fieldwork at Manú was conducted under the permits 041-2010-AG-DGFFSDGEFFS, 008-2011-AG-DGFFS-DGEFFS, 01-C/C-2010SERNANP-JPNM and 01-2011-SERNANP-PNM-JEF and supported by a scholarship from the German Academic Exchange Service to D.M.D. D.M.D. acknowledges funding from the European Research Council (ERC) under the European Union's Horizon 2020 research and innovation programme (grant number 787638) and the Swiss National Science Foundation (grant number 173342), both awarded to C. H. Graham. W.D.K. acknowledges a Global Ecology grant from the University of Amsterdam Faculty Research Cluster. I.D. was funded by the Alexander von Humboldt Foundation and is now supported by the Balearic Government. S.A.F. was funded by the German Research Foundation (DFG; FR 3246/2-2) and the Leibniz Competition of the Leibniz Association (P52/2017)

Global and regional ecological boundaries explain abrupt spatial discontinuities in avian frugivory interactions

Species interactions can propagate disturbances across space via direct and indirect effects, potentially connecting species at a global scale. However, ecological and biogeographic boundaries may mitigate this spread by demarcating the limits of ecological networks. We tested whether large-scale ecological boundaries (ecoregions and biomes) and human disturbance gradients increase dissimilarity among plant-frugivore networks, while accounting for background spatial and elevational gradients and differences in network sampling. We assessed network dissimilarity patterns over a broad spatial scale, using 196 quantitative avian frugivory networks (encompassing 1496 plant and 1004 bird species) distributed across 67 ecoregions, 11 biomes, and 6 continents. We show that dissimilarities in species and interaction composition, but not network structure, are greater across ecoregion and biome boundaries and along different levels of human disturbance. Our findings indicate that biogeographic boundaries delineate the world’s biodiversity of interactions and likely contribute to mitigating the propagation of disturbances at large spatial scales.The authors acknowledge the following funding: University of Canterbury Doctoral Scholarship (L.P.M.); The Marsden Fund grant UOC1705 (J.M.T., L.P.M.); The São Paulo Research Foundation - FAPESP 2014/01986-0 (M.G., C.E.), 2015/15172-7 and 2016/18355-8 (C.E.), 2004/00810-3 and 2008/10154-7 (C.I.D., M.G., M.A.P.); Earthwatch Institute and Conservation International for financial support (C.I.D., M.G., M.A.P.); Carlos Chagas Filho Foundation for Supporting Research in the Rio de Janeiro State – FAPERJ grant E-26/200.610/2022 (C.E.); Brazilian Research Council grants 540481/01-7 and 304742/2019-8 (M.A.P.) and 300970/2015-3 (M.G.); Rufford Small Grants for Nature Conservation No. 22426–1 (J.C.M., I.M.), No. 9163-1 (G.B.J.) and No. 11042-1 (MCM); Universidade Estadual de Santa Cruz (Propp-UESC; No. 00220.1100.1644/10-2018) (J.C.M., I.M.); Fundação de Amparo à Pesquisa do Estado da Bahia - FAPESB (No. 0525/2016) (J.C.M., I.M.); European Research Council under the European Union’s Horizon 2020 research and innovation program (grant 787638) and The Swiss National Science Foundation (grant 173342), both awarded to C. Graham (D.M.D.); ARC SRIEAS grant SR200100005 Securing Antarctica’s Environmental Future (D.M.D.); German Science Foundation—Deutsche Forschungsgemeinschaft PAK 825/1 and FOR 2730 (K.B.G., E.L.N., M.Q., V.S., M.S.), FOR 1246 (K.B.G., M.S., M.G.R.V.) and HE2041/20-1 (F.S., M.S.); Portuguese Foundation for Science and Technology - FCT/MCTES contract CEECIND/00135/2017 and grant UID/BIA/04004/2020 (S.T.) and contract CEECIND/02064/2017 (L.P.S.); National Scientific and Technical Research Council, PIP 592 (P.G.B.); Instituto Venezolano de Investigaciones Científicas - Project 898 (V.S.D.)

TRY plant trait database – enhanced coverage and open access

Plant traits - the morphological, anatomical, physiological, biochemical and phenological characteristics of plants - determine how plants respond to environmental factors, affect other trophic levels, and influence ecosystem properties and their benefits and detriments to people. Plant trait data thus represent the basis for a vast area of research spanning from evolutionary biology, community and functional ecology, to biodiversity conservation, ecosystem and landscape management, restoration, biogeography and earth system modelling. Since its foundation in 2007, the TRY database of plant traits has grown continuously. It now provides unprecedented data coverage under an open access data policy and is the main plant trait database used by the research community worldwide. Increasingly, the TRY database also supports new frontiers of trait‐based plant research, including the identification of data gaps and the subsequent mobilization or measurement of new data. To support this development, in this article we evaluate the extent of the trait data compiled in TRY and analyse emerging patterns of data coverage and representativeness. Best species coverage is achieved for categorical traits - almost complete coverage for ‘plant growth form’. However, most traits relevant for ecology and vegetation modelling are characterized by continuous intraspecific variation and trait–environmental relationships. These traits have to be measured on individual plants in their respective environment. Despite unprecedented data coverage, we observe a humbling lack of completeness and representativeness of these continuous traits in many aspects. We, therefore, conclude that reducing data gaps and biases in the TRY database remains a key challenge and requires a coordinated approach to data mobilization and trait measurements. This can only be achieved in collaboration with other initiatives

AVONET: morphological, ecological and geographical data for all birds

Functional traits offer a rich quantitative framework for developing and testing theories in evolutionary biology, ecology and ecosystem science. However, the potential of functional traits to drive theoretical advances and refine models of global change can only be fully realised when species‐level information is complete. Here we present the AVONET dataset containing comprehensive functional trait data for all birds, including six ecological variables, 11 continuous morphological traits, and information on range size and location. Raw morphological measurements are presented from 90,020 individuals of 11,009 extant bird species sampled from 181 countries. These data are also summarised as species averages in three taxonomic formats, allowing integration with a global phylogeny, geographical range maps, IUCN Red List data and the eBird citizen science database. The AVONET dataset provides the most detailed picture of continuous trait variation for any major radiation of organisms, offering a global template for testing hypotheses and exploring the evolutionary origins, structure and functioning of biodiversity