Common spatial patterns of trees in various tropical forests: Small trees are associated with increased diversity at small spatial scales

Tropical forests are notable for their high species diversity, even on small spatial scales, and right‐skewed species and size abundance distributions. The role of individual species as drivers of the spatial organization of diversity in these forests has been explained by several hypotheses and processes, for example, stochastic dilution, negative density dependence, or gap dynamics. These processes leave a signature in spatial distribution of small trees, particularly in the vicinity of large trees, likely having stronger effects on their neighbors. We are exploring species diversity patterns within the framework of various diversity‐generating hypotheses using individual species–area relationships. We used the data from three tropical forest plots (Wanang—Papua New Guinea, Barro Colorado Island—Panama, and Sinharaja—Sri Lanka) and included also the saplings (DBH ≥ 1 cm). Resulting cross‐size patterns of species richness and evenness reflect the dynamics of saplings affected by the distribution of large trees. When all individuals with DBH ≥1 cm are included, ~50% of all tree species from the 25‐ or 50‐ha plot can be found within 35 m radius of an individual tree. For all trees, 72%–78% of species were identified as species richness accumulators, having more species present in their surroundings than expected by null models. This pattern was driven by small trees as the analysis of DBH >10 cm trees showed much lower proportion of accumulators, 14%–65% of species identified as richness repellers and had low richness of surrounding small trees. Only 11%–26% of species had lower species evenness than was expected by null models. High proportions of species richness accumulators were probably due to gap dynamics and support Janzen–Connell hypothesis driven by competition or top‐down control by pathogens and herbivores. Observed species diversity patterns show the importance of including small tree size classes in analyses of the spatial organization of diversity.     

Functional diversity and coexistence of herbaceous plants in wet,species‐rich savannas

Trait differences among plant species can favor species coexistence. The role that such differences play in the assembly of diverse plant communities maintained by frequent fires remains unresolved. This lack of resolution results in part from the possibility that species with similar traits may coexist because none has a significant fitness advantage and in part from the difficulty of experimental manipulation of highly diverse assemblages dominated by perennial species. We examined a 65‐year chronosequence of losses of herbaceous species following fire suppression (and subsequent encroachment by Pinus elliottii) in three wet longleaf pine savannas. We used cluster analysis, similarity profile permutation tests, and k‐R cluster analysis to identify statistically significant functional groups. We then used randomization tests to determine if the absence of functional groups near pines was greater (or less) than expected by chance. We also tested whether tolerant and sensitive species were less (or more) likely to co‐occur by chance in areas in savannas away from pines in accordance with predictions of modern coexistence theory. Functional group richness near pines was lower than expected from random species extirpations. Wetland perennials with thick rhizomes and high leaf water content, spring‐flowering wetland forbs (including Drosera tracyi), orchids, Polygala spp., and club mosses were more likely to be absent near pines than expected by chance. C3 grasses and sedges with seed banks and tall, fall‐flowering C4 grasses were less likely to be absent near pines than expected by chance. Species sensitive to pine encroachment were more likely to co‐occur with other such species away from pines at two of the three sites. Results suggest that herb species diversity in frequently burned wet savannas is maintained in part by a weak fitness (e.g., competitive) hierarchy among herbs, and not as a result of trait differences among co‐occurring species.     

Response of plant species and life form diversity to variable fire histories and biomass in the jarrah forest of south‐west Australia

Frequent fires reduce the abundance of woody plant species and favour herbaceous species. Plant species richness also tends to increase with decreasing vegetation biomass and cover due to reduced competition for light. We assessed the influence of variable fire histories and site biomass on the following diversity measures: woody and herbaceous species richness, overall species richness and evenness, and life form evenness (i.e. the relative abundance or dominance among six herbaceous and six woody plant life forms), across 16 mixed jarrah (Eucalyptus marginata) and marri (Corymbia calophylla) forest stands in south‐west Australia. Fire frequency was defined as the total number of fires over a 30‐year period. Overall species richness and species evenness did not vary with fire frequency or biomass. However, there were more herbaceous species (particularly rushes, geophytes and herbs) where there were fewer shrubs and low biomass, suggesting that more herbaceous species coexist where dominance by shrubs is low. Frequently burnt plots also had lower number and abundance of shrub species. Life form evenness was also higher at both high fire frequency and low biomass sites. These results suggest that the impact of fire frequency and biomass on vegetation composition is mediated by local interactions among different life forms rather than among individual species. Our results demonstrate that measuring the variation in the relative diversity of different woody and herbaceous life forms is crucial to understanding the compositional response of forests and other structurally complex vegetation communities to changes in disturbance regime such as increased fire frequency.     

Abiotic and biotic responses to woody debris additions in restored old fields in a multi‐site Before‐After‐Control‐Impact experiment

Ecological restoration of former agricultural land can improve soil conditions, recover native vegetation, and provide fauna habitat. However, restoration benefits are often associated with time lags, as many attributes, such as leaf litter and coarse woody debris, need time to accumulate. Here, we experimentally tested whether adding mulch and logs to restoration sites in semi‐arid Western Australia can accelerate restoration benefits. All sites had been cropped and then planted with native trees and shrubs (i.e., Eucalyptus, Melaleuca, and Acacia spp.) 10 years prior to our experiment, to re‐establish the original temperate eucalypt woodland vegetation community. We used a Multi‐site Before‐After‐Control‐Impact (MBACI) design to test the effects on 30 abiotic and biotic response variables over a period of 2 years. Of the 30 response variables, a significant effect was found for just four variables: volumetric water content, decomposition, native herbaceous species cover and species richness of disturbance specialist ants. Mulch addition had a positive effect on soil moisture when compared to controls but suppressed growth of native (but not exotic) herbaceous plants. On plots with log additions, decomposition rates decreased, and species richness of disturbance specialist ants increased. However, we found no effect on total species richness and abundance of other ant species groups. The benefit of mulch to soil moisture was offset by its disbenefit to native herbs in our study. Given time, logs may also provide habitat for ant species that prefer concealed habitats. Indeed, benefits to other soil biophysical properties, vegetation, and ant fauna may require longer time frames to be detected. Further research is needed to determine whether the type, quantity, and context of mulch and log additions may improve their utility for old field restoration and whether effects on native herbs are correlated with idiosyncratic climatic conditions.     

Beneath the veil: plant growth form influences the strength of species richness–productivity relationships in forests

Aim Species richness has been observed to increase with productivity at large spatial scales, though the strength of this relationship varies among functional groups. In forests, canopy trees shade understorey plants, and for this reason we hypothesize that species richness of canopy trees will depend on macroclimate, while species richness of shorter growth forms will additionally be affected by shading from the canopy. In this study we test for differences in species richness–productivity relationships (SRPRs) among growth forms (canopy trees, shrubs, herbaceous species) in small forest plots. Location We analysed 231 plots ranging from 34.0° to 48.3° N latitude and from 75.0° to 124.2° W longitude in the United States. Methods We analysed data collected by the USDA Forest Inventory and Analysis program for plant species richness partitioned into different growth forms, in small plots. We used actual evapotranspiration as a macroclimatic estimate of regional productivity and calculated the area of light‐blocking tissue in the immediate area surrounding plots for an estimate of the intensity of local shading. We estimated and compared SRPRs for different partitions of the species richness dataset using generalized linear models and we incorporated the possible indirect effects of shading using a structural equation model. Results Canopy tree species richness increased strongly with regional productivity, while local shading primarily explained the variation in herbaceous plant richness. Shrub species richness was related to both regional productivity and local shading. Main conclusions The relationship between total forest plant species richness and productivity at large scales belies strong effects of local interactions. Counter to the pattern for overall richness, we found that understorey herbaceous plant species richness does not respond to regional productivity gradients, and instead is strongly influenced by canopy density, while shrub species richness is under multivariate control.     

Disturbance of the herbaceous layer after invasion of an eutrophic temperate forest by wild boar

Disturbances of the soil and the tree canopy are crucial factors determining the diversity, composition and biomass of the herbaceous layer in forests. This study presents a detailed account of ground vegetation in permanent plots surveyed before and after invasion of wild boar (Sus scrofa) to a temperate deciduous broadleaf forest. Specifically, we aimed to quantify the effect of wild boar rooting on cover, richness and composition of spring ephemerals, summer green herbs and saplings of woody species in relation to tree canopy cover. Rooting frequency in sample plots increased from 0% in 2010 to 61% in 2013. In heavily rooted plots, the mean cover of spring ephemeral geophytes (mainly Anemone nemorosa, A. ranunculoides and Ranunculus ficaria) decreased from 75% to 39% between 2010 and 2013. Species richness of summer green herbs generally increased between 2010 and 2013 and was additionally positively affected by heavy rooting and low canopy cover. Rooting also caused heterogenization of the herbaceous layer and amplified ongoing compositional changes induced by changing light conditions. Frequency and richness of spring ephemeral and woody species remained unchanged. We conclude that overall species richness of the herbaceous layer may increase in the short‐term as a result of increased plant recruitment and seed dispersal. However, wild boar rooting can greatly reduce the ground cover of spring ephemerals in eutrophic broadleaf forests, thereby threatening their important ecological function. To avoid long‐term losses of characteristic spring flora elements, local population control of wild boar is necessary to reduce abundance and frequency of soil rooting.     

Are native and non‐native pollinator friendly plants equally valuable for native wild bee communities?

Bees rely on floral pollen and nectar for food. Therefore, pollinator friendly plantings are often used to enrich habitats in bee conservation efforts. As part of these plantings, non‐native plants may provide valuable floral resources, but their effects on native bee communities have not been assessed in direct comparison with native pollinator friendly plantings. In this study, we performed a common garden experiment by seeding mixes of 20 native and 20 non‐native pollinator friendly plant species at separate neighboring plots at three sites in Maryland, USA, and recorded flower visitors for 2 years. A total of 3,744 bees (120 species) were collected. Bee abundance and species richness were either similar across plant types (midseason and for abundance also late season) or lower at native than at non‐native plots (early season and for richness also late season). The overall bee community composition differed significantly between native and non‐native plots, with 11 and 23 bee species being found exclusively at one plot type or the other, respectively. Additionally, some species were more abundant at native plant plots, while others were more abundant at non‐natives. Native plants hosted more specialized plant–bee visitation networks than non‐native plants. Three species out of the five most abundant bee species were more specialized when foraging on native plants than on non‐native plants. Overall, visitation networks were more specialized in the early season than in late seasons. Our findings suggest that non‐native plants can benefit native pollinators, but may alter foraging patterns, bee community assemblage, and bee–plant network structures.     

Response of plant and gastropod species to knotweed invasion

Invasive species of the knotweed complex (Fallopia sp.) have repeatedly been shown to decrease diversity of native local biota. While effects on plant species richness are well established, effects on invertebrate and in particular gastropod species richness are less well understood. We recorded cover of plant species and diversity and abundance of gastropod species in four plots (1 m × 1 m) with Fallopia japonica and compared these to paired control plots without F. japonica at 15 sites along the river Birs (Switzerland) in early summer (June) and autumn (September). Knotweed and control plots did not differ in site characteristics and soil parameters.Average plant species richness in F. japonica plots was 50% lower compared to control plots. This reduction was significant for woody species as well as for herbaceous species. However, species richness of early flowering annuals did not differ significantly. Among the species most affected by knotweed were hop (Humulus lupulus) and European spindle tree (Euonymus europaeus) but also stand-forming species such as nettle (Urtica dioica) or ground elder (Aegopodium podagraria).Average snail species richness was significantly reduced in F. japonica plots. The reduction was pronounced in large (≥5 mm shell size) and long-lived (>2 years) snail species but not in slugs or small and short-lived snails. For example, large snails such as the Roman snail (Helix pomatia, ?85%) or the red-listed species Bradybaena fruticum (?93%), showed reduced abundances in F. japonica compared to control plots. In contrast, the red-listed but small Vertigo pusilla (+92%) had higher abundances in F. japoinca plots. Principal component analyses revealed little overlap in plant communities or community composition of large snail species between F. japonica and control plots. Taken together, knotweed invasion decreased the cover of most plant species and abundance of large and long-lived gastropods.     

Both landscape and local factors influence plant and hexapod communities of industrial water‐abstraction sites

1. At the landscape level, intensification of agriculture, fragmentation, and destruction of natural habitats are major causes of biodiversity loss that can be mitigated at small spatial scales. However, the complex relationships between human activities, landscapes, and biodiversity are poorly known. Yet, this knowledge could help private stakeholders managing seminatural areas to play a positive role in biodiversity conservation.
2. We investigated how water‐abstraction sites could sustain species diversity in vascular‐plant communities and two taxonomic groups of insect communities in a fragmented agricultural landscape.
3. Landscape‐scale variables (connectivity indices and surrounding levels of herbicide use), as well as site‐specific variables (soil type for vascular plants, floral availability for Rhopalocera, and low herbaceous cover for Orthoptera), were correlated to structural and functional metrics of species community diversity for these taxonomic groups, measured on 35 industrial sites in the Ile‐de‐France region in 2018–2019.
4. Rhopalocera and Orthoptera consisted essentially of species with a high degree of dispersal and low specialization, able to reach the habitat patches of the fragmented landscape of the study area. Sandy soil harbored more diverse vascular‐plant communities. Plant diversity was correlated to a greater abundance of Rhopalocera and a lower richness of Orthoptera.
5. Increasing landscape connectivity was related to higher abundance of plants and Rhopalocera, and a higher evenness index for Orthoptera communities. Higher levels of herbicide use were related to a decrease in the biodiversity of plants and Rhopalocera abundance. High levels of herbicide favored high‐dispersal generalist plants, while high levels of connectivity favored low‐dispersal plants. Specialist Orthoptera species were associated with low herbaceous cover and connectivity.
6. Water‐abstraction sites are valuable seminatural habitats for biodiversity. Changing intensive agricultural practices in surrounding areas would better contribute to conserving and restoring biodiversity on these sites.

     

The influence of vines on an oligohaline marsh community: results of a removal and fertilization study

The effects of competitive suppression by vines on the non-vine plant community have received little attention in temperate habitats. This study investigated the impact vines have on their herbaceous hosts in a wetland community at two soil fertility levels. Plots in an oligohaline marsh were treated in a 2 × 2 factorial design with vine removal and fertilization over two growing seasons. There was no significant interaction between removal and fertilization treatments on any of the measured variables. Vine removal initially caused an increase in light penetration through the canopy, but by the end of the study, plots with vines removed had less light due to a 25% increase in biomass by the plants released from competition with vines. For plots with vines removed, species richness was higher during a brief period in the spring of the second year, but by the end of the study, richness in removal plots decreased relative to controls. Fertilization caused a 40% increase in biomass overall, although only two species, Sagittaria lancifolia L. and Polygonum punctatum Ell., showed dramatic increases. Despite fertilization causing a 40% decrease in light penetration to the ground, no change in species richness was observed. Overall, these results show that vine cover in this wetland suppresses non-vine species and reduces community biomass. Removal of vines increased biomass of non-vine dominants but resulted in only an ephemeral change in species richness. Fertilization did not increase the effects of vines on the non-vine community. Received: 14 November 1996 / Accepted: 10 June 1997     

A dominant dwarf shrub increases diversity of herbaceous plant communities in a Trans-Himalayan rangeland

Plant communities are structured by both competition and facilitation. The interplay between the two interactions can vary depending on environmental factors, nature of stress, and plant traits. However, whether positive or negative interactions dominate in regions of high biotic and abiotic stress remains unclear. We studied herbaceous plant communities associated with a dwarf shrub Caragana versicolor in semi-arid, high altitude Trans-Himalayan rangelands of Spiti, India. We surveyed 120 pairs of plots (within and outside shrub canopies) across four watersheds differing in altitude, aspect, and dominant herbivores. Herbaceous communities within shrub canopies had 25% higher species richness, but similar abundance when compared to communities outside the canopy, with the shrub edge having higher diversity than the centre of the canopy. Grasses and erect forbs showed positive associations with the shrub, while prostrate plants occurred at much lower abundance within the canopy. Rare species showed stronger positive associations with Caragana than abundant species. Experimental removal of herbaceous vegetation from within shrub canopies led to 42% increase in flowering in Caragana, indicating a cost to the host shrubs. Our study indicates a robust pattern of a dwarf shrub facilitating local community diversity across this alpine landscape, increasing diversity at the plot level, facilitating rare species, and yet incurring a cost to hosts from the presence of herbaceous plants. Given these large influences of this shrub on the vegetation of these high altitude rangelands, we suggest that the shrub microhabitat be explicitly considered in any analyses of ecosystem health in such rangelands.     

Survival,growth, and photosynthesis of tree seedlings competing with herbaceous vegetation along a water-light-nitrogen gradient

In herbaceous dominated patches and ecosystems, tree establishment is influenced partly by the ability of woody seedlings to survive and grow in direct competition with herbaceous vegetation. We studied the importance of season long wet and dry spells on the competitive interactions between herbaceous vegetation and oak seedlings along a light and nitrogen gradient in an infertile secondary successional grassland in central North America. We conducted a field experiment in which seedlings of bur oak (Quercus macrocarpa) and northern pin oak (Q. ellipsoidalis) were exposed to two levels of light (full sun and 80% shade), three levels of nitrogen input (0, 5, 15 g m^–1 yr^–1), and three levels of water input (low, medium and high). In addition, seedlings were grown with and without the presence of surrounding herbaceous vegetation under both light and all three water levels. Seedling survival, growth, and rate of photosynthesis were significantly affected by competition with herbaceous vegetation and these effects varied along the multiple resource gradient. Overall, seedling survival of both species was significantly greater in wetter and shaded plots and when surrounding herbaceous vegetation was removed and was lower in nitrogen enriched plots. We found that soil water was significantly affected by varying inputs of water, light, and the presence or absence of herbaceous vegetation, and that seedling survival and rate of photosynthesis were highly correlated with available soil water. Our findings show that the impact of season long wet and dry spells on tree seedling success in grasslands can be affected by light and soil nitrogen availability.     

Spatial patterns of light‐demanding tree species in the Yangambi rainforest (Democratic Republic of Congo)

Most Central African rainforests are characterized by a remarkable abundance of light‐demanding canopy species: long‐lived pioneers (LLP) and non‐pioneer light demanders (NPLD). A popular explanation is that these forests are still recovering from intense slash‐and‐burn farming activities, which abruptly ended in the 19th century. This “human disturbance” hypothesis has never been tested against spatial distribution patterns of these light demanders. Here, we focus on the 28 most abundant LLP and NPLD from 250 one‐ha plots distributed along eight parallel transects (~50 km) in the Yangambi forest. Four species of short‐lived pioneers (SLP) and a single abundant shade‐tolerant species (Gilbertiodendron dewevrei) were used as reference because they are known to be strongly aggregated in recently disturbed patches (SLP) or along watercourses (G. dewevrei). Results show that SLP species are strongly aggregated with clear spatial autocorrelation of their diameter. This confirms that they colonized the patch following a one‐time disturbance event. In contrast, LLP and NPLD species have random or weakly aggregated distribution, mostly without spatial autocorrelation of their diameter. This does not unambiguously confirm the “human disturbance” hypothesis. Alternatively, their abundance might be explained by their deciduousness, which gave them a competitive advantage during long‐term drying of the late Holocene. Additionally, a canonical correspondence analysis showed that the observed LLP and NPLD distributions are not explained by environmental variables, strongly contrasting with the results for the reference species G. dewevrei, which is clearly aggregated along watercourses. We conclude that the abundance of LLP and NPLD species in Yangambi cannot be unambiguously attributed to past human disturbances or environmental variables. An alternative explanation is that present‐day forest composition is a result of adaptation to late‐Holocene drying. However, results are inconclusive and additional data are needed to confirm this alternative hypothesis.     

Ungulates increase forest plant species richness to the benefit of non‐forest specialists

Large wild ungulates are a major biotic factor shaping plant communities. They influence species abundance and occurrence directly by herbivory and plant dispersal, or indirectly by modifying plant‐plant interactions and through soil disturbance. In forest ecosystems, researchers’ attention has been mainly focused on deer overabundance. Far less is known about the effects on understory plant dynamics and diversity of wild ungulates where their abundance is maintained at lower levels to mitigate impacts on tree regeneration. We used vegetation data collected over 10 years on 82 pairs of exclosure (excluding ungulates) and control plots located in a nation‐wide forest monitoring network (Renecofor). We report the effects of ungulate exclusion on (i) plant species richness and ecological characteristics, (ii) and cover percentage of herbaceous and shrub layers. We also analyzed the response of these variables along gradients of ungulate abundance, based on hunting statistics, for wild boar (Sus scrofa), red deer (Cervus elaphus) and roe deer (Capreolus capreolus). Outside the exclosures, forest ungulates maintained higher species richness in the herbaceous layer (+15%), while the shrub layer was 17% less rich, and the plant communities became more light‐demanding. Inside the exclosures, shrub cover increased, often to the benefit of bramble (Rubus fruticosus agg.). Ungulates tend to favour ruderal, hemerobic, epizoochorous and non‐forest species. Among plots, the magnitude of vegetation changes was proportional to deer abundance. We conclude that ungulates, through the control of the shrub layer, indirectly increase herbaceous plant species richness by increasing light reaching the ground. However, this increase is detrimental to the peculiarity of forest plant communities and contributes to a landscape‐level biotic homogenization. Even at population density levels considered to be harmless for overall plant species richness, ungulates remain a conservation issue for plant community composition.     

Impact of habitat disturbance in the wetland forests of East Usambara,Tanzania

We evaluated habitat characteristics of East Usambara wetland forests. The abundance and species composition in the tree, shrub and herbaceous layers were enumerated in two sets of nested plots, one set in a natural wetland forest and the second in a wetland forest that had been disturbed by small‐scale gold mining activities. Each set had thirty‐six 1 m × 1 m plots for herbs, inside nine 5 m × 5 m plots for shrubs, in three 20 m × 20 m plots for trees. The habitat profile of herbaceous – shrub – tree layers was found to be sharply different from one obtained in previous studies at the surrounding nonwetland forests as were species composition and abundance. Unlike in the nonwetland forests, the herbaceous layer was thick, the shrub layer very thin and the woody species density and richness much lower. Disturbance significantly reduced woody cover and changed species composition in the herbaceous layer. Recovery of the woody vegetation was low. Wetland forests in the East Usambaras form a small fraction of the total area, but they are a unique biodiversity repository, they appear to be an important carbon dioxide sink and to reserve and purify water. They are sensitive to disturbance and need protection.     

Restored and remnant Banksia woodlands elicit different foraging behavior in avian pollinators

Pollinators and the pollination services they provide are critical for seed set and self‐sustainability of most flowering plants. Despite this, pollinators are rarely assessed in restored plant communities, where their services are largely assumed to re‐establish. Bird–pollinator richness, foraging, and interaction behavior were compared between natural and restored Banksia woodland sites in Western Australia to assess their re‐establishment in restored sites. These parameters were measured for natural communities of varying size and degree of fragmentation, and restored plant communities of high and low complexity for three years, in the summer and winter flowering of Banksia attenuata and B. menziesii, respectively. Bird visitor communities varied in composition, richness, foraging movement distances, and aggression among sites. Bird richness and abundance were lowest in fragmented remnants. Differences in the composition were associated with the size and degree of fragmentation in natural sites, but this did not differ between seasons. Restored sites and their adjacent natural sites had similar species composition, suggesting proximity supports pollinator re‐establishment. Pollinator foraging movements were influenced by the territorial behavior of different species. Using a network analysis approach, we found foraging behavior varied, with more frequent aggressive chases observed in restored sites, resulting in more movements out of the survey areas, than observed in natural sites. Aggressors were larger‐bodied Western Wattlebirds (Anthochaera chrysoptera) and New Holland Honeyeaters (Phylidonyris novaehollandiae) that dominated nectar resources, particularly in winter. Restored sites had re‐established pollination services, albeit with clear differences, as the degree of variability in the composition and behavior of bird pollinators for Banksias in the natural sites created a broad completion target against which restored sites were assessed. The abundance, diversity, and behavior of pollinator services to remnant and restored Banksia woodland sites were impacted by the size and degree of fragmentation, which in turn influenced bird–pollinator composition, and were further influenced by seasonal changes between summer and winter. Consideration of the spatial and temporal landscape context of restored sites, along with plant community diversity, is needed to ensure the maintenance of the effective movement of pollinators between natural remnant woodlands and restored sites.     

Reduced herbivory during simulated ENSO rainy events increases native herbaceous plants in semiarid Chile

El Niño Southern Oscillation (ENSO) events have profound consequences for the dynamics of terrestrial ecosystems. Since increased climate variability is expected to favour the invasive success of exotic species, we conducted a field experiment to study the effects that simulated rainy ENSO events in combination with herbivores and shade have on the composition of a semiarid herbaceous community in north-central Chile. We hypothesized that water pulses, such as those associated with rainy ENSO events could trigger significant changes in the relative abundance of exotic and native herbaceous species. Specifically, we predicted an increase in native grasses and a reduction in the abundance of exotic species, especially prostrate forbs, if water pulses were combined with reduced herbivory. We found that herbivory by small mammals, especially introduced European rabbits (Oryctolagus cuniculus) and hares (Lepus europaeus), have an overwhelming effect on species abundance and composition in this semiarid herbaceous community. Herbivore exclusion produced an overall increase in herb density and biomass mostly due to the extraordinary growth of tall native grasses (especially Bromus berterianus) that outcompeted small prostrate forbs (both native and exotic ones), and small exotic grasses (Koeleria pleoides, Schismus arabicus). Our results suggest that it might be possible to enhance the recovery of native grasses by applying efficient herbivore control during rainy years such as those associated with ENSO events although a negative consequence would be the loss of small native forbs, which greatly contribute to the richness of herbaceous communities in semiarid ecosystems.     

Conversion of rainforest into oil palm and rubber plantations affects the functional composition of litter and soil Collembola

Rainforest conversion and expansion of plantations in tropical regions are associated with changes in animal communities and biodiversity decline. In soil, Collembola are one of the most numerous invertebrate groups that affect the functioning of microbial communities and support arthropod predators. Despite that, information on the impact of changes in land use in the tropics on species and trait composition of Collembola communities is very limited. We investigated the response of Collembola to the conversion of rainforest into rubber agroforestry (“jungle rubber”), rubber, and oil palm plantations in Jambi Province (Sumatra, Indonesia), a region which experienced one of the strongest recent deforestation globally. Collembola were sampled in 2013 and 2016 from the litter and soil layer using heat extraction, and environmental factors were measured (litter C/N ratio, pH, water content, composition of microbial community and predator abundance). In the litter layer, density and species richness in plantation systems were 25%–38% and 30%–40% lower, respectively, than in rainforest. However, in the soil layer, density, species richness, and trait diversity of Collembola were only slightly affected by land‐use change, contrasting the response of many other animal groups. Species and trait composition of Collembola communities in litter and soil differed between each of the land‐use systems. Water content and pH were identified as main factors related to the differences in species and trait composition in both litter and soil, followed by the density of micro‐ and macropredators. Dominant species of Collembola in rainforest and jungle rubber were characterized by small body size, absence of furca, and absence of intense pigmentation, while in plantations, larger species with long furca and diffuse or patterned pigmentation were more abundant. Overall, land‐use change negatively affected Collembola communities in the litter layer, but its impact was lower in the soil layer. Several pantropical genera of Collembola (i.e., Isotomiella, Pseudosinella, and Folsomides) dominated across land‐use systems, reflecting their high environmental adaptability and/or efficient dispersal, calling for studies on their ecology and genetic diversity. The decline in species richness and density of litter‐dwelling Collembola with the conversion of rainforest into plantation systems calls for management practices mitigating negative effects of the deterioration of the litter layer in rubber plantations, but even more in oil palm plantations.     

Tree species richness and density affect parasitoid diversity in cacao agroforestry

In Brazil, cacao is mostly planted beneath shade trees. The diversity of shade trees varies from monospecific to highly diverse canopies, characteristic of pristine Atlantic Forest. This study evaluates the relationships between family richness of Hymenoptera-Parasitica and Chrysidoidea, and tree species richness and density, the species richness of herbaceous understorey, and the area and age of the cacao agroforestry system. We sampled 16 cacao agroforestry systems, with canopy diversity ranging from one to 22 tree species per hectare, in three seasons: summer (March), winter (August) and spring (November). Parasitoids were sampled using eight Malaise-Townes traps per site. Tree species richness and density were enumerated within 1 ha at each site, and herbaceous plant species richness was calculated in eight 1 m² plots, within the hectare. The number of parasitoid families increased with tree species richness and density in spring and summer, but decreased in winter. Neither species richness of herbaceous plants nor area and age of the system affected parasitoid family richness. We suggest that the increase of parasitoid diversity with tree species richness and density in warmer seasons reflects increasing heterogeneity and availability of resources. The decrease in parasitoid family number with tree density in winter may be due to local impoverishment of resources, leading to parasitoid emigration to neighbouring forest remnants. This result implies that a higher diversity of shade trees will help to maintain high parasitoid levels and, in consequence, higher levels of natural enemies of cacao pests, particularly in the warmer seasons. This prediction is borne out in the experience of cacao producers. The proper management of shade tree diversity will play a vital role in maintaining the sustainability of cacao agroforestry production systems in the tropics and, concurrently, will maintain high biodiversity values in these locations.     

Disentangling the links between habitat complexity and biodiversity in a kelp‐dominated subantarctic community

Habitat complexity is one of the most important factors modulating species diversity. This feature comprises several interrelated attributes, such as number, size, and spatial arrangement of complexity‐forming elements. However, the separate and joint effects of these attributes on diversity and community structure are still not well understood. Here, we assess the relationships between several structural‐complexity attributes of the subantarctic kelp Lessonia flavicans and species richness, total abundance, and structure of kelp‐associated macrobenthic communities. We predicted that longer thalli and larger holdfasts favor greater species richness and total abundance of invertebrate organisms. To test the prediction, an observational sampling program was established in two sites of the Strait of Magellan. Uni‐ and multivariate analyses revealed both positive and negative effects of kelp structural‐complexity attributes on diversity. Holdfast diameter and maximum frond length, followed by thallus wet weight, had the strongest positive fits to species richness and total abundance; the number of stipes, on the other hand, was negatively associated with both response variables. Longer fronds were associated with greater abundances of spirorbid polychaetes. Larger holdfasts supported larger abundances of Nereididae and Terebelidae polychaetes and the limpet Nacella mytilina. Contrarily, kelps with longer fronds and more stipes supported fewer amphipods. In this way, we demonstrate that different dimensions of habitat complexity can have contrasting effects on diversity and community structure, highlighting the fundamental role of multiple dimensions of kelp habitat complexity for local biodiversity.