What drives the species richness patterns of non‐volant small mammals along a subtropical elevational gradient?

The biodiversity of non‐volant small mammals along an extensive subtropical elevational gradient was studied for the first time on Gongga Mountain, the highest mountain in Hengduan Mountain ranges in China, located in one of the 25 global biodiversity hotspots. Non‐volant small mammals were replicate sampled in two seasons at eight sampling sites between 1000 and 4200 m elevation on the eastern slope of Gongga Mountain. In all, 726 individual small mammals representing 25 species were documented in 28 800 trap nights. The species richness pattern for non‐volant small mammals along the elevational gradients was hump‐shaped with highest richness at mid‐elevations. However, different richness patterns emerged between endemic and non‐endemic species, between larger‐ranged and smaller‐ranged species and between rodents and insectivores. Temperature, precipitation, plant species richness and geometric constraints (mid‐ domain effect) were most significant in explaining species richness patterns. Based on the analysis of simple ordinary least squares (OLS) and stepwise multiple regressions, the overall richness pattern, as well as the pattern of insectivores, endemic species and larger‐ranged species showed strong correlation with geometric constraint predictions. However, non‐endemic species richness was more strongly correlated with temperature, while rodent richness was correlated with plant species richness. Our study shows that no single key factor can explain all richness patterns of non‐volant small mammals. We need to be cautious in summarizing a general richness pattern of large species groups (e.g. small mammals or mammals) from species in smaller groups having different ecological distributions and life histories. Elevational richness patterns and their driving factors for small mammals are more likely dependent on what kind of species we study.     

What causes latitudinal gradients in species diversity? Evolutionary processes and ecological constraints on swallowtail biodiversity

The latitudinal diversity gradient (LDG) is one of the most striking ecological patterns on our planet. Determining the evolutionary causes of this pattern remains a challenging task. To address this issue, previous LDG studies have usually relied on correlations between environmental variables and species richness, only considering evolutionary processes indirectly. Instead, we use a phylogenetically integrated approach to investigate the ecological and evolutionary processes responsible for the global LDG observed in swallowtail butterflies (Papilionidae). We find evidence for the 'diversification rate hypothesis' with different diversification rates between two similarly aged tropical and temperate clades. We conclude that the LDG is caused by (1) climatically driven changes in both clades based on evidence of responses to cooling and warming events, and (2) distinct biogeographical histories constrained by tropical niche conservatism and niche evolution. This multidisciplinary approach provides new findings that allow better understanding of the factors that shape LDGs.     

What drives invasibility? A multi‐model inference test and spatial modelling of alien plant species richness patterns in northern Portugal

Understanding and predicting biological invasions can focus either on traits that favour species invasiveness or on features of the receiving communities, habitats or landscapes that promote their invasibility. Here, we address invasibility at the regional scale, testing whether some habitats and landscapes are more invasible than others by fitting models that relate alien plant species richness to various environmental predictors. We use a multi‐model information‐theoretic approach to assess invasibility by modelling spatial and ecological patterns of alien invasion in landscape mosaics, and by testing competing hypotheses of environmental factors that may control invasibility. Because invasibility may be mediated by particular characteristics of invasiveness, we classified alien species according to their C‐S‐R plant strategies. We illustrate this approach with a set of 86 alien species in northern Portugal. We first focus on predictors influencing species richness and expressing invasibility, and then evaluate whether distinct plant strategies respond to the same or different groups of environmental predictors. We confirmed climate as a primary determinant of alien invasions, and as a primary environmental gradient determining landscape invasibility. The effects of secondary gradients were detected only when the area was sub‐sampled according to predictions based on the primary gradient. Then, multiple predictor types influenced patterns of alien species richness, with some types (landscape composition, topography and fire regime) prevailing over others. Alien species richness responded most strongly to extreme land management regimes, suggesting that intermediate disturbance induces biotic resistance by favouring native species richness. Land‐use intensification facilitated alien invasion, whereas conservation areas hosted few invaders, highlighting the importance of ecosystem stability in preventing invasions. Plants with different strategies exhibited different responses to environmental gradients, particularly when the variations of the primary gradient were narrowed by sub‐sampling. Such differential responses of plant strategies suggest using distinct control and eradication approaches for different areas and alien plant groups.     

Do local processes scale to global patterns? The role of drought and the species pool in determining treehole insect diversity

Global patterns in community species richness may represent limitations operating at the local scale, such as competitive exclusion and habitat suitability, or, in the case of unsaturated communities, limitations to the species pool of biogeographic regions. Separating the effects of local and regional processes on community richness requires combining small-scale experiments with broad surveys. I examine limitations to the number of aquatic insect species per treehole at the scale of a single woodland, and between eight biogeographic regions. Variation in species richness at the woodland scale can largely be explained by small-scale differences between treeholes in drought disturbance, as shown for British treehole insects for 2 years. At the global scale, drought disturbance is a relatively poor predictor of patterns in the local richness of treehole mosquitoes. Instead, regional differences in the species pool explain most of the variation in local mosquito richness. Treeholes in at least the most species-poor regions appear to be unsaturated with mosquito species. In this system, therefore, local processes do not necessarily scale to global patterns.     

Mountain invasions on the way: are there climatic constraints for the expansion of alien woody species along an elevation gradient in Argentina?

Aims Increasing evidence worldwide indicates that high mountain regions are not immune to invasion by alien plants. Here, we aimed to address whether three major woody invaders of low-mountain areas are constrained to lower altitudes due to climatic restrictions or just by low propagule pressure. We hypothesize that the increasing climatic harshness towards higher altitudes restricts seedling establishment and survival of these woody invaders and thus their potential for altitudinal expansion.Methods The study was carried out in the Sierras Grandes Mountains, Córdoba, central Argentina (32o50′S, 64o90′W). We addressed the hypothesis with an experimental approach, dissociating the effect of the environmental gradient from that of propagule pressure, by discarding the influence of seed sources. We translocated seeds of Gleditsia triacanthos L., Ligustrum lucidum W.T. Aiton. and Pyracantha angustifolia (Franch.) C.K.Schneid. along the altitudinal gradient (from 900 to 2700 m asl). Seven sites were selected along the altitudinal gradient, spaced every 200–400 m. Three plots (4×4 m) were selected within each altitudinal site and excluded from livestock; 100 seeds of each species were sown within each plot (6300 seeds in total). Seedling emergence and survival was monitored during two growing seasons and soil temperature and moisture was recorded as well. The field experiment was complemented with lab assays.Important findings Climate along this altitudinal gradient might be least restrictive at mid elevations, as a result of the intersection between opposite soil temperature and moisture gradients. Sown seeds germinated along the whole altitudinal gradient and seedlings successfully established and over-wintered well above their current altitudinal distribution (up to 2200 m for Ligustrum and 2400 m for Gleditsia and Pyracantha). Additional lab assays confirmed field patterns and gave some insights into contrasting regeneration strategies between these invaders that might help to overcome stochastic environmental constraints in the germination stage. Overall, seedlings of three major woody invaders of low-mountain woodlands can establish and survive at higher elevations than their current distribution. In contrast to natives, they seem to be broad climate tolerators, rather than specialized stress tolerators, capable of germinating and growing across a wide elevational range. While long-term experiments might be necessary to fully address the potentials for altitudinal expansion, out results on early lifecycle stage suggest that the invaders studied here would have mostly a dispersal barrier rather than a climate barrier to establish in the upper belt of the Sierras Grandes.     

Spatial ecology and conservation of two sloth species in a cacao landscape in limón,Costa Rica

The spatial ecology of sloths was studied in an agricultural landscape in Limón Province, Costa Rica. Two sloth species, the three-toed sloth (Bradypus variegatus) and the two-toed sloth (Choloepus hoffmanni), actively used and traveled through a cacao agroforest and its contiguous living fence rows and riparian forests. This agroecosystem was embedded in an agricultural landscape dominated by banana and pineapple plantations and pastures with dispersed trees. The two-toed sloth (C. hoffmanni) was found in 101 tree species and used 34 for food; the three-toed sloth (B. variegatus) was found in 71 tree species and used 15 for food. Choice of preferred species differed between the two sloth species. Trees commonly used by sloths for food and/or refuge in the cacao agroforest included Erythrina poeppigiana, Cecropia obtusifolia, Leucaena leucocephala; in the living fence rows, Cordia alliodora, Erythrina poeppigiana, Ocotea sinuata and Trophis racemosa; in the riparian forests, Coussapoa villosa, Cecropia obtusifolia, Hura crepitans, Pterocarpus officinalis and Spondias mombin; and in the pastures with dispersed trees, Cordia alliodora, Coussapoa villosa, Erythrina poeppigiana, Ocotea sinuata and Hura crepitans. This study demonstrates the importance of the cacao agroforest as well as arboreal elements in other land uses in providing resources for sloth conservation in a larger agricultural landscape.     

Using the Zeta (ζ) diversity approach for understanding the effects of variations in climatic covariates,vegetation patterns,and human influence on species composition of arboreal frogs in India

The biodiversity patterns of an ecosystem provide an opportunity for understanding the drivers of the community structure and, together, interpreting better conservation strategies for the species. Here we map the patterns in compositional turnover of the arboreal frog species (Anura: Rhacophoridae) in India and explore the impacts of climatic covariates, vegetation patterns, and human influence on driving their species composition. Using the previously published data, we compiled the occurrence matrix for the species of nine genera of the family Rhacophoridae from India. The species turnover trends were analyzed using the zeta diversity decline and retention-rate analysis. We further used the multisite generalized dissimilarity modeling (MS-GDM) to test the variations in climatic, vegetational, and anthropogenic drivers that better explain the species turnover. The results suggested a low nestedness among the genera of arboreal frogs barring Polypedates and Raorchestes. The difference in the mean temperature of the warmest quarter in the colder temperature regime is the primary driver of the compositional turnover of arboreal frog species. Also, differences in the precipitation seasonality of the lower and higher precipitation regime emerged as a driver of the compositional turnover for some genera. The differences in diversity patterns and their corresponding drivers are ideally only species-specific or group-specific. We suggest their utility while designing conservation efforts for co-occurring groups in a region.     

Reconsidering null models of diversity: Do geometric constraints on species ranges necessarily cause a mid‐domain effect?

Recent null models that place species ranges randomly within a bounded domain have produced controversial results. Many such geometric constraint models predict a peak in species richness in the centre of domains in the absence of underlying environmental gradients or interspecific interactions. We used two-dimensional simulation models to explore different ways that species ranges could interact with the domain boundary. In the rejection model, a randomly generated range that overlaps a domain boundary is removed from the simulation. In the reshaping model, a range that overlaps the domain boundary is reshaped so that the entire range is placed within the domain. The truncation model allows potential ranges to extend across the boundary, but only that portion of the range within the domain is included in the realized range. Both rejection and reshaping models produced a drop in species richness near domain boundaries, though the effect was less pronounced in the reshaping model. Our truncation model did not produce any spatial pattern in species richness. Thus the random placement of species ranges within a bounded domain does not necessarily lead to a mid-domain effect.
  Range truncation is consistent with bioclimate envelope models, which can successfully predict a species range in response to the availability of appropriate climate conditions. We argue that such flexible range sizes are more realistic than the assumption that range size is an unvarying characteristic of a species. Other range characteristics, including size and shape, can change near domain boundaries in the null models, including the truncation model. A broader consideration of range characteristics near domain boundaries could be productive.     

A survey of homopteran species (Auchenorrhyncha) from coffee shrubs and poró and laurel trees in shaded coffee plantations,in Turrialba,Costa Rica

A survey of homopteran species (Auchenorryncha) was conducted in coffee plantations with no shade (C), and in those with shade of either poró (Erythrina poeppigiana) (CP) or poró plus laurel (Cordia alliodora) (CPL), in Turrialba, Costa Rica. A total of 130 species in ten families were collected, dominated by Cicadellidae (82 species). Species richness was highest in the CP system (88), followed by CPL (74) and C systems (60). Five most common species for all systems were Fusigonalia lativittata, Hebralebra nicaraguensis, Neocoelidia sp., Oliarus sp. and Clastoptera sp. Diversification of the coffee agroecosystem favors some species while limiting others, and have no effect on the majority of species. Thus, only F. lativittata, Neocoelidia sp. and Scaphytopius ca. latidens were well represented in all systems, but were more abundant in coffee shrubs. Additionally, the following were the dominant species in each system: Graphocephala sp. 1 (C), F. lativittata (CP) and H. nicaraguensis (CPL). Four species abundant on laurel trees, including H. nicaraguensis, appeared almost exclusively on these tree species. Species similarity was highest on the CP and CPL systems (51% of the species in common), followed by the C and CP (39%) and the C and CPL systems (38%). These findings show that even disturbed systems can harbor many insect species, so that they deserve attention from conservation advocates and biologists.     

Small-scale species patterns and associations in an alvar limestone grassland on Öland, Sweden

This paper reports on the results of a transect analysis carried out in order to detect small-scale patterns ofAvenetum species in a limestone grassland on the Great Alvar on the Baltic island of Öland, Sweden. Three transects consisting of 500 10×10 cm quadrats were used. The cover and number of flowering stalks (if any) of the 21 most common species were recorded. In addition the total cover of vascular plants, cover of mosses and lichens, cover of cow dung and soil depth in each quadrat were recorded. Ellenberg indicator values for light, moisture, reaction and nitrogen were calculated for each quadrat as weighted averages of the values for the species included in the analysis. Patch size frequency analysis, PASFRAN, a new method for detecting patches and their sizes, revealed that 42% of all significant species patches detected had mean patch sizes (diameter) in the range of 26–50 cm, which matched the size of dung patches—49 cm in transect 1 and 30 cm in transect 2. Results of association analysis revealed correlations for several species pairs and between species and environmental factors with similar patch sizes. Combination of PASFRAN and association analysis is recommended as an effective method for pattern detection.     

Does climate determine broad‐scale patterns of species richness? A test of the causal link by natural experiment

Aim Broad‐scale spatial patterns of species richness are very strongly correlated with climatic variables. If there is a causal link, i.e. if climate directly or indirectly determines patterns of richness, then when the climatic variables change, richness should change in the manner that spatial correlations between richness and climate would predict. The present study tests this prediction using seasonal changes in climatic variables and bird richness. Location We used a grid of equal area quadrats (37 000 km²) covering North and Central America as far south as Nicaragua. Methods Summer and winter bird distribution data were drawn from monographs and field guides. Climatic data came from published sources. We also used remotely sensed NDVI (normalized difference vegetation index — a measure of greenness). Results Bird species richness changes temporally (between summer and winter) in a manner that is close to, but statistically distinguishable from, the change one would predict from models relating the spatial variation in richness at a single time to climatic variables. If one further takes into account the seasonal changes in NDVI and within‐season variability of temperature and precipitation, then winter and summer richness follow congruent, statistically indistinguishable patterns. Main conclusions Our results are consistent with the hypothesis that climatic variables (temperature and precipitation) and vegetation cover directly or indirectly influence patterns of bird species richness.     

Cryptogamic epiphytes in primary and recovering upper montane oak forests of Costa Rica – species richness,community composition and ecology

Species richness, community composition and ecology of cryptogamic epiphytes (bryophytes, macrolichens) were studied in upper montane primary, early secondary and late secondary oak forests of the Cordillera de Talamanca, Costa Rica. Canopy trees of Quercus copeyensis were sampled with the aim of getting insight in patterns and processes of epiphyte succession and recovery of diversity in secondary forest following forest clearing. Species richness of cryptogamic epiphytes in secondary and primary forests were nearly the same, showing that primary forests are not necessarily more diverse than secondary forests. High species richness of secondary forests was presumed due to the closed canopy, resulting in permanently high atmospheric humidity in these forests. Similarity in species composition of secondary and primary forests increases with forest age, but after 40 years of succession one third (46 species) of primary forest species had not re-established in the secondary forest. Community composition in primary and secondary forests differed markedly and indicates that a long time is needed for the re-establishment of microhabitats and re-invasion of species and communities adapted to differentiated niches. Genera and species exclusive to primary forests are relevant as indicator taxa and conservation targets. Forty percent (68 species) of all species recorded are restricted to secondary forests, indicating the important contribution of secondary forest diversity to total species richness of the oak forests of Costa Rica.     

Restoration of beech forest for saproxylic beetles—effects of habitat fragmentation and substrate density on species diversity and distribution

The influence of spatial location and density of beech snags on species diversity and distribution patterns of saproxylic beetles was studied in a 2,400 ha forest landscape in southern Sweden. Complete snag surveys were combined with a beetle survey using small window traps directly attached to the beech snags. The density of beech snags ≥30 cm dbh varied between one and seven snags per ha within the study area, corresponding to 1.1–5.1 m³/ha. A total of 2,610 specimens of 180 saproxylic beetles species were trapped, of which 19 species were red-listed. Within the study area, the number of red-listed and formerly red-listed species was highest around traps in old-growth stands, intermediate in managed stands contiguous with old-growth and lowest in managed stands isolated from old-growth by a two km-wide zone without beech forest. Logistic regressions revealed negative relationships between distance to old-growth forest and occurrence of eleven species, among them six red-listed or formerly red-listed species. The number of non red-listed species was not correlated with isolation from old-growth forest. The number of red-listed species also increased with snag density within 200–300 m around the traps. Our results suggest that red-listed species generally have a lower dispersal capacity than other saproxylic beetles. We conclude that retention of dead wood close to existing populations is more beneficial for red-listed species than an even distribution of snags across the forest landscape.     

What determines species composition and diversity of hypogeous fungi in the diet of small mammals? A comparison across mammal species,habitat types and seasons in Central European mountains

Interactions between diverse groups of organisms influence the functioning and diversity of ecosystems. Salient examples of such relationships are those among hypogeous fungi, trees and mycophagous mammals. To investigate the role of small mammals in transporting fungal spores within and outside forests as well as the influence of seasons, habitats and species on small mammal mycophagy, we set up a study in the Pieniny Mts, Western Carpathians (Southern Poland). The droppings of small mammals were collected during live trapping in July and September 2016 and 2017, to analyze richness, composition and frequency of fungal spores present in faeces. The yellow-necked mouse Apodemus flavicollis, the bank vole Myodes glareolus and the common vole Microtus arvalis were the most frequently trapped. Spores of 27 fungal taxa from 16 genera were retrieved from nearly 70% of faecal samples of rodents and shrews, with up to 9 spore taxa recorded per sample. Spore diversity in samples was higher in September than in July, although seasonal variation was year and animal dependent. The highest mean number of fungal taxa per sample was recorded for the bank vole and the yellow-necked mouse, with the former species showing a higher degree of mycophagy. The two rodents differed in the average frequencies of consumed fungi in samples, which could result from some degree of specialization in the choice of particular fungal species, as shown by the laboratory-based experiment. Within particular animal species, differences in the fungal diet were found between seasons. The spores of hypogeous fungi were transported from forests to meadows mostly by the yellow-necked mouse and, to a lesser extent, by the common vole. However, both, the diversity and the number of transported spores diminished with distance from the forest edge.     

Does spatial heterogeneity blur the signature of dispersal syndromes on spatial patterns of woody species? A test in a tropical dry forest

Spatial patterns of adult plants are a consequence of several ecological processes related to seed dispersal and recruitment. Dispersal limitation, mediated by dispersal syndrome, is considered a key factor in the formation of adult plant spatial patterns. Although this initial pattern determined by dispersal has been thoroughly studied, the subsequently modification by the effect of additional ecological factors, such as habitat heterogeneity is less understood. We explored the relative importance of dispersal syndrome and spatial heterogeneity on the realization of spatial patterns of adult trees in an Ecuadorian tropical dry forest. The spatial distribution of 28 species was modeled with four different spatial point processes each: homogeneous Poisson (HPP), inhomogeneous Poisson (IPP), homogeneous Poisson cluster (HPCP), and inhomogeneous Poisson cluster process (IPCP). These models allowed us to discern between effects of random processes, habitat heterogeneity, limited dispersal, and joint effects of habitat heterogeneity and limited dispersal. We employed Akaike's information criterion (AIC) to select the model which best fit the spatial pattern of each species. The best model of each species was used to analyze differences in cluster size and degree of aggregation, between dispersal syndromes. Seventy‐five percent of the species showed inhomogeneous patterns. IPCP yielded the best fit for the spatial distribution of 50% of species in the studied forest and was the prevalent model for the three dispersal syndromes. Thus, the effect of spatial heterogeneity was prevalent in the distribution of most species in this dry tropical forest. Only 21% of species had spatial patterns compatible with random mechanisms associated to limited dispersal around parent sources. Clearly, ignoring habitat heterogeneity could bias the analysis of relationships between dispersal syndrome and species patterns.     

What is the relevance of smallholders’ agroforestry systems for conserving tropical tree species and genetic diversity in circa situm, in situ and ex situ settings? A review

Smallholders’ agroforests may be valuable for conserving tropical trees through three main mechanisms. First, trees planted and/or retained by farmers in agricultural landscapes where wild stands were once found may be circa situm reservoirs of biodiversity. Second, farmland trees may support conservation in situ by providing an alternative source of product to reduce extraction from forest, and by acting as ‘corridors’ or ‘stepping stones’ that connect fragmented wild stands. Third, the additional value that planting assigns to trees may result in greater interest in including them in seed collections, field trials and field ‘genebanks’ that support ex situ conservation. Here, we critically review the evidence for these mechanisms, and highlight areas for research and for intervention so that agroforestry practices can better support conservation in each setting, with an emphasis on often neglected genetic-level considerations. Based on current global challenges to diversity, conservation will need to rely increasingly on a smallholder-farm circa situm approach, but concerns on long-term effectiveness need to be properly quantified and addressed. Connectivity between widely dispersed, low density trees in agricultural landscapes is an important factor determining the success of the circa situm approach, while improving farmers’ access to a diversity of tree germplasm that they are interested in planting is required. The circumstances in which agroforestry plantings can support in situ conservation need to be better defined, and research on the stability of active tree seed collections (how long are species and populations retained in them?) as ex situ reservoirs of biodiversity is needed.