3. What is Biodiversity?

AimComponents of scale, such as grain, focus and extent, influence the spatial patterns of alpha and gamma diversity and the relationships between them. We explored these scale relations by testing whether the gamma diversity and alpha diversity along an elevation gradient were related independent of scale and whether the elevational patterns of herbaceous and woody species richness were dependent on scale.LocationLangtang National Park, Nepal.MethodsWe estimated alpha diversity (plot richness) for woody and herbaceous plant species along an alpine elevation gradient (3,900–5,000 m a.s.l.) in nested plots of 1 m2, 16 m2 and 100 m2 and gamma diversity (regional richness) from published sources. Generalized linear modelling was used to analyse alpha and gamma diversity and their correspondence at different grain sizes.ResultsElevational trends of gamma and alpha diversity were significantly correlated for both woody and herbaceous species at all grain sizes. The concordance increased with increasing grain size and area for gamma diversity estimation, particularly for the monotonously decreasing elevational gamma and alpha diversity patterns of woody species. The hump‐shaped patterns of elevational gamma and alpha diversity for herbaceous species were also significantly correlated, but the concordance between the alpha diversity of herbaceous species and local gamma diversity was stronger. Elevational patterns of alpha diversity were coarsely consistent across grain sizes, although the patterns became more pronounced at larger grain sizes.Main conclusionsThe correspondence of elevational gamma and alpha diversity was largely scale invariant, implying that elevational and possibly other geographical diversity patterns can reliably be studied at different spatial scales. Nonetheless, the alpha diversity pattern was the least pronounced at fine grain size, particularly for woody life‐forms. This finding suggests that for large‐scale patterns such as elevational gradients at regional or continental scales, coarse grain sizes and large areas for gamma estimation are more appropriate.

Speciation is the process that ultimately generates species richness. However, the time required for speciation to build up diversity in a region is rarely considered as an explanation for patterns of species richness. We explored this "time-for-speciation effect" on patterns of species richness in emydid turtles. Emydids show a striking pattern of high species richness in eastern North America (especially the southeast) and low diversity in other regions. At the continental scale, species richness is positively correlated with the amount of time emydids have been present and speciating in each region, with eastern North America being the ancestral region. Within eastern North America, higher regional species richness in the southeast is associated with smaller geographic range sizes and not greater local species richness in southern communities. We suggest that these patterns of geographic range size variation and local and regional species richness in eastern North America are caused by glaciation, allopatric speciation, and the time-for-speciation effect. We propose that allopatric speciation can simultaneously decrease geographic range size and increase regional diversity without increasing local diversity and that geographic range size can determine the relationship between alpha, beta, and gamma diversity. The time-for-speciation effect may act through a variety of processes at different spatial scales to determine diverse patterns of species richness.

To understand patterns of alpha, beta and gamma diversities in fragmented landscapes we need to explore the three scale components in relation to potential drivers in a scale-dependent manner. Often, the drivers themselves can be partitioned to alpha, beta and gamma diversities. Thus, one can hypothesize that the scale-components of species diversity and drivers’ diversity match, i.e., that species alpha diversity is mainly explained by drivers’ alpha diversity, beta by beta and gamma by gamma. Here, we explore this ‘scale-matching’ hypothesis for spiders in two fragmented agricultural landscapes. In each landscape, we sampled spiders and their potential prey in 12 patches. Then, we sub-sampled pseudo-landscapes in which we calculated spider alpha, beta and gamma diversities using multiplicative diversity-partitioning. Next, we used variance partitioning analysis to explore the relative contribution of eleven explanatory variables from five thematic groups (sampling intensity, area, connectivity, habitat diversity and prey diversity), while further partitioning the habitat and prey diversities to their corresponding alpha, beta and gamma diversities. We found considerable evidence for scale-matching, with spiders’ alpha and beta diversities explained mostly by the corresponding alpha and beta diversities (respectively) of prey and/or habitat. We further found a strong effect of connectivity on spider beta diversity, but not on alpha and gamma diversities. For spiders gamma diversity, a cross-scale effect was observed. Our results suggest that multiple drivers from multiple scales interact in structuring patterns of spider alpha, beta and gamma diversities in agro-ecosystems, yet the strongest effects are of those drivers that match in scale.

Summary 1. Local and regional patterns of plant species richness in tropical rain forests, as well as their possible drivers, remain largely unexplored. The main hypotheses for local species richness (alpha diversity) are (i) local environmental determinism with species-saturated communities, and (ii) regional control, in which the immigration of species from the regional species pool (gamma diversity) determines how many species coexist locally. The species pool hypothesis suggests a combined influence of local and regional drivers on alpha diversity. Differences in gamma diversity may arise from divergent environmental conditions or biogeographic histories. 2. We investigated the cross-scale determinants of palm alpha and gamma diversity across the western Amazon using a large field-based data set: a census of all palm individuals in 312 transects, totalling 98 species. We used regression-based variation partitioning to understand how habitat, topography and region influence alpha diversity, and correlations to assess the importance of the present environment (climate, soil, regional topography) and history (long-term habitat stability) for average regional alpha diversity and gamma diversity, including the link between these two diversity measures (species pool effect). 3. Variation in alpha diversity was primarily explained by region (36%) and habitat (18%), whereas the effect of topography was negligible (1%). Within habitats, region was even more important (up to 69% explained variation). Within regions, habitat and topography covaried and had a variable but an important influence. The pure effect of topography remained of minor importance (up to 13%). 4. Average regional alpha diversity was related to gamma diversity, precipitation seasonality and possibly long-term habitat stability. Gamma diversity was related to long-term habitat stability, and possibly current climate. 5. Synthesis. Gamma diversity strongly influenced alpha diversity, although a clear influence of local environment was also evident, notably habitat type, with a minor, more geographically variable effect of small-scale topography. Apart from gamma diversity, the factor most strongly related to regional alpha diversity was precipitation seasonality, while gamma diversity itself was strongly linked to long-term habitat stability. These results imply that plant species richness is contingent on both contemporary and historical factors with a strong link between local species richness and the regional species pool.

Patterns of Bryophyte Diversity in Peatlands of Continental Western Canada

BackgroundBiodiversity, crucial for understanding ecosystems, encompasses species richness, composition, and distribution. Ecological and environmental factors, such as habitat type, resource availability, and climate conditions, play pivotal roles in shaping species diversity within and among communities, categorized into alpha (within habitat), beta (between habitats), and gamma (total regional) diversity. Hummingbird communities are influenced by habitat, elevation, and seasonality, making them an ideal system for studying these diversities, shedding light on mutualistic community dynamics and conservation strategies.MethodsOver a year-long period, monthly surveys were conducted to record hummingbird species and their visited flowering plants across four habitat types (oak forest, juniper forest, pine forest, and xerophytic shrubland) in Tlaxcala, Mexico. Three locations per habitat type were selected based on conservation status and distance from urban areas. True diversity measures were used to assess alpha, beta, and gamma diversity of hummingbirds and their floral resources. Environmental factors such as altitude and bioclimatic variables were explored for their influence on beta diversity.ResultsFor flowering plants, gamma diversity encompassed 34 species, with oak forests exhibiting the highest richness, while xerophytic shrublands had the highest alpha diversity. In contrast, for hummingbirds, 11 species comprised the gamma diversity, with xerophytic shrublands having the highest richness and alpha diversity. Our data reveal high heterogeneity in species abundance among habitats. Notably, certain floral resources like Loeselia mexicana and Bouvardia ternifolia emerge as key species in multiple habitats, while hummingbirds such as Basilinna leucotis, Selasphorus platycercus, and Calothorax lucifer exhibit varying levels of abundance and habitat preferences. Beta diversity analyses unveil habitat-specific patterns, with species turnover predominantly driving dissimilarity in composition. Moreover, our study explores the relationships between these diversity components and environmental factors such as altitude and climate variables. Climate variables, in particular, emerge as significant contributors to dissimilarity in floral resource and hummingbird communities, highlighting the influence of environmental conditions on species distribution.ConclusionsOur results shed light on the complex dynamics of hummingbird-flower mutualistic communities within diverse habitats and underscore the importance of understanding how habitat-driven shifts impact alpha, beta, and gamma diversity. Such insights are crucial for conservation strategies aimed at preserving the delicate ecological relationships that underpin biodiversity in these communities.

AimOur aim was to test whether species richness patterns are best explained by the effect of the total amount of habitat within the landscape, or instead by a combination of patch size and patch isolation effects. To this end, we jointly contrast the habitat amount hypothesis and countryside biogeography with patch size and isolation concepts from island biogeography.LocationThree multi‐habitat landscapes in Peneda‐Gerês National Park, NW Portugal.TaxonMacro‐moths (Lepidoptera).MethodsLight‐trapping using a semi‐nested design at 84 fixed sites which were each repeatedly sampled six times.ResultsAutocovariate models show that sampling sites with a higher number of forest and meadow macro‐moth species (alpha diversity) were surrounded by a higher amount of forest and meadow habitat, respectively within a 160 and 320 m radius (scale of effect). These top‐ranked models, containing only habitat amount as a significant variable, had lower Akaike's information criteria (AIC) than models (only) containing patch size and/or isolation. Complementary to this, the countryside species–area relationship (SAR) model outperforms the classic SAR model, so that the effective area of habitat explains landscape species richness (gamma diversity) across spatial scales (beta diversity) better than the classic SAR. Specifically, we show that forest macro‐moths have a higher spatial turnover than meadow macro‐moths and that, on average, there are more species in forest than in meadow habitat.Main conclusionsThe habitat amount hypothesis predicts alpha species richness in multi‐habitat landscapes better than do patch size and isolation while the countryside SAR predicts beta and gamma diversity better than the classic SAR. We suggest that evidence is mounting to revise the application of the classical approaches of island biogeography and metapopulation theory to conservation biogeography.

The latitudinal diversity gradient predicts that tropical regions should have higher alpha, beta, and gamma diversity than temperate areas. However, only a few studies have assessed the temporal variability of the different components of diversity across climatic regions. In this study, we compare, using a spatial and temporal approach, the diversity of multiple taxa inhabiting tropical and Mediterranean temporary ponds. We sampled the biological communities of each set of ponds on three occasions during the same hydrological year. Under a spatial framework, we analyzed, alpha, beta, and gamma diversities. With a temporal approach, we compared the coefficients of variation in alpha diversity for each local community, and temporal beta diversity. Differences between regions and sampling periods were tested using generalized linear mixed models. We found higher gamma and alpha diversity in the tropical ponds, as expected given the latitudinal differences between them. However, phytoplankton and microinvertebrates from the Mediterranean region, matched or even exceeded tropical alpha diversity on some occasions. Spatial beta diversity did not differ between regions, and it showed lower values at the middle or the end of the hydroperiod in bacteria, micro‐ and macroinvertebrates and amphibians. Thus, processes homogenizing and heterogenising pond metacommunities must be balanced in both studied regions. Temporal variation in alpha and beta diversity was similar for ponds in both regions, except for macroinvertebrates and amphibians, suggesting differential effects on community variation observable only in animals with longer life‐spans, at our temporal scale of analysis.

Strong anthropogenic pressures on global forests necessitate that managed forests be evaluated as habitat for biodiversity. The complex pattern of habitat types created in forestry systems is ideal for analyses through the theoretical framework of alpha (local), gamma (total) and beta (compositional) diversity. Here I use saproxylic beetles, a species-rich threatened group, to compare four Norway spruce-dominated habitats representative of the boreal forest landscape of northern Europe: unmanaged semi-natural stands, nature reserves, unthinned middle-aged production stands and commercially thinned production stands. The beetles (in total 38 085 individuals of 312 species), including red-listed ones and three feeding guilds (wood consumers, fungivores and predators) were studied in 53 stands in central-southern Sweden, in two regions with differing amounts of conservation forest. Alpha diversity of saproxylic, but not red-listed, beetles was higher in the thinned stands than in the semi-natural stands, and did not differ for the other forest types. Beta diversity of saproxylic beetles was higher in unmanaged semi-natural stands than in the other forest types, but species composition did not differ noticeably. Furthermore, red-listed saproxylic beetles had higher gamma diversity in unmanaged semi-natural stands in the region with more conservation forest, but not in the one with less such forest. The local factors dead wood volume and dead wood diversity did not influence alpha diversity of beetles, but increasing canopy openness had a minor negative influence on saproxylic and red-listed beetles. While the local scale (alpha diversity) indicates the potential for managed forests to house many saproxylic beetle species associated with spruce forests in this boreal landscape, the larger scales (beta and gamma diversity) indicate the value of unmanaged forests for the conservation of the entire saproxylic beetle fauna. These results show the importance of analyses at multiple levels of diversity (alpha, beta, gamma) for identifying patterns relevant to conservation.

Strong anthropogenic pressures on global forests necessitate that managed forests be evaluated as habitat for biodiversity. The complex pattern of habitat types created in forestry systems is ideal for analyses through the theoretical framework of alpha (local), gamma (total) and beta (compositional) diversity. Here I use saproxylic beetles, a species-rich threatened group, to compare four Norway spruce-dominated habitats representative of the boreal forest landscape of northern Europe: unmanaged semi-natural stands, nature reserves, unthinned middle-aged production stands and commercially thinned production stands. The beetles (in total 38 085 individuals of 312 species), including red-listed ones and three feeding guilds (wood consumers, fungivores and predators) were studied in 53 stands in central-southern Sweden, in two regions with differing amounts of conservation forest. Alpha diversity of saproxylic, but not red-listed, beetles was higher in the thinned stands than in the semi-natural stands, and did not differ for the other forest types. Beta diversity of saproxylic beetles was higher in unmanaged semi-natural stands than in the other forest types, but species composition did not differ noticeably. Furthermore, red-listed saproxylic beetles had higher gamma diversity in unmanaged semi-natural stands in the region with more conservation forest, but not in the one with less such forest. The local factors dead wood volume and dead wood diversity did not influence alpha diversity of beetles, but increasing canopy openness had a minor negative influence on saproxylic and red-listed beetles. While the local scale (alpha diversity) indicates the potential for managed forests to house many saproxylic beetle species associated with spruce forests in this boreal landscape, the larger scales (beta and gamma diversity) indicate the value of unmanaged forests for the conservation of the entire saproxylic beetle fauna. These results show the importance of analyses at multiple levels of diversity (alpha, beta, gamma) for identifying patterns relevant to conservation.

Simple SummaryWe studied the diversity of arthropods in native forests along a 1000 m elevation gradient on Terceira Island, Azores (Portugal). These forests form an isolated and threatened habitat with unique endemic species. We analysed the change in alpha and beta diversity of arthropod species with elevation and if the diversity of endemic, native non-endemic and introduced species responds differently to elevation. Resident arthropods were sampled using SLAM (Sea, Land and Air Malaise) traps between 2014 and 2018. Spiders (Araneae), beetles (Coleoptera), true bugs (Hemiptera) and barklice (Psocoptera), as well as endemic, native and introduced species, were analysed separately. Total species richness decreases with elevation for all species, Coleoptera and Psocoptera, and particularly so for introduced species, but peaks at mid-high elevation for Araneae and endemic species. These patterns are probably driven by unfavourable climatic conditions at higher elevations while being influenced by human disturbance at lower elevations. Total species diversity along the whole elevation gradient is shaped by this decreasing richness as well as the replacement of species at different elevations.We present an analysis of arthropod diversity patterns in native forest communities along the small elevation gradient (0–1021 m a.s.l.) of Terceira island, Azores (Portugal). We analysed (1) how the alpha diversity of Azorean arthropods responds to increasing elevation and (2) differs between endemic, native non-endemic and introduced (alien) species, and (3) the contributions of species replacement and richness difference to beta diversity. Arthropods were sampled using SLAM traps between 2014 and 2018. We analysed species richness indicators, the Hill series and beta diversity partitioning (species replacement and species richness differences). Selected orders (Araneae, Coleoptera, Hemiptera and Psocoptera) and endemic, native non-endemic and introduced species were analysed separately. Total species richness shows a monotonic decrease with elevation for all species and Coleoptera and Psocoptera, but peaks at mid-high elevation for Araneae and endemic species. Introduced species richness decreases strongly with elevation especially. These patterns are most likely driven by climatic factors but also influenced by human disturbance. Beta diversity is, for most groups, the main component of total (gamma) diversity along the gradient but shows no relation with elevation. It results from a combined effect of richness decrease with elevation and species replacement in groups with many narrow-ranged species.

Summary A plant that causes specific changes to soil biota may either positively or negatively affect the performance of the plant that subsequently grows in that location. These effects, known as plant–soil feedback, can affect plant species diversity at multiple spatial scales. It has been hypothesized that positive plant–soil feedback reduces alpha (local) diversity by allowing dominance by early‐arriving species, but increases gamma (regional) diversity by promoting community divergence (increased beta diversity) through the emergence of alternative stable states. In contrast, negative plant–soil feedback has been thought to increase alpha diversity by allowing local species coexistence, but to reduce gamma diversity by promoting community convergence (reduced beta diversity). Although widely accepted, these hypotheses do not consider the possibility that plant species differ in their effect on, and their response to, a given other species via soil biota. In reality, plant–soil interactions can be complex, with the strength of the interactions variable between plant species. Using a basic simulation model of plant community assembly, we investigated how complex plant–soil interactions might affect plant diversity during succession. When we included only positive or negative intraspecific plant–soil feedback in the model, with no variation in the strength of interspecific plant–soil interactions, results were consistent with the conventional hypotheses. When we allowed the strength of plant–soil interactions to differ between species, plant–soil interactions enhanced alpha diversity initially and beta and gamma diversity subsequently. Diversity enhancement occurred not necessarily because alternative stable states emerged, but primarily because complex plant–soil interactions lengthened the time during which local species composition changed. Due to the longer time for changes in species composition, the high level of beta and gamma diversity at the early stage of succession was maintained for a long time despite eventual community convergence. Thus, diversity enhancement was often transient, though long‐lasting, making the conventional concept of alternative stable states inadequate for explaining diversity. Synthesis. Based on these findings, we propose the new hypothesis that complex plant–soil interactions enhance plant species diversity by delaying community convergence. This hypothesis highlights the role of plant–soil interactions as a driver of long‐lasting transient dynamics of community assembly.

The partitioning of gamma diversity into its alpha and beta components has been used to quantify the effects of natural habitat reduction on species diversity and distribution. Ant assemblages respond to anthropogenic changes, allowing their use as an indicator of biodiversity conservation in human‐modified landscapes. We investigated the responses of gamma, alpha, and beta diversity of ant assemblages to a forest cover gradient in southwestern Brazilian Amazon. Our prediction was that increasing forest cover would increase gamma diversity, which in turn is driven more by increasing beta diversity than alpha diversity along the forest cover gradient. Additionally, we expected that increasing beta diversity would be mainly driven by species replacement. We sampled ants within 12 circular areas (radius: 500 m) with different forest cover percentages in Chico Mendes Extractive Reserve, state of Acre, southwestern Brazilian Amazon. Gamma and beta diversity, but not alpha diversity, responded positively to the forest cover gradient. Additionally, species replacement was the main driver of beta diversity along the forest cover gradient. Areas with low levels of forest cover probably offer limited variation in habitats, resources, and conditions and, thus, harbor a low number of ant species that can easily disperse spatially and leading to species composition simplification. We recommend the use of diversity partitioning for indicator groups as an efficient way to monitor biodiversity and control the process of biota simplification at the human‐modified landscapes, showing the response of species diversity and distribution to human impacts.Abstract in Portuguese is available with online material.

Response of beta diversity to pulses of Ordovician-Silurian mass extinction.

Species richness – Energy relationships and dung beetle diversity across an aridity and trophic resource gradient