Plant species richness–productivity relationships in a low-productive boreal region

Local species richness–productivity (SR–P) relationship is usually reported as unimodal if long productivity gradients are sampled. However, it tends to be monotonically increasing in low-productive environments due to the decreasing part of the SR–P curve being truncated. Previous work indicated that this can hold true for forest herb layers, because of an upper bound on productivity caused mainly by canopy shading. Here, we ask whether the same pattern exists in a region with an upper bound on productivity caused by a harsh climate. We sampled herbaceous vegetation of boreal forests and grasslands in a low-productive region of central Yakutia (NE Siberia) with dry and winter-cool continental climate. We collected data on species composition, herb-layer productivity (aboveground herbaceous biomass), soil chemistry and light availability. We applied regression models to discriminate between monotonically increasing, decreasing and unimodal responses of herb-layer species richness to measured variables and analysed trends in the species-pool size and beta diversity along the productivity gradient. Our expectation of the monotonically increasing SR–P relationship was confirmed for neither forest herb layers nor grasslands. In the forest herb layers, no relationship was detected. In grasslands, the relationship was unimodal with species richness decline starting at much lower productivity levels than in more productive temperate grasslands. Potential causes for this decline are either limitation of local species richness by the species pool, which contains few species adapted to more productive habitats, or competitive exclusion, which can become an important control of species richness under lower levels of productivity than is the case in temperate grasslands.     

Does environmental heterogeneity affect species co‐occurrence in ecological guilds across stream macroinvertebrate metacommunities?

Null model analyses have greatly improved our understanding of species co‐occurrence. Null model analyses have shown, for example, that cold‐blooded animals show less segregated distributions than warm‐blooded animals. This topic has rarely been studied simultaneously across multiple metacommunities. We analysed data on 10 stream metacommunities (with 10 communities in each metacommunity) of a cold‐blooded animal group, benthic macroinvertebrates, and examined co‐occurrence within five ecological guilds. We found that the segregated species co‐occurrence was not the rule in stream invertebrate guilds. This was evidenced by the finding that only 10% of the 50 guild matrices we analyzed showed significant segregation and no matrices showed significant aggregation in the within‐stream analyses. However, in the across‐streams analysis, all guilds showed significant segregation. We neither found differences in the degree of segregation among the guilds, the degree of species segregation did not increase with overall environmental heterogeneity, and there were no differences in the relationships between species segregation and overall environmental heterogeneity among the guilds. Expanding the spatial extent from single stream metacommunities (i.e. within each stream) to the whole study region (i.e. across the streams) increased significantly segregation in all guilds. Because environmental heterogeneity across streams was much higher than within single streams, overall environmental heterogeneity may nevertheless have effects on species segregation. It also seems that the effects of overall heterogeneity on species segregation were masked by mass effects in the within‐stream analyses.     

Does soil moisture availability explain liana seedling distribution across a tropical rainfall gradient?

Liana density tends to increase with decreasing rainfall and increasing seasonality. However, the pattern of liana distribution may be due to differences in soil water retention capacity, not rainfall and seasonality per se. We tested the effect of rainfall and soil substrate with respect to the distribution of liana seedlings in six sites across a rainfall gradient from the wet Atlantic to the dry Pacific in central Panama. Soils were either limestone, with low water‐holding capacity, or laterite, with higher water‐holding capacity. We sampled liana seedlings at each site using three 1 × 100 m transect. We found that relative liana seedling density was higher on limestone soils compared to laterite soils regardless of the amount of rainfall. Furthermore, liana community composition on limestone soils was more similar to dry forest sites than to adjacent wet and moist forest sites. Liana seedling species diversity relative to trees was significantly higher in a low‐fertility dry forest site compared to a high‐fertility forest, but did not differ from the other sites. Thus, liana seedling density and community structure may be driven more by soil type and thus by soil moisture availability than strictly by mean annual rainfall and the seasonality of rainfall.     

Does energy availability influence classical patterns of spatial variation in exotic species richness?

Aim At macroecological scales, exotic species richness is frequently positively correlated with human population density. Such patterns are typically thought to arise because high human densities are associated with increased introduction effort and/or habitat modification and disturbance. Exotic and native species richness are also frequently positively correlated, although the causal mechanisms remain unclear. Energy availability frequently explains much of the variation in species richness and we test whether such species–energy relationships may influence the relationships of exotic species richness with human population density and native species richness. Location Great Britain. Methods We first investigate how spatial variation in the distributions of the 10 exotic bird species is related to energy availability. We then model exotic species richness using native avian species richness, human population density and energy availability as predictors. Species richness is modelled using two sets of models: one assumes independent errors and the other takes spatial correlation into account. Results The probability of each exotic species occurring, in a 10‐km quadrat, increases with energy availability. Exotic species richness is positively correlated with energy availability, human population density and native species richness in univariate tests. When taking energy availability into account, exotic species richness is negligibly influenced by human population density, but remains positively associated with native species richness. Main conclusions We provide one of the few demonstrations that energy availability exerts a strong positive influence on exotic species richness. Within our data, the positive relationship between exotic species richness and human population density probably arises because both variables increase with energy availability, and may be independent of the influence of human density on the probability of establishment. Positive correlations between exotic and native species richness remain when controlling for the influence of energy on species richness. The relevance of such a finding to the debate on the relationship between diversity and invasibility is discussed.     

Does habitat use explain large scale species richness patterns of aquatic beetles in Europe?

Regularities in species richness are widely observed but controversy continues over its mechanistic explanation. Because richness patterns are usually a compound measure derived from taxonomically diverse species with different ecological requirements, these analyses may confound diverse causes of species numbers. Here we investigate species richness in the aquatic beetle fauna of Europe, separating major taxonomic groups and two major ecological types, species occurring in standing and running water bodies. We collated species distributions for 800+ species of water beetles in 15 regions across western Europe. Species number in any of these regions was related to three variables: total area size, geographic connectedness of the area, and latitude. Pooled species numbers were accurately predicted, but correlations were different for species associated with either running or standing water. The former were mostly correlated with latitude, while the latter were only correlated with the measure of connectedness or with area size. These differences were generally also observed in each of the four phylogenetically independent lineages of aquatic Coleoptera when analysed separately. We propose that effects of habitat, in this case possibly mediated by different long term persistence of running and standing water bodies, impose constraints at the population or local level which, if effective over larger temporal and spatial scales, determine global patterns of species richness.     

Does coevolution promote species richness in parasitic cuckoos?

Why some lineages have diversified into larger numbers of species than others is a fundamental but still relatively poorly understood aspect of the evolutionary process. Coevolution has been recognized as a potentially important engine of speciation, but has rarely been tested in a comparative framework. We use a comparative approach based on a complete phylogeny of all living cuckoos to test whether parasite–host coevolution is associated with patterns of cuckoo species richness. There are no clear differences between parental and parasitic cuckoos in the number of species per genus. However, a cladogenesis test shows that brood parasitism is associated with both significantly higher speciation and extinction rates. Furthermore, subspecies diversification rate estimates were over twice as high in parasitic cuckoos as in parental cuckoos. Among parasitic cuckoos, there is marked variation in the severity of the detrimental effects on host fitness; chicks of some cuckoo species are raised alongside the young of the host and others are more virulent, with the cuckoo chick ejecting or killing the eggs/young of the host. We show that cuckoos with a more virulent parasitic strategy have more recognized subspecies. In addition, cuckoo species with more recognized subspecies have more hosts. These results hold after controlling for confounding geographical effects such as range size and isolation in archipelagos. Although the power of our analyses is limited by the fact that brood parasitism evolved independently only three times in cuckoos, our results suggest that coevolutionary arms races with hosts have contributed to higher speciation and extinction rates in parasitic cuckoos.     

How does forest landscape structure explain tree species richness in a Mediterranean context?

Although the strong relationship between vegetation and climatic factors is widely accepted, other landscape composition and configuration characteristics could be significantly related with vegetation diversity patterns at different scales. Variation partitioning was conducted in order to analyse to what degree forest landscape structure, compared to other spatial and environmental factors, explained forest tree species richness in 278 UTM 10 × 10 km cells in the Mediterranean region of Catalonia (NE Spain). Tree species richness variation was decomposed through linear regression into three groups of explanatory variables: forest landscape (composition and configuration), environmental (topography and climate) and spatial variables. Additionally, the forest landscape characteristics which significantly contributed to explain richness variation were identified through a multiple regression model. About 60% of tree species richness variation was explained by the whole set of variables, while their joint effects explained nearly 28%. Forest landscape variables were those with a greater pure explanatory power for tree species richness (about 15% of total variation), much larger than the pure effect of environmental or spatial variables (about 2% each). Forest canopy cover, forest area and land cover diversity were the most significant composition variables in the regression model. Landscape configuration metrics had a minor effect on forest tree species richness, with the exception of some shape complexity indices, as indicators of land use intensity and edge effects. Our results highlight the importance of considering the forest landscape structure in order to understand the distribution of vegetation diversity in strongly human-modified regions like the Mediterranean.     

Wild bee species abundance and richness across an urban–rural gradient

Long-term and widespread monitoring programs are essential to understanding the role of human-dominated landscapes in supporting wild bee populations. Urbanization results in increased impervious surfaces throughout the landscape, fragmentation of green space, and a loss of naturally occurring floral vegetation. All of these changes have a negative impact on pollinator diversity. The objective of this study was to assess the abundance and richness of wild bee species throughout a small city in northwest Pennsylvania and identify how management of land throughout the city may influence bee communities. Seventeen sites across a land use gradient, moving from areas with large open spaces and mainly permeable surfaces, to sites in the city center consisting of mainly impermeable surfaces, were sampled over a 2-year period. During this time, 106 known species were identified with four state records and 1 undescribed species. Bee species richness was greatest at sites with the largest amount of permeable surface and naturally-occurring, native vegetation. Richness decreased on the college campus and city center where landscapes were highly managed and impermeable surfaces were most abundant. While floral richness was not related to bee abundance and richness, the number of open blooms near traps did have a positive impact on bee species richness. Overall, this survey revealed considerable richness never before recorded for northwest Pennsylvania, suggesting the importance of conservation management in homeowner and community yard space.     

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.     

Rapoport's rule: an explanation or a byproduct of the latitudinal gradient in species richness?

A recent explanation of the declining species richness gradient with increasing latitude away from the tropics implicated broad scale habitat variability, an associated range expansion, and a resulting increase in niche breadth. The niche breadth in turn was thought to affect richness by competition and rescue effect. While all three factors appear to be correlated, neither the postulated nor alternative causal mechanisms have been tested. We conduct such a test using a system which has all the attributes of the large scale pattern but which, in contrast to continental scale observations, allows for estimation or control of crucial variables such as taxonomic composition, habitat heterogeneity, habitat variability, exact species distribution, and local richness. Specifically, we test the alternative that the correlation between the geographical range of species and local diversity is a function of differential species survival and link this phenomenon to habitat variability. We use 40 species of aquatic invertebrates inhabiting a landscape of 49 miniature rock pools on the coast of Jamaica. The system we examined exhibits a gradient of increasing richness with decreasing habitat variability, analogous to the broad scale latitudinal pattern. Furthermore, species with broader ecological ranges are also broadly distributed. Superficially, this appears to be in agreement with the older explanations but two facts suggest different causes. First, there is no evidence of a rescue effect maintaining high richness in many habitats despite their proximity to species sources. Second, ecologically broad species coexist with habitat specialists without reducing richness in jointly occupied habitats.     

Does species diversity limit productivity in natural grassland communities?

Theoretical analyses and experimental studies of synthesized assemblages indicate that under particular circumstances species diversity can enhance community productivity through niche complementarity. It remains unclear whether this process has important effects in mature natural ecosystems where competitive feedbacks and complex environmental influences affect diversity–productivity relationships. In this study, we evaluated diversity–productivity relationships while statistically controlling for environmental influences in 12 natural grassland ecosystems. Because diversity–productivity relationships are conspicuously nonlinear, we developed a nonlinear structural equation modeling (SEM) methodology to separate the effects of diversity on productivity from the effects of productivity on diversity. Meta-analysis was used to summarize the SEM findings across studies. While competitive effects were readily detected, enhancement of production by diversity was not. These results suggest that the influence of small-scale diversity on productivity in mature natural systems is a weak force, both in absolute terms and relative to the effects of other controls on productivity.     

Does genomic variation in a foundation species predict arthropod community structure in a riparian forest?

Understanding how genetic variation within a foundation species determines the structure of associated communities and ecosystem processes has been an emerging frontier in ecology. Previous studies in common gardens identified close links between intraspecific variation and multispecies community structure, and these findings are now being evaluated directly in the complex natural ecosystem. In this study, we examined to what extent genomic variation in a foundation tree species explains the structure of associated arthropod communities in the field, comparing with spatial, temporal and environmental factors. In a continuous mixed forest, arthropods were surveyed on 85 mature alders (Alnus hirsuta) in 2 years. Moreover, we estimated Nei's genetic distance among the alders based on 1,077 single nucleotide polymorphisms obtained from restricted‐site‐associated DNA sequencing of the alders’ genome. In both years, we detected significant correlations between genetic distance and dissimilarity of arthropod communities. A generalized dissimilarity modelling indicated that the genetic distance of alder populations was the most important predictor to explain the variance of arthropod communities. Among arthropod functional groups, carnivores were consistently correlated with genetic distance of the foundation species in both years. Furthermore, the extent of year‐to‐year changes in arthropod communities was more similar between more genetically closed alder populations. This study demonstrates that the genetic similarity rule would be primarily prominent in community assembly of plant‐associated arthropods under temporally and spatially variable environments in the field.     

Can spatial variation and inter-annual variation in precipitation explain the seed density and species richness of the germinable soil seed bank in a tropical dry forest in north-eastern Brazil?

The interaction between microhabitat and inter-annual variation in precipitation has an important role on the dynamics of the seed bank and can play a crucial role in survival and maintenance of plant populations in semi-arid environments. We hypothesized that the type of microhabitat and the inter-annual variability in precipitation can explain the richness and density of the seed bank in a semi-arid region in Brazil. The study was conducted in an area of tropical dry forest with shrub-tree physiognomy, locally called caatinga. We collected 35 soil samples in three distinct microhabitats, at the end of rainy and dry seasons, respectively, over three years, totalling 630 samples. The seed bank (richness and seed density) were determined by seedling emergence method. Over the three years, 79 species emerged from the seed bank, 64, 45 and 42 in riparian, non-riparian and rocky microhabitats, respectively. We recorded differences in species richness and average density between microhabitats and between years, with significant statistical interaction between them. Inter-annual precipitation explained 48% and 5% of the variation in richness and seed density, respectively. Spatial variation explained 7% of the species richness and 31% of the density. Our results show that the interaction between spatial variation and precipitation has an important role on the spatial and temporal heterogeneity of the richness and density of seed banks in dry environments.     

Does scale‐dependent feedback explain spatial complexity in salt‐marsh ecosystems?

Complexity theory highlights scale-dependent feedback mechanisms as an explanation for regular spatial patterning in ecosystems. To what extent scale-dependent feedback clarifies spatial structure in more complex, non-regular systems remains unexplored so far. We report on a scale-dependent feedback process generating patchy landscapes at the interface of intertidal flats and salt marshes. Here, vegetation was characterized by Spartina anglica tussocks, surrounded by erosion gullies. To demonstrate the presence of a scale-dependent feedback, we determined if vegetation induced habitat modification resulted in local facilitation and large scale-inhibition of plant growth. Field surveys revealed that larger tussocks have deeper gullies, suggesting that gully erosion is caused by increased water flow around tussocks. This was confirmed by flume experiments, showing that feedback effects vary with current velocity and water depth. Transplantation of small Spartina units inside and just outside present tussocks revealed that the growth of Spartina transplants compared to transplant growth on bare sediment was higher within the raised Spartina tussocks, but lower in the gully just outside Spartina tussocks, providing clear evidence of scale-dependent feedback. Our results emphasize that scale-dependent feedback is a more general explanation for spatial complexity in ecosystems than previously considered.     

Does anti‐parasitoid defense explain host‐plant selection by a polyphagous caterpillar?

While studies of tri-trophic interactions have uncovered a variety of mechanisms influencing the dietary specialization of insect herbivores, such studies have neglected host-plant selection by generalists. Here, we report an initial investigation on how host-plant quality and a tachinid parasitoid interact to affect the survival and host-plant selection by a polyphagous herbivore. This herbivore, Grammia geneura (Strecker) (Lepidoptera: Arctiidae), is a food-mixing caterpillar that feeds preferentially on forbs. A previous study suggested that G. geneura might eat certain host species for reasons other than benefits of physiological utilization. We hypothesized that host-plant mediated defenses could act against parasitoids, the major mortality agents of late instar G. geneura . Field observations indicated that caterpillars sometimes survived an attack by the parasitoid Exorista mella Walker (Diptera: Tachinidae) in nature. Laboratory experiments showed that the survival of parasitized caterpillars increased on acceptable but nutritionally inferior host-plant species, indicating that anti-parasitoid defense may explain host-plant selection in this dietary generalist. We found no indication that host-plant selection changed according to the parasitism status of individual caterpillars.     

Can host body size explain the parasite species richness in tropical freshwater fishes?

Summary The variability of monogenean gill ectoparasite species richness in 19 West African cyprinid species was analyzed using the following seven predictor variables: host size, number of drainage basins, number of sympatric cyprinid species, host diversity, association with mainland forest, host ecology, and monogenean biological labelling. The size of the host species accounted for 77% of the variation in the number of parasite species per host, and host ecology an additional 8%. Together the effects of host size and host ecology accounted for 85% of the variation in monogenean species richness. This study shows that the deciding factors for explaining monogenean species richness in West African cyprinid fishes are host species size and host ecology. These results were compared with main factors responsible for parasite species richness in fish communities. Other possible explanations of monogenean community structure in west African cyprinids are discussed.     

Can the mass effect explain the mid-altitudinal peak in vascular plant species richness?

A mid-altitudinal peak in species richness is commonly observed and the mass effect (or source–sink effect) has been suggested as a possible cause. We test the importance of the mass effect for generating altitudinal patterns of plant species richness at two grain sizes using a simple estimate of sterility/fertility to indicate sinks and sources. To do this we identified species with fertile specimens (fertile species) and species with only sterile specimens (sterile species) in each sampling unit along altitudinal transects and assumed that the number of sterile species indicated the relative number of sink species, correspondingly that the number of fertile species indicated the relative number of source species when looking at the overall pattern of species richness along a transect. To evaluate this approach, we investigated the distribution of sterility and fertility of each species along the altitudinal transects. We found that sterile species are found more often at the edges and fertile species more often in the centre of the species altitudinal ranges than expected by chance. Using a fine grain, sterile species richness had a humped altitudinal pattern on all transects investigated at this scale, whereas using a coarse grain two of the three transects investigated had a humped pattern. At the fine grain, sterile species richness had a more pronounced peak than fertile species richness in two of the three transects investigated supporting the hypothesis of the mass effect, but this pattern did not persist at coarser grain. The observations at the fine grain are in accordance with the idea that the mass effect is important in shaping the mid-altitudinal peak in species richness, whereas the observations from the coarser grain are ambiguous.     

Plant species richness and composition of heathland relics in north‐western Belgium: evidence for a rescue‐effect?

Aim To study the effects of the degree of fragmentation of heathland patches on their species richness and species composition, and to infer the ecological mechanisms behind the observed patterns. Location The heathland patches of the north‐western part of Flanders, Belgium. During the last 200 years, the heathland area in this study area has been reduced from c. 10,000 to c. 40 ha, yielding c. 150 small and highly isolated relic fragments. Methods Different isolation measures were calculated for each of the 153 inventoried heathland patches. The influence of isolation, area and habitat diversity on species richness was investigated using correlation techniques. Community composition of the patches was tested for nestedness, and the mechanisms potentially underlying this pattern were determined. Results and main conclusions Both the analyses at the species richness and community composition level yielded evidence for a positive species–area relation. This relation was not caused by higher habitat heterogeneity in larger patches. Patch isolation, measured in different ways, however seemed much more important in explaining species richness and community composition than fragment area. Our results indicate that area effects are overcompensated by the rescue‐effect: if a patch is close to other patches, species can disperse between them and prevent the species from going extinct. Species having a short living seed bank were also more sensitive to isolation than species with a long living seed bank, indicating that the latter most probably depend on the seed bank to survive periods when environmental conditions are harsh. Analogously to the spatial rescue‐effect, the existence of a persistent seed bank may lead to a sort of temporal rescue‐effect, where the extinction of a plant species is prevented through survival in the seed bank of a patch.