Decomposing the process of species accumulation into area dependent and time dependent parts

In 1960, Preston predicted that the process of species accumulation in time (species–time relationship, STR) should be similar to the species–area relationship (SAR) and follow a power function with a slope of about 0.26. Here these two conjectures are tested using data of the spatiotemporal species accumulation in a local community of beech forest Hymenoptera. A power function species–area–time model of the form S = S₀ A^z t gave better fits to observed species numbers than a simple power function SAR model, and was able to predict similar species turnover rates (about 9% per year) to those inferred by other methods. The STR was well fitted by a power function, although due the limited time span (8 years) a logarithmic STR pattern cannot be ruled out. STR slopes ranged between 0.01 and 0.23 and were lower than predicted by Preston. Temporal species turnover appeared to be negatively correlated to species densities and positively correlated to species body weights. Ecological guild and taxon membership did not significantly influence temporal species turnover.     

Allometric ecological distributions in a local community of Hymenoptera

The present paper describes basic ecological distributions in a community of beech forest Hymenoptera. It shows that the species diversity–body weight and the density–body weight distributions give rise to a new distribution that relates total community biomass to species diversity. For Hymenoptera this distribution follows a power function with a slope of 1.3. Combining this relation with the species–area and the individuals–area relations resulted in two other distributions that relate community biomass to area and individual numbers. It appeared that population densities decrease when computed over larger areas. The biomass–species diversity relation offers a new and simple way to estimate total community biomass from samples. The possible implications of this distribution to the productivity–diversity debate are discussed.     

Species turnover at small scales in dune slack plant communities

Patterns of both species accumulation with increasing area and of individual species occurrences depend on the scale level considered. This study investigated community diversity and individual species turnover patterns between four scale levels within 2×2 m² nested plots situated in a dune slack plant community. The number of species increased with plot area following a log–log function, with a slope of 0.23. However, species turnover was higher between the lowest scale levels, indicating limitations on species occurrences at the 25×25 cm² scale level. Alpha diversity in rectangular plots was significantly higher than in square plots of the same area. There were strong differences between individual species turnover patterns. Most species occurrence patterns had a box-counting fractal dimension value between 0.8 and 1.6, which is rather low compared with other studies on larger scale levels. Analyses of occurrence probabilities and scale area plots showed that there is a systematic deviation from self-similarity at the smallest scale level. Species had a lower frequency than expected from a fractal distribution, suggesting a higher level of species aggregation. The higher species diversity turnover at the smallest scale level can be linked to a higher spatial aggregation of individual species, due to biotic or abiotic limitations on their occurrence. These results confirm the general nature of the pattern of break-down of self-similarity at the smallest scale level considered.

Zusammenfassung

Sowohl das Muster des Artenanstiegs mit zunehmender Fläche als auch das Muster des Auftretens einzelner Arten hängen vom betrachteten Skalenlevel ab. Diese Studie untersuchte die Diversität der Lebensgemeinschaft und die Muster der Fluktuationen einzelner Arten auf vier Skalenlevels innerhalb von 2×2 m² ineinander geschachtelten Versuchsflächen in einer Pflanzengemeinschaft der Dünentäler. Die Zahl der Arten nahm mit der Versuchsfläche entsprechend einer log–log Funktion mit einer Steigung von 0.23 zu. Die Artenfluktuation zwischen den niedrigsten Skalenlevels war jedoch größer und weist darauf hin, dass es Limitierungen für das Auftreten der Arten auf dem 25×25 cm² Skalenlevel gibt. Die Alpha-Diversität war in rechteckigen Versuchsflächen signifikant größer als in quadratischen Versuchsflächen der gleichen Größe. Es gab größe Unterschiede in den Mustern der Fluktuation einzelner Arten. Die meisten Muster des Auftretens der Arten hatten fraktale Box-Counting-Dimensions-Werte zwischen 0.8 und 1.6, was relativ gering im Vergleich zu Studien auf größeren Skalenlevels ist. Die Analysen der Auftretenswahrscheinlichkeit und der Probefläche der Skalenlevels zeigten, dass es eine systematische Abweichung von der Selbstähnlichkeit auf dem kleinsten Skalenlevel gibt. Die Arten hatten eine geringere Häufigkeit als die aufgrund einer fraktalen Verteilung erwartete, was einen höheren Level der Artaggregation vermuten lässt. Die größere Fluktuation der Artendiversität auf dem kleinsten Skalenlevel kann mit einer größeren räumlichen Aggregation einzelner Arten aufgrund von biotischen und abiotischen Beschränkungen ihres Vorkommens in Verbindung gebracht werden. Diese Ergebnisse bestätigen die generelle Natur des Musters des Zusammenbruchs der Selbstähnlichkeit auf den kleinsten betrachteten Skalenlevels.     

Detecting biodiversity hotspots using species–area and endemics–area relationships: the case of butterflies

Using data of the Red Data Book of European Butterflies we establish the species–area relationship (S = 8.5 A^0.23) and the endemics–area relationship (S = 0.5 A^0.18) of European butterflies. Applying confidence limits as tools for the identification of hotspot countries we show that in the case of butterflies hotspots of endemism and hotspots of overall species richness do not coincide. We introduce plots of residuals from species–area relationships shifted upwards by the 95% confidence limits of the intercept (α-values) as a new tool for identifying and ranking of hotspots.     

Human population, grasshopper and plant species richness in European countries

Surprisingly, several studies over large scales have reported a positive spatial correlation of people and biodiversity. This pattern has important implications for conservation and has been documented for well studied taxa such as plants, amphibians, reptiles, birds and mammals. However, it is unknown whether the pattern applies also to invertebrates other than butterflies and more work is needed to establish whether the species–people relationship is explained by both variables correlating with other environmental factors. We studied whether grasshopper species richness (Orthoptera, suborder Caelifera) is related to human population size in European countries. As expected, the number of Caelifera species increases significantly with increasing human population size. But this is not the case when controlling for country area, latitude and number of plant species. Variations in Caelifera species richness are primarily associated with variations in plant species richness. Caelifera species richness also increases with decreasing mean annual precipitation, Gross Domestic Product per capita (used as an indicator for economic development) and net fertility rate of the human population. Our analysis confirms the hypothesis that the broad-scale human population–biodiversity correlations can be explained by concurrent variations in factors other than human population size such as plant species richness, environmental productivity, or habitat heterogeneity. Nonetheless, more populated countries in Europe still have more Caelifera species than less populated countries and this poses a particular challenge for conservation.     

The theory of the nested species–area relationship: geometric foundations of biodiversity scaling

The relationship between sampled area and the number of species within that area, the species–area relationship (SAR), is a major biodiversity pattern and one of a few law‐like regularities in ecology. While the SAR for isolated units (islands or continents) is assumed to result from the dynamics of species colonization, speciation and extinction, the SAR for contiguous areas in which smaller plots are nested within larger sample areas can be attributed to spatial patterns in the distribution of individuals. The nested SAR is typically triphasic in logarithmic space, so that it increases steeply at smaller scales, decelerates at intermediate scales and increases steeply again at continental scales. I will review current theory for this pattern, showing that all three phases of the SAR can be derived from simple geometric considerations. The increase of species richness with area in logarithmic space is generally determined by overall species rarity, so that the rarer the species are on average, the higher is the local slope z. Rarity is scale‐dependent: species occupy only a minor proportion of area at broad spatial scales, leading to upward accelerating shape of the SAR at continental scales. Similarly, species are represented by only a few individuals at fine spatial scales, leading to high SAR slope also at small areas. Geometric considerations reveal links of the SAR to other macroecological patterns, namely patterns of β‐diversity, the species–abundance distribution, and the relationship between energy availability (or productivity) and species richness. Knowledge of the regularities concerning nested SARs may be used for standardizing unequal areas, upscaling species richness and estimating species loss due to area loss, but all these applications have their limits, which also follow from the geometric considerations.     

A flexible multi-scale approach for standardised recording of plant species richness patterns

A sound monitoring of appropriate biodiversity indicators is necessary in order to assess the progress towards the internationally agreed target of halting the loss of biodiversity by 2010. However, existing monitoring schemes often do not address species richness as a key component of biodiversity directly or do so with insufficient methods. I provide an overview and assessment of the large variety of different sampling approaches for small-scale plant species richness. Major shortcomings of many of these are (i) non-uniform plot sizes or shapes; (ii) analysis of only one spatial scale despite the scale dependence of nearly all biodiversity parameters; (iii) lack of replication of smaller subplots; and (iv) exclusion of bryophytes and lichens despite their often large contribution to total plant diversity. Based on this review, I propose a new standardised sampling approach for plant diversity patterns at small scales that is applicable for a multitude of purposes and in any biome. In its basic variant, species composition is recorded on nested squares of 0.01 m², 0.1 m², 1 m², 10 m², and 100 m², with all smaller subplots being replicated at least 3-fold and evenly spaced within the next larger plot. Not only terricolous vascular plants, but also bryophytes, lichens, macro-algae as well as non-terricolous taxa should be recorded with the any-part system, i.e. those plants are counted within a plot whose superficial parts reach over it. This approach can be used to assess plant diversity patterns (i) of individual plots of interest, (ii) along environmental gradients, (iii) within specific vegetation types, or (iv) for landscape sectors. In the latter case, the series of nested plots must be placed randomly or systematically, but irrespective of plot homogeneity. The proposed approach allows the calculation of many meaningful biodiversity indicators, while being well compatible with a range of other sampling schemes, but avoiding their shortcomings. As this approach is not very time-consuming in its basic variant, but can easily be extended for specific purposes, I suggest its use for any kind of biodiversity studies and particularly for monitoring.     

Speciation-rate dependence in species–area relationships

The general tendency for species number (S) to increase with sampled area (A) constitutes one of the most robust empirical laws of ecology, quantified by species–area relationships (SAR). In many ecosystems, SAR curves display a power-law dependence, S∝A^z. The exponent z is always less than one but shows significant variation in different ecosystems. We study the multitype voter model as one of the simplest models able to reproduce SAR similar to those observed in real ecosystems in terms of basic ecological processes such as birth, dispersal and speciation. Within the model, the species–area exponent z depends on the dimensionless speciation rate ν, even though the detailed dependence is still matter of controversy. We present extensive numerical simulations in a broad range of speciation rates from ν=10^-3 down to ν=10^-11, where the model reproduces values of the exponent observed in nature. In particular, we show that the inverse of the species–area exponent linearly depends on the logarithm of ν. Further, we compare the model outcomes with field data collected from previous studies, for which we separate the effect of the speciation rate from that of the different species lifespans. We find a good linear relationship between inverse exponents and logarithm of species lifespans. However, the slope sets bounds on the speciation rates that can hardly be justified on evolutionary basis, suggesting that additional effects should be taken into account to consistently interpret the observed exponents.     

sars: an R package for fitting,evaluating and comparing species–area relationship models

The species–area relationship (SAR) constitutes one of the most general ecological patterns globally. A number of different SAR models have been proposed. Recent work has shown that no single model universally provides the best fit to empirical SAR datasets: multiple models may be of practical and theoretical interest. However, there are no software packages available that a) allow users to fit the full range of published SAR models, or b) provide functions to undertake a range of additional SAR‐related analyses. To address these needs, we have developed the R package ‘sars’ that provides a wide variety of SAR‐related functionality. The package provides functions to: a) fit 20 SAR models using non‐linear and linear regression, b) calculate multi‐model averaged curves using various information criteria, and c) generate confidence intervals using bootstrapping. Plotting functions allow users to depict and scrutinize the fits of individual models and multi‐model averaged curves. The package also provides additional SAR functionality, including functions to fit, plot and evaluate the random placement model using a species–sites abundance matrix, and to fit the general dynamic model of oceanic island biogeography. The ‘sars’ R package will aid future SAR research by providing a comprehensive set of simple to use tools that enable in‐depth exploration of SARs and SAR‐related patterns. The package has been designed to allow other researchers to add new functions and models in the future and thus the package represents a resource for future SAR work that can be built on and expanded by workers in the field.     

Why larger nations have disproportionate threat rates: area increases endemism and human population size

Global Red List data on mammals, birds and plants for over 100non-island nations are used to identify the impact of area, endemism, humanpopulation, and many other social variables (urbanized population,human-dominated land, national wealth, % land protected) on proportions ofthreatened species among nations. Human population size and, especially,proportion of endemic species emerge as the strongest correlates of proportionof threatened species in nations. Area tends to increase both human populationand proportion of endemics and thus increases the proportion of threatenedspecies. Increasing wealth is associated with increased relative threat inmammals and plants. Proportion of land protected is significantly associatedwith decreased relative threat in mammals and birds.     

Habitat reduction and patterns of species loss

This paper uses data of The Distribution Atlas of Polish Butterflies to simulate the effect of four different types of area loss (aggregated, fractal, random, and nested) on species diversity and species–area relationships (SARs). We found that power function and logarithmic SAR models overestimated species loss in the case of aggregated, fractal, and random patterns of area reduction. Instead, the modification of the power function by Plotkin et al. (Proc. Natl. Acad. Sci. USA 97 (2000b) 10850) (S=S₀A^ze^−kA) with k being a shape-adjusting parameter worked better and gave sufficient predictions of species loss. The net effects of the aggregated, fractal, and random types of area loss on species diversity were very similar with an aggregated pattern of area loss leading to slightly higher rates of species loss than both other processes. We conclude that SARs might be useful tools for biodiversity forecasting if they are constructed in a case-specific manner. The use of standard models instead might be misleading.     

Aphid biodiversity is positively correlated with human population in European countries

At large spatial scales, high numbers of people tend to be located in regions rich in biodiversity. This pattern has been reported for plants, some invertebrate groups, amphibians, reptiles, birds and mammals, but little is known about whether aphids conform to it. Aphids originated from temperate regions of the boreal hemisphere and are thus exceptionally species-poor in the tropics. Here, we test whether aphid species richness is related to human population variation in European countries. The number of aphid species increases significantly with increasing human population size. This happens also when controlling for country area, latitude and plant species richness, which are not factors significantly affecting the response variable in the multivariate model. Given that the species–area and species–people relationships have a slope shallower than 1, small countries have a higher aphid species density relative to area and to people than large ones. There is no evidence that the species-people correlation for aphids in European countries arises because both variables are related to increasing temperature or precipitation. Potential mechanisms underlying the findings could thus be a sampling artefact or an influence of habitat heterogeneity. There is a need for an increase in research, public awareness and conservation of large-scale aphid biodiversity.     

Untangling ecological complexity on different scales of space and time

Ecological systems are complex and essentially unpredictable, because of the multitude of interactions among their constituents. However, there are general statistical patterns emerging on particular spatial and temporal scales, which indicate the existence of some universal principles behind many ecological phenomena, and which can even be used for the prediction of phenomena occurring on finer scales of resolution. These generalities comprise regular frequency distributions of particular macroscopic variables within higher taxa (body size, abundance, range size), relationships between such variables, and general patterns in species richness. All the patterns are closely related to each other and although there are only a few major explanatory principles, there are plenty of alternative explanations. Reconciliation of different approaches cannot be obtained without careful formulation of testable hypotheses and rigorous quantitative empirical research. Two especially promising ways of untangling ecological complexity comprise: (1) analysis of invariances, i.e. universal quantitative relationships observed within many different systems, and (2) detailed analysis of the anatomy of macroecological phenomena, i.e. explorations of how emergent multispecies patterns are related to regular patterns concerning individual species.

Zusammenfassung

Ökologische Systeme sind komplex und im Wesentlichen aufgrund der Vielzahl von Interaktionen zwischen ihren Bestandteilen nicht vorhersagbar. Dennoch gibt es allgemeine statistische Muster, die in bestimmten räumlichen und zeitlichen Skalen auftreten. Dies weist auf die Existenz von einigen universellen Prinzipien hinter diesen ökologischen Phänomenen hin, die sogar für die Vorhersage von Phänomenen genutzt werden können, die auf kleineren Skalen auftreten. Diese Allgemeingültigkeiten bestehen aus Häufigkeitsverteilungen von bestimmten makroskopischen Variablen innerhalb höherer Taxa (Körpergröße, Abundanz, Arealgröße), den Beziehungen zwischen diesen Variablen und allgemeinen Mustern des Artenreichtums. Alle Muster stehen in enger Beziehung zueinander und obwohl es nur wenige bedeutende Erklärungsprinzipien gibt, existieren viele alternative Erklärungen. Die Abstimmung zwischen verschiedenen Ansätzen kann ohne eine sorgfältige Formulierung von testbaren Hypothesen und rigorose quantitative empirische Forschung nicht erreicht werden. Zwei besonders vielversprechende Wege ökologische Komplexität zu entwirren beinhalten (1) die Analyse von Invarianten, d.h. universellen quantitativen Beziehungen, die innerhalb verschiedener Systeme beobachtet werden, und (2) detaillierte Analysen der Anatomie von makroökologischen Phänomenen, d.h. Untersuchungen darüber, in welcher Beziehung die auftauchenden Muster von Multi-Arten-Systemen zu regulären Mustern individueller Arten stehen.     

Boreal carabid-beetle (Coleoptera, Carabidae) assemblages along the clear-cut originated succession gradient

We examined the occurrence of carabid beetles along a forest successiongradient in central Finland (forest age classes: 5, 10, 20, 30 and 60years since clear-cutting). Species richness of carabids was higherin the two youngest age classes, while no clear differences were detected incarabid abundance. The high species richness in the young, open sites was due toinvasion of open-habitat species. Many forest species were absent from or scarcein the young sites and became gradually more abundant towards the older forestage classes. The catches indicated a drastic decrease and assemblage-levelchange in concert with canopy closure, i.e. 20–30 years afterclear-cutting. Some forest specialists with poor dispersal ability may facelocal extinction, if the proportion of mature forest decreases further and theremaining mature stands become more isolated. We recommend that, whileharvesting timber, connectivity between mature stands is ensured, mature standsare maintained close (a few tens of metres) to each other and the matrix qualityis improved for forest species by green tree retention.     

The relationships between the structure of paddy levees and the plant species diversity in cultural landscapes on the west side of Lake Biwa,Shiga, Japan

Paddy levees form networks of narrow linear habitats and play various roles in cultural landscapes. Traditional landscapes on the west side of Lake Biwa consist of paddy field terraces and both stone and soil levees that have been maintained by paddy field management using local resources. Paddy levees in this study site are principally classified into five different types. Our study points out how differences in paddy levee structure as well as in management practices influence the plant species. Seventeen paddy levee transects were split into four habitat types based on their species components by TWINSPAN. Spatial characteristics and physical structures of paddy levees depended on natural conditions and human activities. The species–area curves of each levee type showed a clear distinction: the soil, stone and abandoned curves were steep, while the concrete and consolidated ones were gentle. The vegetation on consolidated levees was utterly different from the vegetation on traditional levee types from the aspect of species richness and species components. Soil type levees contained various woody plant species and included more diverse and indigenous plant species than abandoned type levees.     

Effects of habitat area, isolation, and landscape diversity on plant species richness of calcareous grasslands

Calcareous grasslands harbour a high biodiversity, but are highly fragmented and endangered in central Europe. We tested the relative importance of habitat area, habitat isolation, and landscape diversity for species richness of vascular plants. Plants were recorded on 31 calcareous grasslands in the vicinity of the city of Göttingen (Germany) and were divided into habitat specialist and generalist species. We expected that habitat specialists were more affected by area and isolation, and habitat generalists more by landscape diversity. In multiple regression analysis, the species richness of habitat specialists (n = 66 species) and habitat generalists (n = 242) increased with habitat area, while habitat isolation or landscape diversity did not have significant effects. Contrary to predictions, habitat specialists were not more affected by reduced habitat area than generalists. This may have been caused by delayed extinction of long-living plant specialists in small grasslands. Additionally, non-specialists may profit more from high habitat heterogeneity in large grasslands compared to habitat specialists. Although habitat isolation and landscape diversity revealed no significant effect on local plant diversity, only an average of 54% of habitat specialists of the total species pool were found within one study site. In conclusion, habitat area was important for plant species conservation, but regional variation between habitats contributed also an important 46% of total species richness.