Regional patterns of diversity and estimates of global insect species richness

The total number of insect species in the world is an important if elusive figure. We use a fresh approach to estimate global insect species richness, based on biogeographic patterns of diversity of well or better documented taxa. Estimates generated by various calculations, all variations on a theme, largely serve to substantiate suggestions that insect species are likely to number around 10 million or less.     

Priorities for conserving global species richness and endemism

The Convention on Biological Diversity aims to encourage and enable countries to conserve biological diversity, to use its components sustainably and to share benefits equitably. Species richness and endemism are two key attributes of biodiversity that reflect the complexity and uniqueness of natural ecosystems. National data on vertebrates and higher plants indicate global concentrations of biodiversity and can assist in defining priorities for action. Projections indicate that species and ecosystems will be at maximum risk from human activities during the next few decades. Prompt action by the world community can minimise the eventual loss of species. Highest priorities should be to: (i) strengthen the management of ecosystems containing a large proportion of global biodiversity; (ii) help developing countries complete their biodiversity strategies and action plans, monitor their own biodiversity, and establish and maintain adequate national systems of conservation areas; (iii) support actions at the global level, providing benefit to all countries in managing their own biodiversity. Generally, resources will best be spent in safeguarding ecosystems and habitats that are viable and important for global biodiversity, and which are threatened by factors that can be controlled cost-effectively. Other important criteria are representativeness, complementarity and insurance.     

Patterns of species richness hotspots and estimates of their protection are sensitive to spatial resolution

Aim

Species richness is a measure of biodiversity often used in spatial conservation assessments and mapped by summing species distribution maps. Commission errors inherent those maps influence richness patterns and conservation assessments. We sought to further the understanding of the sensitivity of hotspot delineation methods and conservation assessments to commission errors, and choice of threshold for hotspot delineation.

Location

United States.

Methods

We created range maps and 30‐m and 1‐km resolution habitat maps for terrestrial vertebrates in the United States and generated species richness maps with each dataset. With the richness maps and the GAP Protected Areas Dataset, we created species richness hotspot maps and calculated the proportion of hotspots within protected areas; calculating protection under a range of thresholds for defining hotspots. Our method allowed us to identify the influence of commission errors by comparing hotspot maps.

Results

Commission errors from coarse spatial grain data and lack of porosity in the range data inflated richness estimates and altered their spatial patterns. Coincidence of hotspots from different data types was low. The 30‐m hotspots were spatially dispersed, and some were very long distances from the hotspots mapped with coarser data. Estimates of protection were low for each of the taxa. The relationship between estimates of hotspot protection and threshold choice was nonlinear and inconsistent among data types (habitat and range) and grain size (30‐m and 1‐km).

Main conclusions

Coarse mapping methods and grain sizes can introduce commission errors into species distribution data that could result in misidentifications of the regions where hotspots occur and affect estimates of hotspot protection. Hotspot conservation assessments are also sensitive to choice of threshold for hotspot delineation. There is value in developing species distribution maps with high resolution and low rates of commission error for conservation assessments.     

Climate induced increases in species richness of marine fishes

Climate change has been predicted to lead to changes in local and regional species richness through species extinctions and latitudinal ranges shifts. Here, we show that species richness of fish in the North Sea, a group of ecological and socio-economical importance, has increased over a 22-year period and that this rise is related to higher water temperatures. Over eight times more fish species displayed increased distribution ranges in the North Sea (mainly small-sized species of southerly origin) compared with those whose range decreased (primarily large and northerly species). This increase in species richness can be explained from the fact that fish species richness in general decreases with latitude. This observation confirms that the interaction between large-scale biogeographical patterns and climate change may lead to increasing species richness at temperate latitudes.     

Contribution of oribatid and mesostigmatid soil mites in ecologically based estimates of global species richness

Erwin’s method for estimating total global species richness assumes some host‐specificity among the canopy arthropods. This study examined possible host habitat specialization in two major groups of soil arthropods, the oribatid and mesostigmatid mites, by sampling beneath three tree species: Eucalyptus pilularis Smith, Eucalyptus propinqua Deane and Maiden and Allocasuarina torulosa (Aiton) L. Johnson. The sample sites were in the Lansdowne State Forest, New South Wales, Australia and the three tree species were selected on the basis of their known differential effects on soil. Sampling was conducted over three seasons, and 79 oribatid and 34 mesostigmatid species were identified from 25 196 and 3634 individuals, respectively. Tree species had little effect on mite species composition with only three oribatid species and no mesostigmatid species identified as host‐habitat specialists using a niche breadth measure. Of mite species found under E. pilularis, E. propinqua and A. torulosa trees, 2%, 1% and 0% were defined as host‐habitat specialists, respectively. In contrast, tree species had significant and consistent effects on mite community structure, which differed in relative abundance of the oribatid species, their size class distributions and species rankings. In the mesostigmatid communities, there was a difference in the ranking of the mite species among tree species. Although it was demonstrated that tree species have an impact on the soil environment, the differences between tree species were insufficient to change species composition. The low degree of host‐habitat specialization suggested that other factors were more important for determining mite species composition at a site, and soil mite host‐habitat specialization may not make a large contribution to estimates of total global species richness using methods such as those proposed by Erwin (1982) .     

Functional group dominance and not productivity drives species richness

Background: There is a lack of consensus about the productivity–richness relationship, with several recent studies suggesting that it is not productivity but other factors that are the important drivers that determine species richness.

Aims: Here, we examine the relationship between productivity, functional group dominance and plant species richness at the plot scale in Tibetan Plateau meadows. These alpine meadows are ideal to examine the species productivity-richness relationship because they have a very high species richness, a large gradient in productivity, and can be dominated by either graminoids (grasses and sedges) or forbs.

Methods: We measured plant species richness and above-ground biomass along a natural gradient of functional group abundance in 44 plots distributed across five natural, winter-grazed but otherwise undisturbed sites in the eastern part of the Qing-Hai Tibetan Plateau, in Gansu province, China in 2008.

Results: Graminoid abundance (i.e. graminoid biomass as percent of the total above-ground biomass) explained 39% of plot differences in species richness while neither productivity nor the biomass of the three most abundant plant species, either individually or combined, were a significant predictor of species richness.

Conclusions: Our results show that within these alpine meadows, a shift from graminoid to forb dominance, rather than the individual dominant species or productivity itself, is strongly correlated with species richness. Thus, differences in functional group abundance can be a strong driver of observed plant species richness patterns.     

基于物种的大尺度生物多样性热点研究方法

生物多样性热点是建立保护区、制定保护决策的依据,是生物保护研究的热点问题之一。基于物种的研究方法是大尺度陆地生物多样性热点的主要研究方法,但数据的缺乏限制了直接根据物种丰富度确定热点的方法,因此研究中经常采用其他方法间接的反映物种情况,介绍了4种主要的基于物种的替代方法:指示种、高级分类单元、环境模型和景观异质性,详细阐述了各种方法存在的利弊,并从数据的可获取性、操作的便捷性和对物种特征的反映3个方面对各种方法进行了评价。任何单一的方法都无法准确反映出生物多样性热点的真实分布。合适的研究方法是权衡研究目的、时间和资金的结果,建议选择优势互补的多种方法。     

Trait-based species richness: ecology and macroevolution

Understanding the origins of species richness patterns is a fundamental goal in ecology and evolutionary biology. Much research has focused on explaining two kinds of species richness patterns: (i) spatial species richness patterns (e.g. the latitudinal diversity gradient), and (ii) clade-based species richness patterns (e.g. the predominance of angiosperm species among plants). Here, I highlight a third kind of richness pattern: trait-based species richness (e.g. the number of species with each state of a character, such as diet or body size). Trait-based richness patterns are relevant to many topics in ecology and evolution, from ecosystem function to adaptive radiation to the paradox of sex. Although many studies have described particular trait-based richness patterns, the origins of these patterns remain far less understood, and trait-based richness has not been emphasised as a general category of richness patterns. Here, I describe a conceptual framework for how trait-based richness patterns arise compared to other richness patterns. A systematic review suggests that trait-based richness patterns are most often explained by when each state originates within a group (i.e. older states generally have higher richness), and not by differences in transition rates among states or faster diversification of species with certain states. This latter result contrasts with the widespread emphasis on diversification rates in species-richness research. I show that many recent studies of spatial richness patterns are actually studies of trait-based richness patterns, potentially confounding the causes of these patterns. Finally, I describe a plethora of unanswered questions related to trait-based richness patterns.     

Coherence of terrestrial vertebrate species richness with external drivers across scales and taxonomic groups

Aim

Understanding connections between environment and biodiversity is crucial for conservation, identifying causes of ecosystem stress, and predicting population responses to changing environments. Explaining biodiversity requires an understanding of how species richness and environment covary across scales. Here, we identify scales and locations at which biodiversity is generated and correlates with environment.

Location

Full latitudinal range per continent.

Time Period

Present day.

Major Taxa Studied

Terrestrial vertebrates: all mammals, carnivorans, bats, songbirds, hummingbirds, amphibians.

Methods

We describe the use of wavelet power spectra, cross-power and coherence for identifying scale-dependent trends across Earth's surface. Spectra reveal scale- and location-dependent coherence between species richness and topography (E), mean annual precipitation (Pn), temperature (Tm) and annual temperature range (ΔT).

Results

>97% of species richness of taxa studied is generated at large scales, that is, wavelengths $\gtrsim {10}^3$ km, with 30%–69% generated at scales $\gtrsim {10}^4$ km. At these scales, richness tends to be highly coherent and anti-correlated with E and ΔT, and positively correlated with Pn and Tm. Coherence between carnivoran richness and ΔT is low across scales, implying insensitivity to seasonal temperature variations. Conversely, amphibian richness is strongly anti-correlated with ΔT at large scales. At scales $\lesssim {10}^3$ km, examined taxa, except carnivorans, show highest richness within the tropics. Terrestrial plateaux exhibit high coherence between carnivorans and E at scales $\sim {10}^3$ km, consistent with contribution of large-scale tectonic processes to biodiversity. Results are similar across different continents and for global latitudinal averages. Spectral admittance permits derivation of rules-of-thumb relating long-wavelength environmental and species richness trends.

Main Conclusions

Sensitivities of mammal, bird and amphibian populations to environment are highly scale dependent. At large scales, carnivoran richness is largely independent of temperature and precipitation, whereas amphibian richness correlates strongly with precipitation and temperature, and anti-correlates with temperature range. These results pave the way for spectral-based calibration of models that predict biodiversity response to climate change scenarios.     

Plot size matters: Toward comparable species richness estimates across plot‐based inventories

To understand the state and trends in biodiversity beyond the scope of monitoring programs, biodiversity indicators must be comparable across inventories. Species richness (SR) is one of the most widely used biodiversity indicators. However, as SR increases with the size of the area sampled, inventories using different plot sizes are hardly comparable. This study aims at producing a methodological framework that enables SR comparisons across plot‐based inventories with differing plot sizes. We used National Forest Inventory (NFI) data from Norway, Slovakia, Spain, and Switzerland to build sample‐based rarefaction curves by randomly incrementally aggregating plots, representing the relationship between SR and sampled area. As aggregated plots can be far apart and subject to different environmental conditions, we estimated the amount of environmental heterogeneity (EH) introduced in the aggregation process. By correcting for this EH, we produced adjusted rarefaction curves mimicking the sampling of environmentally homogeneous forest stands, thus reducing the effect of plot size and enabling reliable SR comparisons between inventories. Models were built using the Conway–Maxell–Poisson distribution to account for the underdispersed SR data. Our method successfully corrected for the EH introduced during the aggregation process in all countries, with better performances in Norway and Switzerland. We further found that SR comparisons across countries based on the country‐specific NFI plot sizes are misleading, and that our approach offers an opportunity to harmonize pan‐European SR monitoring. Our method provides reliable and comparable SR estimates for inventories that use different plot sizes. Our approach can be applied to any plot‐based inventory and count data other than SR, thus allowing a more comprehensive assessment of biodiversity across various scales and ecosystems.     

The role of fire in terrestrial vertebrate richness patterns

Productivity is strongly associated with terrestrial species richness patterns, although the mechanisms underpinning such patterns have long been debated. Despite considerable consumption of primary productivity by fire, its influence on global diversity has received relatively little study. Here we examine the sensitivity of terrestrial vertebrate biodiversity (amphibians, birds and mammals) to fire, while accounting for other drivers. We analyse global data on terrestrial vertebrate richness, net primary productivity, fire occurrence (fraction of productivity consumed) and additional influences unrelated to productivity (i.e., historical phylogenetic and area effects) on species richness. For birds, fire is associated with higher diversity, rivalling the effects of productivity on richness, and for mammals, fire's positive association with diversity is even stronger than productivity; for amphibians, in contrast, there are few clear associations. Our findings suggest an underappreciated role for fire in the generation of animal species richness and the conservation of global biodiversity.     

An assessment of animal species diversity in continental waters

There is a need for monitoring the status and trends of freshwater biodiversity in order to quantify the impacts of human actions on freshwater systems and to improve freshwater biodiversity conservation. Current projects carrying assessment of freshwater biodiversity focus mainly on leading-better-known groups such as fish, or identify keystone species and/or endemic freshwater systems for conservation purposes. Our purpose is to complete these existing projects by providing quantitative estimates of species number for all freshwater groups on each continent and/or major eco-regions. This article present the results of the first implementation phase carried out from September 2002 to June 2003 and which addressed only freshwater animal species. The project consisted of: (1) compiling existing data from literature, web sites and museum collections; (2) contacting scientific experts of each group to provide a ‘to the best of their knowledge, estimates of species numbers. In this study, we consider as true freshwater species, those that complete part or all of their life cycle in freshwater, and water-dependant species those that need freshwater for food or that permanently use freshwater habitats. The current order of magnitude for known freshwater animal species world wide is 100 000, of which half are insects. Among other groups, there are some 20 000 vertebrate species; 10 000 crustacean species and 5000 mollusc species that are either true freshwater or water-dependant species. The study highlighted gaps in the basic knowledge of species richness at continental and global scales: (1) Some groups such as Protozoa, nematodes or annelids have been less studied and data on their diversity and distribution is scarce. Because current richness estimates for these groups are greatly biased by knowledge availability, we can expect that real species numbers might be much higher. (2) Continents are not equal in the face of scientific studies: South America and Asia are especially lacking global estimates of species richness for many groups, even for some usually well-known ones such as molluscs or insects. The second phase of the project will address freshwater plants and algae. The present status should be considered as a first sketch of the global picture of freshwater biodiversity. We hope that this project will initiate interactive exchange of data to complete and update this first assessment.     

Land snail species richness and diversity in Idanre hills,Ondo State,Nigeria

The land snail community of Idanre hills was studied using a combination of direct search and leaf litter‐sieving techniques. In total, 36 species and 2192 individuals in nine molluscan families were collected from 19 plots of 400 m² each. Species richness varied from 8 to 23 and the number of individuals from 21 to 566 per plot. Species richness was dominated by the carnivorous Streptaxidae, while numerical abundance was dominated by the Subulinidae, Streptaxidae and Urocyclidae, contributing to more than 95% of the total number of individuals. The single most abundant species was the urocyclid Trochozonites talcosus, contributing to almost 20% of the total number of individuals. The species richness and high abundance of land snails make Idanre hills a unique site for molluscan conservation in Nigeria.     

Trophic structure, species richness and biodiversity in Danish lakes: changes along a phosphorus gradient

1. Using data from 71, mainly shallow (an average mean depth of 3 m), Danish lakes with contrasting total phosphorus concentrations (summer mean 0.02–1.0 mg P L?l), we describe how species richness, biodiversity and trophic structure change along a total phosphorus (TP) gradient divided into five TP classes (class 1–5: <0.05, 0.05–0.1, 0.1–0.2, 0.2–0.4,> 0.4 mg P L?1).
2. With increasing TP, a significant decline was observed in the species richness of zooplankton and submerged macrophytes, while for fish, phytoplankton and floating‐leaved macrophytes, species richness was unimodally related to TP, all peaking at 0.1–0.4 mg P L?1. The Shannon–Wiener and the Hurlbert probability of inter‐specific encounter (PIE) diversity indices showed significant unimodal relationships to TP for zooplankton, phytoplankton and fish. Mean depth also contributed positively to the relationship for rotifers, phytoplankton and fish.
3. At low nutrient concentrations, piscivorous fish (particularly perch, Perca fluviatilis) were abundant and the biomass ratio of piscivores to plankti‐benthivorous cyprinids was high and the density of cyprinids low. Concurrently, the zooplankton was dominated by large‐bodied forms and the biomass ratio of zooplankton to phytoplankton and the calculated grazing pressure on phytoplankton were high. Phytoplankton biomass was low and submerged macrophyte abundance high.
4. With increasing TP, a major shift occurred in trophic structure. Catches of cyprinids in multiple mesh size gill nets increased 10‐fold from class 1 to class 5 and the weight ratio of piscivores to planktivores decreased from 0.6 in class 1 to 0.10–0.15 in classes 3–5. In addition, the mean body weight of dominant cyprinids (roach, Rutilus rutilus, and bream, Abramis brama) decreased two–threefold. Simultaneously, small cladocerans gradually became more important, and among copepods, a shift occurred from calanoid to cyclopoids. Mean body weight of cladocerans decreased from 5.1 μg in class 1 to 1.5 μg in class 5, and the biomass ratio of zooplankton to phytoplankton from 0.46 in class 1 to 0.08–0.15 in classes 3–5. Conversely, phytoplankton biomass and chlorophyll a increased 15‐fold from class 1 to 5 and submerged macrophytes disappeared from most lakes.
5. The suggestion that fish have a significant structuring role in eutrophic lakes is supported by data from three lakes in which major changes in the abundance of planktivorous fish occurred following fish kill or fish manipulation. In these lakes, studied for 8 years, a reduction in planktivores resulted in a major increase in cladoceran mean size and in the biomass ratio of zooplankton to phytoplankton, while chlorophyll a declined substantially. In comparison, no significant changes were observed in 33 ‘control’ lakes studied during the same period.     

An exploration of factors influencing lotic insect species richness

An understanding of factors that influence species richness of lotic insects is generally lacking. We present comparative data on aquatic insect species richness from several North American and other streams. Factors such as large sample numbers and drainage area (species area relationships) are not significant predictors of species richness across the streams we examined. We explore several hypotheses regarding the origins and maintenance of species richness using Upper Three Runs Creek (UTR), South Carolina, USA, as a reference stream. UTR has the highest species richness of any stream in the Western Hemisphere. Hypotheses examined included historical, regional and local processes such as: (1) Evolutionary time, (2) disturbance regime/environmental variability, (3) temperature/evolutionary-speed, (4) productivity, and (5) habitat heterogeneity. Of these hypotheses, we suggest that productivity and habitat heterogeneity appear to contribute most to the high species richness found in UTR. We believe that multidisciplinary analysis of other streams is necessary because without this crucial information our knowledge of, and desire to protect biodiversity in streams will be wanting.     

Assessing fern diversity: relative species richness and its environmental correlates in Uganda

Techniques for the rapid quantification of tropical biodiversity are of critical importance in deciding where to invest scarce conservation resources. Here we describe a simple survey method for assessing species-level richness of a poorly known plant group, the pteridophytes. We then illustrate the use of a powerful, rarefaction- based technique of controlling for inevitable differences in sampling effort to calculate the relative species richness of our study sites. Lastly, we explore how closely observed patterns of relative species richness of Ugandan forests are correlated with a suite of simple environmental variables. We find that fully 75% of the variance in our estimate of fern diversity can be predicted from just two measures: soil fertility (scored as C/N ratio, itself related to rainfall); and distance from the nearest putative Pleistocene refugium.     

Different evolutionary histories underlie congruent species richness gradients of birds and mammals

Aim The global species richness patterns of birds and mammals are strongly congruent. This could reflect similar evolutionary responses to the Earth’s history, shared responses to current climatic conditions, or both. We compare the geographical and phylogenetic structures of both richness gradients to evaluate these possibilities. Location Global. Methods Gridded bird and mammal distribution databases were used to compare their species richness gradients with the current environment. Phylogenetic trees (resolved to family for birds and to species for mammals) were used to examine underlying phylogenetic structures. Our first prediction is that both groups have responded to the same climatic gradients. Our phylogenetic predictions include: (1) that both groups have similar geographical patterns of mean root distance, a measure of the level of the evolutionary development of faunas, and, more directly, (2) that richness patterns of basal and derived clades will differ, with richness peaking in the tropics for basal clades and in the extra‐tropics for derived clades, and that this difference will hold for both birds and mammals. We also explore whether alternative taxonomic treatments for mammals can generate patterns matching those of birds. Results Both richness gradients are associated with the same current environmental gradients. In contrast, neither of our evolutionary predictions is met: the gradients have different phylogenetic structures, and the richness of birds in the lowland tropics is dominated by many basal species from many basal groups, whereas mammal richness is attributable to many species from both few basal groups and many derived groups. Phylogenetic incongruence is robust to taxonomic delineations for mammals. Main conclusions Contemporary climate can force multiple groups into similar diversity patterns even when evolutionary trajectories differ. Thus, as widely appreciated, our understanding of biodiversity must consider responses to both past and present climates, and our results are consistent with predictions that future climate change will cause major, correlated changes in patterns of diversity across multiple groups irrespective of their evolutionary histories.