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.     

Marine macroalgal biodiversity hotspots: why is there high species richness and endemism in southern Australian marine benthic flora?

The southern Australian marine macroalgal flora has the highest levels of species richness and endemism of any regional macroalgal flora in the world. Analyses of species composition and distributions for the southern Australian flora have identified four different floristic elements, namely the southern Australian endemic element, the widely distributed temperate element, the tropical element and a cold water element. Within the southern Australian endemic element, four species distribution patterns are apparent, thought to largely result from the Jurassic to Oligocene fragmentation of East Gondwana, the subsequent migration of Tethyan ancestors from the west Australian coast and the later invasion of high latitude Pacific species. Climatic deterioration from the late Eocene to the present is thought responsible for the replacement of the previous tropical south coast flora by an endemic temperate flora which has subsequently diversified in response to fluctuating environmental conditions, abundant rocky substrata and substantial habitat heterogeneity. High levels of endemism are attributed to Australia's long isolation and maintained, as is the high species richness, by the lack of recent mass extinction events. The warm water Leeuwin Current has had profound influence in the region since the Eocene, flowing to disperse macroalgal species onto the south coast as well as ameliorating the local environment. It is now evident that the high species richness and endemism we now observe in the southern Australian marine macroalgal flora can be attributed to a complex interaction of biogeographical, ecological and phylogenetic processes over the last 160 million years.     

High levels of cryptic species diversity uncovered in Amazonian frogs

One of the greatest challenges for biodiversity conservation is the poor understanding of species diversity. Molecular methods have dramatically improved our ability to uncover cryptic species, but the magnitude of cryptic diversity remains unknown, particularly in diverse tropical regions such as the Amazon Basin. Uncovering cryptic diversity in amphibians is particularly pressing because amphibians are going extinct globally at an alarming rate. Here, we use an integrative analysis of two independent Amazonian frog clades, Engystomops toadlets and Hypsiboas treefrogs, to test whether species richness is underestimated and, if so, by how much. We sampled intensively in six countries with a focus in Ecuador (Engystomops: 252 individuals from 36 localities; Hypsiboas: 208 individuals from 65 localities) and combined mitochondrial DNA, nuclear DNA, morphological, and bioacoustic data to detect cryptic species. We found that in both clades, species richness was severely underestimated, with more undescribed species than described species. In Engystomops, the two currently recognized species are actually five to seven species (a 150-250% increase in species richness); in Hypsiboas, two recognized species represent six to nine species (a 200-350% increase). Our results suggest that Amazonian frog biodiversity is much more severely underestimated than previously thought.     

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.     

Exotic species richness and native species endemism increase the impact of exotic species on islands

Aim Exotic species pose one of the most significant threats to biodiversity, especially on islands. The impacts of exotic species vary in severity among islands, yet little is known about what makes some islands more susceptible than others. Here we determine which characteristics of an island influence how severely exotic species affect its native biota. Location We studied 65 islands and archipelagos from around the world, ranging from latitude 65° N to 54° S. Methods We compiled a global database of 10 island characteristics for 65 islands and determined the relative importance of each characteristic in predicting the impact of exotic species using multivariate modelling and hierarchical partitioning. We defined the impact of exotic species as the number of bird, amphibian and mammal (BAM) species listed by the International Union for Conservation of Nature (IUCN) as threatened by exotics, relative to the total number of BAM species on that island. Results We found that the impact of exotic species is more severe on islands with more exotic species and a greater proportion of native species that are endemic. Unexpectedly, the level of anthropogenic disturbance did not influence an island's susceptibility to the impacts of exotic species. Main conclusions By coupling our results with studies on the introduction and establishment of exotic species, we conclude that colonization pressure, or invasion opportunities, influences all stages of the invasion process. However, species endemism, the other important factor determining the impact of exotic species, is not known to contribute to introduction and establishment success on islands. This demonstrates that different factors correlate with the initial stages of the invasion process and the subsequent impacts of those invaders, highlighting the importance of studying the impacts of exotic species directly. Our study helps identify islands that are at risk of impact by exotics and where investment should focus on preventing further invasions.     

Collembola,the biological species concept and the underestimation of global species richness

Despite its ancient origin, global distribution and abundance in nearly all habitats, the class Collembola is comprised of only 8000 described species and is estimated to number no more than 50 000. Many morphologically defined species have broad geographical ranges that span continents, and recent molecular work has revealed high genetic diversity within species. However, the evolutionary significance of this genetic diversity is unknown. In this study, we sample five morphological species of the globally distributed genus Lepidocyrtus from 14 Panamanian sampling sites to characterize genetic diversity and test morphospecies against the biological species concept. Mitochondrial and nuclear DNA sequence data were analysed and a total of 58 molecular lineages revealed. Deep lineage diversification was recovered, with 30 molecular lineages estimated to have established more than 10 million years ago, and the origin almost all contemporary lineages preceding the onset of the Pleistocene (~2 Mya). Thirty‐four lineages were sampled in sympatry revealing unambiguous cosegregation of mitochondrial and nuclear DNA sequence variation, consistent with biological species. Species richness within the class Collembola and the geographical structure of this diversity are substantially misrepresented components of terrestrial animal biodiversity. We speculate that global species richness of Collembola could be at least an order of magnitude greater than a previous estimate of 50 000 species.     

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.     

Patterns in species richness and endemism of European freshwater fish

Aim  To analyse the patterns in species richness and endemism of the native European riverine fish fauna, in the light of the Messinian salinity crisis and the Last Glacial Maximum (LGM).
Location  European continent.
Methods  After gathering native fish faunistic lists of 406 hydrographical networks, we defined large biogeographical regions with homogenous fish fauna, based on a hierarchical cluster analysis. Then we analysed and compared the patterns in species richness and endemism among these regions, as well as species–area relationships.
Results  Among the 233 native species present in the data set, the Cyprinidae family was strongly dominant (> 50% of the total number of species). Seven biogeographical regions were defined: Western Peri-Mediterranea, Central Peri-Mediterranea, Eastern Peri-Mediterranea, Ponto-Caspian Europe, Northern Europe, Central Europe and Western Europe. The highest regional species richness was observed for Central Peri-Mediterranea and Ponto-Caspian Europe. The highest endemic richness was found in Central Peri-Mediterranea. Species–area relationships were characterized by high slope values for Peri-Mediterranean Europe and low values for Central and Western Europe.
Main conclusions  The results were in agreement with the 'Lago Mare' hypothesis explaining the specificity of Peri-Mediterranean fish fauna, as well as with the history of recolonization of Central and Western Europe from Ponto-Caspian Europe following the LGM. The results also agreed with the mechanisms of speciation and extinction influencing fish diversity in hydrographical networks. We advise the use of the seven biogeographical regions for further studies, and suggest considering Peri-Mediterranean Europe and Ponto-Caspian Europe as 'biodiversity hotspots' for European riverine fish.     

An assessment of endemism and species richness patterns in the Australian Anura

Aim To assemble a continental‐scale data set of all available anuran records and investigate trends in endemism and species richness for the Anura. Location Continental Australia. Methods 97,338 records were assembled, covering 75% of the continent. A neighbourhood analysis was applied to recorded locations for each species to measure richness and endemism for each half‐degree grid square (c. 50 km) in the continent. This analysis was performed for all anurans, and also for each of the three main anuran families found in Australia. A Monte Carlo simulation was used to test a null hypothesis that observed centres of endemism could result simply from an unstructured overlapping of species ranges of different sizes. Results Eleven main centres of anuran endemism were identified, the most important being the Wet Tropics and the south‐west near Bunbury‐Augusta and near Walpole. With the exception of south‐western Australia, all of the identified significant endemic centres are in the northern half of the continent. The regions identified as significant for endemism differed from those identified for species richness and are more localized. Species richness is greatest in the Wet Tropics and the Border Ranges. High species richness also occurs in several areas not previously identified along the east and northern coasts. Main conclusions Weighted endemism provides a new approach for determining significant areas for anuran conservation in Australia and areas can be identified that could be targeted for beneficial conservation gains. Patterns in endemism were found to vary markedly between the three main anuran families, and south‐eastern Australia was found to be far less significant than indicated by previous studies. The need for further survey work in inland Australia is highlighted and several priority areas suggested. Our results for species richness remain broadly consistent with trends previously observed for the Australian Anura.     

Patterns of species richness and turnover in endemic amphibians of the Guineo-Congolian rain forest

Aim

The African Guineo-Congolian (GC) region is a global biodiversity hotspot with high species endemism, bioclimatic heterogeneity, complex landscape features, and multiple biogeographic barriers. Bioclimatic and geographic variables influence global patterns of species richness and endemism, but their relative importance varies across taxa and regions and is poorly understood for many faunas. Here, we test the hypothesis that turnover in endemic amphibians of the GC biodiversity hotspot is influenced mainly by the geographic distance between grid cells and secondarily by rainfall- and temperature-related variables.

Location

West and Central Africa.

Major Taxa Studied

Amphibians.

Methods

We compiled species-occurrence records via field sampling, online databases, and taxonomic literature. Our study used 1205 unique georeferenced records of 222 amphibian species endemic to the GC region. Patterns of species richness were mapped onto a grid with a spatial resolution of 0.5° × 0.5°. We estimated weighted endemism and tested whether endemism was higher than the expected species richness (randomization test). We quantified species turnover using generalized dissimilarity modelling to evaluate the processes underlying observed patterns of species richness in GC endemic amphibians. We explored bioregionalization using agglomerative hierarchical clustering based on the unweighted pair group method with arithmetic averages.

Results

We identified seven areas within the lower GC region – forests in Cameroon, Gabon, Southern Nigeria, Equatorial Guinea, Republic of Congo, Democratic Republic of Congo, and Cote d'Ivoire – as having high species richness of endemic amphibians. The randomization test returned four major areas of significant weighted endemism: Nigeria-Cameroon mountains, forest regions of the Democratic Republic of Congo, Cote d'Ivoire, and Ghana. Our analysis revealed five bioregions for amphibian endemism, four of which were located within the lower Guineo-Congolian forest. Species turnover was strongly related to the geographic distance between grid cells; contributing bioclimatic variables included precipitation of the warmest quarter, mean temperature of the wettest quarter, and mean diurnal temperature range.

Main Conclusions

Our results indicate that geographic distance between grid cells is the primary determinant of turnover in GC endemic amphibians, with secondary but significant effects of rainfall- and temperature-related variables. Our study identifies key areas of endemic amphibian richness that could be prioritized for conservation actions.     

Determinants of Delphacidae richness and endemism in China

1. Identifying the macro-scale patterns and the underlying mechanisms of species richness are key aspects of biodiversity-related research. In China, previous studies on the mechanisms underlying insect richness have primarily focused on the current ecological conditions. Therefore, the impact of historical climate change on these mechanisms is less well understood. 2. Here, we use members of the Delphacidae family to evaluate the relative impact of the current environmental conditions and that of the Last Glacial Maximum on total species richness and endemism. Total species richness and endemic species richness were summed in 1° × 1° grid cells that the insects occupied. Generalised linear models, simultaneous autoregressive models, and random forest models were used to assess the effects of different environmental factors on total species richness and endemism. 3. The two patterns of species richness are jointly regulated by the current environment and the Last Glacial Maximum, but their key determinants differ. Winter coldness and the temperature annual range strongly affected the total species richness, but temperature variation during the Last Glacial Maximum also played an important role in the development of species richness. The distribution of endemic species was most strongly affected by the Last Glacial Maximum temperature change. 4. The studies confirm that historical climate change contributes to patterns of insect species richness, particularly patterns of endemism. Considering that China was mildly affected by the last glacial period, we propose that the incorporation of historical climate data into such studies will provide a better understanding of the underlying mechanisms.     

Declines in littoral species richness across both spatial and temporal nutrient gradients: a palaeolimnological study of two taxonomic groups

1. Using a palaeolimnological approach in shallow lakes, we quantified the species richness responses of diatoms and Cladocera to phosphorus enrichment. We also examined differences in species richness responses between littoral and pelagic assemblages of our focal communities. To address both spatial and temporal relationships, our study includes an analysis of both surface sediments from 40 lakes and of a lake sediment record spanning c. 120 years. The objective of our study was to determine whether similar species richness patterns occurred across trophic levels, as well as along spatial and temporal gradients. 2. We found that both diatom and Cladocera species richness estimates significantly declined with increasing phosphorus across space and through time. When the assemblages were subdivided according to known habitat preferences, littoral biodiversity maintained a negative trend, whereas pelagic species richness tended to show no relationship with phosphorus. 3. Negative productivity–diversity patterns have been observed across almost all palaeolimnological studies that span large productivity gradients. This congruence in patterns is most likely due to the similarity in data collection methods and in focal communities studied. The contrasting responses between littoral and pelagic assemblages may be explained by the differences in physical habitat and the pool of taxa in each of these environments. Consistent with the literature, we found statistical support for the idea that littoral diversity declines could be explained by an interaction between macrophytes and nutrients along strong trophic gradients. The general lack of a diversity response in our pelagic assemblages could be attributable to the limited pool of subfossil taxa. The response of the pelagic diatom could also be related to their broad range of nutrient tolerances. 4. The observed negative response of species richness to phosphorus enrichment, particularly in the littoral assemblages, has implications for ecosystems functioning because communities with reduced biodiversity often are less resilient to anthropogenic change.     

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.     

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.