Groundwater copepods: diversity patterns over ecological and evolutionary scales

Copepods are common components of the groundwater fauna, and greatly increase the diversity of groundwater communities. With more than 900 species/subspecies known from continental groundwaters, stygobiont copepods inhabit all kinds of aquifers (karstic, fissured, porous), as well as surface/subsurface ecotones (land/water and water/water). The polyhedral and varied structure of the stygohabitats is reflected in the surprising mixture of functional morphologies and habitat exploitations experienced by groundwater copepods. Morphological adaptations and specializations are discussed, as well as the chronology of their appearance in the evolutionary history of several taxa. Diversity patterns of copepod assemblages in groundwater are examined under both structural and functional profiles, as well as across a range of scales. Structure and function operate in an interactive, sometimes hierarchical ways, as well as scales. On the ecological scale, local heterogeneity and patchiness in geomorphic and hydrologic characteristics, as well as biotic interactions, are to be considered causal factors affecting the diversity patterns over a range of spatial and temporal scales. On the evolutionary scale, it is widely accepted that stygobiont copepods evolved from ancestors living in marine, freshwater and semiterrestrial environments. They gained access to the groundwater through major highways represented by the interstitial and the crevicular/karstic corridors. `Phylogenetic diversity' in groundwater copepod taxocoenoses is viewed as a heterogeneous assemblage of species belonging to different phylogenetic lineages, which entered groundwater at different times and by different ways.     

Pairwise versus presence–absence approaches for analysing biodiversity patterns

It has been proposed that the study of co‐occurrence of species, which is traditionally performed using full presence–absence matrices of sets of many species, could benefit from simply testing for random co‐occurrence between pairs of species, and that use of a full presence–absence matrix is tantamount to regarding it as having some real ecological identity. Here I argue that although there are valid questions that can be answered using a pairwise approach, there are many others that naturally require the analysis of entire sets of species in a joint way, as provided for through the use of full presence–absence matrices. Moreover, there are theoretical and mathematical advantages to the use of presence–absence matrices, a few of which are briefly discussed in this short note.     

Regional patterns of biodiversity in New Guinea plants

Regional patterns of biodiversity in seven recently-studied, speciose groups of New Guinea plants (comprising 200 species, or 1–2% of the flora) are analysed with maps showing numbers of species in 1^o grid cells. Patterns are correlated with the tectonic history of New Guinea. The New Guinea orogen involved rocks of the northern margin of the Australian craton as well as the terranes accreted to the margin, and the current axial range is geologically and biologically composite. The southern Nothofagus has a main massing on the Australian craton portion of the New Guinea mountains. In contrast, four typical genera of Malesian rainforest (Parsonsia, Archidendron, Aglaia, Amyema) have centres of biodiversity on the accreted terranes north of the craton. There are 32 distinct tectono-stratigraphic terranes (some composite) which have been accreted to the craton at different times through the Tertiary and these may have travelled hundreds or even thousands of kilometres before docking. Finally, the 'decaisninoid group' of Loranthaceae and the fern Grammitis have centres of diversity on both the craton and the accreted terranes.     

Assessing and conserving groundwater biodiversity: synthesis and perspectives

1. This paper is a synthesis of a special issue on groundwater biodiversity with a focus on obligate subterranean species, the stygobionts. The series of papers constitutes a great leap forward in assessing and understanding biodiversity patterns because of the use of large quantitative data sets obtained over a broad geographic scale. They also represent a conceptual shift, away from a purely taxonomic and phylogenetic focus to the analysis of whole groundwater assemblages.
2. The general patterns emerging for groundwater fauna are: very high levels of endemism, low local diversity relative to regional diversity, a limited number of lineages, occurrence of many relicts, and truncated food webs with very few predators.
3. β-Diversity is at least as important as α-diversity in determining total richness at different scales (aquifer, basin and region) and overall taxa richness increases across spatial scales.
4. Advances in understanding groundwater biodiversity patterns further include identification of several important factors related to geology and hydrology that determine the composition of European stygobiotic assemblages.
5. Important challenges for future research include improving sampling strategies, filling gaps in sampling coverage, intensifying research on theoretical and statistical models, and including functional and genetic diversity components in biodiversity assessments.
6. Strategies are proposed for protecting groundwater biodiversity and an argument is made to integrate biodiversity in groundwater management. Applying principles such as complementarity and flexibility for groundwater biodiversity conservation is a major step toward delineating a reserve network that maximise species representation at the European scale.     

Biodiversity indicators in European ground waters: towards a predictive model of stygobiotic species richness

1. Estimates of species richness obtained from exhaustive field inventories over large spatial scales are expensive and time-consuming. For this reason, efficiency demands the use of indicators as 'surrogates' of species richness. Biodiversity indicators are defined herein as a limited suite of taxonomic groups the species richness of which is correlated with the species richness of all other taxonomic groups present in the survey area.
2. Species richness in ground water was assessed at different spatial scales using data collected from six regions in Europe. In total, 375 stygobiotic species were recorded across 1157 sites and 96 aquifers. The taxonomic groups collected from more than one site and with more than two species (Oligochaeta, Gastropoda, Cyclopoida, Harpacticoida, Ostracoda, Isopoda, Amphipoda, Bathynellacea and Acari) were used to develop nonparametric models to predict stygobiotic biodiversity at the aquifer scale.
3. Pair-wise correlations between taxonomic groups were low, i.e. variation in species richness of a single taxonomic group did not usually reflect variation of the other groups. In contrast, multiple regressions calculated between species richness of any combination of taxa and extra-group species richness along the six regions resulted in a number of significant relationships.
4. These results suggest that some taxonomic groups (mainly Copepoda and Amphipoda and, to a lesser extent, Oligochaeta and Gastropoda) combined in different ways across the regions, were good biodiversity indicators in European groundwater ecosystems. However, the uneven distribution of taxonomic groups prevented selection of a common set of indicators for all six regions. Faunal differences among regions are presumably related to both historical and ecological factors, including palaeogeography, palaeoecology, geology, aquifer fragmentation and isolation, and, less clearly, anthropogenic disturbance.     

Global patterns in peatmoss biodiversity

DNA sequence data from the nuclear ribosomal internal transcribed spacers (ITS) and the trnL-trnF chloroplast DNA regions were used to quantify geographical partitioning of global biodiversity in peatmosses (Sphagnum), and to compare patterns of molecular diversity with patterns of species richness. Molecular diversity was estimated for boreal, tropical, Neotropical, nonboreal (tropical plus Southern Hemisphere), Old World and New World partitions, based on a total of 436 accessions. Diversity was partitioned among geographical regions in terms of combined nuclear and chloroplast sequence data and separately for the ITS and trnL-trnF data sets. Levels of variation were estimated using phylogenetic diversity (PD), which incorporates branch lengths from a phylogenetic tree, and the number of polymorphic nucleotide sites. Estimates of species richness suggest that peatmoss diversity is higher in New World than Old World regions, and that the Neotropics constitute a "hotspot" of diversity. Molecular estimates, in contrast, indicate that peatmoss biodiversity is almost evenly divided between New and Old World regions, and that the Neotropics account for only 20-35% of global peatmoss diversity. In general, levels of tropical and boreal peatmoss molecular diversity were comparable. Two species, S. sericeum from the Old World tropics and S. lapazense from Bolivia, are remarkably divergent in nucleotide sequences from all other Sphagna and together account for almost 20% of all peatmoss diversity, although they are represented by only three of the 436 accessions (0.7%). These species clearly demonstrate the nonequivalence of species biodiversity value.     

The role of ecological knowledge in explaining biogeography and biodiversity in Amazonia

Biogeographical studies in Amazonia have commonly taken a historical, rather than an ecological approach. General patterns have been sought in the distribution maps of different species, and these have been explained in terms of past or present distribution barriers, especially past climates and large rivers. Implicitly, and often also explicitly, it is assumed that Amazonia is ecologically so uniform that present-day ecological conditions are rather insignificant in determining species distribution patterns and speciation. However, this assumption is more based on the lack of relevant data than on actual observations of environmental uniformity or ecological unspecialization of the species. Recent studies have indeed documented ecological heterogeneity and floristic differences among sites that were previously thought similar. In the absence of direct knowledge of the past, more complete ecological and environmental understanding of the present-day Amazonia are needed for evaluating the relative roles of historical and ecological factors in Amazonian biogeography and biodiversity.     

Historical arthropod diversity patterns direct rehabilitation targets for Robben Island,South Africa,a continental island in a biodiversity hotspot

Island species are susceptible to extinction through disturbances such as habitat transformation. Due to the small size and isolation of islands, species have limited options for refuges and recolonization, making their rehabilitation a conservation priority. Robben Island is a continental island, isolated from the mainland ca. 15 000 years ago, and has been degraded by humans and alien species for nearly 400 years. Mainland areas with similar vegetation should be good reference sites for the biological restoration of the island due to historical connectedness. However, very little information exists as to which species were lost. Here we aim to identify the best mainland sites to use as reference sites for Robben Island based on remaining arthropod diversity on the island. Sites found to be most similar in terms of arthropod diversity to Robben Island were sites north of Robben Island (Elandsbaai and Dwarskersbos) rather than the geographically closest locations. These sites therefore represent ideal reference sites for biological restoration of the island. We do not suggest the reintroduction of species from these localities, but rather Robben Island should be restored to match their vegetation height and cover.     

The distribution patterns of reptiles in the Riff region, northern Morocco

A biogeographical classification of reptiles in the Riff region (northern Morocco, Africa) was carried out to look for shared distribution patterns, here termed chorotypes. Baroni‐Urbani & Buser's similarity index was applied to the presence/absence data of reptiles in 10×10 km UTM (Universal Transverse Mercator) squares, and then UPGMA (Unweighted Pair‐Group Method using arithmetic average) was used to classify the species. A probabilistic method was employed to assess the statistical significance of the groups obtained. An ordination method, the Canonical Correspondence Analysis, was also used to study the distribution of the reptiles within a continuous framework.
A gradual longitudinal replacement of reptile species was found throughout the Riff, with no sharp discontinuities for the distributions of most of the species. This may be due to the biogeographical northward movement of the Saharan boundaries, which have not yet reached biogeographical equilibrium. Thus, Saharan reptiles enter the Riff region from the east, through the lower basin of the River Moulouya.
Seven reptile chorotypes were identified in the Riff, and these comprise Mediterranean species and others endemic to the Maghreb (the region that spans most of North‐western Africa, excluding the Sahara). These chorotypes have a western distribution, and are segregated from one another according to altitude. Historical and ecological processes can account for the distributions shared by these species, which have inhabited the Riff for longer than eastern reptiles.     

Old and new challenges in using species diversity for assessing biodiversity

Although the maintenance of diversity of living systems is critical for ecosystem functioning, the accelerating pace of global change is threatening its preservation. Standardized methods for biodiversity assessment and monitoring are needed. Species diversity is one of the most widely adopted metrics for assessing patterns and processes of biodiversity, at both ecological and biogeographic scales. However, those perspectives differ because of the types of data that can be feasibly collected, resulting in differences in the questions that can be addressed. Despite a theoretical consensus on diversity metrics, standardized methods for its measurement are lacking, especially at the scales needed to monitor biodiversity for conservation and management purposes. We review the conceptual framework for species diversity, examine common metrics, and explore their use for biodiversity conservation and management. Key differences in diversity measures at ecological and biogeographic scales are the completeness of species lists and the ability to include information on species abundances. We analyse the major pitfalls and problems with quantitative measurement of species diversity, look at the use of weighting measures by phylogenetic distance, discuss potential solutions and propose a research agenda to solve the major existing problems.     

Present patterns and future prospects for biodiversity in the Western Hemisphere

Creating networks of nature reserves to protect areas rich in biodiversity from the adverse impacts of anthropogenic change is a critical and urgent task. We illustrate the skewed geographical and size distributions of protected areas in the Western Hemisphere. For instance, 811 of 1413 reserves in the Western Hemisphere are smaller than 10 km², and 35% of the total area of these reserves is in Alaska. We compile ranges for all bats in the continental Western Hemisphere and find that 82% of threatened and small‐range species are not protected adequately. Many of the most vulnerable species occur in the areas of highest human density. We provide maps delineating areas where conservation investments may have the greatest impact in preventing biodiversity losses.     

The mud mound nature of the Cassian Platform Margins of the Dolomites A case history: the Cipit boulders from Punta Grohmann (Sasso Piatto Massif, northern Italy)

Summary The sedimentological features and the microbiofacies of the Cassian platforms (Late Ladinian-Carnian) of the Dolomites can be studied only on the basis of the socalled “Cipit boulders”, that are platform-derived olistoliths and clasts fed to the basin and escaped to the extensive dolomitization affecting the buildups. Our paper deals with the Cipit boulders occurring in the Punta Grohmann section (Wengen and S. Cassiano formations, Late Ladinian, Archelaus and Regoledanus Zones). The dominant microfacies are represented by boundstone, consisting of nearly 60% of micritic limestone occurring both as peloidal or aphanitic micrite, mostly organized into stromatolitic laminites of thrombolites. The skeletal organism (Tubiphytes, skeletal cyanobacteria, sphinctozoan sponges, etc.) represent only a minor component of the rock (usually less than 10%). Early cements are widespread and consist both of fan-shaped calcite (replacing former aragonite), bladed isopachous magnesian calcite and radial-fibrous calcite (neomorphic after Mg-calcite). The carbonate platforms from which the olistoliths derive were made up mainly of carbonate mud that underwent early lithification, as witnessed by the considerable amount of early cements: therefore they may be regarded to as mudmounds, and more precisely as microbial mud-mounds, due to the clearly accretionary, organic-controlled nature of most micrites. The micrites, subdivided into auto- and allomicrite on the basis of micromorphological and fabric characteristics, have been tested for epifluorescence. The results confirm the organic control on the deposition of automicrite, also in the cases in which a microbial influence is not obvious (i.e. aphanitic micrite without internal organization).     

Origins of marine patterns of biodiversity: some correlates and applications

Abstract: Marine shelf diversity patterns correlate with macroecological features of basic importance that may play causal roles in macroevolution. We have investigated the global diversity pattern of living Bivalvia, which is dominated by the latitudinal diversity gradient (LDG), maintained by high tropical origination rates. Generic‐level lineages expand poleward, chiefly through speciation, so that species richness within provinces and globally is positively correlated with generic geographical ranges. A gradient in diversity accommodation progressively lowers both immigration and speciation rates in higher latitudes. The LDG correlates with seasonality of trophic resources but not with area; tropical provinces are not diverse because they are large but because they are tropical. A similar dynamic evidently underlays Jurassic and Carboniferous LDGs. Larval developmental modes correlate with the LDG and thus with resource seasonality, with tropical dominance of planktotrophs offset by increasing nonplanktotrophy to poleward. The acquisition of planktotrophy in several early Palaeozoic clades indicates a change in macroecological relationships during Cambrian and Ordovician radiations.