On the distinctness of clusters

A common problem in classification is the assignment of objects to meaningful clusters given their relative positions in reduced ordination space. We examine the distinctiveness of such putative clusters.Clusters are enlarged by simulation and then the relative frequency distributions of interpoint distances in the “enlarged” clusters are compared with the distribution of the original points in the original space. This procedure may be repeated for all reasonable clustering hypotheses until the best fit is found.We suggest that one way of looking at the distinctness of clusters is to look at the distribution of interpoint distances for various hypothetical clusters from which the data might have been sampled. The best clustering, given the data, is the one that best matches the original distribution of interpoint distances.     

Patterns in freshwater diatom taxonomic distinctness along an eutrophication gradient

1. A variety of species richness measures have been used to assess the effects of environmental degradation on biodiversity. Such measures can be highly influenced by sample size, sampling effort, habitat type or complexity, however, and typically do not show monotonic responses to human impact. In addition to being independent of the degree of sampling effort involved in data acquisition, effective measures of biodiversity should reflect the degree of taxonomical relatedness among species within ecological assemblages and provide a basis for understanding observed diversity for a particular habitat type. Taxonomic diversity or distinctness indices emphasize the average taxonomic relatedness (i.e. degree of taxonomical closeness) between species in a community. 2. Eutrophication of freshwater ecosystems, mainly due to the increased availability of nutrients, notably phosphorus, has become a major environmental problem. Two measures of taxonomic distinctness (Average Taxonomic Distinctness and Variation in Taxonomic Distinctness) were applied to surface sediment diatoms from 45 lakes across the island of Ireland to examine whether taxonomic distinctness and nutrient enrichment were significantly related at a regional scale. The lakes span a range of concentrations of epilimnic total phosphorus (TP) and were grouped into six different types, based on depth and alkalinity levels, and three different categories according to trophic state (ultra‐oligotrophic and oligotrophic; mesotrophic; and eutrophic and hyper‐eutrophic). 3. The taxonomic distinctness measures revealed significant differences among lakes in the three different classes of trophic state, with nutrient‐rich lakes generally more taxonomically diverse than nutrient‐poor lakes. This implies that enrichment of oligotrophic lakes does not necessarily lead to a reduction in taxonomic diversity, at least as expressed by the indices used here. Furthermore, taxonomic distinctness was highly variable across the six different lake types regardless of nutrient level. 4. Results indicate that habitat availability and physical structure within the study lakes also exert a strong influence on the pattern of taxonomic diversity. Overall the results highlight problems with the use of taxonomic diversity measures for detecting impacts of freshwater eutrophication based on diatom assemblages.     

Seasonal variations in macrobenthic taxonomic diversity and the application of taxonomic distinctness indices in Bohai Bay,northern China

Taxonomic diversity indices have a number of desirable properties as indicators in the assessment of environmental quality, and have become an important measure in biodiversity studies. Macrobenthic taxonomic variations were studied in Bohai Bay, northern China, an area under threat from rapid human development. Four seasonal cruise datasets were collected between 2006 and 2007. Environmental conditions exhibited large fluctuations due to human development; nitrogen and phosphorus were the main environmental stressors. A total of 97 macrofauna taxa were identified belonging to 88 genera, 72 families, 36 orders, 14 classes, and nine phyla. Analysis of similarity indicated that there were significant assemblage differences across sampling stations as well as seasons. Four taxonomic indices, taxonomic diversity (Δ), taxonomic distinctness (Δ*), average taxonomic distinctness (Δ⁺), and variation in taxonomic distinctness (Λ⁺) were calculated using abundance data. Among the stations and seasons, there were greater variations in both Δ and Λ⁺ than in Δ* and Δ⁺. The funnel plot of Δ⁺ could identify disturbed stations to some extent, but was not always a strong indicator of disturbance. The Δ⁺ performance was better in autumn than in spring, but could not identify a disturbed station in autumn due to a low number of species. The efficiency of taxonomic distinctness may depend on taxa or the pollution indicators. Taxonomic distinctness indices can be effective at assessing environmental degradation when correctly applied; however, they are unsuitable for directly assessing environmental quality in a new area prior to efficiency testing.     

The taxonomic distinctness of coastal bottom-dwelling fish communities of the North-east Atlantic

1. New techniques for identifying the average taxonomic range of species assemblages were applied to an extensive dataset of bottom-dwelling fish in the coastal waters of NW Europe. These taxonomic distinctness indices provided much greater resolution than traditional diversity indices as they incorporated information on taxonomic relationships into an index which measures species dominance. Unlike standard measures of species richness and diversity, the mean value of these statistics is independent of sampling effort, and this allows objective comparisons to be made between samples from studies where sampling effort is not standardized.
2. A reduction in the average taxonomic range between the fauna of western waters of the UK and that of the southern North Sea was consistent with the general decline in species richness observed between these regions, and suggests that these two factors may be spatially positively correlated. Indices calculated for individual samples of fish on a local scale, however, did not all fit this trend.
3. Much of the variability in taxonomic diversity within the coastal waters of NW Europe was caused by the variable geographical distribution of the elasmobranchs. Of all the families which comprise the fish communities, this group has life-history characteristics which make it most susceptible to impact by commercial trawl fisheries.
4. The use of taxonomic distinctness measures provided additional insights, of relevance to biodiversity assessment, suggesting that they might usefully be applied to other aquatic and terrestrial fauna.     

Quantifying the effects of nutrient addition on the taxonomic distinctness of serpentine vegetation

Traditional diversity indices summarize the information about the relative abundances of species within a community without regard to differences between species. However, intuitively, a community composed of dissimilar taxa is more diverse than a community composed of more similar taxa. Therefore, useful indices of diversity should account for taxonomic relations among species. In this paper, a new parametric diversity index that combines species relative abundances and their taxonomic distinctiveness is used to quantify the way in which soil fertilization affects the diversity of a garigue community on ultramafic soils of Tuscany (central Italy). Results show that, while ultramafic soils generally host plant communities of limited taxonomic diversity with respect to similar communities on other substrates, fertilization significantly enhances the biomass production of species that are not exclusive to ultramafic soils. As a consequence, if diversity is measured combining species relative abundances with their taxonomic distinctiveness, nutrient addition tends to increase the diversity of ultramafic communities.     

An evaluation of the taxonomic distinctness of two Geonoma macrostachys (Arecaceae) varieties based on intersimple sequence repeat (ISSR) variation

The latest taxonomic account for the palms of the Amazon recognizes two common and widespread varieties of Geonoma macrostachys (Arecaceae) in western Amazonia: acaulis and macrostachys . These varieties are joined by intermediates, which obscure their taxonomic boundaries. An evaluation of the genetic distinctness between G. macrostachys varieties at a local and regional scale is presented. Thirty-one G. macrostachys individuals were randomly sampled from four Peruvian tropical moist forests. Twenty-seven intersimple sequence repeat (ISSR) primers were used, and all clear and reproducible bands were scored and analysed. Of the 99 ISSR bands produced, 51.52% were polymorphic. Nei's gene diversity value ( H ) was 0.2274 and Shannon's information index ( I ) was 0.3237. Clustering, ordination, and analyses of molecular variance (AMOVAs) suggested a lack of genetic distinctness between varieties at the regional level. Individuals clustered by geographical provenance, and two main groups were identified. A significant genetic differentiation between sympatric varieties occurred only in the northern locality (Φ_ST = 0.359, P  = 0.000). A hierarchical AMOVA revealed that the genetic diversity in G. macrostachys mainly lies between localities (76.58%). Mantel's test revealed no significant correlation between the geographical and genetic distances between individuals. We do not support the recognition of the two taxonomic varieties studied.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 381–392.     

Species richness and taxonomic distinctness of lake macrophytes along environmental gradients in two continents

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Spatial variation in taxonomic distinctness of ciliated protozoan communities at genus-level resolution and relationships to marine water quality in Jiaozhou Bay,northern China

Taxonomic distinctness of ciliated protozoan communities at genus-level resolution was studied over a 12-month period (June 2007–May 2008) in Jiaozhou Bay, northern China. Samples were collected biweekly from three different depths at each of five sites. A range of physico-chemical parameters were also measured in order to determine water quality. Multivariate and univariate analyses showed that (1) spatial patterns of ciliated protozoan communities were significantly different among the five sampling sites (P < 0.05); (2) there were no significant differences among the three depths at each site (P > 0.05); (3) the taxonomic patterns of ciliate communities at genus-level resolution were significantly correlated with the spatial variation of environmental variables, in particular nitrate nitrogen (NO₃-N), sum of NO₃-N and nitrite nitrogen (NO _n -N), and soluble reactive phosphate (SRP); (4) the Margalef’s index did not show a significant correlation with chemical parameters although it was higher for the less eutrophic than the more eutrophic sites (P > 0.05); (5) the average taxonomic distinctness (Δ⁺) and variation in taxonomic distinctness (Λ⁺) of the ciliate communities at genus-level resolution were significantly negatively correlated with the combination of NO _n -N and SRP; (6) the paired taxonomic biodiversity indices (Δ⁺ and Λ⁺) showed a clear decreasing trend of departure from the expected taxonomic breadth in response to water quality. These results suggest that spatial patterns and taxonomic distinctness (especially the paired Δ⁺ and Λ⁺) of ciliated protozoan communities at genus-level resolution can be used as potential indicators for assessing marine water quality while minimizing costs in terms of the time and resources needed for sample analysis.     

Vision in the deep sea

The deep sea is the largest habitat on earth. Its three great faunal environments--the twilight mesopelagic zone, the dark bathypelagic zone and the vast flat expanses of the benthic habitat--are home to a rich fauna of vertebrates and invertebrates. In the mesopelagic zone (150-1000 m), the down-welling daylight creates an extended scene that becomes increasingly dimmer and bluer with depth. The available daylight also originates increasingly from vertically above, and bioluminescent point-source flashes, well contrasted against the dim background daylight, become increasingly visible. In the bathypelagic zone below 1000 m no daylight remains, and the scene becomes entirely dominated by point-like bioluminescence. This changing nature of visual scenes with depth--from extended source to point source--has had a profound effect on the designs of deep-sea eyes, both optically and neurally, a fact that until recently was not fully appreciated. Recent measurements of the sensitivity and spatial resolution of deep-sea eyes--particularly from the camera eyes of fishes and cephalopods and the compound eyes of crustaceans--reveal that ocular designs are well matched to the nature of the visual scene at any given depth. This match between eye design and visual scene is the subject of this review. The greatest variation in eye design is found in the mesopelagic zone, where dim down-welling daylight and bio-luminescent point sources may be visible simultaneously. Some mesopelagic eyes rely on spatial and temporal summation to increase sensitivity to a dim extended scene, while others sacrifice this sensitivity to localise pinpoints of bright bioluminescence. Yet other eyes have retinal regions separately specialised for each type of light. In the bathypelagic zone, eyes generally get smaller and therefore less sensitive to point sources with increasing depth. In fishes, this insensitivity, combined with surprisingly high spatial resolution, is very well adapted to the detection and localisation of point-source bioluminescence at ecologically meaningful distances. At all depths, the eyes of animals active on and over the nutrient-rich sea floor are generally larger than the eyes of pelagic species. In fishes, the retinal ganglion cells are also frequently arranged in a horizontal visual streak, an adaptation for viewing the wide flat horizon of the sea floor, and all animals living there. These and many other aspects of light and vision in the deep sea are reviewed in support of the following conclusion: it is not only the intensity of light at different depths, but also its distribution in space, which has been a major force in the evolution of deep-sea vision.     

Ocean-scale patterns of 'biodiversity' of Atlantic asteroids determined from taxonomic distinctness and other measures

We examine patterns of ‘gamma’ (within-rcgion) and ‘beta’ (between-region) diversity from analysis of a presence/absence dataset for species of asteroids encompassing the whole Atlantic Ocean partitioned into 26 regions. Absolute species numbers (a poor measure of biodiversity) in shallow coastal areas and the deep sea are the same, although species richness per area for two well-sampled regions suggests, qualitatively, that coastal areas may be more speciose. Taxonomic distinctness (A*), an index which is markedly less sample-size dependent than other common diversity measures, shows no significant association with geographic area and no clear pattern with depth, suggesting an absence of latitudinal and coastal/deep trends. Cluster analysis shows that distinctive faunal assemblages are most evident in shelf/ shallow waters, where six groupings separate recognizably acccording to geographical location. Three of these arc the southernmost regions of the Atlantic (southeast of S. America, S. Angola/S. Africa & Tristan da Cunha/Gough Island) and arc characterized by their isolation and high levels of endemism. As depth increases so does the amount of faunal similarity between regions. This indicates that beta diversity is highest in shelf regions and lowest in lower bathyal/abyssal regions. Our results may support the contention which questions the emerging paradigm that the deep sea has exceptionally high diversity. It is evident, however, that comparisons (e.g. between coasts and the deep sea) are problematic and can depend very much on the element(s) of biodiversity measured, sampling methods and the spatial scales (e.g. alpha, beta or gamma diversity) over which assessment is made. Any wider conclusions should therefore be drawn cautiously, particularly since assessment is made of only one faunal group. Other findings include significant correlation between the depth range of asteroids and their geographical range. The utility of low-resolution datasets is reviewed. It is concluded that within limitations they can be of value for determining broad (e.g. regional, ocean-scale and global-scale) patterns of diversity and community structure, especially when analysed using measures relatively uninfluenced by sample size.     

Cross-species amplification of microsatellites reveals incongruence in the molecular variation and taxonomic limits of the Pilosocereus aurisetus group (Cactaceae)

The Pilosocereus aurisetus group contains eight cactus species restricted to xeric habitats in eastern and central Brazil that have an archipelago-like distribution. In this study, 5–11 microsatellite markers previously designed for Pilosocereus machrisii were evaluated for cross-amplification and polymorphisms in ten populations from six species of the P. aurisetus group. The genotypic information was subsequently used to investigate the genetic relationships between the individuals, populations, and species analyzed. Only the Pmac101 locus failed to amplify in all of the six analyzed species, resulting in an 88?% success rate. The number of alleles per polymorphic locus ranged from 2 to 12, and the most successfully amplified loci showed at least one population with a larger number of alleles than were reported in the source species. The population relationships revealed clear genetic clustering in a neighbor-joining tree that was partially incongruent with the taxonomic limits between the P. aurisetus and P. machrisii species, a fact which parallels the problematic taxonomy of the P. aurisetus group. A Bayesian clustering analysis of the individual genotypes confirmed the observed taxonomic incongruence. These microsatellite markers provide a valuable resource for facilitating large-scale genetic studies on population structures, systematics and evolutionary history in this group.     

The use of taxonomic distinctness to assess environmental disturbance of insect communities from running water

1. Taxonomic distinctness (or average taxonomic breadth) of a sample is a measure of diversity that has the great advantage of not being sensitive to variations in sampling effort, unlike measures of species richness. It can also be tested for departures from expectation and can thus be used to determine whether a site has suffered from a genuine loss of diversity. 2. Taxonomic distinctness or delta⁺ was measured using all insect species from three datasets obtained from running water habitats: reference or least disturbed sites from channel and bank habitats on rivers from the Australian state of Victoria; test sites distributed along a water quality gradient in Victoria and sites immediately below a series of dams in Victoria and New South Wales. The last two datasets comprised sites subject to a range of obvious disturbances. 3. At reference sites delta⁺ was generally within the range of expected values (95% confidence limits) for both habitats as would be anticipated for these essentially undisturbed sites. However, 13–14% of sites fell below the lower confidence limits. Sites in river basins with less cover of natural habitat had slightly but significantly lower delta⁺ values than those with more cover and these sites contributed to this anomaly. 4. Delta⁺ was inversely correlated with water quality (r = −0.54, P = 0.002) as measured by the first axis of a PCA of water quality variables. It was also inversely correlated with individual variables, such as turbidity and total phosphorous. 5. Delta⁺ was below expectation for 18 of the 19 dam sites and at 13 of these sites delta⁺ values were below the lower confidence limit. 6. Delta⁺ was a sensitive index of diversity that generally distinguished sites suffering from environmental disturbance. Only a few previous studies of river ecosystems have used delta⁺, which did not respond consistently to disturbance because either too few taxa at species or higher taxonomic levels were available or because longitudinal changes in species composition swamped the signal from delta⁺. Neither of these problems occurred with the current datasets. 7. Delta⁺ responded more readily to disturbance than number of species, perhaps because taxonomic distinctness reflects some of the multivariate nature of community samples, whereas number of species is blind to this aspect.     

Biofilm transplantation in the deep sea

A gold rush is currently going on in microbial ecology, which is powered by the possibility to determine the full complexity of microbial communities through next‐generation sequencing. Accordingly, enormous efforts are underway to describe microbiomes worldwide, in humans, animals, plants, soil, air and the ocean. While much can be learned from these studies, only experiments will finally unravel mechanisms. One of the key questions is how a microbial community is assembled from a pool of bacteria in the environment, and how it responds to change – be it the increase in CO₂ concentration in the ocean, or antibiotic treatment of the gut microbiome. The study by Zhang et al. ( 2016 ) in this issue is one of the very few that approaches this problem experimentally in the natural environment. The authors selected a habitat which is both extremely interesting and difficult to access. They studied the Thuwal Seep in the Red Sea at 850 m depth and used a remotely operated vehicle (ROV) to place a steel frame carrying substrata for biofilm growth into the brine pool and into the adjacent normal bottom water (NBW). Biofilms were allowed to develop for 3 days, and then those that had been growing in the brine pool were transported to normal bottom water and stayed there for another 3 days, and vice versa. The ‘switched’ biofilms were then compared with their source communities by metagenome sequencing. Strikingly, both ‘switched’ biofilms were now dominated by the same two species. These species were able to cope with conditions in both source ecosystems, as shown by assembly of their genomes and detection of expression of key genes. The biofilms had adapted to environmental change, rather than to brine pools or NBW. The study shows both the resilience and adaptability of biofilm communities and has implications for microbial ecology in general and even for therapeutic approaches such as transplantation of faecal microbiomes.     

Increasing numbers of bird species result from taxonomic progress,not taxonomic inflation

The impact and significance of modern taxonomy on other fields in biology have been subjects of much debate. It has been proposed that increasing numbers of vertebrate species are largely owing to ‘taxonomic inflation’. According to this hypothesis, newly recognized species result from reinterpretations of species limits based on phylogenetic species concepts (PSCs) rather than from new discoveries. Here, I examine 747 proposals to change the taxonomic rank of birds in the period 1950–2007. The trend to recognize more species of birds started at least two decades before the introduction of PSCs. Most (84.6%) newly recognized species were supported by new taxonomic data. Proposals to recognize more species resulted from application of all six major taxonomic criteria. Many newly recognized species (63.4%) were not based exclusively on PSC-based criteria (diagnosability, monophyly and exclusive coalescence of gene trees). Therefore, this study finds no empirical support for the idea that the increase in species is primarily epistemological rather than data-driven. This study shows that previous claims about the causes and effects of taxonomic inflation lack empirical support. I argue that a more appropriate term for the increase in species is ‘taxonomic progress’.     

Responses of taxonomic distinctness and species diversity indices to anthropogenic impacts and natural environmental gradients in stream macroinvertebrates

1. Many studies have shown traditional species diversity indices to perform poorly in discriminating anthropogenic influences on biodiversity. By contrast, in marine systems, taxonomic distinctness indices that take into account the taxonomic relatedness of species have been shown to discriminate anthropogenic effects. However, few studies have examined the performance of taxonomic distinctness indices in freshwater systems. 2. We studied the performance of four species diversity indices and four taxonomic distinctness indices for detecting anthropogenic effects on stream macroinvertebrate assemblages. Further, we examined the effects of catchment type and area, as well as two variables (pH and total phosphorus) potentially describing anthropogenic perturbation on biodiversity. 3. We found no indications of degraded biodiversity at the putatively disturbed sites. However, species density, rarefied species richness, Shannon's diversity and taxonomic diversity showed higher index values in streams draining mineral as opposed to peatland catchments. 4. Of the major environmental gradients analysed, biodiversity indices showed the strongest relationships with catchment area, lending further support to the importance of stream size for macroinvertebrate biodiversity. Some of the indices also showed weak linear and quadratic relationships to pH and total phosphorus, and residuals from the biodiversity index‐catchment area regressions (i.e. area effect standardized) were more weakly related to pH and total phosphorus than the original index values. 5. There are a number of reasons why the biodiversity indices did not respond to anthropogenic perturbation. First, some natural environmental gradients may mask the effects of perturbation on biodiversity. Secondly, perturbations of riverine ecosystems in our study area may not be strong enough to cause drastic changes in biodiversity. Thirdly, multiple anthropogenic stressors may either increase or decrease biodiversity, and thus the coarse division of sites into reference and altered streams may be an oversimplification. 6. Although neither species diversity nor taxonomic distinctness indices revealed anthropogenic degradation of macroinvertebrate assemblages in this study, the traditional species diversity and taxonomic distinctness indices were very weakly correlated. Therefore, we urge that biodiversity assessment and conservation planning should utilize a number of different indices, as they may provide complementary information about biotic assemblages.     

A test of traditional diversity measures and taxonomic distinctness indices on benthic diatoms of soda pans in the Carpathian basin

Saline lakes are threatened all over the world and their conservation has been a key issue. Various diversity indices are available for ecological status assessments, however, with poorly explored relevance and applicability in saline, alkaline pans. Therefore, traditional diversity measures (species richness and Shannon diversity) and taxonomic distinctness indices (Average [AvTD] and Variance of Taxonomic Distinctness [VarTD]) were tested in more than 100 sampling sites of 39 soda pans in Central-Europe to find sufficient indicators of the ecological condition and simultaneously to facilitate their preservation according to the modern conservation practices. Results of the analyses showed that healthy soda pan ecosystems with high level of natural stress and reduced habitat heterogeneity are characterized by low diversity diatom assemblages. In soda pans where the stress can be extremely high from natural reasons, oligopoly of closely related species can develop: the average taxonomic distinctness appeared between genus and family level. The non-DNA-sequence based phylogenetic diversity measures (AvTD and VarTD), were generally sensitive to the trophic state of the lakes, in contrast to traditional diversity metrics, which were unequivocally indicative for the special physical and chemical parameters (e.g. conductivity, pH) of the soda pans. In some cases, when the response of the diversity measures for a given environmental variable (pH, temperature) overlapped, the AvTD was found to be a more precise indicator of the environmental changes (pH) than traditional ones. The decreasing tendency of the AvTD along the intensified natural impact may be explained by the long available time for the species to adapt to these special environments.     

Nutrient dynamics in the deep blue sea

For more than a century, oceanographers have studied the interactions between the photosynthetic production of organic matter and nutrient dynamics in the sea. This research has been field-oriented and transdisciplinary, occurring at the intersections of research in microbiology, physics, analytical chemistry, cell physiology and ecology. The global database derived from this collective effort established a sound scientific understanding of nutrient dynamics and the vital role of microorganisms, both autotrophic and heterotrophic, in the coupled organic-matter production and decomposition cycles in the sea. However, novel approaches used over the past two decades, including new designs for field experiments, repeat field observations and remote-sensing capabilities, together with updated methods of sample analysis, have led to a revolution in our thinking about the mechanisms and controls of nutrient dynamics in the deep blue sea. Contemporary paradigms bear only partial resemblance to the dogma of the past, and are likely to evolve further as new data and new ideas are presented for open discussion and debate.     

Genetic variation and taxonomic analysis of the subgenus Profundulus

The current taxonomy within the subgenus Profundulus was reviewed and at least five different species were recognized on the basis of unique genetic characters. Two of them are considered to be new species not yet described: one is known only from a single spring in the Tehuantepec basin (Mexico); the second extends mainly through springs in the Mixteca region of Mexico. The maximum Nei distances were found between P. oaxacae and the rest of the species belonging to this subgenus. Compared with the other Profundulus species, the new species from Mixteca also had high Nei values, which suggest separation of these species probably during the Miocene period. Therefore, the previously postulated old Central American origin of Profundulus is supported by our data. The mountain uplift that gave rise to the current division of Pacific and Atlantic rivers may be the reason for the current distribution pattern of the subgenus Profundulus . The sporadic presence of this subgenus in the headwaters of some Atlantic rivers has been interpreted as the result of fluvial captures during the Pleistocene period.