Elevational diversity of terrestrial rainforest herbs: when the whole is less than the sum of its parts

We studied the species richness of herbaceous terrestrial plant species along an elevational gradient at 250–2425 m a.s.l. in evergreen tropical forest in Central Sulawesi, Indonesia. We recorded 302 species belonging to 51 families. Ferns and lycophytes contributed 62% of the species, followed by monocots with 24% and dicots with 14%. Overall herb species richness did not show any particular relation with elevation, while the richness of ferns increased significantly with elevation, monocots did not show a pattern, and dicots showed a hump-shaped pattern with maximum richness at 1800 m. These patterns in turn were only partly reflected in the patterns of the individual plant families making up each group. The independence of different taxa was also reflected in their relationships to environmental factors (temperature, precipitation, and area): although, each single family was related to one or several factors, at the group level and at the overall level these trends were lost. These results show that interpreting diversity at higher taxonomic level may overlook important information at the family level and raises the biologically intriguing question whether overall patterns of diversity result from a random accumulation of group-specific patterns or if there is some interaction between groups (e.g., via competition and niche-pre-emption).     

Angiosperm growth habit,dispersal and diversification reconsidered

Summary Previous studies have sought to elucidate the relationship between dispersal mode (biotic versus abiotic) and the taxonomic diversification of angiosperm families, but with ambiguous results. In this study, we propose the hypothesis that the combination of (1) the large seed size required of plants germinating in closed, light-poor environments and (2) the necessity to move disseminules away from the maternal plant in order to avoid intraspecific competition, predation and pathogens should favour biotically-dispersed relative to abiotically-dispersed woody arborescent angiosperms, resulting in higher diversification of the former. In this paper, we seek patterns of diversification that support this hypothesis. We examine the association between dispersal mode, growth habit and taxonomic richness of monocotyledon and dicotyledon families using (1) contingency table analyses to detect the effect of dispersal mode on the relative abundances and diversification of woody versus herbaceous taxa and (2) non-parametric analyses of variance to detect the statistical effect of dispersal mode on taxonomic diversification (mean number of species per genus, genera per family and species per family) in monocot and dicot families dominated by biotic or abiotic dispersal. We found a clear statistical effect of dispersal mode on diversification. Among families of woody dicots, dispersal by vertebrates is associated with significantly higher levels of species per genus, genera per family and species per family than is abiotic dispersal. The same pattern is observed among woody monocots, but is not significant at the 0.05 level. Among families of herbaceous monocots and dicots, the situation is reversed, with abiotically-dispersed families exhibiting higher levels of diversification than vertebrate-dispersed families. When woody and herbaceous families are pooled, there is no association between dispersal mode and diversification. These data coincide with evidence from the fossil record to suggest vertebrate dispersal has positively contributed to the diversification of woody angiosperms. We suggest that vertebrate dispersal may have promoted the diversity of extant taxa by reducing the probability of extinction over evolutionary time, rather than by elevating speciation rates. Our results suggest vertebrate dispersal has contributed to, but does not explainin toto, the diversity of living angiosperms.     

Genus age,provincial area and the taxonomic structure of marine faunas

Species are unevenly distributed among genera within clades and regions, with most genera species-poor and few species-rich. At regional scales, this structure to taxonomic diversity is generated via speciation, extinction and geographical range dynamics. Here, we use a global database of extant marine bivalves to characterize the taxonomic structure of climate zones and provinces. Our analyses reveal a general, Zipf–Mandelbrot form to the distribution of species among genera, with faunas from similar climate zones exhibiting similar taxonomic structure. Provinces that contain older taxa and/or encompass larger areas are expected to be more species-rich. Although both median genus age and provincial area correlate with measures of taxonomic structure, these relationships are interdependent, nonlinear and driven primarily by contrasts between tropical and extra-tropical faunas. Provincial area and taxonomic structure are largely decoupled within climate zones. Counter to the expectation that genus age and species richness should positively covary, diverse and highly structured provincial faunas are dominated by young genera. The marked differences between tropical and temperate faunas suggest strong spatial variation in evolutionary rates and invasion frequencies. Such variation contradicts biogeographic models that scale taxonomic diversity to geographical area.     

Taxonomic diversity of the terrestrial bird and mammal fauna in temperate and boreal biomes of the northern hemisphere

Abstract. Using comprehensive range information of northern Hemisphere birds and mammals, we assessed the taxonomic diversity of these two groups in four different regions: Europe, east Asia, and western and eastern North America. East Asia is the richest region in the number of bird and mammal species, genera, families and orders, except that mammal species richness is highest in western North America. Eastern North America is taxonomically the poorest region, but when only forest-associated taxa were considered in mammals taxonomic diversity is equally low in Europe and in eastern North America, and in birds, Europe is the least diverse region. Patterns in endemic taxa follow overall taxonomic diversity. The proportion of shared taxa between regions is higher among boreal species and genera than among all taxa. A comparison with tree species diversity underpins the role of east Asia as the most diverse of all northern biota. Largely congruent patterns at different taxonomic levels emphasizes the role of historical processes, such as differential extinction rate in response to paleoenvironmental fluctuations, in producing these patterns, but we stress the need for more research on the coevolution of species diversity and habitat diversity.     

Patterns of taxonomic diversity among terrestrial isopods

The publication of the world catalog of terrestrial isopods some ten years ago by Schmalfuss has facilitated research on isopod diversity patterns at a global scale. Furthermore, even though we still lack a comprehensive and robust phylogeny of Oniscidea, we do have some useful approaches to phylogenetic relationships among major clades which can offer additional insights into isopod evolutionary dynamics. Taxonomic diversity is one of many approaches to biodiversity and, despite its sensitiveness to biases in taxonomic practice, has proved useful in exploring diversification dynamics of various taxa. In the present work, we attempt an analysis of taxonomic diversity patterns among Oniscidea based on an updated world list of species containing 3,710 species belonging to 527 genera and 37 families (data till April 2014). The analysis explores species diversity at the genus and family level, as well as the relationships between species per genera, species per families, and genera per families. In addition, we consider the structure of isopod taxonomic system under the fractal perspective that has been proposed as a measure of a taxon’s diversification. Finally, we check whether there is any phylogenetic signal behind taxonomic diversity patterns. The results can be useful in a more detailed elaboration of Oniscidea systematics.     

Regional variation in the historical components of global avian species richness

Aim Using a global data base of the distribution of extant bird species, we examine the evidence for spatial variation in the evolutionary origins of contemporary avian diversity. In particular, we assess the possible role of the timing of mountain uplift in promoting diversification in different regions. Location Global. Methods We mapped the distribution of avian richness at four taxonomic levels on an equal‐area 1° grid. We examined the relationships between richness at successive taxonomic levels (e.g. species richness vs. genus richness). We mapped the residuals from linear regressions of these relationships to identify areas that are exceptional in the number of lower taxa relative to the number of higher taxa. We use generalized least squares models to test the influence of elevation range and temperature on lower‐taxon richness relative to higher‐taxon richness. Results Peaks of species richness in the Neotropics were congruent with patterns of generic richness, whilst peaks in Australia and the Himalayas were congruent with patterns of both genus and family richness. Hotspots in the Afrotropics did not reflect higher‐taxon patterns. Regional differences in the relationship between richness at successive taxonomic levels revealed variation in patterns of taxon co‐occurrence. Species and genus co‐occurrence was positively associated with elevational range across much of the world. Taxon occurrence in the Neotropics was associated with a positive interaction between elevational range and temperature. Conclusions These results demonstrate that contemporary patterns of richness show different associations with higher‐taxon richness in different regions, which implies that the timing of historical effects on these contemporary patterns varies across regions. We suggest that this is due to dispersal limitation and phylogenetic constraints on physiological tolerance limits promoting diversification. We speculate that diversification rates respond to long‐term changes in the Earth's topography, and that the role of tropical mountain ranges is implicated as a correlate of contemporary diversity, and a source of diversification across avian evolutionary history.     

POLLEN ENERGETICS AND POLLINATION MODES

The caloric content of pollen for several different species was determined by using a semimicro oxygen bomb calorimeter. The species studied were divided into subsets for statistical comparisons. The subsets considered were wind-pollinated dicots, monocots and gymnosperms; insect-pollinated dicots; and taxonomic subclasses of the wind-pollinated angiosperms (Asteridae, Hamamelidae, Caryophyllidae and Commelinidae). The results indicated a statistically significant caloric content difference (0.05 or higher) between wind-pollinated dicots vs. monocots and gymnosperms. There was no statistical difference between wind-pollinated dicots and insect-pollinated dicots. All subclasses were significantly different from one another. A statistically significant difference was found between two tribes in the Asteraceae which have shifted from insect to wind pollination, providing additional evidence for an independent evolution of the two. The need for future research in this area is discussed.     

Phylogenetic balance and ecological evenness

The frequency distribution of numbers of species in taxonomic groups, where many species belong to a few very diverse higher taxa, is mirrored by that of species in most communities, where many individuals belong to a few very abundant species. Various hypotheses mechanistically link a species' community abundance with the diversity of the higher level taxon (genus, family, order) to which it belongs, but empirical data are equivocal about general trends in the relation between rank-taxon diversity and mean abundance. One reason for this inconclusive result may be the effect of the semisubjective nature of rank-based classification. We assessed the relationship between clade diversity and mean species abundance for two diverse tropical tree communities, using both traditional rank-based analysis and two new phylogenetic analyses (based on the ratio of individuals to taxa at each node in the phylogeny). Both rank-based and phylogenetic analyses using taxonomic ranks above the species level as terminal taxa detected a trend associating common species with species-rich families. In contrast, phylogenetic analyses using species as terminal taxa could not distinguish the observed distribution of species abundances from a random distribution with respect to clade diversity. The difference between these results might be due to (1) the absence of a real phylogeny-wide relationship between clade abundance and diversity, (2) the influence of poor phylogenetic resolution within families in our phylogenies, or (3) insufficient sensitivity of our metrics to subtle tree-wide effects. Further development and application of phylogeny-based methods for testing abundance-diversity relationships is needed.     

Floristic composition and endemism pattern of vascular plants in Ethiopia and Eritrea

Traditional attempts to delineate floristic regions are typically based on the qualitative analysis of species distribution, often ignoring the phylogenetic relationships among their taxa. Ethiopia and Eritrea are in the Horn of Africa, known as one of the world's biodiversity hotspots. We quantitatively classified the flora of Ethiopia and Eritrea into meaningful geographical units by analyzing the taxonomic and phylogenetic β‐diversity at genera, total species, and endemic species levels at a scale of 0.5° × 0.5° grid cells. Hierarchical clustering was used to quantitatively delimitate the flora and analysis of similarities was used to test the significant difference between the derived groups in taxonomic composition and phylogenetic relatedness. In total, two floristic subprovinces, five floristic districts, and 13 floristic subdistricts, as well as three centers of species endemism associated with three floristic subdistricts were identified. Our results also showed that the species diversity, endemism, and turnover of the highlands in Ethiopia and Eritrea were much higher than the lowlands, indicating that the floristic differences are closely related to the topography of the East African Rift. In this study, we provided a scientific framework for the composition and relationships of the floristic units in the Horn of Africa, and similarly provided a scientific basis for better conservation of the diversity in this region.     

Taxonomic surrogacy in biodiversity assessments,and the meaning of Linnaean ranks

The majority of biodiversity assessments use species as the base unit. Recently, a series of studies have suggested replacing numbers of species with higher ranked taxa (genera, families, etc.); a method known as taxonomic surrogacy that has an important potential to save time and resources in assesments of biological diversity. We examine the relationships between taxa and ranks, and suggest that species/higher taxon exchanges are founded on misconceptions about the properties of Linnaean classification. Rank allocations in current classifications constitute a heterogeneous mixture of various historical and contemporary views. Even if all taxa were monophyletic, those referred to the same rank would simply denote separate clades without further equivalence. We conclude that they are no more comparable than any other, non‐nested taxa, such as, for example, the genus Rattus and the phylum Arthropoda, and that taxonomic surrogacy lacks justification. These problems are also illustrated with data of polychaetous annelid worms from a broad‐scale study of benthic biodiversity and species distributions in the Irish Sea. A recent consensus phylogeny for polychaetes is used to provide three different family‐level classifications of polychaetes. We use families as a surrogate for species, and present Shannon‐Wiener diversity indices for the different sites and the three different classifications, showing how the diversity measures rely on subjective rank allocations.     

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.     

Phylogeny of North American amblemines (Bivalvia, Unionoida): prodigious polyphyly proves pervasive across genera

Abstract. The subfamily Ambleminae is the most diverse subfamily of fresh‐water mussels (order Unionoida), a globally diverse and ecologically prominent group of bivalves. About 250 amblemine species occur in North America; however, this diversity is highly imperiled, with the majority of species at risk. Assessing and protecting this diversity has been hampered by the uncertain systematics of this group. This study sought to provide an improved phylogenetic framework for the Ambleminae. Currently, 37 North American genera are recognized in Ambleminae. Previous phylogenetic studies of amblemines highlighted the need for more extensive sampling due to the uncertainties arising from polyphyly of many currently recognized taxa. The present study incorporated all amblemine genera occurring in North America north of the Rio Grande, with multiple species of most genera, including the type species for all but seven genera. A total of 192 new DNA sequences were obtained for three mitochondrial gene regions: COI, 16S, and ND1. In combination with published data, this produced a data matrix incorporating 357 gene sequences for 143 operational taxonomic units, representing 107 currently recognized species. Inclusion of published data provides additional taxa and a summary of present molecular evidence on amblemine phylogeny, if at the cost of increasing the amount of missing data. Parsimony and Bayesian analyses suggest that most amblemine genera, as currently defined, are polyphyletic. At higher taxonomic levels, the tribes Quadrulini, Lampsilini, and Pleurobemini were supported; the extent of Amblemini and the relationships of some genera previously assigned to that tribe remain unclear. The eastern North American amblemines appear monophyletic. Gonidea and some Eurasian taxa place as probable sister taxa for the eastern North American Ambleminae. The results also highlight problematic taxa of particular interest for further work.     

Local, among-site, and regional diversity patterns of benthic macroinvertebrates in high altitude waterbodies: do ponds differ from lakes?

In this study we aimed at comparing invertebrate diversity of high altitude lakes and ponds along hierarchical spatial scales. We compared local, among-site, and regional diversity of benthic macroinvertebrates in 25 ponds and 34 lakes in the Tatra Mountains, central Europe. The ponds showed significantly lower local diversity, higher among-site diversity and similar regional diversity than the lakes. The species–area relationships (SAR), habitat heterogeneity, and environmental harshness are assumed as drivers for the local diversity patterns. An ecological threshold separating pond and lake systems emerged at an area of 2 ha, where the SAR pattern changed significantly. Differences in species turnover between these systems were likely driven by greater environmental variability and isolation of the ponds. High altitude ponds neither significantly support greater regional diversity nor higher number of unique taxa than lakes. The higher among-site diversity of ponds relative to lakes highlights the relevance of ponds for regional diversity in mountain areas.     

Three reasons to re-evaluate fungal diversity ‘on Earth and in the ocean’

Attempts to assess fungal global species richness are confounded by several problems: uncertainty about the number of described species, incomplete fungal inventories even at a high taxonomic level, high diversity of unknown, often small and elusive taxa, high levels of morphological conservation, and incomplete knowledge of their ecological and biogeographical distributions. The two main bases for estimating total fungal diversity are (1) the number of described species and their taxonomic structure, and (2) extrapolating species-area relationships. We argue that knowledge of fungal taxonomy and environmental sampling of fungi are both too incomplete for either approach to be reliable. However, it is likely that the true number of fungal species on the planet is a seven-digit number, and may even be an order of magnitude higher.     

Three reasons to re-evaluate fungal diversity ‘on Earth and in the ocean’

Attempts to assess fungal global species richness are confounded by several problems: uncertainty about the number of described species, incomplete fungal inventories even at a high taxonomic level, high diversity of unknown, often small and elusive taxa, high levels of morphological conservation, and incomplete knowledge of their ecological and biogeographical distributions. The two main bases for estimating total fungal diversity are (1) the number of described species and their taxonomic structure, and (2) extrapolating species-area relationships. We argue that knowledge of fungal taxonomy and environmental sampling of fungi are both too incomplete for either approach to be reliable. However, it is likely that the true number of fungal species on the planet is a seven-digit number, and may even be an order of magnitude higher.     

Are there general rules governing parasite diversity? Small mammalian hosts and gamasid mite assemblages

Parasite biodiversity varies on several scales, and in particular among different host species. Previous attempts at finding relationships between host features and the diversity of the parasite assemblages they harbour have yielded inconsistent results, suggesting strongly that any patterns might be taxon-specific. Here, we examined the potential of three host characteristics (host body mass, basal metabolic rate, and area of the geographical range) as determinants of parasite diversity in one group of ectoparasites, gamasid mites (superfamily Dermanyssoidea), using data from 63 species of small mammalian hosts. Our analyses used three measures of parasite diversity (species richness, the Shannon diversity index, and average taxonomic distinctness), and controlled for sampling effort and phylogenetic influences. Although several significant relationships were observed, they depended entirely on which diversity measure was used, or on which host taxon was investigated (insectivores vs. rodents and lagomorphs). In addition, the present results on patterns of mite diversity were not consistent with those of an earlier study involving roughly the same host taxa and the same biogeographical area, but a different group of ectoparasites, i.e. fleas. Thus, there appears to be no universal determinant of parasite diversity, and associations between host features and parasite diversity probably evolve independently in different host–parasite systems.     

Soil fungal communities of grasslands are environmentally structured at a regional scale in the Alps

Studying patterns of species distributions along elevation gradients is frequently used to identify the primary factors that determine the distribution, diversity and assembly of species. However, despite their crucial role in ecosystem functioning, our understanding of the distribution of below‐ground fungi is still limited, calling for more comprehensive studies of fungal biogeography along environmental gradients at various scales (from regional to global). Here, we investigated the richness of taxa of soil fungi and their phylogenetic diversity across a wide range of grassland types along a 2800 m elevation gradient at a large number of sites (213), stratified across a region of the Western Swiss Alps (700 km²). We used 454 pyrosequencing to obtain fungal sequences that were clustered into operational taxonomic units (OTUs). The OTU diversity–area relationship revealed uneven distribution of fungal taxa across the study area (i.e. not all taxa are everywhere) and fine‐scale spatial clustering. Fungal richness and phylogenetic diversity were found to be higher in lower temperatures and higher moisture conditions. Climatic and soil characteristics as well as plant community composition were related to OTU alpha, beta and phylogenetic diversity, with distinct fungal lineages suggesting distinct ecological tolerances. Soil fungi, thus, show lineage‐specific biogeographic patterns, even at a regional scale, and follow environmental determinism, mediated by interactions with plants.     

The genetic signature of ecologically different grassland Lepidopterans

The level of genetic diversity found for species is strongly influenced by properties of the species’ ecology, abundance and behaviour (as dispersal). To address this coherence, we selected twenty-two grassland butterfly and burnet moth species, which were previously analysed by allozyme electrophoresis (using 15–25 loci per species) over a study area in western Germany with adjoining areas of Luxembourg and north-eastern France. For this study area, we calculated the species’ specific climatic niche breadths and derived various ecological parameters from literature and own field observations. The obtained parameters of genetic diversity (heterozygosity, number of alleles and percentage of polymorphic loci), genetic differentiation (D _est as well as F _ST and F _IS values as proxis for genetic differentiation among populations and inbreeding within populations), as well as ecological and climatic niche dimensions did not show significant differences among the different Lepidoptera families; therefore taxonomic assignment apparently has a negligible influence on the genetic structure of taxa. Genetic diversity and differentiation showed a significant correlation with the ecological and climatic niche-breadth of species in many cases: generalistic species with rather unspecific ecological characteristics and climatic niche had higher genetic diversities and tend to have lower differentiation and inbreeding, whereas specialist taxa (i.e. with narrow ecological and climatic niches) have lower genetic diversities and higher differentiation and inbreeding. The results might reflect contrasting population structures of specialist species with lower abundances compared with the more common generalists. The more restricted and isolated occurrence of specialists might consequence a reduction in genetic diversity and an increase in genetic differentiation among local populations. In contrast, generalists with unspecific habitat requirements occur in higher abundances and in consequence show a more homogenous genetic structure with higher diversities.     

Diversity and evolution of African Grass Rats (Muridae: Arvicanthis)—From radiation in East Africa to repeated colonization of northwestern and southeastern savannas

African Grass Rats of the genus Arvicanthis Lesson, 1842, are one of the most important groups of rodents in sub‐Saharan Africa. They are abundant in a variety of open habitats, they are major agricultural pests, and they became a popular model in physiological research because of their diurnal activity. Despite this importance, information about their taxonomy and distribution is unsatisfactory, especially in eastern Africa. In this study, we collected the most comprehensive multilocus DNA dataset to date across the geographic and taxonomic range of the genus (229 genotyped specimens from 130 localities in 16 countries belonging to all currently recognized species). We reconstructed phylogenetic relationships, mapped the distribution of major genetic clades, and used the combination of cytogenetic, nuclear, and mitochondrial markers for species delimitations and taxonomic suggestions. The genus is composed of two major evolutionary groups, called here the ANSORGEI and NILOTICUS groups. The former contains four presumed species, while the latter is more diverse and we recognized nine species. Most relationships among species are not resolved, which suggests a rapid radiation (dated to early–middle Pleistocene). Further, there is an indication of reticulate evolution in Ethiopia, that is, the region of the highest Arvicanthis diversity. The distribution of genetic diversity suggests diversification in eastern Africa, followed by repeated dispersals to the west (Sudano‐Guinean savannas) and to the south (Masai steppe). We propose nomenclatural changes for Ethiopian taxa and provide suggestions for future steps toward solving remaining taxonomic questions in the genus.