Global Leptocheliidae diversity: environmental drivers and richness hotspots

Polychaeta and Sipuncula are abundant inhabitants of benthic marine habitats and have been increasingly sampled in the Northwest Pacific (NWP). However, polychaete and sipunculan species richness, composition, and distribution patterns still require further investigation, despite previous studies due to increasing deep-sea data flow. Using occurrence records for Polychaeta and Sipuncula from the Ocean Biodiversity Information System (OBIS) and the Global Biodiversity Information Facility (GBIF), we analyzed sampling effort (the number of distribution records), alpha (the number of species per 700,000 km2 hexagon cells) and gamma (the number of species per 5° latitudinal band) species diversity, and estimated species richness along latitudinal and bathymetric gradients. The species richness estimations were also correlated with multiple environmental and topographic variables, including depth, temperature, dissolved oxygen, chlorophyll, primary production, phytoplankton, current velocity, light, iron, nitrate, phosphate, silicate, and salinity. The dataset included over 30,000 distribution records belonging to polychaete (31,114 records, 98%) and sipunculan (690 records, 2%) species. Half of the distribution records were reported at a species level. The area around the island of Hainan and South Korea showed the highest alpha species richness (the number of species per 700,000 km2 hexagon cell), yet the estimated species richness (ES50) indicated that there might be many unknown, unsampled, or non-digitized species throughout the whole NWP. Correspondingly, most distribution records (sampling effort) and gamma species richness were found between latitudes 20 and 40° and decreased towards higher latitudes. Sipuncula were reported relatively more frequently from the deep sea than Polychaeta (62.8% vs. 12%). Overall, the number of species and records decreased with increasing depth, with a peak at about 5000 m. The alpha species richness had the strongest positive correlations with temperature, chlorophyll, primary production, and phytoplankton concentration. Here, we provide an overview of the species richness and distribution of Polychaeta in comparison with Sipuncula in the NWP, in both shallow and deep environments. This study demonstrates where further sampling efforts are needed to fill our knowledge gaps on annelids’ distribution and diversity along the NWP. This could improve the analyses of the distribution and diversity of annelids to better understand the current environmental drivers of biodiversity, as well as predicting potential future drivers. The outcome of the environmental correlation provides thus valuable knowledge for predicting the future impacts of global warming on potential distribution shifts of annelids into new environments such as the Arctic Sea, possibly resulting in biological invasions.

Crustaceans of the Northwest Pacific Ocean: Species richness and distribution patterns

Global patterns of taxonomic and phylogenetic diversity of flowering plants: Biodiversity hotspots and coldspots

AimPatterns of benthic biodiversity at the macroecological scale remain poorly characterised throughout the Chilean latitudinal gradient, in part due to the lack of integrated databases, uneven sampling effort, and the use of species richness alone to quantify biodiversity. Different diversity measures, encompassing taxonomic and functional components, may give us extra information on biodiversity relevant to conservation planning and management. Thus, evaluating the spatial complementarity of these measures is essential.LocationCoast and continental shelf of Chile.MethodsThe latitudinal gradient of Chile was divided into five ecoregions according to the Marine Ecosystems of the World classification. Using a 55 × 55 km equal area grid, we estimated the incidence coverage‐based estimator (ICE), taxonomic distinctness (Δ+) and three measures of functional diversity: functional richness (FRic), functional evenness (FEve) and functional divergence (FDiv). For each measure, we described spatial patterns, identified hotspots, evaluated hotspot congruence and evaluated complementarity between measures.ResultsDiversity patterns varied between ecoregions and over the latitudinal gradient. ICE and Δ+ peaked in the Chiloense and Channels and Fjords ecoregions. Δ+ and FRic present a similar pattern at mid‐latitudes. FEve showed a contrary pattern, principally with FRic. Areas with high numbers of hotspots differed spatially according to each metric, and three latitudinal bands were observed. ICE, Δ+ and FRic were positively correlated, but the hotspot overlap at the grid cell level was more limited.Main ConclusionsThe complementarity between taxonomic and functional diversity measures is limited when we observe the overlap between grid cells representing hotspots. However, some regions are consistently identified as highly diverse, with the Magellanic Province (Chiloense and Channels and Fjords ecoregions) being the most important for the richness, taxonomic and functional diversity of benthos. Confirmation of the importance of this region can help prioritise conservation efforts.

Biodiversity is a multifaceted concept covering different levels of organization from genes to ecosystems. Biodiversity has at least three dimensions: (a) Taxonomic diversity (TD): a measure that is sensitive to the number and abundances of species. (b) Phylogenetic diversity (PD): a measure that incorporates not only species abundances but also species evolutionary histories. (c) Functional diversity (FD): a measure that considers not only species abundances but also species' traits. We integrate the three dimensions of diversity under a unified framework of Hill numbers and their generalizations. Our TD quantifies the effective number of equally abundant species, PD quantifies the effective total branch length, mean‐PD (PD divided by tree depth) quantifies the effective number of equally divergent lineages, and FD quantifies the effective number of equally distinct virtual functional groups (or functional ‘species’). Thus, TD, mean‐PD and FD are all in the same units of species/lineage equivalents and can be meaningfully compared. Like species richness, empirical TD, PD and FD based on sampling data depend on sampling effort and sample completeness. For TD (Hill numbers), the iNEXT (interpolation and extrapolation) standardization was developed for standardizing sample size or sample completeness (as measured by sample coverage, the fraction of individuals that belong to the observed species) to make objective comparisons across studies. This paper extends the iNEXT method to the iNEXT.3D standardization to encompass all three dimensions of diversity via sample size‐ and sample coverage‐based rarefaction and extrapolation under the unified framework. The asymptotic diversity estimates (i.e. sample size tends to infinity and sample coverage tends to unity) are also derived. In addition to individual‐based abundance data, the proposed iNEXT.3D standardization is adapted to deal with incidence‐based occurrence data. We apply the integrative framework and the proposed iNEXT.3D standardization to measure temporal alpha‐diversity changes for estuarine fish assemblage data spanning four decades. The influence of environmental drivers on diversity change are also assessed. Our analysis informs a mechanistic interpretation of biodiversity change in the three dimensions of diversity. The accompanying freeware, iNEXT.3D, developed during this project, facilitates all computation and graphics.

Patterns of foraminiferal species diversity were examined along the eastern margin of North America by utilizing the number of species, S, the information function, H(S), and species equitability, E. The 350 modern samples we studied extended from the Arctic to the Gulf of Mexico at depths ranging from a few meters to more than 5,000 m. In addition, 29 samples from Miocene strata of the Atlantic Coastal Plain and continental shelf were studied. Modern species diversity as measured by S and H(S) generally increases as depth increases and latitude decreases. Some notable exceptions occur, however, which are difficult to explain. For example, species diversity in the Arctic depth interval of 0 to 100 m is as high or higher than that found immediately south of Nova Scotia, in the Gulf of Maine, on Browns and Georges Banks, and even off the Gulf of Mexico deltas. At the moderate depth interval of 100 to 1,000 m, however, the entire margin north of Browns and Georges Banks has lower diversities than that to the south. The highest diversity by far in this depth interval occurs in the northeastern Gulf of Mexico. At the depth interval greater than 1,000 m, the more southern areas studied generally have a higher species diversity than the more northern Cape Cod to Maryland area. An exception to this is the northwestern Gulf of Mexico; this area is also an exception in that species diversity is significantly lower in the deeper waters than in the shallower waters in the same area. The measure of species equitability, E, showed no clear pattern with depth or latitude. This may be so because no simple pattern of species proportions exists or because the sampling was inadequate to measure it. Samples from the Miocene strata show a striking resemblance in species diversity to modern samples at similar depths and latitudes. Our observations indicate that species diversity and equitability have not increased during the last 15 × 106 yrs. The fossil and modern data indicate that each environment has its own carrying capacity and that this capacity is reached rather quickly. Although time and environmental stability are undoubtedly important in determining species diversity, as presently defined they are inadequate to explain all observed patterns. Long-term observations in various environments will be required to determine the relative importance of variables that affect species diversity.

Frameworks exclusively considering functional diversity are gaining popularity, as they complement and extend the information provided by taxonomic diversity metrics, particularly in response to disturbance. Taxonomic diversity should be included in functional diversity frameworks to uncover the functional mechanisms causing species loss following disturbance events. We present and test a predictive framework that considers temporal functional and taxonomic diversity responses along disturbance gradients. Our proposed framework allows us to test different multidimensional metrics of taxonomic diversity that can be directly compared to calculated multidimensional functional diversity metrics. It builds on existing functional diversity-disturbance frameworks both by using a gradient approach and by jointly considering taxonomic and functional diversity. We used previously unpublished stream insect community data collected prior to, and for the two years following, an extreme flood event that occurred in 2013. Using 14 northern Colorado mountain streams, we tested our framework and determined that taxonomic diversity metrics calculated using multidimensional methods resulted in concordance between taxonomic and functional diversity responses. By considering functional and taxonomic diversity together and using a gradient approach, we were able to identify some of the mechanisms driving species losses following this extreme disturbance event.

Conserving and restoring biodiversity are key objectives for an ecosystem approach to management in the North Sea, but ecological quality objectives for the groundfish community instead concentrate on restoring size structure. Species richness and diversity estimates are strongly influenced by sampling effort. Failure to account for this has led to the belief that species richness and diversity indices are not adequate indicators of 'state' for the groundfish community. However, ad- herence to a standard procedure that is robust within respect to sampling effort influence should allow these metrics to perform a state indicator role. The Arrhenius power and Gleason semi-log spe- cies-area relationships are examined to determine whether they can provide modelled estimates of species richness at the ICES (International Council for the Exploration of the Sea) rectangle scale. Of these, the Gleason semi-log appears most reliable, particularly when a randomised aggregation pro- cess is followed. Aggregation of at least 20 trawl samples is required to provide empirically derived index values that are representative of the communities sampled, and therefore sensitive to drivers of change in these communities. However, given current groundfish survey sampling levels, combining 20 half-hour trawl samples to provide single estimates of species richness and diversity will require considerable aggregation over time and/or space. This can lead to estimates of α or local richness/ diversity becoming inflated through the inclusion of elements of β or regional richness/diversity. For the North Sea groundfish assemblage, this occurs when the distance between the focal position and the location of the most distant sample exceeds 49 km.

The study describes an analysis of the impact of plantation forestry on the taxonomic diversity of plants in south-central Chile. In this biodiversity hotspot, plantations of non-native species like Pinus radiata D. Don, Eucalyptus globules Labill. and Populus nigra L. have largely replaced native deciduous and sclerophyllous forests. The study compares taxonomic diversity of commercial plantations and native forests using taxonomic distinctness and diversity and the Simpson diversity index. Most of these indexes attest a considerably reduced taxonomic diversity to plantations. However, taxonomic distinctness values for P. radiata plantations seem to contradict other indexes at first glance. It is shown that the higher values of taxonomic distinctness of P. radiata plantations come from taxonomic dominance. Taxonomic dominance describes the fact that P. radiate—a member of the infradivison of Gymnospermae-bears only little taxonomic resemblance to other plants, which are Angiospermae. Thus, it strongly dominates the taxonomic distinctness index and the high taxonomic resemblance of other plant within its plots is neglected. Indexes are developed that identify such dominant species and adjust for taxonomic dominance in taxonomic diversity analyses. After this adjustment, all indexes provide a coherent image on taxonomic diversity. Plantation forestry produces a considerable decline of taxonomic diversity. Taxonomic diversity analysis provides valuable insights in biodiversity impacts and complements standard analyses.

Local and regional species diversity of benthic Isopoda (Crustacea) in the deep Gulf of Mexico

Human disturbances are greatly threatening to the biodiversity of vascular plants. Compared to seed plants, the diversity patterns of ferns have been poorly studied along disturbance gradients, including aspects of their taxonomic, phylogenetic, and functional diversity. Longnan County, a biodiversity hotspot in the subtropical zone in South China, was selected to obtain a more thorough picture of the fern–disturbance relationship, in particular, the taxonomic, phylogenetic, and functional diversity of ferns at different levels of disturbance. In 90 sample plots of 5 × 5 m2 along roadsides at three sites, we recorded a total of 20 families, 50 genera, and 99 species of ferns, as well as 9759 individual ferns. The sample coverage curve indicated that the sampling effort was sufficient for biodiversity analysis. In general, the taxonomic, phylogenetic, and functional diversity measured by Hill numbers of order q = 0–3 indicated that the fern diversity in Longnan County was largely influenced by the level of human disturbance, which supports the ‘increasing disturbance hypothesis’. Many functional traits of ferns at the most disturbed site were adaptive to the disturbance. There were also some indicators of fern species responding to the different disturbance levels. Hence, ferns may be considered as a good indicator group for environmental stress.

Biodiversity hotspots are increasingly recognized as areas of high taxonomic and functional diversity. These hotspots are dynamic and shift geographically over time in response to environmental change. To identify drivers of the origin, evolution, and persistence of diversity hotspots, we investigated the diversity patterns of reef-building corals (Scleractinia) in the Central Indo-Pacific, a marine biodiversity hotspot for the last 25 Myr. We used the scleractinian fossil record (based on literature and a newly acquired fossil collection) to examine the taxonomic and functional diversity of corals from the Eocene to Pliocene. Our data identify potential drivers of coral diversity through time (and space) in the Central Indo-Pacific by constraining the timing of taxonomic turnover events and correlating them with known environmental changes. Increases in taxonomic diversity, high origination rates, and changes in abundance of functional character states indicate that the origin of the Central Indo-Pacific hotspot occurred during the Oligocene, initially through proliferation of pre-existing taxa and then by origination of new taxa. In contrast to taxonomic diversity, overall functional diversity of Central Indo-Pacific reef-building corals remained constant from the Eocene to the Pliocene. Our results identify global sea level as a main driver of diversity increase that, together with local tectonics, regulates availability of suitable habitats. Moreover, marine biodiversity hotspots develop from both the accumulation of taxa from older biodiversity hotspots and origination of new taxa. Our study demonstrates the utility of a combined literature-based and new collection approach for gaining new insights into the timing, drivers, and development of tropical biodiversity hotspots.

In the semiarid areas of Ghana, the interactive effects of various environmental drivers, their relative importance, and their direct and indirect effects on plant species composition and diversity are still poorly understood, hence affecting effective ecosystem management. Using a combined gradient approach, the study investigated the predictors of species diversity of both the woody and the herbaceous layers of a steep land-use and climatic gradient from the forest-savanna transition to the Sudan savanna of Ghana. Species richness and the Shannon-Weiner Index are the response variables. Two-way ANOVA was performed to test the interaction effects of climate and land-use on species diversity; linear mixed-effect models were used to test the relationships between multiple environmental variables, and structural equation modelling was used to determine the direct and indirect effects of climate and land-use on species diversity. We found significant effects of climate and land use, and their interactions on species diversity for both vegetation layers. We also found differential responses of the herbaceous and woody layers to environmental drivers. Land use (Grazing pressure) was the most important predictor of the woody layer while climatic aridity was the most important for the herbaceous layer. Climatic aridity and fire were only directly important for herbaceous vegetation but not the woody layer, although their indirect effects cannot be discounted. For soil properties, organic matter was important for both vegetation layers. Synthesis: The marked differences in species composition for various land uses along the climatic gradient imply that climate change will indeed affect vegetation. The observed importance of grazing for all response variables implies that land use could override climate effects and that appropriate land management strategies could mitigate potential negative effects of climate change.

Environmental filtering (EF) and dispersal filtering (DF) are widely known to shape plant community assembly. Particularly in arid and semi-arid mountainous regions, however, it remains unclear whether EF or DF dominate in the community assembly of different life forms or how they interact along elevational gradients. This research aims to reveal how different ecological processes influence herbaceous and woody community assembly and how they respond to various environmental drivers and elevational gradients. Here we integrated taxonomic diversity (TD), phylogenetic diversity (PD), and ecological drivers across an elevational gradient of 1,420 m in the Helan Mountain Nature Reserve, in typical arid and semi-arid areas of China. This study showed that the TD and PD of herbaceous communities significantly increase linearly with changing elevation gradients, while woody ‘TD’ showed a unimodal pattern, and there was little relationship between woody ‘PD’ and elevation. Herbaceous species exhibited significant phylogenetic clustering at low elevations, where they were influenced by climate, aspect, and tree cover. However, woody species exhibited random patterns across elevations. Herbaceous and woody species’ taxonomic and phylogenetic beta diversity is governed primarily by spatial turnover rather than nestedness. Spatial turnover is caused primarily by EF and DF’s combined influence, but their relative importance differs between herbaceous and woody communities. Therefore, we conclude that the responses of herbaceous and woody plants along elevation gradients in the Helan Mountains are decoupled due to their different adaptation strategies to climate factors in the drylands. These findings are important for understanding the assembly mechanisms driving plant communities in dryland under the context of dramatic increases in drought driven by climate warming.

(introduction to Special Issue: The roles of habitat heterogeneity in generating and maintaining biodiversity on continental margins A Contribution to the Census of Marine Life)