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Showing papers on "Species richness published in 2012"


Journal ArticleDOI
07 Jun 2012-Nature
TL;DR: The analyses clearly show that the ecosystem consequences of local species loss are as quantitatively significant as the direct effects of several global change stressors that have mobilized major international concern and remediation efforts.
Abstract: Evidence is mounting that extinctions are altering key processes important to the productivity and sustainability of Earth’s ecosystems 1–4 . Further species loss will accelerate change in ecosystem processes 5–8 , but it is unclear how these effects compare to the direct effects of other forms of environmental change that are both driving diversity loss and altering ecosystem function. Here we use a suite of meta-analyses of published data to show that the effects of species loss on productivity and decomposition—two processes important in all ecosystems—are of comparable magnitude to the effects of many other global environmental changes. In experiments, intermediate levels of species loss (21–40%) reduced plant production by 5–10%, comparable to previously documented effects of ultraviolet radiation and climate warming. Higher levels of extinction (41–60%) had effects rivalling those of ozone, acidification, elevated CO2 and nutrient pollution. At intermediate levels, species loss generally had equal or greater effects on decomposition than did elevated CO2 and nitrogen addition. The identity of species lost also had a large effect on changes in productivity and decomposition, generating a wide range of plausible outcomes for extinction. Despite the need for more studies on interactive effects of diversity loss and environmental changes, our analyses clearly show that the ecosystem consequences of local species loss are as quantitatively significant as the direct effects of several global change stressors that have mobilized major international concern and remediation efforts 9 .

1,858 citations


Journal ArticleDOI
TL;DR: In this paper, the authors provide new unconditional variance estimators for classical, individual-based rarefaction and for Coleman Rarefaction under two sampling models: sampling-theoretic predictors for the number of species in a larger sample (multinomial model), a larger area (Poisson model) or a larger number of sampling units (Bernoulli product model), based on an estimate of asymptotic species richness.
Abstract: Aims In ecology and conservation biology, the number of species counted in a biodiversity study is a key metric but is usually a biased underestimate of total species richness because many rare species are not detected. Moreover, comparing species richness among sites or samples is a statistical challenge because the observed number of species is sensitive to the number of individuals counted or the area sampled. For individual-based data, we treat a single, empirical sample of species abundances from an investigator-defined species assemblage or community as a reference point for two estimation objectives under two sampling models: estimating the expected number of species (and its unconditional variance) in a random sample of (i) a smaller number of individuals (multinomial model) or a smaller area sampled (Poisson model) and (ii) a larger number of individuals or a larger area sampled. For sample-based incidence (presence–absence) data, under a Bernoulli product model, we treat a single set of species incidence frequencies as the reference point to estimate richness for smaller and larger numbers of sampling units. Methods The first objective is a problem in interpolation that we address with classical rarefaction (multinomial model) and Coleman rarefaction (Poisson model) for individual-based data and with sample-based rarefaction (Bernoulli product model) for incidence frequencies. The second is a problem in extrapolation that we address with sampling-theoretic predictors for the number of species in a larger sample (multinomial model), a larger area (Poisson model) or a larger number of sampling units (Bernoulli product model), based on an estimate of asymptotic species richness. Although published methods exist for many of these objectives, we bring them together here with some new estimators under a unified statistical and notational framework. This novel integration of mathematically distinct approaches allowed us to link interpolated (rarefaction) curves and extrapolated curves to plot a unified species accumulation curve for empirical examples. We provide new, unconditional variance estimators for classical, individual-based rarefaction and for Coleman rarefaction, long missing from the toolkit of biodiversity measurement. We illustrate these methods with datasets for tropical beetles, tropical trees and tropical ants.

1,445 citations


Journal ArticleDOI
TL;DR: It is shown that there is no universal measure of impact and the pattern observed depends on the ecological measure examined, and some species traits, especially life form, stature and pollination syndrome, may provide a means to predict impact, regardless of the particular habitat and geographical region invaded.
Abstract: With the growing body of literature assessing the impact of invasive alien plants on resident species and ecosystems, a comprehensive assessment of the relationship between invasive species traits and environmental settings of invasion on the characteristics of impacts is needed. Based on 287 publications with 1551 individual cases that addressed the impact of 167 invasive plant species belonging to 49 families, we present the first global overview of frequencies of significant and non-significant ecological impacts and their directions on 15 outcomes related to the responses of resident populations, species, communities and ecosystems. Species and community outcomes tend to decline following invasions, especially those for plants, but the abundance and richness of the soil biota, as well as concentrations of soil nutrients and water, more often increase than decrease following invasion. Data mining tools revealed that invasive plants exert consistent significant impacts on some outcomes (survival of resident biota, activity of resident animals, resident community productivity, mineral and nutrient content in plant tissues, and fire frequency and intensity), whereas for outcomes at the community level, such as species richness, diversity and soil resources, the significance of impacts is determined by interactions between species traits and the biome invaded. The latter outcomes are most likely to be impacted by annual grasses, and by wind pollinated trees invading mediterranean or tropical biomes. One of the clearest signals in this analysis is that invasive plants are far more likely to cause significant impacts on resident plant and animal richness on islands rather than mainland. This study shows that there is no universal measure of impact and the pattern observed depends on the ecological measure examined. Although impact is strongly context dependent, some species traits, especially life form, stature and pollination syndrome, may provide a means to predict impact, regardless of the particular habitat and geographical region invaded.

1,067 citations


Journal ArticleDOI
TL;DR: In this article, the authors used 867 vegetation samples above the treeline from 60 summit sites in all major European mountain systems to show that ongoing climate change gradually transforms mountain plant communities.
Abstract: Climate impact studies have indicated ecological fingerprints of recent global warming across a wide range of habitats1, 2. Although these studies have shown responses from various local case studies, a coherent large-scale account on temperature-driven changes of biotic communities has been lacking3, 4. Here we use 867 vegetation samples above the treeline from 60 summit sites in all major European mountain systems to show that ongoing climate change gradually transforms mountain plant communities. We provide evidence that the more cold-adapted species decline and the more warm-adapted species increase, a process described here as thermophilization. At the scale of individual mountains this general trend may not be apparent, but at the larger, continental scale we observed a significantly higher abundance of thermophilic species in 2008, compared with 2001. Thermophilization of mountain plant communities mirrors the degree of recent warming and is more pronounced in areas where the temperature increase has been higher. In view of the projected climate change5, 6 the observed transformation suggests a progressive decline of cold mountain habitats and their biota.

943 citations


Journal ArticleDOI
13 Jan 2012-Science
TL;DR: A global empirical study relating plant species richness and abiotic factors to multifunctionality in drylands, which collectively cover 41% of Earth’s land surface and support over 38% of the human population, suggests that the preservation of plant biodiversity is crucial to buffer negative effects of climate change and desertification in dryland.
Abstract: Experiments suggest that biodiversity enhances the ability of ecosystems to maintain multiple functions, such as carbon storage, productivity, and the buildup of nutrient pools (multifunctionality). However, the relationship between biodiversity and multifunctionality has never been assessed globally in natural ecosystems. We report here on a global empirical study relating plant species richness and abiotic factors to multifunctionality in drylands, which collectively cover 41% of Earth’s land surface and support over 38% of the human population. Multifunctionality was positively and significantly related to species richness. The best-fitting models accounted for over 55% of the variation in multifunctionality and always included species richness as a predictor variable. Our results suggest that the preservation of plant biodiversity is crucial to buffer negative effects of climate change and desertification in drylands.

941 citations


Journal ArticleDOI
Ward Appeltans1, Shane T. Ahyong2, Shane T. Ahyong3, Gary L. Anderson4, Martin V. Angel5, Tom Artois6, Nicolas Bailly7, Roger N. Bamber, Anthony Barber, Ilse Bartsch8, Annalisa Berta9, Magdalena Błażewicz-Paszkowycz, Phil Bock10, Geoff A. Boxshall11, Christopher B. Boyko12, Simone N. Brandão13, R. A. Bray11, Niel L. Bruce14, Niel L. Bruce15, Stephen D. Cairns16, Tin-Yam Chan17, Lanna Cheng18, Allen Gilbert Collins19, Thomas H. Cribb20, Marco Curini-Galletti21, Farid Dahdouh-Guebas22, Farid Dahdouh-Guebas23, Peter J. F. Davie24, Michael N Dawson25, Olivier De Clerck26, Wim Decock1, Sammy De Grave8, Nicole J. de Voogd27, Daryl P. Domning28, Christian C. Emig, Christer Erséus29, William N. Eschmeyer30, William N. Eschmeyer31, Kristian Fauchald16, Daphne G. Fautin8, Stephen W. Feist32, Charles H. J. M. Fransen27, Hidetaka Furuya33, Óscar García-Álvarez34, Sarah Gerken35, David I. Gibson11, Arjan Gittenberger27, Serge Gofas36, Liza Gómez-Daglio25, Dennis P. Gordon37, Michael D. Guiry38, Francisco Hernandez1, Bert W. Hoeksema27, Russell R. Hopcroft39, Damià Jaume40, Paul M. Kirk41, Nico Koedam23, Stefan Koenemann42, Jürgen B. Kolb43, Reinhardt Møbjerg Kristensen44, Andreas Kroh45, Gretchen Lambert46, David Lazarus47, Rafael Lemaitre16, Matt Longshaw32, Jim Lowry3, Enrique Macpherson40, Laurence P. Madin48, Christopher L. Mah16, Gill Mapstone11, Patsy A. McLaughlin49, Jan Mees1, Jan Mees26, Kenneth Meland50, Charles G. Messing51, Claudia E. Mills46, Tina N. Molodtsova52, Rich Mooi30, Birger Neuhaus47, Peter K. L. Ng53, Claus Nielsen44, Jon L. Norenburg16, Dennis M. Opresko16, Masayuki Osawa54, Gustav Paulay31, William F. Perrin19, John F. Pilger55, Gary C. B. Poore10, P.R. Pugh5, Geoffrey B. Read37, James Davis Reimer56, Marc Rius57, Rosana M. Rocha58, J.I. Saiz-Salinas59, Victor Scarabino, Bernd Schierwater60, Andreas Schmidt-Rhaesa13, Kareen E. Schnabel37, Marilyn Schotte16, Peter Schuchert, Enrico Schwabe, Hendrik Segers61, Caryn Self-Sullivan51, Noa Shenkar62, Volker Siegel, Wolfgang Sterrer8, Sabine Stöhr63, Billie J. Swalla46, Mark L. Tasker64, Erik V. Thuesen65, Tarmo Timm66, M. Antonio Todaro, Xavier Turon40, Seth Tyler67, Peter Uetz68, Jacob van der Land27, Bart Vanhoorne1, Leen van Ofwegen27, Rob W. M. Van Soest27, Jan Vanaverbeke26, Genefor Walker-Smith10, T. Chad Walter16, Alan Warren11, Gary C. Williams30, Simon P. Wilson69, Mark J. Costello70 
Flanders Marine Institute1, University of New South Wales2, Australian Museum3, University of Southern Mississippi4, National Oceanography Centre, Southampton5, University of Hasselt6, WorldFish7, American Museum of Natural History8, San Diego State University9, Museum Victoria10, Natural History Museum11, Dowling College12, University of Hamburg13, James Cook University14, University of Johannesburg15, National Museum of Natural History16, National Taiwan Ocean University17, Scripps Institution of Oceanography18, National Oceanic and Atmospheric Administration19, University of Queensland20, University of Sassari21, Université libre de Bruxelles22, Vrije Universiteit Brussel23, Queensland Museum24, University of California, Merced25, Ghent University26, Naturalis27, Howard University28, University of Gothenburg29, California Academy of Sciences30, Florida Museum of Natural History31, Centre for Environment, Fisheries and Aquaculture Science32, Osaka University33, University of Santiago de Compostela34, University of Alaska Anchorage35, University of Málaga36, National Institute of Water and Atmospheric Research37, National University of Ireland, Galway38, University of Alaska Fairbanks39, Spanish National Research Council40, CABI41, University of Siegen42, Massey University43, University of Copenhagen44, Naturhistorisches Museum45, University of Washington46, Museum für Naturkunde47, Woods Hole Oceanographic Institution48, Western Washington University49, University of Bergen50, Nova Southeastern University51, Shirshov Institute of Oceanology52, National University of Singapore53, Shimane University54, Agnes Scott College55, University of the Ryukyus56, University of California, Davis57, Federal University of Paraná58, University of the Basque Country59, University of Veterinary Medicine Hanover60, Royal Belgian Institute of Natural Sciences61, Tel Aviv University62, Swedish Museum of Natural History63, Joint Nature Conservation Committee64, The Evergreen State College65, Estonian University of Life Sciences66, University of Maine67, Virginia Commonwealth University68, Trinity College, Dublin69, University of Auckland70
TL;DR: The first register of the marine species of the world is compiled and it is estimated that between one-third and two-thirds of marine species may be undescribed, and previous estimates of there being well over one million marine species appear highly unlikely.

822 citations


Journal ArticleDOI
TL;DR: The results suggested that seasonal changes in environmental variables are more important than trophic interactions, and correlations in abundance were stronger within bacterial taxa rather than between bacteria and eukaryotes, or betweenacteria and environmental variables.
Abstract: Here we describe, the longest microbial time-series analyzed to date using high-resolution 16S rRNA tag pyrosequencing of samples taken monthly over 6 years at a temperate marine coastal site off Plymouth, UK. Data treatment effected the estimation of community richness over a 6-year period, whereby 8794 operational taxonomic units (OTUs) were identified using single-linkage preclustering and 21 130 OTUs were identified by denoising the data. The Alphaproteobacteria were the most abundant Class, and the most frequently recorded OTUs were members of the Rickettsiales (SAR 11) and Rhodobacteriales. This near-surface ocean bacterial community showed strong repeatable seasonal patterns, which were defined by winter peaks in diversity across all years. Environmental variables explained far more variation in seasonally predictable bacteria than did data on protists or metazoan biomass. Change in day length alone explains >65% of the variance in community diversity. The results suggested that seasonal changes in environmental variables are more important than trophic interactions. Interestingly, microbial association network analysis showed that correlations in abundance were stronger within bacterial taxa rather than between bacteria and eukaryotes, or between bacteria and environmental variables.

800 citations


Journal ArticleDOI
20 Apr 2012-Science
TL;DR: Recent changes in vascular plant species richness observed in a standardized monitoring network across Europe’s major mountain ranges are presented and indicate that high-altitude species, and in particular the rich endemic alpine flora of many Mediterranean mountain ranges, will come under increasing pressure in the predicted warmer and drier climates in this region.
Abstract: In mountainous regions, climate warming is expected to shift species' ranges to higher altitudes. Evidence for such shifts is still mostly from revisitations of historical sites. We present recent (2001 to 2008) changes in vascular plant species richness observed in a standardized monitoring network across Europe's major mountain ranges. Species have moved upslope on average. However, these shifts had opposite effects on the summit floras' species richness in boreal-temperate mountain regions (+3.9 species on average) and Mediterranean mountain regions (-1.4 species), probably because recent climatic trends have decreased the availability of water in the European south. Because Mediterranean mountains are particularly rich in endemic species, a continuation of these trends might shrink the European mountain flora, despite an average increase in summit species richness across the region.

669 citations


Journal ArticleDOI
TL;DR: Very high richness at any spatial grain is found only in two particular habitat/community types, and these high richness values form a very strong, consistent pattern, not greatly affected by the method of sampling, and this pattern extrapolates amazingly well.
Abstract: Questions The co-existence of high numbers of species has always fascinated ecologists, but what and where are the communities with the world records for plant species richness? The species–area relationship is among the best-known patterns in community ecology, but does it give a consistent global pattern for the most saturated communities, the global maxima? Location The world. Methods We assembled the maximum values recorded for vascular plant species richness for contiguous areas from 1 mm2 up to 1 ha. We applied the power function to relate maximal richness to area and to make extrapolations to the whole Earth. Results Only two community types contain global plant species maxima. The maxima at smaller spatial grain were from oligo- to meso-trophic, managed, semi-natural, temperate grasslands (e.g. 89 species on 1 m2), those at larger grains were from tropical rain forests (e.g. 942 species on 1 ha). The maximum richness values closely followed a power function with z = 0.250: close to Preston's ‘canonical’ value of 0.262. There was no discernable difference between maxima using rooted presence (i.e. including only plants rooted in the plot) vs shoot presence (i.e. including any plant with physical cover over the plot). However, shoot presence values must logically be greater, with the curves flattening out at very small grain, and there is evidence of this from point quadrats. Extrapolating the curve to the terrestrial surface of the Earth gave a prediction of 219 204 vascular plant species, surprisingly close to a recent estimate of 275 000 actual species. Conclusions Very high richness at any spatial grain is found only in two particular habitat/community types. Nevertheless, these high richness values form a very strong, consistent pattern, not greatly affected by the method of sampling, and this pattern extrapolates amazingly well. The records challenge ecologists to consider mechanisms of species co-existence, answers to the ‘Paradox of the Plankton’.

621 citations


Journal ArticleDOI
TL;DR: This analysis is the first to demonstrate the critical role of species evenness, richness and the importance of contrasting traits in defining net diversity effects in forest polycultures and should motivate future studies to link richness,Evenness, contrasting traits and life-history stage to the mechanisms that are expected to produce positive net biodiversity effects.
Abstract: Summary 1. Although there is ample support for positive species richness–productivity relationships in planted grassland experiments, a recent 48-site study found no diversity–productivity relationship (DPR) in herbaceous communities. Thus, debate persists about diversity effects in natural versus planted systems. Additionally, current knowledge is weak regarding the influence of evenness on the DPRs, how DPRs are affected by the variation in life-history traits among constituent species in polycultures and how DPRs differ among biomes. The impacts of these factors on DPRs in forest ecosystems are even more poorly understood. 2. We performed a meta-analysis of 54 studies to reconcile DPRs in forest ecosystems. We quantified the net diversity effect as log effect size [ln(ES)], the log ratio of the productivity in polycultures to the average of those in monocultures within the same type of mixture, site condition and stand age of each study. The first use of a boosted regression tree model in meta-analysis, a useful method to partition the effects of multiple predictors rather than relying on vote-counting of individual studies, unveiled the relative influences of individual predictors. 3. Global average ln(ES) was 0.2128, indicating 23.7% higher productivity in polycultures than monocultures. The final model explained 21% of the variation in ln(ES). The predictors that substantially accounted for the explained variation included evenness (34%), heterogeneity of shade tolerance (29%), richness (13%) and stand age (15%). In contrast, heterogeneity of nitrogen fixation and growth habits, biome and stand origin (naturally established versus planted) contributed negligibly (each £ 4%). Log effect size strongly increased with evenness from 0.6 to 1 and with richness from 2 to 6. Furthermore, it was higher with heterogeneity of shade tolerance and generally increased with stand age. 4. Synthesis. Our analysis is, to our knowledge, the first to demonstrate the critical role of species evenness, richness and the importance of contrasting traits in defining net diversity effects in forest polycultures. While testing the specific mechanisms is beyond the scope of our analysis, our results should motivate future studies to link richness, evenness, contrasting traits and life-history stage to the mechanisms that are expected to produce positive net biodiversity effects such as niche differentiation, facilitation and reduced Janzen–Connell effects.

613 citations


Journal ArticleDOI
TL;DR: In this paper, the lack of a consistent relationship between actual plant, butterfly, and bird species richness and the psychological well-being of urban greenspace visitors was investigated, showing a positive relationship with the richness that the greenspace users perceived to be present.
Abstract: Over half of the world's human population lives in cities, and for many, urban greenspaces are the only places where they encounter biodiversity. This is of particular concern because there is growing evidence that human well-being is enhanced by exposure to nature. However, the specific qualities of greenspaces that offer the greatest benefits remain poorly understood. One possibility is that humans respond positively to increased levels of biodiversity. Here, we demonstrate the lack of a consistent relationship between actual plant, butterfly, and bird species richness and the psychological well-being of urban greenspace visitors. Instead, well-being shows a positive relationship with the richness that the greenspace users perceived to be present. One plausible explanation for this discrepancy, which we investigate, is that people generally have poor biodiversity-identification skills. The apparent importance of perceived species richness and the mismatch between reality and perception pose a serious challenge for aligning conservation and human well-being agendas.

Journal ArticleDOI
TL;DR: The results reveal that body size and dispersal mode are important drivers of metacommunity structure and these traits should therefore be considered when developing a predictive framework for metacomunity dynamics.
Abstract: Relationships between traits of organisms and the structure of their metacommunities have so far mainly been explored with meta-analyses. We compared metacommunities of a wide variety of aquatic organism groups (12 groups, ranging from bacteria to fish) in the same set of 99 ponds to minimise biases inherent to meta-analyses. In the category of passive dispersers, large-bodied groups showed stronger spatial patterning than small-bodied groups suggesting an increasing impact of dispersal limitation with increasing body size. Metacommunities of organisms with the ability to fly (i.e. insect groups) showed a weaker imprint of dispersal limitation than passive dispersers with similar body size. In contrast, dispersal movements of vertebrate groups (fish and amphibians) seemed to be mainly confined to local connectivity patterns. Our results reveal that body size and dispersal mode are important drivers of metacommunity structure and these traits should therefore be considered when developing a predictive framework for metacommunity dynamics.

Journal ArticleDOI
13 Jun 2012-PLOS ONE
TL;DR: It is found that commonalities between samples based on taxonomy could sometimes belie variability at the sub-genus OTU level, and even OTUs present in nearly every subject, or that dominate in some samples, showed orders of magnitude variation in relative abundance emphasizing the highly variable nature across individuals.
Abstract: We explore the microbiota of 18 body sites in over 200 individuals using sequences amplified V1–V3 and the V3–V5 small subunit ribosomal RNA (16S) hypervariable regions as part of the NIH Common Fund Human Microbiome Project. The body sites with the greatest number of core OTUs, defined as OTUs shared amongst 95% or more of the individuals, were the oral sites (saliva, tongue, cheek, gums, and throat) followed by the nose, stool, and skin, while the vaginal sites had the fewest number of OTUs shared across subjects. We found that commonalities between samples based on taxonomy could sometimes belie variability at the sub-genus OTU level. This was particularly apparent in the mouth where a given genus can be present in many different oral sites, but the sub-genus OTUs show very distinct site selection, and in the vaginal sites, which are consistently dominated by the Lactobacillus genus but have distinctly different sub-genus V1–V3 OTU populations across subjects. Different body sites show approximately a ten-fold difference in estimated microbial richness, with stool samples having the highest estimated richness, followed by the mouth, throat and gums, then by the skin, nasal and vaginal sites. Richness as measured by the V1–V3 primers was consistently higher than richness measured by V3–V5. We also show that when such a large cohort is analyzed at the genus level, most subjects fit the stool “enterotype” profile, but other subjects are intermediate, blurring the distinction between the enterotypes. When analyzed at the finer-scale, OTU level, there was little or no segregation into stool enterotypes, but in the vagina distinct biotypes were apparent. Finally, we note that even OTUs present in nearly every subject, or that dominate in some samples, showed orders of magnitude variation in relative abundance emphasizing the highly variable nature across individuals.

Journal ArticleDOI
27 Apr 2012-PLOS ONE
TL;DR: Results obtained demonstrate that regional approaches to analytical biogeography are at present more likely to achieve insights into the biogeographic history of sponges than a global perspective, which appears currently too ambitious.
Abstract: With the completion of a single unified classification, the Systema Porifera (SP) and subsequent development of an online species database, the World Porifera Database (WPD), we are now equipped to provide a first comprehensive picture of the global biodiversity of the Porifera. An introductory overview of the four classes of the Porifera is followed by a description of the structure of our main source of data for this paper, the WPD. From this we extracted numbers of all 'known' sponges to date: the number of valid Recent sponges is established at 8,553, with the vast majority, 83%, belonging to the class Demospongiae. We also mapped for the first time the species richness of a comprehensive set of marine ecoregions of the world, data also extracted from the WPD. Perhaps not surprisingly, these distributions appear to show a strong bias towards collection and taxonomy efforts. Only when species richness is accumulated into large marine realms does a pattern emerge that is also recognized in many other marine animal groups: high numbers in tropical regions, lesser numbers in the colder parts of the world oceans. Preliminary similarity analysis of a matrix of species and marine ecoregions extracted from the WPD failed to yield a consistent hierarchical pattern of ecoregions into marine provinces. Global sponge diversity information is mostly generated in regional projects and resources: results obtained demonstrate that regional approaches to analytical biogeography are at present more likely to achieve insights into the biogeographic history of sponges than a global perspective, which appears currently too ambitious. We also review information on invasive sponges that might well have some influence on distribution patterns of the future.

Journal ArticleDOI
TL;DR: There is an increasing trend in new introductions via the Suez Canal and via shipping, while Lessepsian species decline westwards, while the reverse pattern is evident for ship-mediated species and for those introduced with aquaculture.
Abstract: More than 60 marine non-indigenous species (NIS) have been removed from previous lists and 84 species have been added, bringing the total to 986 alien species in the Mediterranean [775 in the eastern Mediterranean (EMED), 249 in the central Mediterranean (CMED), 190 in the Adriatic Sea (ADRIA) and 308 in the western Mediterranean (WMED)]. There were 48 new entries since 2011 which can be interpreted as approximately one new entry every two weeks. The number of alien species continues to increase, by 2-3 species per year for macrophytes, molluscs and polychaetes, 3-4 species per year for crustaceans, and 6 species per year for fish. The dominant group among alien species is molluscs (with 215 species), followed by crustaceans (159) and polychaetes (132). Macrophytes are the leading group of NIS in the ADRIA and the WMED, reaching 26-30% of all aliens, whereas in the EMED they barely constitute 10% of the introductions. In the EMED, molluscs are the most species-rich group, followed by crustaceans, fish and polychaetes. More than half (54%) of the marine alien species in the Mediterranean were probably introduced by corridors (mainly Suez). Shipping is blamed directly for the introduction of only 12 species, whereas it is assumed to be the most likely pathway of introduction (via ballasts or fouling) of another 300 species. For approximately 100 species shipping is a probable pathway along with the Suez Canal and/or aquaculture. Approximately 20 species have been introduced with certainty via aquaculture, while >50 species (mostly macroalgae), occurring in the vicinity of oyster farms, are assumed to be introduced accidentally as contaminants of imported species. A total of 18 species are assumed to have been introduced by the aquarium trade. Lessepsian species decline westwards, while the reverse pattern is evident for ship-mediated species and for those introduced with aquaculture. There is an increasing trend in new introductions via the Suez Canal and via shipping.

Journal ArticleDOI
14 Dec 2012-Science
TL;DR: This work sampled the phylogenetic breadth of arthropod taxa from the soil to the forest canopy in the San Lorenzo forest, Panama using a comprehensive range of structured protocols and found that models based on plant diversity fitted the accumulated species richness of both herbivore and nonherbivore taxa exceptionally well.
Abstract: Most eukaryotic organisms are arthropods. Yet, their diversity in rich terrestrial ecosystems is still unknown. Here we produce tangible estimates of the total species richness of arthropods in a tropical rainforest. Using a comprehensive range of structured protocols, we sampled the phylogenetic breadth of arthropod taxa from the soil to the forest canopy in the San Lorenzo forest, Panama. We collected 6144 arthropod species from 0.48 hectare and extrapolated total species richness to larger areas on the basis of competing models. The whole 6000-hectare forest reserve most likely sustains 25,000 arthropod species. Notably, just 1 hectare of rainforest yields >60% of the arthropod biodiversity held in the wider landscape. Models based on plant diversity fitted the accumulated species richness of both herbivore and nonherbivore taxa exceptionally well. This lends credence to global estimates of arthropod biodiversity developed from plant models.

Journal ArticleDOI
01 Mar 2012-Ecology
TL;DR: A system of diversity measurement is arrived at that should lay much of this suspicion about diversity indices to rest, by dropping the naive assumption that distinct species have nothing in common, working with effective numbers, and using diversity profiles.
Abstract: Realistic measures of biodiversity should reflect not only the relative abundances of species, but also the differences between them. We present a natural family of diversity measures taking both factors into account. This is not just another addition to the already long list of diversity indices. Instead, a single formula subsumes many of the most popular indices, including Shannon's, Simpson's, species richness, and Rao's quadratic entropy. These popular indices can then be used and understood in a unified way, and the relationships between them are made plain. The new measures are, moreover, effective numbers, so that percentage changes and ratio comparisons of diversity value are meaningful. We advocate the use of diversity profiles, which provide a faithful graphical representation of the shape of a community; they show how the perceived diversity changes as the emphasis shifts from rare to common species. Communities can usefully be compared by comparing their diversity profiles. We show by example that this is a far more subtle method than any relying on a single statistic. Some ecologists view diversity indices with suspicion, questioning whether they are biologically meaningful. By dropping the naive assumption that distinct species have nothing in common, working with effective numbers, and using diversity profiles, we arrive at a system of diversity measurement that should lay much of this suspicion to rest.

Journal ArticleDOI
TL;DR: Evidence is found to support the hypotheses that host diet and taxonomy structure insect gut bacterial communities (P < 0.001) were important in hymenopteran and termite gut community structure, and diet was an important community structuring factor particularly for insect hosts that ingest lignocellulose‐derived substances.
Abstract: Many insects contain diverse gut microbial communities. While several studies have focused on a single or small group of species, comparative studies of phylogenetically diverse hosts can illuminate general patterns of host–microbiota associations. In this study, we tested the hypotheses that (i) host diet and (ii) host taxonomy structure intestinal bacterial community composition among insects. We used published 16S rRNA gene sequence data for 58 insect species in addition to four beetle species sampled from the Sevilleta National Wildlife Refuge to test these hypotheses. Overall, gut bacterial species richness in these insects was low. Decaying wood xylophagous insects harboured the richest bacterial gut flora (102.8 species level operational taxonomic units (OTUs)/sample ± 71.7, 11.8 ± 5.9 phylogenetic diversity (PD)/sample), while bees and wasps harboured the least rich bacterial communities (11.0 species level OTUs/sample ± 5.4, 2.6 ± 0.8 PD/sample). We found evidence to support our hypotheses that host diet and taxonomy structure insect gut bacterial communities (P < 0.001 for both). However, while host taxonomy was important in hymenopteran and termite gut community structure, diet was an important community structuring factor particularly for insect hosts that ingest lignocellulose-derived substances. Our analysis provides a baseline comparison of insect gut bacterial communities from which to test further hypotheses concerning proximate and ultimate causes of these associations.

Journal ArticleDOI
01 Apr 2012-Ecology
TL;DR: The observed relationship between species richness and ANPP provides support for the theoretical expectation that a trade-off occurs between richness and productivity in herbaceous communities.
Abstract: Woody encroachment is a widespread and acute phenomenon affecting grasslands and savannas worldwide. We performed a meta-analysis of 29 studies from 13 different grassland/savanna communities in North America to determine the consequences of woody encroachment on plant species richness. In all 13 communities, species richness declined with woody plant encroachment (average decline = 45%). Species richness declined more in communities with higher precipitation (r2 = 0.81) and where encroachment was associated with a greater change in annual net primary productivity (ANPP; r2 = 0.69). Based on the strong positive correlation between precipitation and ANPP following encroachment (r2 = 0.87), we hypothesize that these relationships occur because water-limited woody plants experience a greater physiological and demographic release as precipitation increases. The observed relationship between species richness and ANPP provides support for the theoretical expectation that a trade-off occurs between richness and productivity in herbaceous communities. We conclude that woody plant encroachment leads to significant declines in species richness in North American grassland/savanna communities.

Journal ArticleDOI
TL;DR: It is predicted that climate change will most threaten communities of species that have narrow niches, vary in dispersal (most communities) and compete strongly, and current forecasts probably underestimate climate change impacts on biodiversity by neglecting competition and dispersal differences.
Abstract: Most climate change predictions omit species interactions and interspecific variation in dispersal. Here, we develop a model of multiple competing species along a warming climatic gradient that includes temperature-dependent competition, differences in niche breadth and interspecific differences in dispersal ability. Competition and dispersal differences decreased diversity and produced so-called ‘no-analogue’ communities, defined as a novel combination of species that does not currently co-occur. Climate change altered community richness the most when species had narrow niches, when mean community-wide dispersal rates were low and when species differed in dispersal abilities. With high interspecific dispersal variance, the best dispersers tracked climate change, out-competed slower dispersers and caused their extinction. Overall, competition slowed the advance of colonists into newly suitable habitats, creating lags in climate tracking. We predict that climate change will most threaten communities of species that have narrow niches (e.g. tropics), vary in dispersal (most communities) and compete strongly. Current forecasts probably underestimate climate change impacts on biodiversity by neglecting competition and dispersal differences.

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TL;DR: The life-history traits that cause some hosts to be widespread and resilient might be correlated with those that promote infection and transmission by some pathogens, supporting the notion that the dilution effect might be widespread among disease systems.
Abstract: The dynamics of infectious diseases can be affected by genetic diversity within host populations, species diversity within host communities, and diversity among communities. In principle, diversity can either increase or decrease pathogen transmission and disease risk. Theoretical models and laboratory experiments have demonstrated that a dilution effect (decreased disease risk with increasing diversity) can occur under a wide range of conditions. Field studies of plants, aquatic invertebrates, amphibians, birds, and mammals demonstrate that the phenomenon indeed does occur in many natural systems. A dilution effect is expected when (a) hosts differ in quality for pathogens or vectors; (b) higher quality hosts tend to occur in species-poor communities, whereas lower quality hosts tend to occur in more diverse communities; and (c) lower quality hosts regulate abundance of high-quality hosts or of vectors, or reduce encounter rates between these hosts and pathogens or vectors. Although these conditions characterize many disease systems, our ability to predict when and where the dilution effect occurs remains poor. The life-history traits that cause some hosts to be widespread and resilient might be correlated with those that promote infection and transmission by some pathogens, supporting the notion that the dilution effect might be widespread among disease systems. Criticisms of the dilution effect have focused on whether species richness or species composition (both being metrics of biodiversity) drives disease risk. It is well established, however, that changes in species composition correlate with changes in species richness, and this correlation could explain why the dilution effect appears to be a general phenomenon.

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TL;DR: A global analysis to disentangle the global determinants of diversity and community composition for ectomycorrhizal fungi-microbial symbionts that play key roles in plant nutrition in most temperate and many tropical forest ecosystems provides useful biogeographic and ecological hypotheses for explaining the distribution of fungi.
Abstract: Global species richness patterns of soil micro-organisms remain poorly understood compared to macro-organisms. We use a global analysis to disentangle the global determinants of diversity and community composition for ectomycorrhizal (EcM) fungi—microbial symbionts that play key roles in plant nutrition in most temperate and many tropical forest ecosystems. Host plant family has the strongest effect on the phylogenetic community composition of fungi, whereas temperature and precipitation mostly affect EcM fungal richness that peaks in the temperate and boreal forest biomes, contrasting with latitudinal patterns of macro-organisms. Tropical ecosystems experience rapid turnover of organic material and have weak soil stratification, suggesting that poor habitat conditions may contribute to the relatively low richness of EcM fungi, and perhaps other soil biota, in most tropical ecosystems. For EcM fungi, greater evolutionary age and larger total area of EcM host vegetation may also contribute to the higher diversity in temperate ecosystems. Our results provide useful biogeographic and ecological hypotheses for explaining the distribution of fungi that remain to be tested by involving next-generation sequencing techniques and relevant soil metadata.

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TL;DR: The use of molecular tools to measure whole community dispersal provides a direct confirmation for a key mechanism underlying island biogeography theory and has the potential to make microbial systems a model for understanding the role of dispersal in ecological theory.
Abstract: Dispersal plays a prominent role in most conceptual models of community assembly. However, direct measurement of dispersal across a whole community is difficult at ecologically relevant spatial scales. For cryptic organisms, such as fungi and bacteria, the scale and importance of dispersal limitation has become a major point of debate. We use an experimental island biogeographic approach to measure the effects of dispersal limitation on the ecological dynamics of an important group of plant symbionts, ectomycorrhizal fungi. We manipulated the isolation of uncolonized host seedlings across a natural landscape and used a range of molecular techniques to measure the dispersal rates of ectomycorrhizal propagules and host colonization. Some species were prolific dispersers, producing annual spore loads on the order of trillions of spores per km 2 . However, fungal propagules reaching host seedlings decreased rapidly with increasing distance from potential spore sources, causing a concomitant reduction in ectomycorrhizal species richness, host colonization and host biomass. There were also strong differences in dispersal ability across species, which correlated well with the predictable composition of ectomycorrhizal communities associated with establishing pine forest. The use of molecular tools to measure whole community dispersal provides a direct confirmation for a key mechanism underlying island biogeography theory and has the potential to make microbial systems a model for understanding the role of dispersal in ecological theory.

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TL;DR: The distribution of small-ranged species is concentrated near tropical coasts, where moderation of the climate in topographically complex areas creates cloud forests and stable local conditions, and thereby the role of these places as cradles of biodiversity.
Abstract: Avian faunas vary greatly among montane areas; those at high latitudes are biologically impoverished, whereas those of some low-latitude mountains are biologically very complex. Their high level of species richness is caused by the aggregation of many small-ranged species, which has been difficult to explain from purely macroecological models focusing on contemporary ecological processes. Because the individual mountain tracts harbor species that represent different evolutionary trajectories, it seems plausible to relate these species assemblages to high persistence (or absence of extinction) in addition to high levels of speciation. The distribution of small-ranged species is concentrated near tropical coasts, where moderation of the climate in topographically complex areas creates cloud forests and stable local conditions. The stability underpins specialization and resilience of local populations, and thereby the role of these places as cradles of biodiversity.

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TL;DR: Survey of endophytic fungal communities in leaves of a single tree species on Mauna Loa Volcano suggests that microbial diversity may be even greater than has been assumed and that broad-scale environmental controls such as temperature and rainfall can structure eukaryotic microbial diversity.
Abstract: We surveyed endophytic fungal communities in leaves of a single tree species (Metrosideros polymorpha) across wide environmental gradients (500–5,500 mm of rain/y; 10–22 °C mean annual temperature) spanning short geographic distances on Mauna Loa Volcano, Hawai’i. Using barcoded amplicon pyrosequencing at 13 sites (10 trees/site; 10 leaves/tree), we found very high levels of diversity within sites (a mean of 551 ± 134 taxonomic units per site). However, among-site diversity contributed even more than did within-site diversity to the overall richness of more than 4,200 taxonomic units observed in M. polymorpha, and this among-site variation in endophyte community composition correlated strongly with temperature and rainfall. These results are consistent with suggestions that foliar endophytic fungi are hyperdiverse. They further suggest that microbial diversity may be even greater than has been assumed and that broad-scale environmental controls such as temperature and rainfall can structure eukaryotic microbial diversity. Appropriately constrained study systems across strong environmental gradients present a useful means to understand the environmental factors that structure the diversity of microbial communities.

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TL;DR: In this paper, an acoustic analysis of choruses produced by animal communities was carried out to assess α and β diversity in three woodland habitats: a mature forest, a young forest and a forest-cropland ecotone within the Parc Naturel Regional of Haute-Vallee de Chevreuse (France).

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TL;DR: The novel methodology allows the discrimination of different causes of beta-diversity patterns along latitudinal, biogeographic or ecological gradients, by estimating correctly the relative contributions of replacement and richness differences.
Abstract: Aim To determine the relative contribution of species replacement and species richness differences to the emergence of beta-diversity patterns. Innovation A novel method that disentangles all compositional differences (βcc, overall beta diversity) in its two components, species replacement (β-3) and species richness differences (βrich) is proposed. The performance of the method was studied with ternary plots, which allow visualization of the influence of the relative proportions of shared and unique species of two sites over each metric. The method was also tested in different hypothetical gradients and with real datasets. The novel method was compared with a previous proposal based on the partitioning of overall compositional differences (βsor) in replacement (βsim) and nestedness (βnes). The linear response of βcc contrasts with the curvilinear response of βsor to linear gradients of dissimilarity. When two sites did not share any species, βsim was always 1 and β-3 only reached 1 when the number of exclusive species of both sites was equal. β-3 remained constant along gradients of richness differences with constant replacement, while βsim decreased. βrich had a linear response to a linear gradient of richness differences with constant species replacement, whereas βnes exhibited a hump-shaped response. Moreover, βsim > βnes when clearly almost all species of one site were lost, whereas β-3 < βrich in the same circumstances. Main conclusions The behaviour of the partition of βcc into β-3 and βrich is consistent with the variation of replacement and richness differences. The partitioning of βsor into βsim and βnes overestimates the replacement component and underestimates richness differences. The novel methodology allows the discrimination of different causes of beta-diversity patterns along latitudinal, biogeographic or ecological gradients, by estimating correctly the relative contributions of replacement and richness differences.

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TL;DR: It is shown that meiofaunal diversity has been greatly underestimated by 18S eDNA surveys and that the use of COI provides a better estimate of diversity, while estimates of species richness using COI were robust among three of four commonly used delimitation metrics, whereas estimates using 18S varied widely with the different metrics.
Abstract: Molecular tools have revolutionized the exploration of biodiversity, especially in organisms for which traditional taxonomy is difficult, such as for microscopic animals (meiofauna). Environmental (eDNA) metabarcode surveys of DNA extracted from sediment samples are increasingly popular for surveying biodiversity. Most eDNA surveys use the nuclear gene-encoding small-subunit rDNA gene (18S) as a marker; however, different markers and metrics used for delimiting species have not yet been evaluated against each other or against morphologically defined species (morphospecies). We assessed more than 12,000 meiofaunal sequences of 18S and of the main alternatively used marker [Cytochrome c oxidase subunit I (COI) mtDNA] belonging to 55 datasets covering three taxonomic ranks. Our results show that 18S reduced diversity estimates by a factor of 0.4 relative to morphospecies, whereas COI increased diversity estimates by a factor of 7.6. Moreover, estimates of species richness using COI were robust among three of four commonly used delimitation metrics, whereas estimates using 18S varied widely with the different metrics. We show that meiofaunal diversity has been greatly underestimated by 18S eDNA surveys and that the use of COI provides a better estimate of diversity. The suitability of COI is supported by cross-mating experiments in the literature and evolutionary analyses of discreteness in patterns of genetic variation. Furthermore its splitting of morphospecies is expected from documented levels of cryptic taxa in exemplar meiofauna. We recommend against using 18S as a marker for biodiversity surveys and suggest that use of COI for eDNA surveys could provide more accurate estimates of species richness in the future.

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TL;DR: The intermediate disturbance hypothesis (IDH) predicts a hump-shaped pattern between community diversity and disturbance, and is central to understanding patterns of species diversity, which is examined in the context of alien plant invasions to suggest a range of strategies can be used to manage diversity.
Abstract: The intermediate disturbance hypothesis (IDH) predicts a hump-shaped pattern between community diversity and disturbance, and is central to understanding patterns of species diversity. Here, we examine IDH in the context of alien plant invasions. IDH can offer insight into the role of disturbance in facilitating plant invasions and the effect of these invasions on floristic diversity. Early stages of succession are most susceptible to invasion because resources and colonisation opportunities are elevated after disturbance. This trend is accentuated by human-mediated dispersal, a bias towards early successional species in the alien species pool, the tendency for fast-growing species to profit most from enemy release, and increased disturbance levels in human-modified habitats. Human disturbance, coupled with plant introductions, extends the diversity–disturbance curve and shifts peak diversity towards higher disturbance levels. However, invasive aliens can reduce native diversity at the community scale, especially in mid succession where competitive interactions structure communities. Certain invasive plants may have higher impacts because they overcome some life history tradeoffs as a result of their association with humans or novel evolutionary histories, e.g. enemy release. This may directly or indirectly (e.g. through plastic reallocation of resources from defence into growth) enable invasive plants to colonise earlier or persist into later stages of succession. By modifying disturbance regimes, invaders that transform the environment may also interfere with succession and precipitate low diversity communities. Low introduction rates of late successional species may currently limit impacts of aliens under infrequent disturbance. IDH is a useful framework for understanding ecological communities. However, because of the novel evolutionary histories of alien species and the anthropogenic context in which they invade, disturbance levels that maximise total diversity in invaded communities can differ from those that maximise native diversity. Joint consideration of IDH and alien invasion patterns suggests a range of strategies can be used to manage diversity.

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TL;DR: Using a 10-year data set on Costa Rican birds, it is found that low-intensity agriculture sustained β-diversity across large scales on a par with forest, and point the way towards multi-functional agricultural systems that maintain agricultural productivity while simultaneously conserving biodiversity.
Abstract: Biodiversity is declining from unprecedented land conversions that replace diverse, low-intensity agriculture with vast expanses under homogeneous, intensive production. Despite documented losses of species richness, consequences for β-diversity, changes in community composition between sites, are largely unknown, especially in the tropics. Using a 10-year data set on Costa Rican birds, we find that low-intensity agriculture sustained β-diversity across large scales on a par with forest. In high-intensity agriculture, low local (α) diversity inflated β-diversity as a statistical artefact. Therefore, at small spatial scales, intensive agriculture appeared to retain β-diversity. Unlike in forest or low-intensity systems, however, high-intensity agriculture also homogenised vegetation structure over large distances, thereby decoupling the fundamental ecological pattern of bird communities changing with geographical distance. This ~40% decline in species turnover indicates a significant decline in β-diversity at large spatial scales. These findings point the way towards multi-functional agricultural systems that maintain agricultural productivity while simultaneously conserving biodiversity.