A review of methods for analysing spatial and temporal patterns in coastal water quality

Coastal environments contain some of the marine world's most important ecosystems and represent significant resources for human industry and recreation. Water quality in the coastal environment is extremely important for a number of reasons from the protection of marine organisms and the well being of marine ecosystems to the health of people in the region and the safety of industries such as aquaculture. As a result it is essential that environmental health in coastal environments is monitored. Traditional monitoring methods include assessment of biological indices or direct measurements of water quality, which are based on in situ data collection and hence are often spatially or temporally limited. Remote sensing imagery is increasingly used as a rich source of spatial information, providing more detailed coverage then other methods. But the complexity of information in the imagery requires new analysis techniques that allow us to identify the components and possible causes of spatial and temporal variability.This paper presents a review of methods to analyse spatial and temporal variations in remote sensing data of coastal water quality and discusses and compares these methods and the outcomes they achieve. Selected techniques are illustrated by using a sample dataset of MODIS chlorophyll-a imagery. We consider classification methods (cluster analysis, discriminant analysis) that may be used in exploratory, confirmatory and predictive ways, methods that summarize and identify patterns within complex datasets (factor analysis, principal components analysis, self-organizing maps), and techniques that explicitly analyse spatial relationships (the semivariogram and geographically weighted regression). Each technique has a different purpose and addresses different questions. This review identifies how these methods can be utilized to address water quality variability in order to foster a wider application of such techniques for coastal water quality assessment and monitoring.     

How does dune morphology shape coastal EC habitats occurrence? A remote sensing approach using airborne LiDAR on the Mediterranean coast

We examined the relationship between coastal habitats (sensu European Union Habitats Directive) and local dune morphology along a Mediterranean coastal dune system by integrating field collected vegetation data and remotely sensed imagery. Specifically, we described the morphological profile of each EC habitat based on the morphological variables that are most likely to affect their occurrence, including elevation, slope, curvature, northness, eastness and sea distance. In addition, we assessed the role and strength of each morphological variable in determining the occurrence of EC habitats.We used 394 random vegetation plots representative of six EC habitats (Habitat 1210: “Annual vegetation of drift lines”; Habitat 2110: “Embryonic shifting dunes”; Habitat 2120: “Shifting dunes along the shoreline with Ammophila arenaria”; Habitat 2210 and 2230: “Crucianellion maritimae fixed beach dunes” and “Malcolmietalia dune grasslands”; Habitat 2250: “Coastal dunes with Juniperus spp.”; Habitat 2260: “Cisto-Lavanduletalia dune sclerophyllous scrubs”) found along the Tyrrhenian coast of central Italy. We derived each morphological variable from a DTM (Digital Terrain Model) obtained from 2-m resolution LiDAR (Light Detection And Range) images. The mean value of each variable was calculated at different spatial scales using buffer areas of increasing radius (2 m, 4 m, 8 m) around each vegetation plot. Mean morphological values for each EC habitat were compared using Kruskal-Wallis rank test. The role and strength of the relationship between habitat type and the morphological variables were assessed using Generalized Linear Models.EC habitats occur differentially across dune morphology, and the role and strength of each morphological variable define habitat specificity. Dune elevation and sea distance were determined to be the key factors in shaping EC habitat occurrence along this section of the Mediterranean coast. Identification of the close relationship between habitat type and morphological variables deriving from airborne LiDAR imagery points to the high potential of such remote sensing tool for analyzing and monitoring the integrity of coastal dune ecosystems. As airborne LiDAR enables the rapid collection of extremely accurate topographic data over large areas, it also offers useful information for the management of these threatened and fragile ecosystems.     

Natural history museum collections provide information on phenological change in British butterflies since the late-nineteenth century

Museum collections have the potential to provide valuable information on the phenological response of organisms to climate change. This is particularly useful for those species for which few data otherwise exist, but also to extend time series to the period before other observational data are available. To test this potential, we analysed data from 2,630 specimens of four species of British butterflies (Anthocharis cardamines, Hamearis lucina, Polyommatus bellargus and Pyrgus malvae), collected from 1876 to 1999 and stored in the Natural History Museum, London, UK (NHM). In A. cardamines, first-generation P. bellargus and P. malvae, we found that there was a strong significant negative relationship between spring temperature and 10th percentile collection dates, which approximates mean first appearance date, and median collection date, which approximates mean flight date. In all four species, there was a significant negative relationship between the 10th percentile collection date and the length of the collection period, which approximates flight period. In second-generation P. bellargus, these phenological measurements were correlated with summer temperature. We found that the rates of phenological response to temperature, based on NHM data, were similar to, or somewhat greater than, those reported for other organisms based on observational data covering the last 40 years. The lower rate of phenological response, and the significant influence of February rather than March or April temperatures, in recent decades compared with data from earlier in the twentieth century may indicate that early emerging British butterfly species are currently approaching the limits of phenological advancement in response to recent climate warming.     

Application of specific exergy to macrophytes as an integrated index of environmental quality for coastal lagoons

This work aims to explore the application of specific exergy on macrophytes as an integrated index to assess ecosystem health in coastal lagoons, and to analyse its possible application in view of the implementation of the Water Framework Directive of the European Union (European Commission, 2000). Exergy and specific exergy are calculated as a function of the biomass multiplied by a weighting factor (β), which express the quantity of information embedded in the biomass. Until now only few general values of β have been published. Therefore, we have extended the calculation of β-values for 244 seaweed and seagrass species which are common in Mediterranean coastal lagoons. The estimated β-values have been then compared with macrophyte-based indicators which have been recently developed, namely the Ecological Evaluation Index and the IFREMER classification scheme. Overall, we found a good agreement between β-values and macrophyte indicators. Furthermore, the specific exergy was calculated for 71 sites in coastal lagoons of Southern France and compared with existing classification schemes, finding a good agreement between them. As a result, we argue that specific exergy may be used as an integrated index that is able to synthesize and complement existing approaches.     

Birds of Antioquia: Georeferenced database of specimens from the Colección de Ciencias Naturales del Museo Universitario de la Universidad de Antioquia (MUA)

The department of Antioquia, Colombia, lies in the northwestern corner of South America and provides a biogeographical link among divergent faunas, including Caribbean, Andean, Pacific and Amazonian. Information about the distribution of biodiversity in this area is of relevance for academic, practical and social purposes. This data paper describes the dataset containing all bird specimens deposited in the Colección de Ciencias Naturales del Museo Universitario de la Universidad de Antioquia (MUA). We curated all the information associated with the bird specimens, including the georeferences and taxonomy, and published the database through the Global Biodiversity Information Facility network. During this process we checked the species identification and existing georeferences and completed the information when possible. The collection holds 663 bird specimens collected between 1940 and 2011. Even though most specimens are from Antioquia (70%), the collection includes material from several other departments and one specimen from the United States. The collection holds specimens from three endemic and endangered species (Coeligena orina, Diglossa gloriossisima, and Hypopirrhus pyrohipogaster), and includes localities poorly represented in other collections. The information contained in the collection has been used for biodiversity modeling, conservation planning and management, and we expect to further facilitate these activities by making it publicly available.     

Stranding Events of Kogia Whales along the Brazilian Coast

The genus Kogia, which comprises only two extant species, Kogia sima and Kogia breviceps, represents one of the least known groups of cetaceans in the global ocean. In some coastal regions, however, stranding events of these species have been relatively common over the last decades. Stranding provides the opportunity to investigate the biology of these cetaceans and to explore the epidemiological aspects associated with the mortality of the organisms found on the beach. A number of disturbances (including pelagic fisheries, chemical pollution, boat strikes, and noise pollution) have been confirmed to pose a particular threat to the Kogia species. However, no study has yet investigated potential relationships between environmental conditions and stranding events. Here we analyse how a collection of environmental, physical, and biological variables, such as wind, sea surface temperature (SST), water depth, and chlorophyll-a, correlate to Kogia stranding events along the Brazilian coast. The results of our statistical analyses suggest that K. sima is more likely found in warm tropical waters, which provide an explanation for the high frequency of stranding in northeastern Brazilian coast. In contrast, K. breviceps appears to have a preference for temperate and productive waters. Wind speed results to be also an important factor for predicting Kogia strandings in Brazilian coast. Additionally, literature information in combination with our own data and analyses of stomach contents confirms that oceanic cephalopods constitute the primary nutritional source of both Kogia species. By using the available information as a qualitative proxy for habitat preference and feeding ecology, our study provides a novel and comprehensive assessment of Kogia stranding data in relation to environmental conditions along the Brazilian coast.     

Genetic Structure of Wild Chinook Salmon Populations of Southeast Alaska and Northern British Columbia

Allozyme variation was used to examine population genetic structure of adult chinook salmon, Oncorhynchus tshawytscha, collected between 1988 and 1993 from 22 spawning locations in Southeast Alaska and northern British Columbia. Thirty-five loci and two pairs of isoloci were variable, and of these, 25 loci and one pair of isoloci expressed the most abundant allele with a frequency of less than or equal to 0.95 in at least one collection. A neighbor-joining (NJ) tree of genetic distances defined five regional groups: (1) King Salmon River (the only island collection), which has large allelic frequency differences from other populations in this study; (2) heterogeneous coastal populations from southern southeast Alaska; (3) transmountain collections from the Taku and Stikine Rivers on the eastern side of the coastal mountain range; (4) Chilkat River in northern Southeast Alaska; and (5) northern coastal Southeast Alaska, which consists of the Situk River and the Klukshu River, a tributary of the Alsek River. A second NJ tree that included collections from the Yukon River and British Columbia did not reveal any strong genetic similarity between Southeast Alaska and the Yukon River. The data suggest that Southeast Alaska may have been colonized from both northern and southern refugia following the last glaciation — a period of sufficient time to allow for isolation by distance to occur.     

Demographic and Genetic Patterns of Variation among Populations of Arabidopsis thaliana from Contrasting Native Environments

Background

Understanding the relationship between environment and genetics requires the integration of knowledge on the demographic behavior of natural populations. However, the demographic performance and genetic composition of Arabidopsis thaliana populations in the species'' native environments remain largely uncharacterized. This information, in combination with the advances on the study of gene function, will improve our understanding on the genetic mechanisms underlying adaptive evolution in A. thaliana.

Methodology/Principal Findings

We report the extent of environmental, demographic, and genetic variation among 10 A. thaliana populations from Mediterranean (coastal) and Pyrenean (montane) native environments in northeast Spain. Geographic, climatic, landscape, and soil data were compared. Demographic traits, including the dynamics of the soil seed bank and the attributes of aboveground individuals followed over a complete season, were also analyzed. Genetic data based on genome-wide SNP markers were used to describe genetic diversity, differentiation, and structure. Coastal and montane populations significantly differed in terms of environmental, demographic, and genetic characteristics. Montane populations, at higher altitude and farther from the sea, are exposed to colder winters and prolonged spring moisture compared to coastal populations. Montane populations showed stronger secondary seed dormancy, higher seedling/juvenile mortality in winter, and initiated flowering later than coastal populations. Montane and coastal regions were genetically differentiated, montane populations bearing lower genetic diversity than coastal ones. No significant isolation-by-distance pattern and no shared multilocus genotypes among populations were detected.

Conclusions/Significance

Between-region variation in climatic patterns can account for differences in demographic traits, such as secondary seed dormancy, plant mortality, and recruitment, between coastal and montane A. thaliana populations. In addition, differences in plant mortality can partly account for differences in the genetic composition of coastal and montane populations. This study shows how the interplay between variation in environmental, demographic, and genetic parameters may operate in natural A. thaliana populations.     

On the Potential of Surfers to Monitor Environmental Indicators in the Coastal Zone

The social and economic benefits of the coastal zone make it one of the most treasured environments on our planet. Yet it is vulnerable to increasing anthropogenic pressure and climate change. Coastal management aims to mitigate these pressures while augmenting the socio-economic benefits the coastal region has to offer. However, coastal management is challenged by inadequate sampling of key environmental indicators, partly due to issues relating to cost of data collection. Here, we investigate the use of recreational surfers as platforms to improve sampling coverage of environmental indicators in the coastal zone. We equipped a recreational surfer, based in the south west United Kingdom (UK), with a temperature sensor and Global Positioning System (GPS) device that they used when surfing for a period of one year (85 surfing sessions). The temperature sensor was used to derive estimates of sea-surface temperature (SST), an important environmental indicator, and the GPS device used to provide sample location and to extract information on surfer performance. SST data acquired by the surfer were compared with data from an oceanographic station in the south west UK and with satellite observations. Our results demonstrate: (i) high-quality SST data can be acquired by surfers using low cost sensors; and (ii) GPS data can provide information on surfing performance that may help motivate data collection by surfers. Using recent estimates of the UK surfing population, and frequency of surfer participation, we speculate around 40 million measurements on environmental indicators per year could be acquired at the UK coastline by surfers. This quantity of data is likely to enhance coastal monitoring and aid UK coastal management. Considering surfing is a world-wide sport, our results have global implications and the approach could be expanded to other popular marine recreational activities for coastal monitoring of environmental indicators.     

Addition of Escherichia coli K-12 Growth Observation and Gene Essentiality Data to the EcoCyc Database

The sets of compounds that can support growth of an organism are defined by the presence of transporters and metabolic pathways that convert nutrient sources into cellular components and energy for growth. A collection of known nutrient sources can therefore serve both as an impetus for investigating new metabolic pathways and transporters and as a reference for computational modeling of known metabolic pathways. To establish such a collection for Escherichia coli K-12, we have integrated data on the growth or nongrowth of E. coli K-12 obtained from published observations using a variety of individual media and from high-throughput phenotype microarrays into the EcoCyc database. The assembled collection revealed a substantial number of discrepancies between the high-throughput data sets, which we investigated where possible using low-throughput growth assays on soft agar and in liquid culture. We also integrated six data sets describing 16,119 observations of the growth of single-gene knockout mutants of E. coli K-12 into EcoCyc, which are relevant to antimicrobial drug design, provide clues regarding the roles of genes of unknown function, and are useful for validating metabolic models. To make this information easily accessible to EcoCyc users, we developed software for capturing, querying, and visualizing cellular growth assays and gene essentiality data.     

Geographic range and structure of cryptic genetic diversity among Pacific North American populations of the non-native amphipod Grandidierella japonica

Reconstructing the invasion history of aquatic invasive species can enhance understanding of invasion risks by recognizing areas most susceptible to invasion and forecasting future spread based on past patterns of population expansion. Here we reconstruct the invasion history of the Japanese amphipod Grandidierella japonica Stephensen 1938 combining information from historical collection data with molecular genetic data to better understand post-invasion range expansion and anthropogenic connectivity across the Pacific coast of North America. Compilation of collection data from bays and estuaries of the Pacific North American coast show many new localities have been colonized in the last two decades, moving outward from harbors and bays with high commercial traffic into smaller coastal locations dominated by local recreational traffic. DNA barcode sequence data for G. japonica reveals two distinct clades: one found in San Francisco Bay and sites to the north, and one also found in San Francisco Bay and sites to the south. The two clades differ by an average 7.28 % genetic distance, large enough to consider these invasive amphipods two separate species. Both northern and southern clades exhibit low levels of genetic diversity, suggesting a single introduction event for each. The presence of cryptic diversity within this invasive amphipod highlights the need for more extensive study of the invasive and native populations of aquatic invasive invertebrates to address questions of taxonomy, diversity, and invasion history.     

Comparison of low salinity tolerance in Callinectes sapidus Rathbun and Callinectes similis Williams postlarvae upon entry into an estuary

Planktonic larvae of estuarine crabs are commonly exported to the continental shelf for development and then return to coastal and estuarine areas as postlarvae (megalopae). Megalopae returning to estuaries must be adapted to survive in brackish water whereas those of coastally distributed species should not need such adaptations. We investigated 1) whether megalopae of the estuarine crab Callinectes sapidus and the coastal crab Callinectes similis undergo changes in salinity tolerance upon entry into an estuary and 2) what factors induce those changes. Megalopae were collected at a coastal site and a nearby estuarine site and exposed to a range of salinities (5, 10, 15, 20 and 30) for 6 h. Percent survival was determined after 24 h reintroduction to the collection site water. We also investigated 1) whether increased salinity tolerance was induced by reduced salinity or estuarine chemical cues, 2) the time to acclimation and 3) the salinity necessary for acclimation. C. sapidus megalopae from the estuarine site were more likely to survive exposure to low salinities than those from the coastal site. C. sapidus megalopae from the coastal site exhibited increased survival after acclimation to salinities of 27 and 23 for 12 h. Estuarine chemical cues had no effect on salinity tolerance. C. similis megalopae were less likely to survive at low salinities and did not exhibit an acclimation response upon exposure to reduced salinities. These results suggest that megalopae of C. sapidus are physiologically adapted to recruit to estuaries whereas megalopae of C. similis are unable to acclimate to low salinity conditions.     

DNA repair defects ascribed to pby1 are caused by disruption of Holliday junction resolvase Mus81-Mms4

PBY1 continues to be linked with DNA repair through functional genomics studies in yeast. Using the yeast knockout (YKO) strain collection, high-throughput genetic interaction screens have identified a large set of negative interactions between PBY1 and genes involved in genome stability. In drug sensitivity screens, the YKO collection pby1Δ strain exhibits a sensitivity profile typical for genes involved in DNA replication and repair. We show that these findings are not related to loss of Pby1. On the basis of genetic interaction profile similarity, we pinpoint disruption of Holliday junction resolvase Mus81-Mms4 as the mutation responsible for DNA repair phenotypes currently ascribed to pby1. The finding that Pby1 is not a DNA repair factor reconciles discrepancies in the data available for PBY1, and indirectly supports a role for Pby1 in mRNA metabolism. Data that has been collected using the YKO collection pby1Δ strain confirms and expands the chemical-genetic interactome of MUS81-MMS4.     

Invasive moss alters patterns in life-history traits and functional diversity of spiders and carabids

Invasive plants can modify terrestrial habitats and affect the natural faunal composition. In acidic coastal dunes the invasive moss Campylopus introflexus can form dense carpets that largely replace native vegetation. As shown in a previous study, moss invasion affects habitat structure and ground-dwelling arthropod diversity. We suggested that including the functional diversity concept in the analysis of moss invasion impacts may offer further insights. We used pitfall trap data to compare trait composition and functional diversity of spiders and carabids in (a) invaded, moss-rich (C. introflexus) and (b) native, lichen-rich (Cladonia spp.) acidic coastal dunes. Moss invasion induced shifts in the trait values body size and feeding preference (carabids) and hunting mode (spiders): Species were smaller in native sites, and the percentages of web-building spiders and phytophagous beetles were reduced in invaded sites. Furthermore, moss invasion led to a more heterogeneous trait composition for spiders, and changed functional diversity of both arthropod groups, although with the opposite effects: While spiders were functionally more diverse in invaded sites, moss invasion reduced carabid beetles’ functional diversity. We also observed changes in the relationship between species richness and functional diversity that indicate a high functional similarity for spiders but a lower one for carabid beetles in native grey dunes. C. introflexus invasion not only alters the arthropod diversity and assemblage structure of endangered acidic coastal dunes but also interferes at a functional level. These results provide further insight into the way plant invasions might alter the structure and function of ecosystems.     

Genetic characterization of the scyphozoan jellyfish Aurelia spp. in Chinese coastal waters using mitochondrial markers

Blooms of the moon jellyfish Aurelia spp. have occurred in the harbors and coastal waters around the world. The phylogenetic relationship and genetic characterization of Aurelia spp. was determined along the Chinese coastal waters based on sequences of the mitochondrial cytochrome c oxidase subunit I (COI) gene. The molecular analysis confirmed that all samples collected in Chinese coastal waters were Aurelia sp.1. We also analyzed the phylogenetic and population genetic structure of Aurelia sp.1 using the newly generated sequences supplemented with existing data from previous studies. The phylogenetic analyses of the COI regions did not support geographically restricted groups among the global samples of Aurelia sp.1. Analysis of molecular variance (AMOVA) indicated a complex genetic population structure and pattern of connectivity. Populations of Aurelia sp.1 were highly structured between most sampling sites over distances as small as 100 km (Rizhao and Qingdao) in certain cases. However, non-significant pairwise F_ST values were also observed between short geographic distances (Yantai, Rongcheng and Qingdao) and relatively distant sampling sites (Caofeidian, Rizhao and Japan). The life-cycle characteristics, together with the prevailing ocean currents in this region and possible anthropogenic introduction, were proposed and discussed as the main factors that determined the genetic patterns of Aurelia sp.1.     

A method to evaluate body length of live aquatic vertebrates using digital images

Traditional methods to measure body lengths of aquatic vertebrates rely on anesthetics, and extended handling times. These procedures can increase stress, potentially affecting the animal''s welfare after its release. We developed a simple procedure using digital images to estimate body lengths of coastal cutthroat trout (Oncorhynchus clarkii clarkii) and larval coastal giant salamander (Dicamptodon tenebrosus). Images were postprocessed using ImageJ2. We measured more than 900 individuals of these two species from 200 pool habitats along 9.6 river kilometers. The percent error (mean ± SE) of our approach compared to the use of a traditional graded measuring board was relatively small for all length metrics of the two species. Total length of trout was −2.2% ± 1.0. Snout–vent length and total length of larval salamanders was 3.5% ± 3.3 and −0.6% ± 1.7, respectively. We cross‐validated our results by two independent observers that followed our protocol to measure the same animals and found no significant differences (p > .7) in body size distributions for all length metrics of the two species. Our procedure provides reliable information of body size reducing stress and handling time in the field. The method is transferable across taxa and the inclusion of multiple animals per image increases sampling efficiency with stored images that can be reviewed multiple times. This practical tool can improve data collection of animal size over large sampling efforts and broad spatiotemporal contexts.     

Morphometric and molecular differences among Calvertius tuberosus (Coleoptera: Curculionidae) populations associated with Andean and coastal populations of Araucaria araucana in the La Araucanía Region,Chile

Calvertius tuberosus (Curculionidae) lives exclusively on Araucaria araucana trees (commonly known as pehuen) in southern Chile. In this study, morphometric and molecular genetic analyses of Andean and coastal populations of C. tuberosus were performed to evaluate evolutionary divergence associated with the discontinuity of the Araucaria forest between the coastal and Andean regions. Specimens of C. tuberosus were collected in Nahuelbuta National Park, Villa Las Araucarias, and Malalcahuello National Reserve and were classified and stored at the Animal Biotechnology Researching Laboratory (LINBA), University of La Frontera, Temuco, Chile. Thirteen morphometric parameters and the expression patterns of ISSR (inter-simple sequence repeat) markers were analyzed. Morphometric data revealed high phenotypic similarity between coastal populations. The genetic analysis revealed a high similarity between coastal populations (genetic identity, 93%), which were differentiated from the Andean population (genetic identity, 84%). This study contributes new genotypic and phenotypic data for the C. tuberosus populations in forest ecosystems of A. araucana, and clarifies the associations between these characteristics and the geographic distributions of populations.     

A comparative approach of assessing ecological status in two coastal areas of Eastern Mediterranean

This work presents an assessment of the ecological quality status of two marine coastal areas in the Aegean Sea (Greece, Eastern Mediterranean) based on the benthic macroinvertebrate quality element. S. Evvoikos and Thessaloniki gulfs, two coastal areas subjected to slight and heavier anthropogenic pressures, were selected to test the biotic index Bentix developed for the assessment of the ecological status. Other ecological indicators, such as the Shannon diversity index (H′), the species richness (S) and the AMBI biotic index were also applied and evaluated comparatively. Faunistic data were also used to interpret results. The resulting classification was validated with the use of physicochemical parameters and pressure information. This work also provides an insight into the structure of the Bentix classification scheme within the scope of its use in Water Framework Directive (WFD) implementation.     

Multiple-Locus Sequence Typing Analysis of Bacillus cereus and Bacillus thuringiensis Reveals Separate Clustering and a Distinct Population Structure of Psychrotrophic Strains

We used multilocus sequence typing (MLST) to characterize phylogenetic relationships for a collection of Bacillus cereus group strains isolated from forest soil in the Paris area during a mild winter. This collection contains multiple strains isolated from the same soil sample and strains isolated from samples from different sites. We characterized 115 strains of this collection and 19 other strains based on the sequences of the clpC, dinB, gdpD, panC, purF, and yhfL loci. The number of alleles ranged from 36 to 53, and a total of 93 allelic profiles or sequence types were distinguished. We identified three major strain clusters—C, T, and W—based on the comparison of individual gene sequences or concatenated sequences. Some less representative clusters and subclusters were also distinguished. Analysis of the MLST data using the concept of clonal complexes led to the identification of two, five, and three such groups in clusters C, T, and W, respectively. Some of the forest isolates were closely related to independently isolated psychrotrophic strains. Systematic testing of the strains of this collection showed that almost all the strains that were able to grow at a low temperature (6°C) belonged to cluster W. Most of these strains, including three independently isolated strains, belong to two clonal complexes and are therefore very closely related genetically. These clonal complexes represent strains corresponding to the previously identified species Bacillus weihenstephanensis. Most of the other strains of our collection, including some from the W cluster, are not psychrotrophic. B. weihenstephanensis (cluster W) strains appear to comprise an effectively sexual population, whereas Bacillus thuringiensis (cluster T) and B. cereus (cluster C) have clonal population structures.