Geometry of biodiversity patterning: assemblages of benthic macroinvertebrates at tributary confluences

We assessed whether tributaries in upland catchments (=watersheds) affected assemblages of benthic macroinvertebrates in mainstems, as has been reported in northern hemisphere systems. Eight confluences of small to medium streams (stream orders 1–4, 2.2–10.8 m wide) were studied in the Acheron River basin in Victoria, Australia. For each confluence, two transects were sampled at each of five zones relative to the confluence: two zones upstream in the mainstem, one zone upstream in the tributary, one zone at the confluence and one zone downstream in the mainstem. Surveys were conducted in both high-flow and low-flow conditions. In mainstems, there was no change in macroinvertebrate density, taxonomic richness or functional feeding group composition downstream relative to upstream of the confluences. While tributaries statistically had distinctive benthic macroinvertebrate assemblages compared to mainstems, these distinctions were small. In low flows, densities in tributaries were substantially lower than in mainstems, but densities during high flows were more similar (albeit only about one-third as high as in low flow) in tributaries and mainstems. An inverse pattern was evident for taxonomic richness, where richness in tributaries and mainstems was similar in low flows but was greater in mainstems than in tributaries in high flows. We found little evidence of tributary effects in macroinvertebrate assemblages in this basin, which is at odds with some previous results from other continents. To explain this divergence, we suggest a conceptual model outlining factors that control variation in effects of tributaries on assemblages of benthic macroinvertebrates in mainstems.     

Estimating bacterial diversity from clone libraries with flat rank abundance distributions

There are a number of parametric and non-parametric methods for estimating diversity. However all such methods employ either the proportional abundance of the most abundant taxon in a sample or require that a specific taxon is sampled more than once. Consequently, the available methods for estimating diversity cannot be applied to samples consisting entirely of singletons, which might be characteristic of some hyperdiverse communities. Here we present a non-parametric method that estimates the probability that a given number of unique taxa would be sampled from a community with a particular diversity. We have applied this approach to a well known data set of 100 unique clones from a sample of Amazonian soil (Borneman and Triplett (1997) Appl Environ Microbiol 63: 2647-2653) and determine the probability that this observation would be made from an environment of a given diversity. On this basis we can state this observation would be very unlikely (P = 0.006) if the soil diversity was less than 10(3), and quite unlikely (P = 0.6) if the diversity was less than 10(4), and probable (P = 0.95) if the diversity was about 10(5). There are essentially no contestable assumptions in our method. Thus we are able to offer almost unequivocal evidence that the bacterial diversity, of at least soils, is very large and a method that may be used to interpret samples consisting entirely of singletons from other hyperdiverse communities.     

Paleodepth determination from Antarctic benthic diatom assemblages

Analysis of modern surface sediments from fjords in the Vestfold Hills (Antarctica) indicates that 58% of the variation in benthic diatom assemblages can be attributed to changes in environmental parameters with water depth. Attenuation of light through the water column is suggested to account for 45% of the variation, and the decrease in substrate grain size with depth possibly accounts for a further 13%. Cluster analysis and principal component analysis were used to objectively circumscribe five floral zones between the surface and 35 m depth. The depth distribution of the benthic diatoms was then used to interpret the paleodepth of relict fjord (Holocene) sediments exposed around Deep Lake in the Death Valley (Vestfold Hills). Paleodepths measured from the sea-ice bench around Deep Lake combined with data from grain size analysis indicate that the relict fjord sediments have no analogue amongst the modern fjord sediments sampled in the Vestfold Hills. Without a comparable modern habitat on which to model the diatom depth zones, however, this study was unable to accurately determine the paleowater depth at Deep Lake using diatoms. Paleodepth determination will be possible using grain size analysis and diatom data when the substrate and light requirements of benthic diatoms are understood.     

Conservation in a cup of water: estimating biodiversity and population abundance from environmental DNA

Three mantras often guide species and ecosystem management: (i) for preventing invasions by harmful species, ‘early detection and rapid response’; (ii) for conserving imperilled native species, ‘protection of biodiversity hotspots’; and (iii) for assessing biosecurity risk, ‘an ounce of prevention equals a pound of cure.’ However, these and other management goals are elusive when traditional sampling tools (e.g. netting, traps, electrofishing, visual surveys) have poor detection limits, are too slow or are not feasible. One visionary solution is to use an organism’s DNA in the environment (eDNA), rather than the organism itself, as the target of detection. In this issue of Molecular Ecology, Thomsen et al. (2012) provide new evidence demonstrating the feasibility of this approach, showing that eDNA is an accurate indicator of the presence of an impressively diverse set of six aquatic or amphibious taxa including invertebrates, amphibians, a fish and a mammal in a wide range of freshwater habitats. They are also the first to demonstrate that the abundance of eDNA, as measured by qPCR, correlates positively with population abundance estimated with traditional tools. Finally, Thomsen et al. (2012) demonstrate that next‐generation sequencing of eDNA can quantify species richness. Overall, Thomsen et al. (2012) provide a revolutionary roadmap for using eDNA for detection of species, estimates of relative abundance and quantification of biodiversity.     

Recent benthic foraminiferal assemblages in deep high-energy environments from the Gulf of Cadiz (Spain)

The benthic environment in the Gulf of Cadiz, north-eastern Atlantic, is strongly affected by the Mediterranean outflow water undercurrent (MOW) which flows northwards along the western Iberian Margin at 500–1500 m water depth. Foraminiferal census counts of living and dead assemblages from 27 surface samples ranging from 103 to 1917 m water depth, and the examination of hard substrates reveal a close correlation of the fauna with the local hydrography and sediment facies. Four different faunal groups are separated by factor analysis of the living fauna. Assemblage 1 contains typical lower slope species and dominates samples from the lower MOW core layer and in the North Atlantic deep water below. Shelf edge foraminifera are common in assemblage 2a which shows the highest proportions in samples from 103 to 272 m. Assemblage 2b is dominated by upper slope species and suspension-feeders that are frequent in the upper MOW core layer and in distal settings between 396 and 901 m. Species from assemblage 3 prefer epibenthic habitats and are recorded with high proportions exclusively in the immediate flow paths of the upper MOW between 496 and 881 m. Colonisation structures and species composition of epibenthic assemblages from the proximal facies largely differ from those in distal settings. In general, epibenthic foraminifers only use elevated substrates under the influence of near-bottom flow. Under high current velocities, epibenthic foraminifers prefer large and heavy objects. They colonise high attachment levels where a maximum yield of advected food particles can be achieved. In distal settings at lower flow velocities, the elevation height does not exceed 20 mm above the surrounding sediment surface. This level is related to a hydrologic transition layer with high concentrations of suspended particles. The comparison of microhabitat preferences and faunal structure under high and low current velocities reveal that substrate stability may be a confining environmental variable for endobenthic and shallow epibenthic foraminifers. The observations also indicate that the preferential settling height of epibenthic foraminifera is related to the highest lateral flux rates of food particles within reach from the sea floor. A dynamic selection of elevated microhabitats is only used by 7.8% of all species recognised in the Gulf of Cadiz area.     

The effect of habitat stability on benthic invertebrate communities: the utility of species abundance distributions

Spatial and temporal patterns in the species abundance distribution of benthic invertebrate communities of 11 freshwater habitats (10 streams and a wind-swept lake shore) were examined with respect to habitat stability. Abundance patterns varied markedly between seasons at most sites. However, mean abundance distributions at 4 of the 5 unstable sites and the 2 most stable sites were dominated by one or two taxa with a large number of rare species, whereas sites of intermediate stability had more equitable distributions. Both the log series and log normal distributions were statistically indistinguishable, at the 5% level, from all the observed mean abundance patterns. In contrast, graphical comparisons of the observed and fitted distributions suggested the log series may be the better fit at most of the unstable sites and the two most stable sites, whereas the more equitable distribution at sites of intermediate stability suggested the log normal distribution was the better fit. If conditions at a site favoured one or two species, either through severe physical conditions, or through competitive superiority in the absence of disturbance then the log series distribution may result. However, if no species in the community was strongly advantaged over others, a log normal distribution should result. Given the discriminating power of the appropriate statistical test it may not, however, be possible to pick up these differences without graphical comparisons as well.     

Pliocene benthic foraminifera from the Blake Plateau: Faunal assemblages and paleocirculation

Relative abundance of benthic foraminifera have been analyzed from core V26-145 from the Blake Plateau. The investigated sequence represents the time interval between 1.8 and 4.6 Ma. In order to determine how different sieve sizes influence the relative abundance patterns, three sediment size fractions were studied separately. It becomes difficult to maintain consistent taxonomic concepts in the fraction 63–125 μm, partly because this fraction contains high abundances of juvenile forms. However, the 63–125 μm fraction holds high abundances of the important small speciesEpistominella exigua. Due to these reasons only the two larger fractions (125–250 μm and >250 μm) were considered meaningful to analyze for relative abundance patterns. An analysis of the two larger fractions (>125 μm; >250 μm) shows no consistency in relative abundance patterns.The relative abundance patterns for the 34 most common species in the size fraction >125 μm were analyzed by means of correspondence analysis. Three benthic foraminiferal assemblages (I, II, and III) were recognized and these can be associated with water masses. Assemblage I is associated with the Florida Current and consists of shallow water species (Amphistegina gibbosa, Compressigerina sp. A,Discorbinella biconcavus, Islandiella teretis, Reussella atlantica, andSiphonina pulchra). Assemblage II contains key species for North Atlantic Deep Water (NADW) (Cibicidoides kullenbergi, Epistominella exigua, Globocassidulina subglobosa, Lenticulina peregrina, Oridorsalis umbonatus, andPlanulina wuellerstorfi). The third assemblage (III) contains species associated with the Antilles Current (Bolivina rhomboidalis, Cassidulina obtusa, Cassidulina vortex, andNuttallides umbonifera). The correspondence analysis reveals an alternation in dominance between Assemblage I and Assemblage II prior to 3.3 Ma, suggesting lateral oscillations between the Florida Current and NADW. At about 3.3 Ma Assemblage I disappears and Assemblage III increases in importance, suggesting an increasing influence of the Antilles Current in the upper part of the record.     

Absolute depths of Silurian benthic assemblages

A variety of evidence can be used to estimate the absolute depth of the well-established depth gradient of Silurian onshore-to-offshore benthic assemblages (BA 1–5); this evidence is reviewed herein. There is a fair degree of consistency between fossil benthic assemblages and the occurrence of certain primary sedimentary structures, particularly those involved in storm deposition. A second, probably stronger, line of evidence for absolute depth of Silurian benthic assemblages involves the distribution of fossils of light-sensitive organisms and of reefs. Among these, the most important are the calcified dasycladacean algae, to which the cyclocrinitids of the Middle Ordovician - Early Silurian may belong. The modem dasycladaceans have a narrowly restricted bathymetric range. Maximum depth for modern calcified dasycladaceans is about 90 m, but the vast majority occur at 30 m or less. Some Silurian occurrences of these algae are abundant, particularly in BA 3 and 4; rarely, small specimens of cyclocrinitids and receptaculitids are found in the lower, outer portions of BA 4 and even into 5. This evidence constrains much of the spectrum (BA 1–5) of Silurian fossil communities to a rather narrow depth range, within the photic zone. Based on taxonomic uniformitarianism, we would place the depth of Silurian BA 1 through 4 between 0 and about 40–60 m. The common coincidence of the lower end of storm wave base and the lower end of the photic zone near the BA 4–5 boundary also suggests that this position may represent water depths on the order of 50 m. Several other lines of evidence -including algal borings, widths of facies belts, separation of communities by basalt flows whose thicknesses are known - also support a relatively shallow depth range for BA 14. The absence of storm-disturbed beds over large parts of several major platforms below about the BA 3–4 boundary region suggests that major storms of the Silurian may have been far weaker than those of the present, possibly owing to a different climatic regime. □Absolute depth, Silurian, photic zone, calcareous algae, Benthic Assemblage, storms.     

Island benthic assemblages: With examples from the Late Ordovician of Eastern Australia

The Benthic Assemblage (B.A.) concept, developed by A.J. Boucot two decades ago principally for continental marine margins, is extended to offshore island settings. Characteristics of modern island biotas, including ecological displacement, and the effects of r and K selection, can be identified in Late Ordovician volcanic islands of central New South Wales. Two variants of B.A.1 are represented, a quiet‐water lingulide biofacies, and a rough‐water rhynchonellide biofacies. The quiet‐water Eodinobolus biofacies occupied a B.A.1–2 position, onshore of and sheltered by a Tetradium wave‐baffle (B.A.2). The offshore shelfal high diversity strophomenide biofacies is equivalent to B.A.3. Remnants of periplatformal B.A.4–5 communities are recognised in allochthonous limestone breccias which were displaced downslope into graptolitic B.A.6 sediments. Steeper offshore gradients, typical of islands, laterally compress the B.A. profiles, and also contribute to downslope slumping. Ecological displacement in island environments results in extension of the habitat range of species into adjacent B.A.s. Reefs complicate the usual B.A. profile by introducing distinct sheltered and turbulent water environments. These characteristics may have applicability in interpretation of islands throughout the Palaeozoic record.     

Filling biodiversity gaps: benthic hydroids from the Bellingshausen Sea (Antarctica)

The Bellingshausen Sea constitutes the third largest sea in the Southern Ocean, though it is widely recognized as one of the less-studied Antarctic areas. To reduce this lack of knowledge, a survey to study the biodiversity of its marine benthic communities was carried out during the Bentart 2003 and Bentart 2006 Spanish Antarctic expeditions. The study of the hydroid collection has provided 27 species, belonging to ten families and 15 genera. Twenty-one out of the 27 species constitute new records for the Bellingshausen Sea, raising the total number of known species to 37, as also do nine out of the 15 genera. Candelabrum penola, Lafoea annulata, and Staurotheca juncea are recorded for the second time. Most species belong to Leptothecata. Sertulariidae with 13 species (48%) is by far the most speciose family, and Symplectoscyphus with seven species (26%), including S. bellingshauseni sp. nov. and S. hesperides sp. nov., the most diverse genus. Considering the whole benthic hydroid fauna of the Bellingshausen Sea, 18 species (69%) are endemic to Antarctic waters, either with a circum-Antarctic (12 species, 46%) or West Antarctic (6 species, 23%) distribution, 23 (88%) are restricted to Antarctic or Antarctic/sub-Antarctic waters, and only three species have a wider distribution. Bellingshausen Sea hydroid fauna is composed of a relatively high diversity of typical representatives of the Antarctic benthic hydroid fauna, though with a surprisingly low representation of some of the most diverse and widespread Antarctic genera (Oswaldella and Schizotricha), what could be related to the fact that its shelf-inhabiting hydroid fauna remains practically unknown.     

Late Quaternary paleoproductivity and deep water circulation in the eastern South Atlantic Ocean: Evidence from benthic foraminifera

Two late Quaternary sediment cores from the northern Cape Basin in the eastern South Atlantic Ocean were analyzed for their benthic foraminiferal content and benthic stable carbon isotope composition. The locations of the cores were selected such that both of them presently are bathed by North Atlantic Deep Water (NADW) and past changes in deep water circulation should be recorded simultaneously at both locations. However, the areas are different in terms of primary production. One core was recovered from the nutrient-depleted Walvis Ridge area, whereas the other one is from the continental slope just below the coastal upwelling mixing area where present day organic matter fluxes are shown to be moderately high. Recent data served as the basis for the interpretation of the late Quaternary faunal fluctuations and the paleoceanographic reconstruction.

During the last 450,000 years, NADW flux into the eastern South Atlantic Ocean has been restricted to interglacial periods, with the strongest dominance of a NADW-driven deep water circulation during interglacial stages 1, 9 and 11. At the continental margin, high productivity faunas and very low epibenthic δ¹³C values indicate enhanced fluxes of organic matter during glacial periods. This can be attributed to a glacial increase and lateral extension of coastal upwelling. The long term glacial-interglacial paleoproductivity cycles are superimposed by high-frequency variations with a period of about 23,000 yr. Enhanced productivity in surface waters above the Walvis Ridge, far from the coast, is indicated during glacial stages 8, 10 and 12. During these periods, cold, nutrient-rich filaments from the mixing area were probably driven as far as to the southeastern flank of the Walvis Ridge.     

Multivariate analysis of periphytic and benthic diatom assemblages from Papua New Guinea

The benthic and periphytic diatom flora is studied in 136 fresh waters in Papua New Guinea. DCA ordination and TWINSPAN are used to describe 20 diatom assemblages. Conductivity, water temperature and altitude are the main sources for the variation in the data. Laing Island Biological Station publication n° 214 Laing Island Biological Station publication n° 214     

Natural succession of macroalgal-dominated epibenthic assemblages at different water depths and after transplantation from deep to shallow water on Spitsbergen

In the current study, we investigated the primary succession of seaweeds over different time periods at different water depths. Furthermore, we followed the succession of field-grown benthic communities of different successional age, developing on ceramic tiles, prior to and after transplantation from 8 to 0.5 m water depth. The transplantation simulated changes associated with the break up of sea-ice cover, e.g. light regime or wave exposure. For this purpose, we transplanted 12 and 21-month old communities, grown at 8 m water depth, together with a set of sterile tiles, onto rafts, floating in 0.5 m water depth. Our results describe for the first time the succession of macroalgal communities in the Arctic and give important insights into the effect of disturbance of differently aged communities. The primary succession at 0.5 m water depth was mainly driven by Bacillariophyta and filamentous green algae like Urospora sp. and Ulothrix implexa. Twelve-month old communities at 8 m water depth are dominated by members of the Ectocarpales (Phaeophyceae), like Pylaiella littoralis, P. varia, and Ectocarpus siliculosus and the green alga U. implexa, whereas the 21-month old community showed a higher cover of the green algal class Ulvophyceae and sessile invertebrates. After transplantation to near surface conditions, species composition of the communities changed, but this effect was differently strong between communities of different age.     

Hydromorphological restoration of running waters: effects on benthic invertebrate assemblages

1. River restoration has received considerable attention, with much recent focus on restoring river hydromorphology to improve impoverished aquatic communities. However, we still lack a clear understanding of the response of aquatic biota to river restoration. 2. We studied the effects of hydromorphological restoration on benthic invertebrate assemblages in 25 river sites in Germany using standardised methods. Restoration efforts were primarily aimed to restore habitat heterogeneity; correspondingly, habitat diversity increased at most sites. 3. Similarity of benthic invertebrate assemblages between restored and unrestored river sections was low (mean similarity was 0.32 for Jaccard and 0.46 for Sørensen). Community‐based metrics, such as the percentage of Ephemeroptera, Plecoptera and Trichoptera taxa, also differed between restored and unrestored sections. 4. Only three of the 25 restored sections were classified as having ‘good ecological quality’ class according to the European Water Framework Directive criteria; hence, poor water quality is probably one factor impeding recolonisation. 5. Our results show that isolated restoration measures do not necessarily result in positive effects on aquatic biota and that better understanding of the interconnectedness within a catchment is required before we can adequately predict biotic responses to structural river restoration.     

Recent benthic foraminiferal assemblages from cold-water coral mounds in the Porcupine Seabight

Cold-water coral ecosystems are characterised by a high diversity and population density. Living and dead foraminiferal assemblages from 20 surface sediment samples from Galway and Propeller Mounds were analysed to describe the distribution patterns of benthic foraminifera on coral mounds in relation to different sedimentary facies. Hard substrates were examined to assess the foraminiferal microhabitats and diversities in the coral framework. We recognised 131 different species, of which 27 prefer an attached lifestyle. Epibenthic species are the main constituents of the living and dead foraminiferal assemblages. The frequent species Discanomalina coronata was associated with coral rubble, Cibicides refulgens showed preference to the off-mound sand veneer, and Uvigerina mediterranea displayed abundance maxima in the main depositional area on the southern flank of Galway Mound, and in the muds around Propeller Mound. The distribution of these species is rather governed by their specific ecological demands and microhabitat availability than by the sedimentary facies. Benthic foraminiferal assemblages from coral mounds fit well into basin-wide-scale distribution patterns of species along the western European continental margin. The diversity of the foraminiferal faunas is not higher on the carbonate mounds as in their vicinity. The living assemblages show a broad mid-slope diversity maximum between 500 and 1,300 m water depth, which is the depth interval of coral mound formation at the Celtic and Amorican Margin. The foraminiferal diversity maximum is about 700 m shallower than comparable maxima of nematodes and bivalves. This suggests that different processes are driving the foraminiferal and metazoan diversity patterns.     

Insects and low temperatures: from molecular biology to distributions and abundance

Insects are the most diverse fauna on earth, with different species occupying a range of terrestrial and aquatic habitats from the tropics to the poles. Species inhabiting extreme low-temperature environments must either tolerate or avoid freezing to survive. While much is now known about the synthesis, biochemistry and function of the main groups of cryoprotectants involved in the seasonal processes of acclimatization and winter cold hardiness (ice-nucleating agents, polyols and antifreeze proteins), studies on the structural biology of these compounds have been more limited. The recent discovery of rapid cold-hardening, ice-interface desiccation and the daily resetting of critical thermal thresholds affecting mortality and mobility have emphasized the role of temperature as the most important abiotic factor, acting through physiological processes to determine ecological outcomes. These relationships are seen in key areas such as species responses to climate warming, forecasting systems for pest outbreaks and the establishment potential of alien species in new environments.     

Spatial scaling of species abundance distributions

Species abundance distributions are an essential tool in describing the biodiversity of ecological communities. We now know that their shape changes as a function of the size of area sampled. Here we analyze the scaling properties of species abundance distributions by using the moments of the logarithmically transformed number of individuals. We find that the moments as a function of area size are well fitted by power laws and we use this pattern to estimate the species abundance distribution for areas larger than those sampled. To reconstruct the species abundance distribution from its moments, we use discrete Tchebichef polynomials. We exemplify the method with data on tree and shrub species from a 50 ha plot of tropical rain forest on Barro Colorado Island, Panama. We test the method within the 50 ha plot, and then we extrapolate the species abundance distribution for areas up to 5 km². Our results project that for areas above 50 ha the species abundance distributions have a bimodal shape with a local maximum occurring for the singleton classes and that this maximum increases with sampled area size.     

Assessment of species diversity from species abundance distributions at different localities

We show how the spatial structure of species diversity can be analyzed using the correlation between the log abundances of the species in the communities, assuming that two communities at different localities can be described by a bivariate lognormal species abundance distribution. A useful property of this approach is that the log abundances of the species at two localities can be considered as samples from a bivariate normal distribution defined by only five parameters. The variances and the correlation can be estimated by maximum likelihood methods even if there is no information about the sampling intensity and the number of unobserved species. This method also enables estimation of over-dispersion in the sampling relative to a Poisson distribution that allows sampling adjustment of the estimate of β-diversity. Furthermore, we also obtain a partitioning of species diversity into additive components of α-, β- and γ-diversity. For instance, if the correlation between the log abundances of the species is close to one, the same species will be common and rare in the two communities and the β-diversity will be low. We illustrate this approach by analysing similarities of communities of rare and endangered species of oak-living beetles in south-eastern Norway. The number of recorded species was estimated to be only 48.1% of the total number of species actually present in these communities. The correlations among communities dropped rather quickly with distance with a scaling of order 200 km. This illustrates large spatial heterogeneity in species composition, which should be accounted for in the design of schemes of such devices for assessing species diversity in these habitat-types.     

Sampling species abundance distributions: resolving the veil-line debate

Preston's classic work on the theory of species abundance distributions (SADs) in ecology has been challenged by Dewdney. Dewdney contends that Preston's veil-line concept, relating to the shape of sample SADs, is flawed. Here, I show that Preston's and Dewdney's theories can be reconciled by considering the differing mathematical properties of the sampling process on logarithmic (Preston) versus linear (Dewdney) abundance scales. I also derive several related results and show, importantly, that one cannot reject the log-normal distribution as a plausible SAD based only on sampling arguments, as Dewdney and others have done.     

Infection paradox: high abundance but low impact of freshwater benthic viruses

The discovery of an abundant and diverse virus community in oceans and lakes has profoundly reshaped ideas about global carbon and nutrient fluxes, food web dynamics, and maintenance of microbial biodiversity. These roles are exerted through massive viral impact on the population dynamics of heterotrophic bacterioplankton and primary producers. We took advantage of a shallow wetland system with contrasting microhabitats in close proximity to demonstrate that in marked contrast to pelagic systems, viral infection, determined directly by transmission electron microscopy, and consequently mortality of prokaryotes were surprisingly low in benthic habitats in all seasons. This was true even though free viruses were abundant throughout the year and bacterial infection and mortality rates were high in surrounding water. The habitats in which we found this pattern include sediment, decomposing plant litter, and biofilms on aquatic vegetation. Overall, we detected viruses in only 4 of a total of approximately 15,000 bacterial cells inspected in these three habitats; for comparison, nearly 300 of approximately 5,000 cells suspended in the water column were infected. The strikingly low incidence of impact of phages in the benthos may have important implications, since a major portion of microbial biodiversity and global carbon and nutrient turnover are associated with surfaces. Therefore, if failure to infect benthic bacteria is a widespread phenomenon, then the global role of viruses in controlling microbial diversity, food web dynamics, and biogeochemical cycles would be greatly diminished compared to predictions based on data from planktonic environments.