Changes In Plankton Communities Research Articles

Models of Plankton Community Changes during a Warm Water Anomaly in Arctic Waters Show Altered Trophic Pathways with Minimal Changes in Carbon Export

Carbon flow through pelagic food webs is an expression of the composition, biomass and activity of phytoplankton as primary producers. In the near future, severe environmental changes in the Arctic Ocean are expected to lead to modifications of phytoplankton communities. Here, we used a combination of linear inverse modeling and ecological network analysis to study changes in food webs before, during, and after an anomalous warm water event in the eastern Fram Strait of the West Spitsbergen Current (WSC) that resulted in a shift from diatoms to flagellates during the summer (June–July). The model predicts substantial differences in the pathways of carbon flow in diatom- vs. Phaeocystis/nanoflagellate-dominated phytoplankton communities, but relatively small differences in carbon export. The model suggests a change in the zooplankton community and activity through increasing microzooplankton abundance and the switching of meso- and macrozooplankton feeding from strict herbivory to omnivory, detritivory and coprophagy. When small cells and flagellates dominated, the phytoplankton carbon pathway through the food web was longer and the microbial loop more active. Furthermore, one step was added in the flow from phytoplankton to mesozooplankton, and phytoplankton carbon to higher trophic levels is available via detritus or microzooplankton. Model results highlight how specific changes in phytoplankton community composition, as expected in a climate change scenario, do not necessarily lead to a reduction in carbon export.

Comparison of zooplankton data collected by a continuous semi-automatic sampler (CALPS) and a traditional vertical ring net

We compared and evaluated the performance of a Continuous Automatic Litter and Plankton Sampler (CALPS) against the traditional ring net vertical haul. CALPS is a custom-made semi-automatic sampler, which collects water using a pump system at a single depth along a predetermined transect as the ship sails. CALPS underestimated species abundance compared to the ring net by a factor 1.61, but both datasets illustrated a similar species composition, community size structure and good agreement in the spatial distribution of abundance. Our analysis suggests that avoidance of the CALPS is likely to be the main factor responsible for the observed difference in sampling efficiency, but other factors, such as depth, area sampled and zooplankton patchiness, are also likely to play their part. We conclude that whilst the CALPS is not suitable for investigations that require accurate measures of abundance, it is an ideal tool to identify and quantify changes in plankton communities and diversity. A particular advantage over more traditional vertical sampling methods is that it can be integrated within existing multidisciplinary surveys at little extra cost, thus making the CALPS particularly valuable as part of integrated monitoring programmes to underpin policy areas such as the EU Marine Strategy Framework Directive.

Carbonate-to-biosilica transition at the Norian–Rhaetian boundary controlled by rift-related subsidence in the western Tethyan Lagonegro Basin (southern Italy)

The Lagonegro Basin, located in the Southern Apennines, Italy, records a transition from carbonate sediments of the Calcari con Selce Fm. (“Cherty Limestone”) to biosiliceous sediments of the Scisti Silicei Fm. (“Siliceous Shale”) around the Norian–Rhaetian boundary (NRB), Upper Triassic. Sedimentologic and chemostratigraphic analysis of three sections (Pignola-Abriola, Monte Volturino, and Madonna del Sirino) representing a proximal-to-distal transect across the basin document marine productivity, redox, and chemical weathering conditions. The basin was characterized by low to moderate productivity and mainly oxic deep waters, with no systematic secular variation throughout the ~10-Myr-long study interval. A weathering proxy, the chemical index of alteration (CIA*), shifts toward higher values (from ~0.70 to ~0.80) in all three study sections, suggesting development of warmer and/or more humid conditions just before the NRB. The lack of evidence for changes in marine productivity or redox conditions and the time-transgressive nature of the carbonate-biosilica transition suggests that the latter was caused not by changes in planktonic communities but, rather, by a rise in the CCD, the most likely cause of which was seafloor subsidence within the actively rifting Ionian Ocean (i.e., the westernmost branch of the Tethys Ocean). The tectonically active character of the Lagonegro Basin is also reflected in evidence of synsedimentary paleoseismic activity, e.g., numerous debris flows around the basin margins. The present study thus documents sedimentologic changes within a marginal basin in a geologically young ocean.

Can a copper sulphate pulse below toxic threshold change plankton communities?

More field relevant ecological assessments, apart from single species tests using standard species, are needed to better predict agrochemical effects at higher ecosystems levels. Therefore, an experiment using a non-target aquatic community was used to test the hypothesis of the negative effect of a single pulse of copper sulfate on plankton abundance, structure, richness and diversity endpoints. Microcosms (20 l volume) were established during 21 days of experimentation, using six replicates for controls and with two concentrations of copper sulfate (High treatment, H: 20 µg Cu l−1; and Low treatment, L: 2 µg Cu l−1), both within the copper legal threshold following the Water Framework Directive (2000). The general lineal model found significant differences at the phytoplankton abundance endpoint at the end of the experiment, with an increase of phytoplankton abundance in L treatments dominated by the smaller cell size class. The principal response curve on zooplankton data, despite being insignificant, pointed out some dissimilarities in abundance and structure between treatments and controls: treatments showed lower abundances and were richer in cladocerans and copepods than the control microcosm where rotifers and nauplii were dominant. This indicated that trends change in community structure due to the effects of copper sulfate, and that even if the copper concentrations under study were within legal limits, they showed potential to induce changes in planktonic communities.

Changes in Planktonic Communities and Environmental Factors between Open Versus Closed Artificial Marine Microcosms

Changes in Planktonic Communities and Environmental Factors between Open Versus Closed Artificial Marine Microcosms

Hexabromocyclododecane affects benthic-pelagic coupling in an experimental ecosystem

Hexabromocyclododecane (HBCDD) is an additive brominated flame retardant and a recognized PBT chemical. However, little is known about its effects on coastal species, and even less on ecosystem effects. We investigated the dose–response effects of HBCDD over 8 months in 1000 L experimental mesocosms assembled from coastal Baltic Sea ecosystem components. HBCDD was added via spiked plankton material and a range of structural and functional endpoints were measured during the experiment. Increasing HBCDD concentration decreased the biomass of large Macoma balthica, resulting in a decreased recirculation of nutrients to the water. Changes in plankton communities were also observed, either due to direct toxic HBCDD effects or indirect via changes in benthic-pelagic coupling of nutrients. Such complex ecosystem responses can only be quantified and understood by using realistic experimental set-ups, and including knowledge of system-specific ecological interactions. This is the first study of HBCDD effects on ecosystem level.

Modifying the PEG model for Mediterranean lakes – no biological winter and strong fish predation

Summary The widely cited Plankton Ecology Group (PEG) model of plankton seasonal succession is often used as a template to explain the seasonal changes in plankton communities outside the cold temperate zone, where it was developed, but this may be inappropriate for lower‐latitude lakes. Lower‐latitude lakes have high light availability in winter and less pronounced seasonal variations in fish predation on zooplankton. We might therefore expect higher phytoplankton crops in winter and much more predation on zooplankton by fish than in colder lakes. This might lead to less grazing in summer and relatively higher phytoplankton crops. We compared data on phytoplankton biovolume, zooplankton biomass and body size from 18 German and 6 Greek lakes to test these hypotheses.

Enhanced greenhouse gas emissions and changes in plankton communities following an experimental increase in organic carbon loading to a humic lake

Organic carbon concentrations in the surface waters of the boreal region have increased during the past two decades. We investigated the impact of elevated dissolved organic carbon (DOC) loading to a humic lake by a whole-lake experiment in which DOC in the form of cane sugar was added monthly during the ice-free period over two consecutive years. The sugar addition represented an increased concentration of 2 mg l−1 of DOC in the epilimnion and led to an increase in CO2 emission and also an apparent increase in CH4 emission to the atmosphere from the lake surface. The composition of the bacterial, phytoplankton and zooplankton communities altered during the study period and the bacterial abundance in the metalimnion and hypolimnion of the lake decreased. No changes were detected in epilimnetic primary production or respiration, but there was an increase in bacterial production in the epilimnion. The nutrient and particulate organic carbon concentrations also suggested possible changes in the activity of heterotrophic bacteria in the metalimnion. Carbon stable isotope analyses indicated transfer of some added sugar carbon through the food web to zooplankton consumers. Overall the results suggest that future increases in organic carbon loading to boreal lakes will increase greenhouse gas emissions, although the magnitude of any change is likely to depend on the availability of nutrients like phosphorus and nitrogen which influence organic matter processing and the development of plankton communities.

Climate and ecosystem linkages explain widespread declines in North American Atlantic salmon populations

North American Atlantic salmon (Salmo salar) populations experienced substantial declines in the early 1990s, and many populations have persisted at low abundances in recent years. Abundance and productivity declined in a coherent manner across major regions of North America, and this coherence points toward a potential shift in marine survivorship, rather than local, river-specific factors. The major declines in Atlantic salmon populations occurred against a backdrop of physical and biological shifts in Northwest Atlantic ecosystems. Analyses of changes in climate, physical, and lower trophic level biological factors provide substantial evidence that climate conditions directly and indirectly influence the abundance and productivity of North American Atlantic salmon populations. A major decline in salmon abundance after 1990 was preceded by a series of changes across multiple levels of the ecosystem, and a subsequent population change in 1997, primarily related to salmon productivity, followed an unusually low NAO event. Pairwise correlations further demonstrate that climate and physical conditions are associated with changes in plankton communities and prey availability, which are ultimately linked to Atlantic salmon populations. Results suggest that poor trophic conditions, likely due to climate-driven environmental factors, and warmer ocean temperatures throughout their marine habitat area are constraining the productivity and recovery of North American Atlantic salmon populations.

Climate Change and Eutrophication Induced Shifts in Northern Summer Plankton Communities

Marine ecosystems are undergoing substantial changes due to human-induced pressures. Analysis of long-term data series is a valuable tool for understanding naturally and anthropogenically induced changes in plankton communities. In the present study, seasonal monitoring data were collected in three sub-basins of the northern Baltic Sea between 1979 and 2011 and statistically analysed for trends and interactions between surface water hydrography, inorganic nutrient concentrations and phyto- and zooplankton community composition. The most conspicuous hydrographic change was a significant increase in late summer surface water temperatures over the study period. In addition, salinity decreased and dissolved inorganic nutrient concentrations increased in some basins. Based on redundancy analysis (RDA), warming was the key environmental factor explaining the observed changes in plankton communities: the general increase in total phytoplankton biomass, Cyanophyceae, Prymnesiophyceae and Chrysophyceae, and decrease in Cryptophyceae throughout the study area, as well as increase in rotifers and decrease in total zooplankton, cladoceran and copepod abundances in some basins. We conclude that the plankton communities in the Baltic Sea have shifted towards a food web structure with smaller sized organisms, leading to decreased energy available for grazing zooplankton and planktivorous fish. The shift is most probably due to complex interactions between warming, eutrophication and increased top-down pressure due to overexploitation of resources, and the resulting trophic cascades.

Bayesian learning and predictability in a stochastic nonlinear dynamical model

Bayesian inference methods are applied within a Bayesian hierarchical modeling framework to the problems of joint state and parameter estimation, and of state forecasting. We explore and demonstrate the ideas in the context of a simple nonlinear marine biogeochemical model. A novel approach is proposed to the formulation of the stochastic process model, in which ecophysiological properties of plankton communities are represented by autoregressive stochastic processes. This approach captures the effects of changes in plankton communities over time, and it allows the incorporation of literature metadata on individual species into prior distributions for process model parameters. The approach is applied to a case study at Ocean Station Papa, using particle Markov chain Monte Carlo computational techniques. The results suggest that, by drawing on objective prior information, it is possible to extract useful information about model state and a subset of parameters, and even to make useful long-term forecasts, based on sparse and noisy observations.

Mismatch revisited: what is trophic mismatching from the perspective of the plankton?

Long-term changes in the seasonal timing of phytoplankton and zooplankton population development (i.e. phenological changes) have been widely reported in freshwater and marine ecosystems, and have been interpreted as a biological response to ongoing climate change. Observed among-species variations in the rates and patterns of these changes have led to concerns that seasonal trophic interactions may become de-synchronized in plankton communities, following the logic of the match– mismatch hypothesis. Herein, the current evidence for long-term mismatching within plankton communities is evaluated and the meaning of this concept for the plankton is explored. Despite the large number of studies documenting phenological change in plankton communities, very few explicitly quantify mismatching among trophic levels within the same study system, and quantify its ecological impacts. There is a clear need to build the evidence base for the mismatching phenomenon. In this endeavour, we must carefully define the seasonality of resource availability based on prior ecological knowledge, and embrace the challenges posed by organisms with more complex life cycles and the ecological network context within which plankton interact. In addition, we must acknowledge that, while mismatching in the plankton and in non-planktonic systems relates to synchrony between resource supply and consumer demand, there are conceptual differences in the phenomenon between these two broad classes of systems. For planktonic systems, mismatching is best visualized as an alteration of the rates and lags of the interaction between dynamically linked consumers and resources, rather than simple and independent phase shifts of seasonal abundance peaks at each trophic level.

Protistan diversity does peak in the horohalinicum of the Baltic Sea: Reply to Ptacnik et al. (2011)

In reply to the Comment by Ptacnik et al. (2011; Mar Ecol Prog Ser 432:291-292), a sub- stantiation is given for the use of the simplest method to conduct a meta-analysis of published plank- ton data sets and annotated checklists aimed at refining the 'Artenminimum' (species-minimum) con- cept (Remane 1934; Zool Anz 7(Suppl):34-74). The applicability of this concept, which originally derived from macrozoobenthos data, was tested by Telesh et al. (2011; Mar Ecol Prog Ser 421:1-11) for plankton species using much larger data sets, and these results backed up the novel 'protistan species-maximum concept' for the horohalinicum of the Baltic Sea. Re-fitting our algorithm to the available data sets after the critical comments of Ptacnik et al. (2011) did not alter the shape of the plankton diversity trends, and the original conclusions that (1) the brackish-water Baltic Sea is not poor in plankton species, (2) Remane's curve is invalid for plankton, and (3) protistan species richness peaks in the horohalinicum at salinities of 5 to 8 remained robust. The suggestion by Ptacnik et al. (2011) to use a more detailed statistical analysis for describing the patterns of overall plankton diver- sity is recognized as a perspective for future research of changes in plankton communities within the Baltic Sea salinity gradient, although it cannot be implemented until spatially representative zoo- plankton data sets become available.

Molecular isotopic evidence of environmental and ecological changes across the Cenomanian–Turonian boundary in the Levant Platform of central Jordan

We evaluated the structure of planktonic communities and paleoenvironmental conditions throughout the Cenomanian–Turonian Oceanic Anoxic Event (OAE2) by studying bulk geochemical properties and the molecular isotopic composition of source-specific hydrocarbons from organic-rich sediments deposited in an intrashelf basin at the Levant Platform, central Jordan. High concentrations of desmethyl and 4-methylsteranes as well as dinosteranes indicate that marine algae including dinoflagellates were the main primary producing organisms. The presence of 2-methylhopanes and 13C-enriched hopanes, in addition to isotopically enriched aryl isoprenoids, evidenced the contribution of cyanobacteria and green sulfur bacteria, respectively. Additionally, variable but fairly low δ 15N values during OAE2 suggest the occurrence of diazotrophy as a likely important process fueling primary production during OAE2 in this stratified/anoxic continental platform. Variations in the relative contribution of biomarkers revealed changes in planktonic communities associated with sea level change and water column stratification. OAE2 was characterized by strong stratification, anoxic bottom waters and a deep chemocline, as evidenced by high gammacerane and homohopane indices and the absence of photic zone euxinia (PZE) markers, respectively. However, the presence of isorenieratane and its derivatives in post-OAE black shales points to a shoaling of the chemocline and to PZE. This interval was also characterized by an exceptionally high abundance of chlorophyll-derived pristane and phytane (up to 2 mg g −1 TOC), likely as a result of highly enhanced primary production and organic matter preservation. Remarkably, this high productivity event co-occurs with an exceptionally high abundance of calcispheres reported elsewhere to be part of a global bio-event.

Biomarker evidence of macrophyte and plankton community changes in Zeekoevlei, a shallow lake in South Africa

Zeekoevlei is the largest freshwater lake in South Africa and has a century-long history of anthropogenic impact that caused hyper-eutrophic conditions. We used biomarkers (alkanes and pigments), stable isotopes (δ 13 C and δ 15 N), rates of primary palaeoproduction and total inorganic carbon (TIC) accumulation rates in the lake sediments to investigate changes in plankton and macrophyte communities in response to anthropogenic activities in this shallow lake. Specific alkanes (ΣC 15,17,19 , pristane, phytane and n-C 29 /n-C 17 ratio) and pigment (chlorophyll a, β,β-carotene, echinenone, fucoxanthin and zeaxanthin) concentrations in lake waters indicated the present-day hyper-eutrophic condition and seasonal fluctuations of cyanobacteria, zooplankton and diatom populations. Eutrophic conditions were initiated in the lake with the start of recreational activities and construction of a sewage treatment plant in the early 1920s. The lake transformed from a eutrophic to a hyper-eutrophic waterbody following damming, pondweed eradication and accelerated catchment-derived nutrient input. The change in lake trophic state was recorded by a sharp decline in the terrestrial to aquatic ratio (TAR) of specific n-alkanes, low carbon preference index (CPI) and increased δ 13 C values in the sediment core. In addition, the aquatic macrophyte n-alkane proxy (P aq ) values (~1) indicated a slow takeover by floating macrophytes after the eradication of submerged pondweeds in 1951. Elevated n-alkane (ΣC 15,17,19 ), total alkane and pigment (chlorophyll a, β,β-carotene, zeaxanthin and zeaxanthin to β,β-carotene ratio) concentrations, low δ 15 N values and low TIC accumulation rates in the upper middle section of the core indicated the beginning of intense cyanobacterial blooms after the dredging in 1983. Although the cyanobacterial population has decreased in recent years, hyper-eutrophic conditions are reflected by low CPI <0.04 and TAR <1 values at the top of the sediment core.

From plankton to top predators: bottom‐up control of a marine food web across four trophic levels

1. Abundant mid-trophic pelagic fish often play a central role in marine ecosystems, both as links between zooplankton and top predators and as important fishery targets. In the North Sea, the lesser sandeel occupies this position, being the main prey of many bird, mammal and fish predators and the target of a major industrial fishery. However, since 2003, sandeel landings have decreased by > 50%, and many sandeel-dependent seabirds experienced breeding failures in 2004. 2. Despite the major economic implications, current understanding of the regulation of key constituents of this ecosystem is poor. Sandeel abundance may be regulated 'bottom-up' by food abundance, often thought to be under climatic control, or 'top-down' by natural or fishery predation. We tested predictions from these two hypotheses by combining unique long-term data sets (1973-2003) on seabird breeding productivity from the Isle of May, SE Scotland, and plankton and fish larvae from the Continuous Plankton Recorder survey. We also tested whether seabird breeding productivity was more tightly linked to sandeel biomass or quality (size) of individual fish. 3. The biomass of larval sandeels increased two- to threefold over the study period and was positively associated with proxies of the abundance of their plankton prey. Breeding productivity of four seabirds bringing multiple prey items to their offspring was positively related to sandeel larval biomass with a 1-year lag, indicating dependence on 1-year-old fish, but in one species bringing individual fish it was strongly associated with the size of adult sandeels. 4. These links are consistent with bottom-up ecosystem regulation and, with evidence from previous studies, indicate how climate-driven changes in plankton communities can affect top predators and potentially human fisheries through the dynamics of key mid-trophic fish. However, the failing recruitment to adult sandeel stocks and the exceptionally low seabird breeding productivity in 2004 were not associated with low sandeel larval biomass in 2003, so other mechanisms (e.g. predation, lack of suitable food after metamorphosis) must have been important in this case. Understanding ecosystem regulation is extremely important for predicting the fate of keystone species, such as sandeels, and their predators.

Overview of key phytoplankton toxins and their recent occurrence in the North and Baltic Seas

The frequency and intensity of harmful algal blooms (HABs) appear to be on the rise globally. There is also evidence of the geographic spreading of toxic strains of these algae. Consequently, methods had to be established and new ones are still needed for the evaluation of possible hazards caused by increased algal toxin production in the marine food chain. Different clinical effects of algae-related poisoning have attracted scientific attention; paralytic shellfish poisoning, diarrhetic shellfish poisoning, and amnesic shellfish poisoning are among the most common. Additionally, cyanobacteria (blue-green algae) in brackish waters often produce neurotoxic and hepatotoxic substances. Bioassays with mice or rats are common methods to determine algal and cyanobacterial toxins. However, biological tests are not really satisfactory because of their low sensitivity. In addition, there is growing public opposition to animal testing. Therefore, there has been increasing effort to determine algal toxins by chemical methods. Plankton samples from different European marine and brackish waters were taken during research cruises and analyzed on board directly. The ship routes covered marine areas in the northwest Atlantic, Orkney Islands, east coast of Scotland, and the North and Baltic seas. The first results on the occurrence and frequency of harmful algal species were obtained in 1997 and 1998. During the 2000 cruise an HPLC/MS coupling was established on board, and algal toxins were measured directly after extraction of the plankton samples. In contrast to earlier cruises, the sampling areas were changed in 2000 to focusing on coastal zones. The occurrence of toxic algae in these areas was compared to toxin formation during HABs in the open sea. It was found that the toxicity of the algal blooms depended on the prevailing local conditions. This observation was also confirmed by monitoring cyanobacterial blooms in the Baltic Sea. Optimal weather conditions, for example, during the summers of 1997 and 2003, favored blooms of cyanobacteria in all regions of the Baltic. The dominant species regarding the HABs in the Baltic was Nodularia spumigena. However, in addition to high concentrations of Nodularia spumigena in coastal zones, other blue-green algae are involved in bloom formation, with changes in plankton communities influencing both toxin profiles and toxicity.

Parasites and phytoplankton, with special emphasis on dinoflagellate infections.

Planktonic members of most algal groups are known to harbor intracellular symbionts, including viruses, bacteria, fungi, and protozoa. Among the dinoflagellates, viral and bacterial associations were recognized a quarter century ago, yet their impact on host populations remains largely unresolved. By contrast, fungal and protozoan infections of dinoflagellates are well documented and generally viewed as playing major roles in host population dynamics. Our understanding of fungal parasites is largely based on studies for freshwater diatoms and dinoflagellates, although fungal infections are known for some marine phytoplankton. In freshwater systems, fungal chytrids have been linked to mass mortalities of host organisms, suppression or retardation of phytoplankton blooms, and selective effects on species composition leading to successional changes in plankton communities. Parasitic dinoflagellates of the genus Amoebophrya and the newly described Perkinsozoa, Parvilucifera infectans, are widely distributed in coastal waters of the world where they commonly infect photosynthetic and heterotrophic dinoflagellates. Recent work indicates that these parasites can have significant impacts on host physiology, behavior, and bloom dynamics. Thus, parasitism needs to be carefully considered in developing concepts about plankton dynamics and the flow of material in marine food webs.

Changes in bacterial community composition and microbial activities along the longitudinal axis of two canyon-shaped reservoirs with different inflow loading

Changes in microbial activities and bacterial community composition (BCC) were studied using fluorescence in situ hybridization (FISH) along the longitudinal axis of two canyon-shaped reservoirs with contrasting retention times and nutrient and organic matter loads. The reservoirs showed different specific longitudinal changes in plankton communities and BCC shifts that were related mainly to allochthonous input and longitudinal patterns of primary production and nutrient availability. The studied branch of the Orlik reservoir is entered by a large river and so had a short retention time (23 days) and a large allochthonous input. Primary production, bacterial abundance, bacterial production and protozoan bacterivory showed a consistent trend, decreasing gradually from the river inflow downstream. Overall, shifts in BCC were minor and statistically insignificant. The relatively large input of the river-borne allochthonous plankton communities and the short retention time of the reservoir probably limited the effect of the autochthonous in-reservoirs processes. In contrast, the Řimov reservoir has a long retention time (∼100 days) and had a limited input of allochthonous organic matter to its lacustrine area. The autochthonous processes of primary and bacterial production were tightly linked and they showed peaks in the middle part of the studied stretch. This together with a trend of downstream decreasing nutrient concentrations and increasing protistan grazing pressure seem to support the development of specific BCC in the lacustrine part of the reservoir that significantly differed from that in river inflow. Apart from the marked differences between the limnological variables of reservoirs we found some commonalties in BCC – in both systems the phylogenetic groups of β-Proteobacteria and Cythophaga/Flavobacterium clearly dominated bacterioplankton assemblages. %

Paleozoic plankton revolution: Evidence from early gastropod ontogeny

A major macroevolutionary trend in gastropod larval shell morphology occurred throughout the middle and late Paleozoic. The majority of Ordovician and Silurian gastropods have openly coiled initial whorls; this morphology was gradually lost. Mesozoic gastropods have exclusively tightly coiled initial whorls. Most protoconchs were formed by planktonic larvae, and this is the first documentation of a supraspecific trend in Paleozoic invertebrate larvae. Open coiling makes shells weak and vulnerable to attack and force. The replacement of weak-shelled plankton by stronger-shelled plankton was probably caused by predatory, selective forces. It coincides with other fundamental changes in plankton communities during the middle and late Paleozoic, including the decline of acritarchs and graptolites. Plankton communities suffered from crises that were probably caused by extrinsic forces. During plankton restructuring in the aftermath, predation could have become an increasingly important selective factor for gastropod larvae.