Limnological survey of the warm monomictic lake Trichonis (central western Greece); II. Seasonal phytoplankton periodicity – a community approach

Phytoplankton assemblages of the warm monomictic lakeTrichoniswere studied during the period April 1985 to February 1986.Speciescomposition and biomass data are presented along withinformationon the seasonal periodicity of dominant taxa of microalgae.Multivariate methods were used to analyze community structureandannual succession. Population succession patterns correspondtochanges in environmental variables. According to thegeneralizedphytoplankton sequences Trichonis is classified as anoligotrophiclake.     

Effects of regular and irregular temporal patterns of disturbance on biomass accrual and species composition of a subtidal hard-bottom assemblage

Assessing patterns of species distribution and abundance is important to understand the driving processes of, and predict future changes in, biodiversity. To this date, ecological studies have been mainly designed to investigate the effects of the mean magnitude of predictor variables, although ecological factors naturally vary in space and time. In a nine month long field experiment, we tested the effects of different temporal patterns (regular, lowly and highly irregular) in biomass removal (=disturbance event) on the diversity, species composition, and biomass accrual of macrobenthic assemblages grown on 15 × 15 cm² PVC-panels. For each pattern of disturbance, disturbance events were timed at three sequences to control for possible confounding effects with recruitment patterns. Disturbance intensity was kept identical among treatments. Assemblages developed in the absence of disturbance for 3 months prior to a 150-day manipulation period, during which the biomass from 20% of the panel area was removed at each of ten disturbance events. Additional undisturbed settlement panels were deployed in the field to assess monthly recruitment rates and species succession over a one year period. Disturbance (i) reduced biomass and total species cover, (ii) changed species composition during the first half of the manipulation period significantly, and (iii) was without effect on species richness and evenness. Irregular disturbance regimes enhanced the abundance of the ascidian Ciona intestinalis, biomass accrual, and total species cover of assemblages relative to the regular disturbance regime, but had either no or only transient effects on diversity and species composition, respectively. Neither the degree of irregularity in disturbance nor the sequence of disturbance events affected any of the response variables significantly. Recruitment of species was strongly seasonal with almost only diatoms recruiting during winter, while recruitment was most intense during summer. Our results suggest that the temporal patterns of predictor variables might be of low explanatory power for the variance of responses in communities with seasonal recruitment patterns that are exposed to a high level of disturbance. Thus the need to include temporal patterns of predictor variables in experimental designs may depend on community dynamics and the characteristics of the process under investigation. S. Wollgast and M. Molis contributed equally to this work.     

Spatio-temporal changes of benthic macroinvertebrates in a cold Arkansas tailwater

This paper investigates spatial, seasonal and long-term changes in benthic macroinvertebrates in riffles of a cold tailwater. Cold tailwaters initially disrupt previously existing macroinvertebrate assemblages, but little is known about the long-term biological effects of a stable cold thermal regime. Assemblages at an upstream and downstream site of the Little Red River, Arkansas were investigated almost 30 years apart (1971 and 1999). Based upon published literature demonstrating the stability of benthic assemblages within unaltered environments, we predicted that the assemblages would be similar for each variable investigated. The benthic macroinvertebrate assemblages can be characterized as low diversity, with a total of 17 taxa identified. Isopods and Diptera comprised ~80% of all individuals. Other than chironomids, insects and particularly EPT taxa were poorly represented. Recent macroinvertebrate densities were significantly greater compared to the historical study period for the downstream site. Assemblage comparisons revealed moderate differences between study periods. Macroinvertebrate density was significantly greater upstream than downstream in the 1971 study period, yet taxa richness was significantly greater downstream for both study periods. Faunal composition was significantly different for upstream and downstream sites. Seasonal differences in numerical standing crop were identified for the 1971 upstream and 1999 downstream data sets. Low to moderate levels of seasonal, spatial and historical variation among benthic macroinvertebrate assemblages were attributed to environmental (temperature and flow) stability. The lack of aquatic insects other than chironomids over a 30-year period is indicative of the extreme constraints placed upon insect development within this cold regulated river.     

Nestedness and successional trajectories of macroinvertebrate assemblages in man-made wetlands

Current successional models, primarily those based on floral succession, propose several distinct trajectories based on the integration of two key hypotheses from succession theory: convergence versus divergence in species composition among successional sites, and progression towards versus deviation from a desired reference state. We applied this framework to faunal succession, including differential colonization between active and passive dispersers, and the nested patterns generated as a consequence of this peculiarity. Nine man-made wetlands located in three different areas, from 0–3 years from wetland creation, were assessed. In addition, 91 wetlands distributed throughout the region were used as references for natural macroinvertebrate communities. We predicted the following: (1) highly nested structures in pioneering assemblages will decrease to lower mid-term values due to a shift from active pioneering taxa to passive disperser ones; (2) passive idiosyncratic taxa will elicit divergent successional trajectories among areas; (3) the divergent trajectories will provoke lower local and higher regional diversity values in the mid-term assemblages than in pioneer assemblages. Our results were largely congruent with hypotheses (1) and (2), diverging from the anticipated patterns only in the case of the temporary wetlands area. However, overall diversity trends based on hypothesis (3) did not follow the expected pattern. The divergent successional trajectories did not compensate for regional biodiversity losses that occurred as a consequence of pioneering colonizer decline over time. Consequently, we suggest reconsidering wetland construction for mitigation purposes within mid-term time frames (≤3 years). Wetlands may not offset, within this temporal scenario, regional biodiversity loss because the ecosystem may not support idiosyncratic taxa from natural wetlands.     

Quantifying temporal variation in heterobranch (Mollusca: Gastropoda) sea slug assemblages: tests of alternate models

Assemblages of heterobranch sea slugs are notoriously variable in space and time which has sometimes led to their exclusion from broader studies of patterns of biodiversity. This variability may also result in a failure to detect underlying spatial and temporal patterns. Assemblage data (species abundances) from three intertidal sites over a period of 13 months, were used to compare the conclusions drawn from three different analytical approaches—a non-specific test for differences among months, and tests of cyclicity, and seriation, using specific model matrices. While no significant difference was detected for the non-specific test (i.e. concluding no significant temporal variation), the data were significantly correlated to both model matrices with the cyclicity model outperforming the seriation model at two of the three sites. The study also highlighted the spatial variability of assemblages over a scale of just a few kilometres. Wider testing using previously published datasets confirmed the utility of these models for exploring specific hypotheses about patterns of temporal change.     

Metabarcoding improves detection of eukaryotes from early biofouling communities: implications for pest monitoring and pathway management

In this experimental study the patterns in early marine biofouling communities and possible implications for surveillance and environmental management were explored using metabarcoding, viz. 18S ribosomal RNA gene barcoding in combination with high-throughput sequencing. The community structure of eukaryotic assemblages and the patterns of initial succession were assessed from settlement plates deployed in a busy port for one, five and 15 days. The metabarcoding results were verified with traditional morphological identification of taxa from selected experimental plates. Metabarcoding analysis identified > 400 taxa at a comparatively low taxonomic level and morphological analysis resulted in the detection of 25 taxa at varying levels of resolution. Despite the differences in resolution, data from both methods were consistent at high taxonomic levels and similar patterns in community shifts were observed. A high percentage of sequences belonging to genera known to contain non-indigenous species (NIS) were detected after exposure for only one day.     

Structure of Benthic Diatom Assemblages from a Mangrove Environment in a Mexican Subtropical Lagoon1

The taxonomic list and the structure of benthic diatom assemblages occurring in fine sediments (silt and sand) from the mangrove forest of the Balandra lagoon in Baja California Sur, Mexico was determined based on seasonal samplings for one year. Assemblage structure was analyzed using several ecological indices for estimating diversity (H'), dominance (REDI), equitability, and similarity. A total of 230 diatom taxa were identified and include 109 new records for the Baja California peninsula coast. Taxa representative of highly productive and hypersaline environments were common. Assemblages were characterized by a few abundant species and many uncommon or rare taxa. High diatom diversity estimates at all sampling sites during all seasons suggest that diatom assemblages in sediments of the Balandra lagoon represent a quasi-pristine environment.     

Distribution patterns of benthic foraminifera across the Ypresian–Lutetian Gorrondatxe section,Northern Spain: Response to sedimentary disturbance

We present a study of benthic foraminiferal assemblages from an Ypresian–Lutetian distal submarine fan system in the lower bathyal Gorrondatxe section (Basque-Cantabrian Basin, northern Spain). The objective of our study is to analyze the benthic foraminiferal distribution patterns and their response to sedimentary disturbance and related factors.Assemblages contain a high percentage of allochthonous taxa, such as asterigerinids and other shallow water taxa, which were transported downslope by turbidity currents.Detailed quantitative analyses, supported by R-mode cluster and Detrended Correspondence Analyses (after removing allochthonous taxa from the foraminiferal counts) allowed us to identify 6 assemblages that are divided into two groups related to the turbidite content in the Gorrondatxe section. Assemblages 1, characteristic of the turbidite-poor intervals with low sedimentary disturbance, include assemblage 1a (with highly diverse common middle–lower bathyal calcareous taxa) assemblage 1b (with common agglutinated taxa, mainly trochamminids), and assemblage 1c (characterized by calcareous taxa that are also common in the turbidite-rich interval).Assemblages 2, characterized by a high dominance, prevail in the turbidite-rich interval, and include assemblage 2a (characterized by the dominance of infaunal bolivinids and epifaunal cibicids), assemblage 2b (typified by moderate to low diversity and dominated by deep-infaunal Globobulimina species), and assemblage 2c (typified by very abundant suspension-feeding astrorhizids). The high abundance of bolivinids and Globobulimina species may be related to an enhanced input of low-quality organic matter transported by turbidity currents to the seafloor, representing different stages of recolonisation after disturbance and different energy regimes. High current activity was probably responsible for the abundance of cibicids, while moderate to low diverse and high dominance assemblages characterize the recolonisation of the substrate after disturbance.We conclude that sedimentary disturbance and other related factors such as current activity, resuspension of sediments at the seafloor, and supply of organic matter (and its quality) played an important role in the distribution of benthic foraminifera in the Gorrondatxe section. The identification of allochthonous taxa emerges as an essential aspect of the study of environments with sedimentary disturbance.     

Seasonal dynamics and composition of cladoceran and copepod assemblages in ponds of a Hungarian cutaway peatland

Seasonal dynamics of cladoceran and copepod zooplankton were studied over a one‐year period in five permanent ponds of a cutaway peatland, situated in the Danube−Tisza Interfluve, Central Hungary. 17 cladoceran, 11 cyclopoid copepod and 6 harpacticoid copepod species were identified and most of them were typical of small lowland ponds. Nevertheless, some taxa like Cyclops insignis, Ceriodaphnia setosa and Macrocyclops distinctus are considered to be rare in Hungary. The microcrustacean assemblages exhibited apparent seasonal succession with typical seasonal species. There appears to be at least two main successional patterns in the five ponds. After general cyclopoid copepod dominance in winter (Cyclops strenuus and Cyclops insignis), at sites with higher proportion of open water and submerged vegetation, spring was characterized by the dominance of the large cladoceran Daphnia curvirostris, which declined during summer, when microcrustacean assemblages composed mainly of smaller, littoral cladocerans. At these sites, species richness and diversity reached their maximum in autumn. In the case of duckweed covered ponds, succession led to less diverse autumn assemblages with fewer species, dominated by Simocephalus exspinosus. Our results draw the attention to the importance of long‐term investigations and the often neglected winter sampling in the accurate evaluation of species richness (© 2012 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Decadal abundance patterns in an isolated urban reptile assemblage: Monitoring under a changing climate

Determine seasonal, annual, and decadal patterns of abundance in reptile species and assemblages occupying central Bold Park (~338 ha), an isolated urban bushland remnant in Perth, Southwestern Australia. Fenced pitfall trapping in four sampling sites, representing different habitats and fire history, over the primary reptile activity period for 35 consecutive years with over 17,000 individuals captured during 3300 days of sampling; the trapping regime was modified for the last 28 years. Sampling occurred in one of 35 global biodiversity hotspots that has a Mediterranean climate experiencing a 15% decline from the century average rainfall over the last 50 years. Twenty‐nine species were recorded, with 16 captured in 32 or more years and accounting for nearly 97% of all captures; the six most common for 81%. Three taxa became locally extinct. Activity predominates in warmer and dryer months (October to April), peaking in November–December. Species richness remained relatively constant between years with around 73% of known taxa captured annually. Assemblages did not change when analyzing the presence/absence data but moved through five statistically significant assemblages analyzing relative abundance data. Over the last 28 years, relative abundance was significantly and positively correlated with annual rainfall residuals, uniquely for the 4 years preceding annual sampling, resulting in significant changes in total assemblages and significantly similar patterns in four sample sites; the presence/absence data indicated only minor assemblage changes across sites. The number of species recorded annually remained relatively constant, but relative abundance illustrated significant temporal changes in assemblages over decades. The modeled relationship between relative abundance and annual rainfall residuals for 4 years preceding annual sampling is supported by known ecological responses and reptile demographics within this Mediterranean climate. Maintenance of urban biodiversity should consider impacts of a significantly drying climate exacerbating the extinction debt already inherent in isolated bushland populations experiencing limited immigration.     

Surface composition and orientation interact to affect subtidal epibiota

Settlement panels were used to evaluate the effects of composition of the substratum (sandstone, concrete, wood) and orientation (vertical, horizontal undersides) on subtidal epibiota. It was predicted that both factors would influence the development of epibiotic assemblages, but that differences due to composition would be less marked on horizontal undersides compared to vertical panels. Differences in assemblages among sandstone, concrete and wooden panels orientated vertically were predicted to be similar to those described previously among vertical surfaces of sandstone rocky reefs and concrete and wooden urban structures (pilings and pontoons). Multivariate analyses indicated that assemblages were influenced greatly by orientation, whereas the effects of surface composition differed for the two orientations and among sites. Assemblages on wood were always significantly different from those on sandstone or concrete - patterns between the latter two surfaces depended on the orientation of the panels. The taxa that dominated these surfaces were not similar in identity nor abundance to those on urban structures of the same composition. The covers of most taxa were influenced by orientation alone or by surface composition for just one orientation. This study demonstrates the need for caution in generalizing about effects of orientation and surface composition because they may interact with each other and/or with other factors and they are certainly quite different for different taxa and among sites.     

Stochastic and deterministic processes jointly structure tropical arthropod communities

The question of whether ecological assemblages are structured by stochastic and deterministic (e.g. interspecific competition) processes is controversial, but it is difficult to design sampling regimes and experiments that can dissect the relative importance of stochastic and deterministic processes in natural assemblages. Using null models, we tested communities of arthropod decomposers in tropical epiphytes for patterns of species co-occurrence, while controlling for habitat gradients, seasonal variations and ecological succession. When environmental conditions were controlled, our analysis showed that the communities were structured stochastically. However, analysing mixed sets of communities that were deliberately created either from two distinct heights or two successional stages revealed that communities were structured deterministically. These results confirm that habitat gradients and dispersal/competition trade-offs are capable of generating non-random patterns within decomposer arthropod communities, but reveal that when such effects are accounted for, species co-occurrence is fundamentally random.     

Broad-Scale Patterns of Late Jurassic Dinosaur Paleoecology

Background

There have been numerous studies on dinosaur biogeographic distribution patterns. However, these distribution data have not yet been applied to ecological questions. Ecological studies of dinosaurs have tended to focus on reconstructing individual taxa, usually through comparisons to modern analogs. Fewer studies have sought to determine if the ecological structure of fossil assemblages is preserved and, if so, how dinosaur communities varied. Climate is a major component driving differences between communities. If the ecological structure of a fossil locality is preserved, we expect that dinosaur assemblages from similar environments will share a similar ecological structure.

Methodology/Principal Findings

This study applies Ecological Structure Analysis (ESA) to a dataset of 100+ dinosaur taxa arranged into twelve composite fossil assemblages from around the world. Each assemblage was assigned a climate zone (biome) based on its location. Dinosaur taxa were placed into ecomorphological categories. The proportion of each category creates an ecological profile for the assemblage, which were compared using cluster and principal components analyses. Assemblages grouped according to biome, with most coming from arid or semi-arid/seasonal climates. Differences between assemblages are tied to the proportion of large high-browsing vs. small ground-foraging herbivores, which separates arid from semi-arid and moister environments, respectively. However, the effects of historical, taphonomic, and other environmental factors are still evident.

Conclusions/Significance

This study is the first to show that the general ecological structure of Late Jurassic dinosaur assemblages is preserved at large scales and can be assessed quantitatively. Despite a broad similarity of climatic conditions, a degree of ecological variation is observed between assemblages, from arid to moist. Taxonomic differences between Asia and the other regions demonstrate at least one case of ecosystem convergence. The proportion of different ecomorphs, which reflects the prevailing climatic and environmental conditions present during fossil deposition, may therefore be used to differentiate Late Jurassic dinosaur fossil assemblages. This method is broadly applicable to different taxa and times, allowing one to address questions of evolutionary, biogeographic, and climatic importance.     

Small-scale temporal variability in assemblages of larval fishes: implications for sampling

Short-term changes in the assemblages of larval fishes in watersover the inner continental shelf off Sydney (Australia) wereexamined.This was done to highlight the potential sampling problemof confounding short-term variability with longer-term and spatialvariability in plankton communities. Assemblages at the surfaceand at 30 m depth were sampled twice a day over consecutivedays at two locations 5 km apart. The assemblages and the densitiesof individual taxa varied greatly between sampling times anddays. The small-scale temporal changes in densities of larvaewere not the same at both locations or between depths. Largeand consistent depth-related differences in assemblages andabundances of most taxa were evident. The consequences of small-scaletemporal variations are discussed relative to the design ofsampling programs for ecological and environmental purposes.Studies that include the double sampling of some stations arerecommended. Greater emphasis needs to be placed on fine-scaleoceanography and larval behaviour to determine causative mechanismsof short-term changes in assemblages of larval fishes.     

Litter Breakdown and Microbial Succession on Two Submerged Leaf Species in a Small Forested Stream

Microbial succession during leaf breakdown was investigated in a small forested stream in west-central Georgia, USA, using multiple culture-independent techniques. Red maple (Acer rubrum) and water oak (Quercus nigra) leaf litter were incubated in situ for 128 days, and litter breakdown was quantified by ash-free dry mass (AFDM) method and microbial assemblage composition using phospholipid fatty acid analysis (PLFA), ribosomal intergenic spacer analysis (RISA), denaturing gradient gel electrophoresis (DGGE), and bar-coded next-generation sequencing of 16S rRNA gene amplicons. Leaf breakdown was faster for red maple than water oak. PLFA revealed a significant time effect on microbial lipid profiles for both leaf species. Microbial assemblages on maple contained a higher relative abundance of bacterial lipids than oak, and oak microbial assemblages contained higher relative abundance of fungal lipids than maple. RISA showed that incubation time was more important in structuring bacterial assemblages than leaf physicochemistry. DGGE profiles revealed high variability in bacterial assemblages over time, and sequencing of DGGE-resolved amplicons indicated several taxa present on degrading litter. Next-generation sequencing revealed temporal shifts in dominant taxa within the phylum Proteobacteria, whereas γ-Proteobacteria dominated pre-immersion and α- and β-Proteobacteria dominated after 1 month of instream incubation; the latter groups contain taxa that are predicted to be capable of using organic material to fuel further breakdown. Our results suggest that incubation time is more important than leaf species physicochemistry in influencing leaf litter microbial assemblage composition, and indicate the need for investigation into seasonal and temporal dynamics of leaf litter microbial assemblage succession.     

Factors influencing macroinvertebrate assemblages in artificial subtropical ponds of Taiwan

Macroinvertebrate assemblages and its association with environmental factors at the 11 artificial subtropical ponds of Taiwan were examined using the multivariate analysis software STATICO. The aims of the study were to determine whether spatial and seasonal variation of macroinvertebrate assemblages changed seasonally, to examine which environmental factors determined the spatial and temporal structure of maroinvertebrate assemblages, and to compare between-pond variations in the taxon composition of macroinvertebrates. Macroinvertebrates were collected seasonally by a corer and a sweep net in 2007, and 13 physical and chemical factors were measured at the same time. A total of 31 macroinvertebrate taxa were collected during the sampling period, and the most dominant taxa were Chironomidae (31.7% of total animal abundance) and Tubificidae (22.4%). STATICO identified pond size, pond depth, sediment depth, and altitude as the major abiotic factors and Bufo melanostictus (Amphibia) as the major biotic factor to influence macroinvertebrate assemblages at these ponds. These factors changed with seasonality. For example, the abundance of B. melanostictus was the most important factor during the spring but became much less important in other seasons. According to the spatial distribution patterns of macroinvertebrate assemblages, macroinvertebrates could be split into two groups based on their dispersal. The active dispersers, such as insect taxa, were strongly associated with pond size and the passive dispersers, such as non-insect taxa, were strongly associated with the pond depth and/or sediment depth. The results of this study suggested that pond size might influence macroinvertebrate assemblages through their dispersal mechanisms and that the environmental factors which influenced the macroinvertebrate assemblages most changed with seasons in this study area.     

Seasonal patterns of horse fly richness and abundance in the Pampa biome of southern Brazil

Fluctuations in seasonal patterns of horse fly populations were examined in rainforests of tropical South America, where the climate is seasonal. These patterns were evaluated with robust analytical models rather than identifying the main factors that influenced the fluctuations. We examined the seasonality of populations of horse flies in fields and lowland areas of the Pampa biome of southern Brazil with generalized linear models. We also investigated the diversity of these flies and the sampling effort of Malaise traps in this biome over two years. All of the 29 species had clear seasonality with regard to occurrence and abundance, but only seven species were identified as being influenced by temperature and humidity. The sampling was sufficient and the estimated diversity was 10% more than observed. Seasonal trends were synchronized across species and the populations were most abundant between September and March and nearly zero in other months. While previous studies demonstrated that seasonal patterns in population fluctuations are correlated with climatic conditions in horse fly assemblages in South America rainforests, we show a clear effect of each factor on richness and abundance and the seasonality in the prevalence of horse fly assemblages in localities of the Pampa biome.     

SEASONAL PERIODICITY OF CHRYSOPHYCEAE AND SYNUROPHYCEAE IN A SMALL NEW ENGLAND LAKE: IMPLICATIONS FOR PALEOLIMNOLOGICAL RESEARCH1

The seasonal periodicity of taxa of Chrysophyceae and Synurophyceae from a small New England lake is described for the period September 1983 through June 1988. We found 51 taxa, including 29 that accounted for over 10% of the total in at least one collection. The taxa were fitted into one of five seasonal patterns. Patterns I and II represented taxa restricted to warm (pattern I) or cold (pattern II) months, respectively. Pattern HI represented organisms that began growth in the summer, persisted through autumn and disappeared with the onset of an ice cover. Pattern IV was an extension of pattern III, in which the taxon remained in the plankton throughout the winter and disappeared soon after ice out. Species without a clear seasonal pattern were grouped as pattern V. The seasonal periodicity of the flora, as examined with ordination analyses, was found to remain remarkedly similar during the 58–month study. Except for episodes of low pH during spring snow melt and unseasonally warm or cold weather, sample scores followed a fairly consistent pattern along the first and second primary axes. Water temperature, specific conductance, and pH were important variables that controlled changes in the species composition during the course of a given year. The flora was used to develop an inference model for water temperature. According to the analyses, the remains of a surface sediment sample represented a flora that grew primarily during the late autumn period at 7.6 ° C. Ways in which seasonal data could be utilized to improve paleolimnological inference work are discussed.     

Spatial and temporal variability in early successional patterns of an invertebrate assemblage at an offshore oil platform

The effects of selected physical and biological factors on the early development of a subtidal invertebrate assemblage were examined at an offshore oil platform in the Santa Barbara Channel (California, USA). The effects of date, year, length, and depth of submersion were investigated by replacing sets of ceramic tiles with new tiles at frequencies of 2, 4, 6, 12, and 24 months at three depths (6, 12, and 18 m) over a period of 24 months. The effects of existing colonists and depth were explored in a second experiment by removing selected early colonists from ceramic tiles deployed at the same three depths over a period of 12 months. More than 40 invertebrate taxa from seven phyla colonized the tiles. Colonial tunicates and encrusting bryozoans appeared early in the successional sequence (∼2 months), in cover ranging from <5% to 80% and from <5% to 55%, respectively. Tubiculous amphipods, barnacles, and sponges could also appear early, but in low cover (<20%). Composition of the assemblage changed over time with barnacles, sponges, and mussels becoming the principal space holders on tiles submerged for 24 months. When potential competitors were removed monthly, variation in the cover of dominant taxa (i.e., bryozoans, amphipods, barnacles) was maintained to 12 months, suggesting that other factors, such as larval availability or post-settlement mortality, were responsible for these patterns. Development of this assemblage appeared to fit a pattern of early succession that was largely predictable in terms of the composition and sequence of occurrence of dominant taxa, but variable in rate of development, depending on date of submersion, year, and depth.