Transoceanic ships as vectors for nonindigenous freshwater bryozoans

Aim The transport of organisms in ships’ ballast tanks is a dominant vector for aquatic invasions worldwide. Until recently, efforts to manage this vector have overlooked the potential transport of invertebrate resting stages in the residual waters and sediments within emptied ballast tanks, i.e. NOBOB (‘No Ballast On Board’) tanks. The resting stages (statoblasts) of freshwater bryozoans are often buoyant and locally abundant and thus can be taken up easily during ballasting operations. They are also resistant to extreme environmental conditions and can generate new colonies after being dormant for decades; as such, they would likely remain viable propagules after lengthy transport in ship ballast tanks. This study quantified the occurrence of freshwater bryozoan statoblasts in ballast tank sediments of transoceanic ships. Location North American Great Lakes. Methods We quantified the frequency of occurrence, abundance and diversity of bryozoans (as statoblasts) in residual sediment samples taken from 51 NOBOB tanks of 33 transoceanic ships visiting the Great Lakes from 2000 to 2002. Results Our study identified 11 species, comprising nearly 12% of the total number of freshwater bryozoans known worldwide. These include two exotic species unrecorded in the Great Lakes (Fredericella sultana and Lophopus crystallinus), an exotic species already established in the region (Lophopodella carteri) and three cosmopolitan species (Plumatella casmiana, P. fungosa and P. repens). Our estimates suggest that a ship with NOBOB tanks may carry up to 10⁶ statoblasts. Main conclusions The discovery of species unrecorded in the Great Lakes and the potentially large numbers of statoblasts being transported in ship ballast tanks indicate a significant risk of new species introductions. Furthermore, the presence of cosmopolitan species and an exotic species already established in the Great Lakes suggests the strong possibility of cryptic invasions via the introduction of exotic genotypes.     

Divergence and species discrimination in freshwater bryozoans (Bryozoa: Phylactolaemata)

We explored the suitability of nuclear and mitochondrial ribosomal markers [small subunit nuclear ribosomal RNA gene, large subunit nuclear ribosomal RNA gene, and a region spanning partial small mitochondrial ribosomal RNA subunit, four transfer RNA genes, and partial large mitochondrial ribosomal RNA subunit (referred to as rrnS‐rrnL)] for resolving patterns of diversification of 27 freshwater bryozoan species (class: Phylactolaemata) and evaluated the utility of statoblast ultrastructural features and molecular phylogenies for species discrimination in the Fredericellidae and Plumatellidae. Molecular data identified Plumatella fruticosa as distinct from the rest of the plumatellids, rendering the latter polyphyletic. rrnS‐rrnL was the most suitable marker for species discrimination and identified two undescribed species of Plumatella and at least two undescribed species of Fredericella. Lack of wide dispersal by fredericellid statoblasts may underlie the observed propensity for cryptic speciation and phylogeographical structure in Fredericella. Conversely, the strong dispersal potential of plumatellid statoblasts may mediate efficient gene flow between distant populations and explain the relatively low intraspecific divergence and lack of evidence for cryptic speciation. We show that species identification based on external features of statoblasts can be problematic in both genera, including for a putatively highly invasive, biofouling species, Plumatella vaihiriae, thereby highlighting the utility of rrnS‐rrnL sequences for species barcoding. © 2013 The Linnean Society of London     

Bryozoans are returning home: recolonization of freshwater ecosystems inferred from phylogenetic relationships

Bryozoans are aquatic invertebrates that inhabit all types of aquatic ecosystems. They are small animals that form large colonies by asexual budding. Colonies can reach the size of several tens of centimeters, while individual units within a colony are the size of a few millimeters. Each individual within a colony works as a separate zooid and is genetically identical to each other individual within the same colony. Most freshwater species of bryozoans belong to the Phylactolaemata class, while several species that tolerate brackish water belong to the Gymnolaemata class. Tissue samples for this study were collected in the rivers of Adriatic and Danube basin and in the wetland areas in the continental part of Croatia (Europe). Freshwater and brackish taxons of bryozoans were genetically analyzed for the purpose of creating phylogenetic relationships between freshwater and brackish taxons of the Phylactolaemata and Gymnolaemata classes and determining the role of brackish species in colonizing freshwater and marine ecosystems. Phylogenetic relationships inferred on the genes for 18S rRNA, 28S rRNA, COI, and ITS2 region confirmed Phylactolaemata bryozoans as radix bryozoan group. Phylogenetic analysis proved Phylactolaemata bryozoan's close relations with taxons from Phoronida phylum as well as the separation of the Lophopodidae family from other families within the Plumatellida genus. Comparative analysis of existing knowledge about the phylogeny of bryozoans and the expansion of known evolutionary hypotheses is proposed with the model of settlement of marine and freshwater ecosystems by the bryozoans group during their evolutionary past. In this case study, brackish bryozoan taxons represent a link for this ecological phylogenetic hypothesis. Comparison of brackish bryozoan species Lophopus crystallinus and Conopeum seurati confirmed a dual colonization of freshwater ecosystems throughout evolution of this group of animals.     

The effects of infection by Tetracapsuloides bryosalmonae (Myxozoa) and temperature on Fredericella sultana (Bryozoa)

The myxozoan, Tetracapsuloides bryosalmonae, exploits freshwater bryozoans as definitive hosts, occurring as cryptic stages in bryozoan colonies during covert infections and as spore-forming sacs during overt infections. Spores released from sacs are infective to salmonid fish, causing the devastating Proliferative Kidney Disease (PKD). We undertook laboratory studies using mesocosm systems running at 10, 14 and 20 °C to determine how infection by T. bryosalmonae and water temperature influence fitness of one of its most important bryozoan hosts, Fredericella sultana, over a period of 4 weeks. The effects of infection were context-dependent and often undetectable. Covert infections appear to pose very low energetic costs. Thus, we found that growth of covertly infected F. sultana colonies was similar to that of uninfected colonies regardless of temperature, as was the propensity to produce dormant resting stages (statoblasts). Production of statoblasts, however, was associated with decreased growth. Overt infections imposed greater effects on correlates of host fitness by: (i) reducing growth rates at the two higher temperatures; (ii) increasing mortality rates at the highest temperature; (iii) inhibiting statoblast production. Our results indicate that parasitism should have a relatively small effect on host fitness in the field as the negative effects of infection were mainly expressed in environmentally extreme conditions (20 °C for 4 weeks). The generally low virulence of T. bryosalmonae is similar to that recently demonstrated for another myxozoan endoparasite of freshwater bryozoans. The unique opportunity for extensive vertical transmission in these colonial invertebrate hosts couples the reproductive interests of host and parasite and may well give rise to the low virulence that characterises these systems. Our study implies that climate change can be expected to exacerbate PKD outbreaks and increase the geographic range of PKD as a result of the combined responses of T. bryosalmonae and its bryozoan hosts to higher temperatures.     

Novel morphological and genetic tools to discriminate species among the family Plumatellidae (Phylactolaemata,Bryozoa)

Some species of the freshwater bryozoans (Bryozoa, Phylactolaemata) belonging to the genus Plumatella are remarkably difficult to identify because of the large similarity of superficial architecture of their statoblasts. The examination of statoblasts by scanning electron microscope (SEM) has in fact resolved only some taxonomic questions. In this article, the authors report on novel morphological and molecular traits to discriminate among ten species of Plumatellidae (P. viganoi, P. repens, P. geimermassardi, P. rugosa, P. reticulata, P. casmiana, P. fungosa, P. emarginata, P. vaihiriae, and Hyalinella punctata). The former traits are based on shape, number, and position of annular chamber pores, whereas the latter reside on amplification and sequence analysis of the Internal Transcribed Spacer (ITS) region of nuclear rDNA. The successful amplification of ITS region from statoblasts and zooidial tubules allowed us to sequence this region on all the species investigated. The ITS sequences showed the presence of sufficient and informative polymorphisms to discriminate among morphologically similar species. It is noteworthy that the resulting ITS phylogenetic tree largely corroborated the distinction of at least two groups of freshwater bryozoans inferred on the basis of the annular chamber pore morphology. This study provides innovative approaches to reliably characterize freshwater bryozoans species and gain more insight into their taxonomy, phylogenetic relationship, and biodiversity.     

Cryptic invaders: nonindigenous and cryptogenic freshwater Bryozoa and Entoprocta in the St. Lawrence River

The distributions of most cosmopolitan invertebrate species are assumed to result from natural processes. Cryptic invertebrates with obscure biogeographic origins are often considered native by default, resulting in potentially severe underestimation of the extent of human-assisted invasions. This problem is exemplified by freshwater Bryozoa (Ectoprocta) and Entoprocta—small and widely distributed invertebrates commonly found in lakes and rivers. A benthic survey of a thermally modified section of the St. Lawrence River revealed the presence of two nonindigenous bryozoans: Carter’s moss animal Lophopodella carteri (Hyatt) and the crystal moss animal Lophopus crystallinus Pallas. Also discovered was a cryptogenic entoproct, the goblet worm Urnatella gracilis Leidy. These species were collected as statoblasts and (in the case of U. gracilis) colonial fragments downstream of the Gentilly-2 nuclear power plant at Bécancoeur, Quebec. Local densities of both U. gracilis and L. carteri increased by an order of magnitude at sites closer to the power plant. The occurrence of Lophopus crystallinus statoblasts in St. Lawrence River sediments is the first documented physical evidence of the species in North America. Contrary to the presumed natural Holarctic distribution of L. crystallinus, our literature review found that published historical records of L. crystallinus in the United States are erroneous or unsubstantiated. We propose that L. crystallinus is a western Palearctic species recently introduced to the St. Lawrence River, most likely as statoblasts discharged with ballast water from transoceanic ships.     

Comparative studies on the statoblasts of higher phylactolaemate bryozoans

Statoblasts of five higher phylactolaemates were compared morphologically. As a result, they were divided into two groups: Group I comprising Lophopus crystallinus, Lophopodella carteri, and Pectinatella gelatinosa, and Group II comprising Pectinatella magnifica and Cristatella mucedo. These two groups are thought to represent independent evolutionary series. In Group I and in P. magnifica, the statoblasts are curved to varying degrees after the manner of a saddle. When the dorsal and ventral valves are flattened, therefore, the contour is different between the two. In Group I, the outermost layer of a mature statoblast is hard-gelatinous and basophilic; it remains intact after the statoblast is set free. The statoblast does not float until it is dry, and the float is similar in size on both valves. In Group II, a mature statoblast is covered by a softgelatinous basophilic layer, which decays after the statoblast is released. The statoblast floats without drying, and the float is better developed on the dorsal valve than on the ventral. Moreover, in the members of Group II, large yolk granules are first formed, followed by much smaller yolk granules. When their statoblasts are treated with KOH, the shell is separated completely into two valves. These characters are common to many lower phylactolaemates. By contrast, in L. carteri and P. gelatinosa, the yolk granules are uniformly small and the capsule proper resists KOH treatment. On these points, L. crystallinus is somewhat different from these two species, suggesting its primitive nature.     

A benign approach to the preparation of freshwater bryozoan statoblasts for scanning electron microscopy (SEM) imaging

Abstract

Several different species of freshwater Bryozoa, belonging to the genera Plumatella, Rumarcanella and Fredericella, were detected within the Northern Mallee Pipeline (NMP) system in Victoria, Australia, that required definitive identification. These organisms produce asexual buds called statoblasts, with valves composed of sclerotised chitin that bear minute micro-ornamentations of considerable taxonomical significance. Imaging and analysis of these distinctive micro-ornamentations using scanning electron microscopy (SEM) is often employed for species identification. Meticulous preparation of statoblast samples is therefore required that necessitates the removal of adhering debris, dehydration and drying—whilst mitigating specimen damage and distortion. This technical note describes an approach whereby each of these three steps have been individually designed to be as benign as possible, using mild detergent/sonication to remove debris, a gradual and gentle dehydration procedure using ethanol, and critical point drying. For the overall process, these methods are chosen to optimise control and to minimise the use of harsh and hazardous chemicals.     

Are plumatellid statoblasts in freshwater bryozoans phylogenetically informative?

Morphological characters of statoblasts (including floatoblasts and sessoblasts) in freshwater bryozoans have been important in phylactolaemate systematics and identification in that older phylogenetic hypotheses relied heavily on statoblast morphology. To assess the reliability of statoblast characters in drawing conclusions about phylogeny, we examined the phylogenetic distribution of metric and proportional floatoblast characters, floatoblast symmetry, and floatoblast and sessoblast microsculpture in Plumatellidae, the largest family of phylactolaemates, in the context of molecular phylogenetic reconstructions based on nucleotide sequences of parts of the mitochondrial 12S rRNA, 16S rRNA, and cytochrome b (cytb) genes. Adding cytb sequences to a previous phylogeny based on 12S and 16S increased support only for a node including Plumatella vaihiriae in the P. repens clade. Characters of gross floatoblast morphology were generally not phylogenetically informative individually, but collectively discriminated among members of three relatively well-supported clades that were analyzed as pre-defined groups in a discriminant analysis. Two characters of floatoblast microsculpture (reticulation and hypertubercles) were restricted to particular clades; other characters (e.g., villi) were clearly convergent. In nine of 11 cases, fenestral microsculpture was identical or partly correlated between the floatoblast and sessoblast of a species. Overall, our results indicate that statoblast morphology is not highly phylogenetically constrained.     

Bryozoan colonization of the marine isopod Glyptonotus antarcticus at Signy Island, Antarctica

Sixty specimens of the giant marine isopod Glyptonotus antarcticus Eights, collected from Borge Bay, Signy Island, Antarctica were examined for epizoans. Ten species of cheilostomatid bryozoans were found on the isopods. The purpose of the study was to quantify the prevalence, intensity, abundance, and spatial distribution of the bryozoans on the isopods. The proportion of isopods colonized was 42%. The larger isopods had both significantly more epizoic bryozoan colonies and species. The greatest density of bryozoans was on the fused pleon and telson. There was no significant difference between the dorsal and ventral abundance of bryozoan colonies. The diversity of epizoic bryozoans on the isopods is higher than on other host organisms from more stable environments. This may be because of active selection by settling larvae. The frequency of local substrata being scoured by ice is high around Signy Island, so there may be a selective advantage in colonizing a motile host. Accepted: 20 June 1998     

Evaluation of malacosporean life cycles through transmission studies

Myxozoans, belonging to the recently described Class Malacosporea, parasitise freshwater bryozoans during at least part of their life cycle, but no complete malacosporean life cycle is known to date. One of the 2 described malacosporeans is Tetracapsuloides bryosalmonae, the causative agent of salmonid proliferative kidney disease. The other is Buddenbrockia plumatellae, so far only found in freshwater bryozoans. Our investigations evaluated malacosporean life cycles, focusing on transmission from fish to bryozoan and from bryozoan to bryozoan. We exposed bryozoans to possible infection from: stages of T. bryosalmonae in fish kidney and released in fish urine; spores of T. bryosalmonae that had developed in bryozoan hosts; and spores and sac stages of B. plumatellae that had developed in bryozoans. Infections were never observed by microscopic examination of post-exposure, cultured bryozoans and none were detected by PCR after culture. Our consistent negative results are compelling: trials incorporated a broad range of parasite stages and potential hosts, and failure of transmission across trials cannot be ascribed to low spore concentrations or immature infective stages. The absence of evidence for bryozoan to bryozoan transmissions for both malacosporeans strongly indicates that such transmission is precluded in malacosporean life cycles. Overall, our results imply that there may be another malacosporean host which remains unidentified, although transmission from fish to bryozoans requires further investigation. However, the highly clonal life history of freshwater bryozoans is likely to allow both long-term persistence and spread of infection within bryozoan populations, precluding the requirement for regular transmission from an alternate host.     

Annual net production and life span of floating leaves in Nymphaea tetragona Georgi: a comparison with other floating-leaved macrophytes

Fredericella sultana (Blumenbach, 1779) has long been considered one of the few freshwater bryozoan species with a truly cosmopolitan distribution. However, chromosome spreads from European material show 2n = 16 compared to 2n = 14 in North American specimens. In laboratory rearing the two forms are morphologically indistinguishable except for the surface texture of their statoblasts. Smooth statoblasts of European colonies match early illustrations of the species, while the densely pitted statoblasts of the North American form resemble those of F. indica Annandale 1909. On the basis of these observations we tentatively designate the North American F. sultana as F. indica. The only known American species with smooth statoblasts is F. australiensis Goddard 1909, in which the 2n = 16 karyotype is similar or identical to European F. sultana; however, despite this karyotypic similarity the two species retain their distinguishing morphology when reared together in the laboratory. Two enzymes from a single specimen of European F. sultana were electrophoretically distinct from the corresponding enzymes present in samples of both F. australiensis and North American F. sultana. Four phosphoglucose isomerase alleles were present in North American F. aultana from four geographically separated collection sites, although only one genotype for this locus was observed in material from any one site. These genetic findings are consistent with a relatively short-range dispersal potential in this species as compared to Plumatella species.     

Fusion of ancestrulae germinated from statoblasts in plumatellid freshwater bryozoans

Upon germination of a statoblast, the shell is split into two valves; a mucous pad which represents the basal portion of the body wall of the incipient zooid or ancestrula then appears from between the valves; lastly, a tiny polypide evaginates at the opposite site. When two or more contiguously located statoblasts (floatoblasts or sessoblasts) of the same species germinate simultaneously, their mucous pads often come into contact with each other. The walls of the mucous pads then disappear in the contact areas, thus uniting the coeloms of the ancestrulae. This type of fusion between mucous pads of statoblast-derived ancestrulae was ascertained in Plumatella emarginata, P. repens, P. casmiana, and Hyalinella punctata. The fusion is clearly species specific, and shows no clone specificity or allogeneic recognition. The fusibility test reported here seems to be a useful method for the examination of conspecificity in plumatellid bryozoans.     

Particle capture in ciliary filter‐feeding gymnolaemate and phylactolaemate bryozoans – a comparative study

Riisgård, H.U., Okamura, B. and Funch, P. 2009. Particle capture in ciliary filter‐feeding gymnolaemate and phylactolaemate bryozoans – a comparative study. —Acta Zoologica (Stockholm) 91 : 416–425. We studied particle capture using video‐microscopy in two gymnolaemates, the marine cheilostome Electra pilosa and the freshwater ctenostome Paludicella articulata, and three phylactolaemates, Fredericella sultana with a circular funnel‐shaped lophophore, and Cristatella mucedo and Lophophus crystallinus, both with a horseshoe‐shaped lophophore. The video‐microscope observations along with studies of lophophore morphology and ultrastructure indicated that phylactolaemate and gymnolaemate bryozoans with a diversity of lophophore shapes rely on the same basic structures and mechanisms for particle capture. Our study also demonstrates that essential features of the particle capture process resemble one another in bryozoans, brachiopods and phoronids.     

Shallow lake sediments provide evidence for metapopulation dynamics: a pilot study

The aim of this pilot study is to test the hypotheses that sediment cores can provide evidence for metapopulation dynamics and that these can be linked with site characteristics. We focus on temporal patterns of incidence and abundance of overwintering stages (statoblasts) produced by the freshwater bryozoan Cristatella mucedo, an organism characterised by a metapopulation ecology, in sediment cores retrieved from 18 UK lakes. Runs and goodness-of-fit tests provided evidence for population instability, periods of low abundance and absence, and of asynchrony—all signatures of metapopulation dynamics. Further hypothesis testing indicated that extinction risk is greater in more isolated sites and in sites of smaller size. Absence of statoblasts from the top sections of spatially separated, replicate cores provided independent evidence for extinction in one site. Our study demonstrates how the abundances of sedimentary-bound propagules may be analysed initially for metapopulation dynamics and subsequently how this may lead to working hypotheses regarding the drivers of such dynamics. The sediment archive represents a unique historical record whose potential for characterising metapopulation dynamics has previously been untapped but is broadly relevant for understanding the population biology of freshwater organisms.     

Dormant stages in freshwater bryozoans--an adaptation to transcend environmental constraints

Freshwater invertebrates often disperse between discrete habitat patches via the production of dormant propagules. Being dispersed passively by animal vectors or wind, certain adaptations for exposures to terrestrial and aerial conditions like desiccation and freezing are required. In the present study, we investigate the mechanisms of survival and physiological adaptations due to desiccation and low temperatures in the statoblasts of two populations of the freshwater bryozoan Cristatella mucedo. Our results show that both sessoblasts and floatoblasts tolerate almost complete desiccation and subzero temperatures. Trehalose, a non-reducing disaccharide which has been related to desiccation tolerance, was detected by amperometric chromatography. However, due to the low concentrations found, it is unlikely that trehalose is playing a major part in desiccation tolerance of bryozoan statoblasts. Vitrification is assumed to be important in the survival of desiccation tolerant organisms. Differential scanning calorimetry revealed thermal transitions (T_g onset around 70 °C) in desiccated statoblasts, indicating that a vitreous matrix is present. During the exposure to subzero temperatures, freeze tolerance of statoblasts was confirmed by the detection of internal ice formation, which took place at a crystallisation temperature of between −6 °C and −12 °C.     

Variation in reproductive characteristics of colonies of the freshwater bryozoan Cristatella mucedo

1. We studied relationships between different forms of reproduction and local variation in the reproductive state of colonies of a common freshwater bryozoan Cristatella mucedo Cuvier (Bryozoa: Phylactolaemata). Four sampling locations in central Finland, including both lotic and lentic habitats, were sampled by scuba diving. The traits studied were occurrence of sexual larvae, colony size, number of resting stages (statoblasts) and number of zooids. 2. While 76.9% of the sampled colonies carried statoblasts at the time of sampling, only 4.5% of the colonies were brooding sexual larvae. Most of the colonies were relatively small with a mean colony size of 16.6 mm. 3. In colonies brooding larvae, the number of statoblasts was positively correlated with the number of larvae. We did not detect a colony size threshold for sexual reproduction or formation of statoblasts. Colonies carrying statoblasts had a lower number of zooids per unit dry weight. 4. We found significant variation in reproductive traits of colonies among the sampling locations, and among lotic and lentic habitats. The observed phenotypic differences may reflect broadly distributed, phenotypically plastic clones. Differences in clonal composition of local populations cannot, however, be ruled out and factors that may provide clonal diversity are discussed.     

Plumatella mukaii,a new phylactolaemate bryozoan from Asia and South America

A new freshwater bryozoan species, Plumatella mukaii, is recognized from eastern Asia and western South America. Colonies and statoblasts both bear a resemblance to P. emarginata, and have often been confused with that species, especially in Japanese studies. Floatoblasts and sessoblasts are enclosed within a tough, wrinkled, membrane which resists removal by mechanical means. Floatoblasts are generally smaller than those of P. emarginata, but display unusually high variability in their overall dimensions. The species has been reported from both lentic and lotic habitats. In Asia, the range includes Japan, Korea, China, India and Indonesia. It has most recently also been found at several sites in Chile. The recognition of P. mukaii narrows the previously reported range of P. emarginata and invites a re-inspection of that species worldwide.     

Grazing impact and phenology of the freshwater sponge Ephydatia muelleri and the bryozoans Plumatella emarginata and Fredericella sultana under experimental warming

1. Both the clearance rates (CR) and abundances of the freshwater sponge Ephydatia muelleri and the bryozoans Plumatella emarginata and Fredericella sultana were investigated from autumn to spring under different temperature regimes. The experiments were performed in bypass channels of the River Rhine (Cologne, Germany) in which temperature could be manipulated. 2. The impact of temperature increase on CRs depends upon the grazer: E. muelleri showed a clear increase in CRs with increasing temperature whereas P. emarginata was not significantly affected by experimental warming. 3. Distinct differences in food preference were found for the sponge (which is an efficient grazer of bacteria and small algae) and for the bryozoan P. emarginata (which feeds primarily on large algae, and with no significant grazing on bacteria). 4. In contrast to their temperature‐related patterns in CR, respiration of both P. emarginata and E. muelleri increased with temperature between 19 and 32 °C, suggesting that the risk of experiencing energy deficiency at high temperatures due to increased metabolic rates is particularly high for the bryozoan. 5. A temperature elevation of 3 °C above the natural Rhine temperature resulted in a delay in the disappearance of active tissue and formation of resting stages for E. muelleri in autumn. This delay ranged from 8 (beginning of gemmulation) to 22 days (termination of gemmulation). In contrast, there was no distinct effect of warming on the disappearance of active zooids of the two bryozoan species in autumn. However, warming can positively affect the maintenance of active zooids during winter in F. sultana. In spring, the appearance of active zooids of P. emarginata was clearly stimulated by temperature elevations, whereas the hatching of both F. sultana and E. muelleri was hardly affected by warming. 6. The study demonstrated different patterns in the thermal ecology of the two freshwater bryozoans and the sponge in comparison to other filter feeders, particularly mussels. Such patterns need to be considered when predicting the impact of temperature on pelagic‐benthic coupling in aquatic habitats.     

Abiotic and biotic factors affecting zooperiphyton development in a waterflow of a cooling water pool

The study is concerned with invertebrates populating riffle stones in a cooling water pool. It was shown that the invertebrate community is dominated by Oligochaeta of the genus Naididae in abundance, and Plumatella emarginata bryozoans, in biomass. The study revealed the following: (1) in communities with biomass >100 g/m² abundance and biomass are affected by roughness and lightness of the substrate; (2) bryozoan biomass and Oligochaeta abundance correlate.