Infection of bryozoans by Tetracapsuloides bryosalmonae at sites endemic for salmonid proliferative kidney disease

Laboratory-reared colonies of the bryozoans Fredericella sultana and Plumatella fungosa were placed upstream of 2 fish farms endemic for salmonid proliferative kidney disease (PKD) to assess rates of infection of bryozoans by Tetracapsuloides bryosalmonae, the causative agent of PKD. Colonies were deployed in the field for 8 trial periods of 2 wk each throughout the summer of 2001. Following each trial, bryozoan colonies were maintained in laboratory culture for 28 d and were regularly monitored for infection by searching for sac stages of T. bryosalmonae. Infections were never identified by observations of sac stages, however positive PCR results and sequencing of cultured material confirmed that cryptic infections were present in colonies of both species deployed at one site. The possibility that PCR results reflected contamination of surfaces of bryozoans can be excluded, given the short period of spore viability of T. bryosalmonae. Highest rates of infection occurred when 4 of 23 colonies of F. sultana and 1 of 12 colonies of P. fungosa were infected during the period 10 to 24 July. No infections were detected from mid-August to late October at this site. None of the colonies at the other site became infected throughout the period of study. Our data provide the first estimates of infection rates of bryozoans by T. bryosalmonae. Additionally, they provide evidence that a cryptic stage can be maintained within bryozoan hosts for a period of 4 to 6 wk.     

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.     

Diversity and systematics of the Malacosporea (Myxozoa)

Abstract. Myxozoans belonging to the recently described class Malacosporea parasitize freshwater bryozoans during at least part of their life cycle. There are at present only two species described in this class: Buddenbrockia plumatellae and Tetracapsuloides bryosalmonae . The former can exist as vermiform and sac-like stages in bryozoan hosts. The latter, in addition to forming sac-like stages in bryozoans, is the causative agent of salmonid proliferative kidney disease (PKD). We undertook molecular and ultrastructural investigations of new malacosporean material to further resolve malacosporean diversity and systematics. Phylogenetic analyses of 18S rDNA sequences provided evidence for two new putative species belonging to the genus Buddenbrockia , revealing a two-fold increase in the diversity of malacosporeans known to date. One new malacosporean is a vermiform parasite infecting the bryozoan Fredericella sultana and the other occurs as sac-like stages in the rare bryozoan, Lophopus crystallinus . Both bryozoans represent new hosts for the genus Buddenbrockia . Our results have established that the malacosporean which infected F. sultana was not a vermiform stage of T. bryosalmonae , although it was collected from a site endemic for PKD. Ultrastructural investigation of new material of B . plumatellae revealed the presence of numerous external tubes associated with developing polar capsules, confirming that the absence of external tubes should no longer be considered as a character of the class Malacosporea.     

Transmission of Tetracapsuloides bryosalmonae (Myxozoa: Malacosporea) to Fredericella sultana (Bryozoa: Phylactolaemata) by various fish species

Tetracapsuloides bryosalmonae is a myxozoan parasite of salmonids and freshwater bryozoans, which causes proliferative kidney disease (PKD) in the fish host. To test which fish species are able to transmit T. bryosalmonae to bryozoans, an infection experiment was conducted with 5 PKD-sensitive fish species from different genera. Rainbow trout Oncorhynchus mykiss, brown trout Salmo trutta, brook trout Salvelinus fontinalis, grayling Thymallus thymallus and northern pike Esox lucius were cohabitated with T. bryosalmonae-infected Fredericella sultana colonies and then subsequently cohabitated with statoblast-reared parasite free Bryozoa. Statoblasts from infected colonies were tested by PCR to detect cryptic stages of T. bryosalmonae, which may indicate vertical transmission of the parasite. In this study, brown trout and brook trout were able to infect Bryozoa, while there was no evidence that rainbow trout and grayling were able to do so. Few interstitial kidney stages of the parasite were detected by immunohistochemistry in brown trout and brook trout, while rainbow trout and grayling showed marked proliferation of renal interstitial tissue and macrophages with numerous parasite cells. Intraluminal stages in the kidney tubules were only detected in brown trout and rainbow trout. In contrast to previous observations, pike was not susceptible to PKD in these trials according to the results of T. bryosalmonae-specific PCR. No DNA of T. bryosalmonae was detected in any statoblast.     

Temperature-driven proliferation of Tetracapsuloides bryosalmonae in bryozoan hosts portends salmonid declines

Proliferative kidney disease (PKD) is an emerging disease of salmonid fishes. It is provoked by temperature and caused by infective spores of the myxozoan parasite Tetracapsuloides bryosalmonae, which develops in freshwater bryozoans. We investigated the link between PKD and temperature by determining whether temperature influences the proliferation of T. bryosalmonae in the bryozoan host Fredericella sultana. Herein we show that increased temperatures drive the proliferation of T. bryosalmonae in bryozoans by provoking, accelerating and prolonging the production of infective spores from cryptic stages. Based on these results we predict that PKD outbreaks will increase further in magnitude and severity in wild and farmed salmonids as a result of climate-driven enhanced proliferation in invertebrate hosts, and urge for early implementation of management strategies to reduce future salmonid declines.     

Evidence that infectious stages of Tetracapsula bryosalmonae for rainbow trout Oncorhynchus mykiss are present throughout the year

Proliferative kidney disease (PKD) is a hyperplastic condition of the lymphoid tissue of salmonids infected with the spores of Tetracapsula bryosalmonae, a myxozoan parasite formerly designated PKX, which has recently been described as a parasite of several species of bryozoans. The occurrence of PKD is generally associated with seasonal increase in water temperature, with research indicating that transmission of the disease does not occur below 12 to 13 degrees C. This suggested that the infectious stages are absent from about November to March/April. Here we document the transmission of PKD at water temperatures and seasons previously considered to be non permissive for PKD infection. The exposure of naive rainbow trout Oncorhynchus mykiss (Walbaum) to PKD-infected water ranging from 8 to 13 degrees C during the Autumn, Winter and early Spring, resulted in the infection of kidney interstitium once the trout were transferred to 16 degrees C. In addition, cohabitation studies were conducted with the bryozoan host Fredericella sultana collected from a river at times of low seasonal temperatures because this bryozoan species overwinters as living colonies. Cohabitation of trout with colonies of F sultana in parasite-free city water at 16 degrees C, also led to renal lymphoid tissue infection with the parasite and even to nephromegaly. Our results provide evidence that the infectious stages of T bryosalmonae for rainbow trout were present in the water throughout the entire year and that the impact of temperature on the development of PKD is primarily a result of the kinetics of Tetracapsula multiplication in bryozoan and fish hosts.     

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.     

Transmission of freshwater myxozoans during the asexual propagation of invertebrate hosts

The phylum Myxozoa contains over 1350 species almost all of which are considered to be obligate parasites of aquatic animals. The phylum is composed of two classes, the Myxosporea and the Malacosporea, species of which are important pathogens responsible for severe economic losses in cultured fisheries. The life cycles of freshwater Myxozoa are believed to involve horizontal, indirect transmission, involving an invertebrate (oligochaetes or bryozoans) and a vertebrate host (fish or amphibians). Here, we describe myxozoan propagation through the fragmentation of invertebrate hosts to form new infected individuals. The two hosts examined are an oligochaete Lumbriculus variegatus infected with an unidentified myxosporean (Triactinomyxon sp.) and the bryozoan Fredericella sultana infected with the malacosporean Tetracapsuloides bryosalmonae which causes proliferative kidney disease, a major constraint of the European rainbow trout industry. Such intra-clonal propagation is a novel form of vertical transmission that is likely to be widespread within the Myxozoa and could form an important method by which some of these parasites maintain and proliferate within the aquatic environment.     

A new class and order of myxozoans to accommodate parasites of bryozoans with ultrastructural observations on Tetracapsula bryosalmonae (PKX organism)

Tetracapsula bryosalmonae, formerly PKX organism, is a myxozoan parasite that causes proliferative kidney disease in salmonid fish. Its primary hosts, in which it undergoes a sexual phase, are phylactolaemate bryozoans. It develops in the bryozoan coelomic cavity as freely floating sacs which contain two types of cells, stellate cells and sporoplasmogenic cells, which become organised as spores. Eight stellate cells differentiate as four capsulogenic cells and four valve cells which surround a single sporoplasmogenic cell. The sporoplasmogenic cell undergoes meiosis and cytoplasmic fission to produce two sporoplasms with haploid nuclei. Sporoplasms contain secondary cells. The unusual development supports previously obtained data from 18S rDNA sequences, indicating that species of Tetracapsula form a clade. It diverged early in the evolution of the Myxozoa, before the radiation that gave rise to the better known genera belonging to the two orders in the single class Myxosporea. The genus Tetracapsula as seen in bryozoans shares some of the characters unique to the myxosporean phase and others typical of the actinosporean phase of genera belonging to the class Myxosporea. However, it exhibits other features which are not found in either phase. A new class Malacosporea and order Malacovalvulida are proposed to accommodate the family Saccosporidae and genus Tetracapsula. Special features of the new class are the sac-like proliferative body, valve cells not covering the exit point of the polar filament, lack of a stopper-like structure sealing the exit, maintenance of valve cell integrity even at spore maturity, absence of hardened spore walls and unique structure of sporoplasmosomes in the sporoplasms.     

Sequential development of Buddenbrockia plumatellae (Myxozoa: Malacosporea) within Plumatella repens (Bryozoa: Phylactolaemata)

Colonies of the freshwater bryozoan Plumatella repens collected from a river in the UK were found to be infected with the myxozoan parasite Buddenbrockia plumatellae following laboratory maintenance. Optimisation of the bryozoan diet allowed maintenance of infected colonies for 90 d, permitting observation by light and electron microscopy of the sequential parasitic developmental cycle. Parasite stages were associated with host peritoneum, identifying the primary developmental phase. The association of B. plumatellae cells with peritoneal basal lamina and morphological similarities between parasite and host suggested that the parasite remodelled host tissue. Progressive expansion and elongation of individual parasites led to the release of freely floating vermiform stages within the host coelomic cavities. Within these 'worms', intraluminal masses developed, resulting in the formation of spores. Upon maturation, the 'worms' ruptured, releasing many spores within the host that were subsequently discharged. Although parasitism led to increased bryozoan fragmentation and lowered statoblast production, some colonies did survive, resulting in repeated waves of infection. Long-term laboratory maintenance of infected bryozoan colonies could provide a means of maintaining B. plumatellae for study until the full life cycle is ascertained.     

Distribution of Kroeyerina elongata (Kroyeriidae: Siphonostomatoida, Copepoda) in the olfactory sacs of the blue shark, Prionace glauca

The infection pattern of Kroeyerina elongata (Kroyeriidae, Copepoda) in the olfactory sacs of the blue shark, Prionace glauca, was investigated using 4,722 copepods from 54 olfactory sacs. Copepod prevalence and mean intensity of infection per olfactory sac were 94.0 and 91.1%, respectively, and the most intensely infected olfactory sac and shark hosted 218 and 409 copepods, respectively. There were significant linear relationships between the number of female and total copepods per left olfactory sac and shark fork length as well as between the numbers of female, male, and total copepods per shark and mean olfactory sac width and cumulative olfactory sac width. Female copepods typically outnumbered males within olfactory sacs (mean intensity = 65.7 and 26.3, respectively), and no statistical differences were detected between the numbers of copepods inhabiting the left and right olfactory sacs. Copepods were not evenly distributed within olfactory sacs. Typically, female copepods occupied olfactory chambers located centrally along the length of the olfactory sac, while males infected lateral olfactory chambers nearest the naris. The orientation of most copepods (84.6%) suggested positive rheotaxis relative to the path of water through the olfactory sac. Within olfactory chambers, most mature females (68.2%) infected the first third of the peripheral excurrent channel and the adjacent fringe of olfactory lamellae, while most males (91.7%) infected the olfactory lamellae, and the 4 larval females collected were attached within the lamellar field and grasped by males. Based on the observed infection patterns and the pattern of water flow throughout the olfactory sac, a hypothesis regarding the life cycle of K. elongata is advanced wherein infective copepodids are swept into the olfactory sac from the surrounding sea and initially colonize the olfactory lamellae. Copepodids feed and mature among the olfactory lamellae, and adult males search for mates and copulate with young females among the olfactory lamellae. Inseminated females move to the peripheral excurrent channels to mature and produce ovisacs. Hatching ovisacs release free-swimming nauplii into the excurrent water flow to be swept into the milieu, where they can molt into infective copepodids that may infect new hosts.     

Severe, chronic proliferative kidney disease (PKD) induced in rainbow trout Oncorhynchus mykiss held at a constant 18 degrees C

Proliferative kidney disease (PKD), caused by the myxozoan parasite Tetracapsuloides bryosalmonae, is well documented as a seasonal disease of rainbow trout Oncorhynchus mykiss. Water temperatures influence the course of the infection both within the fish and the invertebrate host, the recovery of fish from the disease being accelerated with decreasing water temperatures. During this study, groups of rainbow trout were held at a constant temperature (18 degrees C) for a sustained period of time following initial exposure to T. bryosalmonae. While the majority of these fish had recovered from the clinical disease after 9 mo, 10% remained infected, showing clinical signs of disease. A histological study revealed that the majority exhibited very high parasite loads and unusually severe symptoms of PKD. This demonstrates that while most rainbow trout can recover from PKD independent of water temperature, there exists a sub-population that cannot.     

A palaeoecological study of the Upper Coralline Limestone, Terebratula-Aphelesia bed (miocene,malta) based on bryozoan growth-form studies and brachiopod distributions

A prominent bed, containing brachiopods and ectoproct bryozoans, is widely distributed within the Maltese Islands, being associated with a coralline algal rhodolite bioherm. This bed provides a useful stratigraphic marker horizon at the base of the Upper Coralline Limestone Formation.Samples taken from the bed at localities in Malta and Gozo have been examined and their contained biodata studied. Distribution analysis of the bryozoan growth-forms present has permitted recognition of four assemblages, dominated by cellariform, vinculariiform, encrusters, and cellariform and vinculariiform bryozoan growth-forms, respectively. Morphological variations and distributions of brachiopods contained within the samples were also examined and these were compared with the bryozoan growth-form distributions previously obtained. An interpretation of the palaeoecology of the four common brachiopod species, Terebratula terebratula, Aphelesia bipartita, Argyrotheca cf. cordata and Megathiris decollata, is offered partly on the basis of present-day brachiopod ecology but mainly on the basis of information gained for the bryozoans. Interpretations of the palaeoecology of several other common invertebrate genera, also based on the bryozoan growth-form distributions, are also presented.     

The settlement and metamorphosis of the marine bryozoan Bowerbankia gracilis (Ctenostomata: Vesicularioidea)

Summary The settlement and metamorphosis of the marine bryozoan Bowerbankia gracilis has been examined by light and electron microscopy. The period of rapid morphogenesis consists of the following sequence of morphogenetic movements: 1) eversion of the internal sac, 2) retraction of the apical disc, 3) coronal involution and exposure of the pallial epithelium, and 4) closure of the internal coronal cavity. The eversion of the internal sac at the onset of metamorphosis coincides with a sudden reversal of the direction of beat of the coronal cilia. The reversed beating of the coronal cilia wafts the adhesive secreted by the internal sac over the metamorphosing larva, forming the pellicle. The internal sac is subsequently internalized and histolyzed with the corona and the other transitory larval tissues, and the extensive pallial epithelium forms the epidermis of the ancestrular body wall (cystid). Type I mesenchyme cells form an incomplete somatic mesothelium beneath the differentiating cystid epidermis, and Type II mesenchyme cells become mobile phagocytes. The main body cavity develops by the histolytic enlargement of the internal cavity formed during coronal involution. The apical disc degenerates and the polypide develops from rudiments in the oral hemisphere of the larva. The distinctive larval morphology and metamorphosis of vesicularioid ctenostomes are compared with other bryozoans, and possible evolutionary trends are considered.     

Sporogony of Tetracapsuloides bryosalmonae in the brown trout Salmo trutta and the role of the tertiary cell during the vertebrate phase of myxozoan life cycles

Tetracapsuloides bryosalmonae is the myxozoan that causes the commercially and ecologically important proliferative kidney disease of salmonid fish species. Immunohistochemistry and electron microscopy were used to examine the development of this parasite within the kidney of the brown trout Salmo trutta. The main replicative phase of T. bryosalmonae is a cell doublet composed of a primary cell and a single secondary cell. Engulfment of one secondary cell by another to form a secondary-tertiary doublet (S-T doublet) heralded the onset of sporogony whereupon the parasite migrated to the kidney tubule lumen. Within the tubule, the parasite transformed into a pseudoplasmodium and anchored to the tubule epithelial cells via pseudopodial extensions. Within each pseudoplasmodium developed a single spore, composed of 4 valve cells, 2 polar capsules and 1 sporoplasm. The pseudoplasmodia formed clusters suggesting that large numbers of spores develop within the fish. This examination of T. bryosalmonae suggests that the main replicative phase of freshwater myxozoans within vertebrates is via direct replication of cell doublets rather than through the rupturing of extrasporogonic stages, while tertiary cell formation relates only to sporogony. Taken in conjunction with existing phylogenetic data, 5 distinct sporogonial sequences are identified for the Myxozoa.     

Reef-bryozoans and bryozoan-microreefs: Control factor evidence from the philippines and other regions

Summary In this paper, a preliminary concept on the interplay of local, regional and global control factors of bryozoan diversity and distribution pattern is introduced. Recent bryozoans from the Philippines, New Zealand and the Gulf of Aqaba are compared to the selected fossil specimens from the Oxfordian and Santonian. Reef bryozoan skeletons are studied in order to separate local control within the substrate-water interface from regional control. The latter originate mainly from the transport function of the water column (e.g. sediment load, wave energy, vagile predators). This is true especially for erect (tree-like) and massive (multilaminar) bryozoans which are subjected to the dynamics of the water body in the littoral area. This regional control, affects simultaneously several structural and substrate zones of a reef. Early life history of vertically growing reef bryozoans reflect local control, while older zoarial structures reflect the signals of regional influence. Three types of multiserial nodular bryozoans are cited: self-overgrowing sheets (‘S-Nodule’, derived from ‘S-Sheet’), circumlaterally budding colonies (‘C-Nodules’), and fungiform bryozoans. Bryozoan growth form selection allows the separation of two types of regional controls, long range control which favours the selection of specialized sheets, and slow rate control documented in the growth form modification of sheets changing into nodules. In the domain of local control epibiontic microorganisms, microbial mats and biofilms on hard substrata represent probably the most important elements, aside from the limited substrate space. Symbiotic and/or competitive bryozoan-microorganism interrelationships result in the distinct adaptations of bryozoan growth. It is apparent that bryozoan modifications of substrate micro-topography influence the character of epibiontic microbial settlement. The peak occurrence of microbial settlement on prominent external bryozoan skeleton parts is discussed as an effect of feeding currents. In contrast, smooth and unelevated exoskeletons are less favourable for micro-epibiontic colonization. Due to the high level of order in the spatial zonation patterns of epibionts on calcifying bryozoans, the term ‘bryozoan-microreef’ is introduced. Bryozoan-microreefs are locally controlled reefs. They are e.g. characterized by higher competitive abilities than bryozoans without ‘reefdwellers’. Local control has a fast rate of change and is reflected in morphologies of individual zooids and/or single zooid generations. Regional control has a slower rate but a higher range. It is important for growth modifications of the whole zoaria. The very slow rate of global control cannot be recorded within the life span of bryozoan zoaria unless it is expressed through regional control (such as monsoons). Nevertheless, global control is paleoecologically important because it is traceable in bryozoan (paleo-)biogeography. For practical purpose, we suggest to define those control factors of bryozoans as global which affect simultaneously at least one tropical and one non-tropical bryozoan community. This reportis dedicated to the memory of our friend and research colleague Mr. Joselito G. Santiago of the University of San Carlos, Marine Biology Section. Tiago (as we fondly call him) has started within the scope of the Philippine Bryozoan Project an applied research on fish abundance and productivity around a new artificial reef model. This prototype model was conceptualized with him. His promising study remained unfinished; he died May 13, 1993, while sampling inside the reef cave off Marigondon.     

Sacculogenesis of Buddenbrockia plumatellae (Myxozoa) within the invertebrate host Plumatella repens (Bryozoa) with comments on the evolutionary relationships of the Myxozoa

Members of the phylum Myxozoa are obligate parasites, primarily of aquatic organisms. Their phylogeny has remained problematic, with studies placing them within either the Bilateria or Cnidaria. The discovery that the enigmatic Buddenbrockia plumatellae is a myxozoan that possesses distinct bilaterian features appeared to have finally resolved the debate. B. plumatellae is described as a triploblastic 'worm-like' organism, within which typical myxozoan malacospores form. Using EM we examined the early development of the B. plumatellae 'worms' within the bryozoan host Plumatella repens. The initial development involved numerous unicellular, amoeboid pre-saccular stages that were present within the basal lamina of the host's body wall. These stages migrate immediately beneath the peritoneum where a significant host tissue reaction occurs. The stages aggregate, initiating the formation of a 'worm'. The base of a developing 'worm' forms a pseudosyncytium which resolves into an ectoderm surrounding a mesendoderm. The pseudosyncytium is directly anchored into neighbouring host cells via masses of striated fibres. The replication of the ectodermal and mesendodermal cells extends the developing 'worm' into the coelom of the host. The mesendoderm resolves to form a mesoderm and an endoderm. Myogenesis appears to be initiated from the anchored end of the 'worm' and develops along the mesoderm. The aggregation and differentiation of amoeboid pre-saccular stages to initiate the 'worm' draws analogies to the sacculogenesis observed for Tetracapsuloides bryosalmonae, B. plumatellae's sister taxon within the class Malacosporea. The development of a multicellular, spore forming organism, from single cells does not correlate to any bilaterian or cnidarian species. Current phylogenies indicate the Myxozoa are basal bilaterians along with the Acoela and Mesozoa. Comparison with these other basal groups may help to resolve the placement of Myxozoa within the tree of life.     

Severe mortality in wild Atlantic salmon Salmo salar due to proliferative kidney disease (PKD) caused by Tetracapsuloides bryosalmonae (myxozoa)

Extensive mortality in Atlantic salmon fry was reported in the River Aelva from 2002 to 2004. Dead fish were collected in late summer 2006, and live fish were sampled by electrofishing in September the same year. At autopsy and in histological sections, the fish kidneys were found to be pale and considerably enlarged. Proliferative lesions with characteristic PKX cells were seen in a majority of the fish. DNA from kidney samples of diseased fish was subjected to PCR and sequencing, and the amplified sequences matched those of Tetracapsuloides bryosalmonae. We concluded that this myxozoan transmitted from bryozoans was the main cause of the observed mortality in salmon fry in 2006. Results from quantitative electrofishing in 2005 and 2006, combined with the observed fry mortality from 2002 to 2004, show that the smolt production in the river is severely reduced and that T. bryosalmonae is the most likely explanation for this decline. The present study is the first to report a considerable negative population effect in wild Atlantic salmon due to proliferative kidney disease (PKD). It also represents the northernmost PKD outbreak in wild fish. The river is regulated for hydroelectric power purposes, causing reduced water flow and elevated summer temperatures, and the present PKD outbreak may serve as an example of increased disease vulnerability of northern fish populations in a warmer climate.     

Organogenesis during budding and lophophoral morphology of Hislopia malayensis Annandale, 1916 (Bryozoa, Ctenostomata)

Background  

Bryozoans represent a large lophotrochozoan phylum with controversially discussed phylogenetic position and in group relationships. Developmental processes during the budding of bryozoans are in need for revision. Just recently a study on a phylactolaemate bryozoan gave a comprehensive basis for further comparisons among bryozoans. The aim of this study is to gain more insight into developmental patterns during polypide formation in the budding process of bryozoans. Particular focus is laid upon the lophophore, also its condition in adults. For this purpose we studied organogenesis during budding and lophophoral morphology of the ctenostome bryozoan Hislopia malayensis.