Ultrastructure of the body cavities in Phylactolaemata (Bryozoa)

Only species belonging to the bryozoan subtaxon Phylactolaemata possess an epistome. To test whether there is a specific coelomic cavity inside the epistome, Fredericella sultana, Plumatella emarginata, and Lophopus crystallinus were studied on the ultrastructural level. In F. sultana and P. emarginata, the epistome contains a coelomic cavity. The cavity is confluent with the trunk coelom and lined by peritoneal and myoepithelial cells. The lophophore coelom extends into the tentacles and is connected to the trunk coelom by two weakly ciliated coelomic ducts on either side of the rectum. The lophophore coelom passes the epistome coelom on its anterior side. This region has traditionally been called the forked canal and hypothesized to represent the site of excretion. L. crystallinus lacks an epistome. It has a simple ciliated field where an epistome is situated in the other species. Underneath this field, the forked canal is situated. Compared with the other species, it is pronounced and exhibits a dense ciliation. Despite the occurrence of podocytes, which are prerequisites for a selected fluid transfer, there is no indication for an excretory function of the forked canal, especially as no excretory porus was found. J. Morphol. 2009. © 2008 Wiley‐Liss, Inc.     

Bacterial feeding by the freshwater bryozoan Plumatella fungosa (Pallas, 1768).

The present study demonstrates for the first time the capacity of a bryozoan to feed on bacteria. Qualitative feeding tests were performed in laboratory conditions with Plumatella fungosa raised from statoblasts and fed with formazan-coloured Escherichia coli. They establish that bacteria were not only ingested but digested as shown by their transit in the digestive tract and the presence of broken bacteria in the faecal pellets. The ingestion rate of E. coli was determined by measuring bacterial removal in a vessel containing a colony of P. fungosa collected from a pond. It attained 10³ cells per zooid during the first hour of the experiment, for an initial bacterial concentration of 5.5 × 10⁴ cells ml⁻¹. The filtration rate was estimated at 0.0246 ml zooid⁻¹ h⁻¹ . These results indicate the ecological role of bryozoans in water purification.     

Asajirella gelatinosa in Panama: a bryozoan range extension in the Western Hemisphere (Ectoprocta: Phylactolaemata)

Asajirella gelatinosa, a freshwater bryozoan known previously only from eastern Asia is now reported in the Republic of Panama. Colonies were collected during April 1992 in the broad reaches of the lower Río Chagres as it enters Gatun Lake, part of the Panama Canal system. A subsequent collection in 1998 included large colonies and numerous statoblasts from a region upstream of this site in the impounded Lago Alajuela (Madden Lake). Colonies and statoblasts resemble those from Japan in almost every detail. Human transport of statoblasts is a likely cause of this disjunct distribution. This revised version was published online in July 2006 with corrections to the Cover Date.     

Global diversity of bryozoans (Bryozoa or Ectoprocta) in freshwater

The present study considers 88 bryozoan species occurring in freshwater: 69 phylactolaemate and 19 gymnolaemate species. Roughly 49% of these species are confined to one zoogeographical region. The cosmopolitan status of species like Fredericella sultana, Plumatella repens or P. emarginata has to be reconsidered. Among the Phylactolaemata, which are phylogenetically older than the Gymnolaemata, the gelatinous species (Lophopodidae, Pectinatellidae, Cristatellidae) are more primitive than the branching tubular species (Plumatellidae, Fredericellidae). Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Plumatella nitens, a new species of freshwater bryozoan from North America (Ectoprocta: Phylactolaemata), previously misidentified

Plumatella nitens is a newly recognized species of phylactolaemate bryozoan in North America. It has previously been combined with either P. repens or P. fungosa, from which it differs primarily by the floatoblast. In P. nitens the ventral floatoblast annulus is uniformly narrow with no significant widening at the poles. The dorsal floatoblast fenestra is conspicuously larger than in either P. repens or P. fungosa. Reticulated ridges on the dorsal fenestra are weak along the margins, becoming unusually prominent in the center. The sessoblast is densely covered with uneven papillae. Spotty sampling so far shows a distribution only in the northern half of North America, including Massachusetts, Michigan, Minnesota, Wisconsin, Ontario, and the northern regions of Illinois, Indiana, and Ohio.     

Biochemical genetics and taxonomy in Plumatella fungosa and P. repens (Bryozoa: Phylactolaemata)

SUMMARY. Some authors have recently concluded that, using morphological criteria, the freshwater bryozoans Plumatella repens and P. fugosa cannot be separated and should be regarded as conspecific. To test this conclusion, electrophoretic techniques have been used to examine genetic differences between the two nominate species at several enzyme loci. Significant variation at a malate dehydrogenase locus and three aminopeptidase loci establish beyond doubt that P. repens and P. fugosa are separate but related species. No significant variation was found between two populations of P. repens . Morphological characters for the distinction of the two species are discussed and evaluated.     

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.     

Freshwater bryozoans (Bryozoa) of Norway III: distribution and ecology of Plumatella fruticosa

Bryozoans were investigated during field studies of 601 lakes and other surface water bodies throughout Norway from 1960 to 1978. The frequency of occurrence of Plumatella fruticosawas evaluated in relation to 12 environmental variables. Statistically significant deviations from the frequencies expected on the basis of random distribution were described using the categories preference, avoidance, and absence. According to our material P. fruticosais one of the most common bryozoan species in Norwegian fresh water, represented in samples from 251 localities. This species occurred frequently all over the country, north to 71° 06 N (the northernmost record globally). Maximum elevation above sea level was 1179 m (maximum for Northern Europe).In most studies of freshwater bryozoans from the holarctic region, Plumatella fruticosahas been reported as widely distributed, but locally rare. In Norway, however, P. fruticosa is frequently found all over the country, even far to the north. The species apparently finds an optimal climate in the cold temperate and cold regions of Norway. Plumatella fruticosapreferred lakes, avoided slow-flowing rivers and ponds, and was absent in smaller water bodies. The species preferred somewhat higher elevation (400–900 m above sea level), but avoided lakes with the lowest summer temperatures (below 11 °C). P. fruticosa also preferred oligotrophic conditions with poor aquatic vegetation, stony shores, lakes poor in calcium and magnesium (sometimes almost comparable with distilled water), where the water was clear and colourless, and slightly acidic to neutral. The species also preferred lakes surrounded by Sphagnum bogs and was absent when pH was below 5.2. P. fruticosa avoided eutrophic conditions with rich aquatic vegetation, alkaline water (pH above 7.0), lakes with a higher content of calcium and magnesium and those with strongly coloured water (above 100 mg Pt l^–1). In spite of its preference for ion-poor conditions, it was also found in a brackish water lake. For most environmental variables, the species had a wider tolerance range than reported from elsewhere.     

Ganglion ultrastructure in phylactolaemate Bryozoa: evidence for a neuroepithelium

In contrast to other Bryozoa, members of the subtaxon Phylactolaemata bear a subepithelial cerebral ganglion that resembles a hollow vesicle rather than being compact. In older studies this ganglion was said to originate by an invagination of the pharyngeal epithelium. Unfortunately, documentation for this is fragmentary. In chordates the central nervous system also arises by an invagination-like process, but this mode is uncommon among invertebrate phyla. As a first attempt to gather more data about this phenomenon, cerebral ganglia in two phylactolaemate species, Fredericella sultana and Plumatella emarginata, were examined at the ultrastructural level. In both species the ganglion bears a small central lumen. The ganglionic cells are organized in the form of a neuroepithelium. They are polarized and interconnected by adherens junctions on their apical sides and reside on a basal lamina. The nerve cell somata are directed towards the central lumen, whereas the majority of nervous processes are distributed basally. Orientation of the neuroepithelial cells can be best explained by the possibility that they develop by invagination. A comparison with potential outgroups reveals that a neuroepithelial ganglion is at least derived. Since, however, a reliable phylogenetic system of the Bryozoa is missing, a decision on whether such a ganglion is apomorphic for Bryozoa or evolved within this taxon can hardly be made.     

Development and molecular characterisation of the microsporidian Schroedera airthreyi n. sp. in a freshwater bryozoan Plumatella sp. (Bryozoa: Phylactolaemata)

The development of a new species of microsporidian, infecting a freshwater Plumatellid bryozoan, is described. The small-subunit rDNA, internal transcribed spacer region (ITS), and partial large-subunit rDNA genes were sequenced. Phylogenetic analysis demonstrated that the parasite clustered with Schroedera plumatellae. However, while there were morphological affinities with this species, significant differences were also observed. The infection initially appeared as a roughening of the peritoneum lining the metacoelom of the bryozoan. This roughening resolved into meront-infected syncytia, composed of interconnected cells of the body wall that detached to float in the coleomic cavity. Spores were observed to develop within these syncytia. All stages of development were diplokaryotic in contrast to S. plumatellae, which has a distinct monokaryotic merogony preceding sporogony. The infection was pathogenic to the host. Direct bryozoan-bryozoan transmission was not observed. We propose to name the microsporidian Schroedera aithreyi n. sp.     

Freshwater bryozoans (Bryozoa) of Norway IV: Distribution and ecology of four species of Plumatella with notes on Hyalinella punctata

Bryozoans were investigated during field studies of 601 lakes and other surface water bodies throughout Norway from 1960 to 1978. The frequency of occurrence of Plumatella repens was evaluated in relation to 12 environmental variables. Statistically significant deviations from the frequencies expected on the basis of random distribution were described using the categories preference, avoidance and absence. P. repens is one of the most common bryozoan species in Norway. It is represented at 227 localities in our material and by 29 additional records. The species occurred frequently all over the country, north to 71°09N (the northernmost record on the European mainland). Maximum elevation above sea level was 1052 m (maximum for Northern Europe). P. repens preferred lakes, avoided rivers, and was absent from the smallest water bodies. The species preferred (1) low elevation (2–200 m above sea level), (2) medium water temperature, (3) rich aquatic vegetation (eutrophic conditions), (4) small wave action, (5) high content of calcium and magnesium, (6) pH 6.4–9.6, and (7) rather high water colour. Otherwise P. repens avoided (1) habitats above 500 m but occurred up to 1052 m, (2) the lowest water temperature intervals, (3) sites with poor aquatic vegetation and stony shores, (4) medium wave action, (5) sites with a low calcium and magnesium content, (6) lakes of low water colour, and (7) lakes with pH below 6.6: it was absent from lakes with pH below 5.2. For many environmental variables, the species had a wider tolerance range than reported from elsewhere. P. fungosa, P. casmiana, and P. emarginata are all rare in Norway. The total numbers of known localities, with our records in (), are 18 (11), 7 (7), and 7 (5), respectively. These sites are all from lowland areas up to 133 m above sea level. Environmental variables in the habitats are described. P. fungosa was represented by a single floatoblast in each of the four northernmost lakes where it was recorded. One of these lakes was in North Norway some 1000 km away from the nearest known sites with living colonies. The presence of large colonies of P. fungosa in some parts of southern Norway and only `single floatoblast lakes' further north is discussed in relation to the equilibrium theory of island biogeography: birds facilitate immigration by carrying floatoblasts and environmental conditions prevent their germination and the development of living colonies. All records of P. casmiana and P. emarginata were from South Norway. Results for P. fruticosa have been published previously. Notes are given on an old and now missing sample from ca. 1900, identified by Lacourt (1968) as Hyalinella punctata – the only record of this species from Norway.     

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.     

Biochemical genetics and taxonomy in Plumatella emarginata and P. repens (Bryozoa: Phylactolaemata)

SUMMARY. Although morphological distinctions are well documented, many authors have regarded the freshwater bryozoan Plumatella emarginata as a variety of P. repens. Differences between the two nominate species have been studied by the electrophoretic examination of genetic variation at enzyme loci. Genetic identity (Nei, 1972) was 0.286, a value well below that expected for conspecific allopatric populations. The results leave little doubt that specimens of the two species are from quite distinct and separate gene pools, with an unexpectedly high level of genetic differentiation. It is therefore concluded that P. emarginata is a valid species and is not merely a variety of P. repens. Morphological differences between the two species are also discussed.     

Ultrastructural studies on the early cuticular metamorphosis of adult female Lernaeocera branchialis (L.) (Copepoda,Pennellidae)

After its final moult and fertilization an adult female of the marine fish-parasitic copepod, Lernaeocera branchialis, begins an extensive metamorphosis. This commences while the parasite is still on the flounder intermediate host and is completed once the female has established itself on the whiting final host. One early component of the metamorphosis is a considerable elongation of the parasite's abdominal region. S.e.m. and t.e.m. studies have revealed that part of the mechanism of the elongation consists of a straightening out of a highly folded abdominal cuticle. Before fertilization, the epicuticle and outer procuticular layers of this integument are thrown into a series of transverse, 4–6 µm deep pleats or folds with a density of 1–1.2 folds/µm of abdominal length. Straightening these folds can generate an approximately 6-fold length increase. The folds are already present beneath the female chalimus IV cuticle when the epidermis of this development stage starts to secrete the adult cuticle. Immediately before the final moult, the adult cuticle is super-folded with the whole cuticle displaying second-order folds, 8–10 µm deep.The capacity of Lernaeocera to engage in extensive cuticular modifications without recourse to a moult is compared with similar abilities shown by some insect species.     

Freshwater bryozoans (Bryozoa) of Norway V: review and comparative discussion of the distribution and ecology of the 10 species recorded

The paper is based on material presented in five contributions [Hydrobiologia 421:1–24, 2000; 459: 103–123, 2001; 479: 11–22, 2002; 501: 179–198, 2003; Archives de lInstitut grand-ducal de Luxembourg. Section des Sciences naturelles, physiques et mathématiques, Nouvelle Série 44: 127–143, 2002]. Nine species of Bryozoa were collected during field studies of 601 lakes and other surface water types in Norway. Five of the species occur throughout Norway (Plumatella fruticosa, P. repens, Paludicella articulata, Cristatella mucedo and Fredericella sultana) and one has a northerly range (Fredericella indica). With the exception of Plumatella repens, the northernmost records of these species globally are in Norway. All six species have been found at elevations of more than 1000m in South Norway, which is also their maximum elevation above sea level in Northern Europe. Three species (Plumatella fungosa, P. emarginata and P. casmiana) are rare and present only in lowland areas in South Norway. Lacourt [A Monograph of the Freshwater Bryozoa–Phylactolaemata. Zoologische Verhandelingen, Leiden 93: 1–159, 1968] considered an old and now mislaid sample from the same area to be Hyalinella punctata. Up to six species and an average of two were found in lakes. The mean numbers of species in ponds, and slow-flowing and rapid-flowing rivers were 0.6, 0.8 and 1.0, respectively. Co-occurrence of pairs of species in lakes was investigated. For 10pairs, the number of cases of co-occurrence was significantly higher than expected on the basis of random distribution. This was ascribed to some shared ecological requirements. Only one pair (Paludicella articulata–Plumatella fungosa) showed fewer cases of co-occurrence than expected. This was explained by differences in their geographical distribution and tolerance to ecological factors. Tolerance and other ecological responses were described for the six frequent species using two statistical methods. Method 1: frequency deviations, is based on the null hypothesis of random distribution. Ten environmental variables were studied (type of surface water, elevation above sea level, water temperature, aquatic vegetation, sediment, wave action, Ca, Mg, pH and water colour).The concepts absence, avoidance (frequency lower than expected), preference (frequency higher than expected) and presence (no significant deviation from expected frequency) were used. The degree of response of a particular species to a specific environmental variable was measured by the sum of the numerical values of the significant frequency deviations divided by the number of categories with significant deviations. The results can be used as adaptability indices or tolerance indices: the lower the index, the better a species is adapted for surviving with respect to the environmental variable in question. Diagrams showed (1) the relative strength of the response of the six frequent species, (2) the average index for each species for all variables and (3) the effect of each of the 10 environmental variables on the Bryozoa (the species treated as a unit: an ecological guild). Method 2: logistic regression, treated the presence or absence of each of the six frequent species as dependent variables using nine independent environmental variables. Comparison of the results of method 1 and 2, indicated that method 1, which gives details of how a species responds to different categories of a variable, was particularly useful.