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.     

Bryozoan populations reflect nutrient enrichment and productivity gradients in rivers

1. The hypothesis that nutrient enrichment will affect bryozoan abundance was tested using two complementary investigations; a field‐based method determining bryozoan abundance in 20 rivers of different nutrient concentrations by deploying statoblast (dormant propagule) traps and an experimental laboratory microcosm study measuring bryozoan growth and mortality. These two methods confirmed independently that increased nutrient concentrations in water promote increases in the biomass of freshwater bryozoans. 2. Statoblasts of the genus Plumatella were recorded in all rivers, regardless of nutrient concentrations, demonstrating that freshwater bryozoans are widespread. Concentrations of Plumatella statoblasts were high in rivers with high nutrient concentrations relative to those with low to moderate nutrient concentrations. Regression analyses indicated that phosphorus concentrations, in particular, significantly influenced statoblast concentrations. 3. Concentrations of Lophopus crystallinus statoblasts were also higher in sites characterised by high nutrient concentrations. Logistic regression analysis revealed that the presence of L. crystallinus statoblasts was significantly associated with decreasing altitude and increasing phosphorus concentrations. This apparently rare species was found in nine rivers (out of 20), seven of which were new sites for L. crystallinus. 4. Growth rates of Fredericella sultana in laboratory microcosms increased with increasing nutrient concentration and high mortality rates were associated with low nutrient concentrations. 5. Our results indicate that bryozoans respond to increasing nutrient concentrations by increased growth, resulting in higher biomasses in enriched waters. We also found that an important component of bryozoan diets can derive from food items lacking chlorophyll a. Finally, bryozoans may be used as independent proxies for inferring trophic conditions, a feature that may be especially valuable in reconstructing historical environments by assessing the abundance of statoblasts in sediment cores.     

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.     

A fish-based index for the assessment of the ecological quality of temperate lakes

A fish – based index for the assessment of the ecological quality of natural temperate lakes was developed, in accordance to the requirements of the Water Framework Directive (WFD) 2000/60/EC. As a case study, 11 natural lakes located at northern and western Greece were selected. Fish surveys were conducted during mid summer to mid autumn in 2010, 2011 and 2012 using Nordic gillnets and electrofishing. Environmental parameters and anthropogenic pressures were assessed for each lake. Fish species richness, abundance, trophic, reproductive and habitat functional guilds were used for extracting a set of 107 metrics, meeting the requirements of the WFD. All metrics were initially tested as candidates for the index development. A stepwise linear regression of each metric against environmental parameters (lake area, altitude, maximum depth, alkalinity) and anthropogenic pressures (drainage area covered by non-natural land uses – NNLC, water total phosphorus concentrations – TP, Lake Habitat Modification Score – LHMS) was initially conducted for ensuring pressure-response relationships. Reference conditions for each lake were estimated by the hindcasting procedure and the ecological quality for each lake was expressed as the ecological quality ratio (EQR) by a value ranging from 0 (poor quality) to 1 (excellent quality). Two fish fauna metrics, the relative numerical abundance of introduced species (Introduced_a) and the relative biomass of omnivorous species (OMNI_b) were finally extracted as the most significant, responding to LHMS and TP, respectively. The final index was expressed as the mean values of the EQRs of these two metrics. The multimetric fish index presented herein could serve as a tool for assessing the ecological quality of natural lakes at broad geographical scale and generally, in the Mediterranean temperate lakes with similar hydromorphological characteristics.     

Representation of aquatic invertebrate communities in subfossil death assemblages sampled along a salinity gradient of western Uganda crater lakes

We analysed subfossil death assemblages of aquatic invertebrate communities in a salinity series of 35 western Uganda maar-crater lakes to evaluate their potential as biological indicators of past habitat conditions in paleo-environmental research. The study region encompasses the climatological and hydrological gradient between the dry floor and moist shoulders of the Edward-George branch of the East African Rift Valley, and includes mesotrophic to hyper-eutrophic, and shallow unstratified to deep meromictic lakes with a surface-water salinity range between 101 and 135 400 μS/cm. Focusing on non-chironomid aquatic invertebrates with good fossil preservation, we found that fossil larval remains of the Dipteran families Culicidae, Ephydridae, and Stratiomyidae are good indicators of saline environments. Our data further suggest that the abundances of Bryozoan statoblasts and Chaoboridae are indicative of, respectively, the fraction of the littoral zone covered by aquatic macrophytes and of lake trophic state, but a lake reference data set more specifically designed to cover variation in these environmental factors will be needed to determine the strength of these relationships. In these small, simple lake basins, recent death assemblages recovered from a single mid-lake surface-sediment sample provides a more complete inventory of local aquatic invertebrate communities and the distribution of species among lakes than exploratory live sampling of those taxa in a selection of littoral, benthic and pelagic habitats.     

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.     

Lake morphometry predicts the degree of habitat coupling by a mobile predator

Habitat coupling is an ecosystem process whereby semi-discontinuous habitats are connected through the movement of energy and nutrients by chemical, physical or biological processes. One oft-cited example is that of littoral–pelagic coupling in lakes. Theory has argued that such habitat coupling may be critical to food web dynamics, yet there have been few empirical studies that have quantified ecological factors that affect the degree of habitat coupling in ecosystems. Specifically, the degree to which habitat coupling occurs across important physical gradients has largely been ignored. To address this, we investigate the degree of littoral habitat coupling (i.e. the degree to which a top predator lake trout, Salvelinus namaycush, derives energy from the littoral zone) along a gradient of lake shape, where lake shape modifies the relative quantity of coupled epilimnetic benthic and pelagic habitats within each lake. Herein we demonstrate that littoral habitat coupling is intensified in simple circular lakes compared to their reticulate counterparts in seven Canadian Shield lakes. Although the more reticulate lakes had larger areas of epilimnetic benthic habitat, littoral food sources comprised 11% compared to 24% of lake trout diet in reticulate and circular lakes, respectively. This heightened interaction in circular lakes also appears to translate into increased omnivory in more circular lakes compared to reticulate lakes such that lake trout of circular lakes have a significantly lower trophic position than lake trout of reticulate lakes (F_1,5=6.71 p=0.05). These results suggest that it is the accessibility of littoral production via thermal refugia, and not the amount of littoral production, that determines the degree to which lake trout couple littoral and pelagic habitats in lakes.     

Composition and dispersal of riverine and lake phytoplankton communities in connected systems with different water retention times

1. Lake phytoplankton community structure may be influenced by both internal factors (predation, competition, resource constraints) and external ones, such as dispersal of materials and cells between connected habitats. However, little is known about the importance of cell dispersal for phytoplankton community structure in lakes. 2. We investigated the abundance and dispersal of phytoplankton cells between connected rivers and lakes, and analysed whether similarities in phytoplankton community composition between rivers and lakes were primarily related to cell import rates or to characteristics of the local habitat. We focused on lakes along a gradient of theoretical water retention times (TWRT). Two data sets from Swedish lakes were used; a seasonal study of two connected boreal forest lakes, differing in TWRT, and a multi‐lake study of 13 lakes with a continuous range of TWRTs. 3. Phytoplankton cells were transported and dispersed in all investigated rivers. In the seasonal study, cell import rates and similarities in phytoplankton community composition between the lake and its inlet(s) were much higher in the lake with a shorter TWRT. Phytoplankton community structure in different habitats was associated with total organic carbon (TOC). This indicates that local habitat characteristics may be important in determining lake phytoplankton community composition, even in the presence of substantial cell import. 4. The multi‐lake study also showed a negative relationship between TWRT and similarities in phytoplankton community composition between inlets and lakes. Moreover, similarity in community structure was related to both cell import rates from inlet to lake and differences in habitat characteristics between inlet and lake. However, the variable most strongly correlated with community structure was TOC, indicating that species sorting rather than a mass effect was the most important mechanism underlying the correlation between community structure and retention time. 5. Overall, our data suggest that local habitat characteristics may play a key role in determining community similarity in this set of lakes covering a large range of habitat connectedness. Due to the strong co‐variations between cell dispersal and TOC, it was hard to unequivocally disentangle the different mechanisms; hence, there is a need for further studies of the role of dispersal for phytoplankton community structures.     

A spatially explicit computer model for immature distributions of Glyptotendipes paripes (Diptera: Chironomidae) in central Florida lakes

Density and spatial distribution data of immature Glyptotendipes paripes Edwards (Diptera: Chironomidae), in relation to selected water and sediment characteristics prevailing in three central Florida lakes, were used to develop and calibrate a qualitative model of G. paripes distributions with the purpose of developing more efficient sampling plans for research or population management. For the model, each lake habitat was defined by a two-dimensional matrix, while a three-dimensional matrix was used to simulate the life stage-structured population. A Lefkovitch population matrix was used to project survival and development of the population in each spatial unit. The model incorporated lake bathymetry, sediment dry weight, water level, food availability (as a function of Secchi disk transparency) and strength of density dependence as influences on immature survival and adult fecundity. Temperature-dependent development data for G. paripes were incorporated to estimate development time. A simple redistribution function was used to simulate the dispersal of adults over the simulated habitat for oviposition. Immatures and relevant environmental data taken for eight dates, at each of two other central Florida lakes were used to validate the model. The mean correct prediction rate of the model for field density distributions within 0.5 log(n+1) immatures/m² density in spatial habitat cells was 0.64 and 0.66 for the validation lakes. Presence/absence correct prediction rates were 0.61 and 0.76, while matching to a proposed sampling stratification based on a treatment threshold was 0.85 and 0.88. These values indicate that the model is efficient for preparing stratified immature G. paripes sampling plans for central Florida lakes.     

Distribution of macroinvertebrates in relation to physical and biological variables in the littoral zone of nine New Zealand lakes

Macroinvertebrates play a key role in the littoral zone of lakes. Macroinvertebrate community composition is closely linked to habitat conditions. To date, there have been few attempts to relate macroinvertebrates to habitat factors in lakes. In this study, nine mainly oligotrophic lakes from throughout New Zealand were surveyed for macroinvertebrates. The lakes were selected to represent a range of suspended sediment loading and lake level regimes. Within each lake, several sites were selected to provide a range of exposure to wave action. A multiple regression approach was taken to relate macroinvertebrate community composition and habitat characteristics. The results of the analysis suggest that the littoral zone of the lakes we studied could be divided into four general habitats. The first is the wave wash zone characterised by coarse substrates and macroinvertebrate taxa usually associated with lotic environments, such as Ephemeroptera and Plecoptera. The second habitat is associated with macrophytes and is limited at the top by wave action and at depth by light attenuation. In this zone, the snail Potamopyrgus antipodarum is dominant, along with Trichoptera and Odonata. At the base of the macrophytes is the detrital habitat characterised by fine, organic rich sediments and dominated by chironomids, oligochaetes and Trichoptera. At depths below the macrophyte zone, fine sediments are found, and bivalves such as the freshwater mussel Hyridella menziesi are common. While macroinvertebrate abundance can be highly variable, some general predictions of community structure can be made based on a few key environmental factors. Abundance of snails Odonata and Trichoptera was positively related to macrophyte biomass. Some macroinvertebrate groups such as oligochaetes, chironomids, snails and bivalves were more common in fine substrates, while Ephemeroptera were characteristic of coarse substrates. Detrital biomass was important for most of the macroinvertebrate groups studied showing a positive relationship for oligochaetes and Trichoptera and a negative relationship for Ephemeroptera and Plecoptera.     

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.     

Soluble reactive phosphorus release from bryozoan dominated periphyton

Net release of soluble reactive phosphorus from mass developments of bryozoans, grown on artificial substrata in Mikolajskie Lake, Poland, was measured in depth profiles from 0 to 4 m. Release rates up to 94 g P dm^-2 h^-1 were recorded. P uptake by algae in the bryozoan community was a minor component of the P balance. The P exchange was several times higher in this community than typical values for other periphyton communities in the same lake. It is concluded that well developed bryozoan communities, when present, can contribute considerably to the recycling of phosphorus in the littoral of smaller lakes.     

Impacts of space,local environment and habitat connectivity on macrophyte communities in conservation lakes

Aim  To assess the relative impacts of spatial, local environmental and habitat connectivity on the structure of aquatic macrophyte communities in lakes designated for their conservation value. Location  Selected lakes of conservation importance all over Scotland, representing a wide variety of lake habitat types and associated macrophyte communities. Methods  Local environmental variables and species occurrence were measured in the field. Spatial variables were generated using principal coordinates of neighbour matrices (PCNM) analysis. Connectivity between each lake and its neighbours was defined as either (i) all lakes within a radius of 5, 10, 25, 50, 75 or 100 km; (ii) all lakes in same river system; or (iii) all lakes in the same catchment and upstream of the lake. Using variance partitioning within canonical correspondence analysis, the relative impact of E = local environment, S = space and C = lake connectivity was compared on submerged (n = 119 lakes) and emergent (n = 96 lakes) macrophyte assemblages. Results  Local environmental conditions, such as total phosphorus, alkalinity/conductivity and the presence of invasive species, as well as spatial gradients were key drivers of observed variation in macrophyte communities; e.g., for submerged macrophytes, a combination of local to moderate factors relating to water chemistry and broad‐scale gradients reflecting elevation and climate are important. Spatially structured environmental variables explained a large portion of observed variation. Main conclusions  Our findings confirmed the need to manage local environmental pressures such as eutrophication, but suggested that the traditional catchment approach was insufficient. The spatial aggregation of environmental and connectivity factors indicated that a landscape scale approach should be used in lake management to augment the risk assessment to conservation species from the deterioration of suitable lake sites over broad biogeographic areas.     

Experimental cultivation of the invasive freshwater bryozoan <Emphasis Type="Italic">Pectinatella magnifica</Emphasis>

The freshwater bryozoan Pectinatella magnifica (Leidy, 1851) is an invasive species in many countries all over the world. Although native to North America, it has been found in many countries of Europe and Asia. Pectinatella magnifica forms the largest colonial masses from all recently known bryozoan species. Culturing this organism in an aquarium has never been achieved for more than few days so far. Colonies from laboratory culture are important for various studies on its biology and life cycle of this species in experimental conditions. Young colonies successfully hatched from germinating statoblasts of P. magnifica in the laboratory and were maintained over eight weeks. Moreover, this was the first time when the compound colonies of this species were carried from its natural habitat to the laboratory, into a special aquarium system, and kept alive for more than three weeks. In both experiments the physicochemical parameters of the water (temperature, concentration of dissolved oxygen, electrolytic conductivity and pH) and changes in weight of compound colonies of P. magnifica in laboratory conditions were checked. The results found in this study are essential for understanding the invasiveness of this species and identifying methods for elimination of its ecological risks because these are closely resembling those of other invasive species.     

Landsat TM inventory and assessment of waterbird habitat in the southern <Emphasis Type="Italic">altiplano</Emphasis> of South America

The diverse set of wetlands in southern altiplano of South America supports a number of endemic and migratory waterbirds. These species include endangered endemic flamingos and shorebirds that nest in North America and winter in the altiplano. This research developed maps from nine Landsat Thematic Mapper (TM) images (254,300 km²) to provide an inventory of aquatic waterbird habitats. Image processing software was used to produce a map with a classification of wetlands according to the habitat requirements of different types of waterbirds. A hierarchical procedure was used to, first, isolate the bodies of water within the TM image; second, execute an unsupervised classification on the subsetted image to produce 300 signatures of cover types, which were further subdivided as necessary. Third, each of the classifications was examined in the light of field data and personal experience for relevance to the determination of the various habitat types. Finally, the signatures were applied to the entire image and other adjacent images to yield a map depicting the location of the various waterbird habitats in the southern altiplano. The data sets referenced with a global positioning system receiver were used to test the classification system. Multivariate analysis of the bird communities censused at each lake by individual habitats indicated a salinity gradient, and then the depth of the water separated the birds. Multivariate analysis of the chemical and physical data from the lakes showed that the variation in lakes were significantly associated with difference in depth, transparency, latitude, elevation, and pH. The presence of gravel bottoms was also one of the qualities distinguishing a group of lakes. This information will be directly useful to the Flamingo Census Project and serve as an element for risk assessment for future development.This revised version was published online in March 2005 with corrections to the issue cover date.     

Phylogenetic conservation of freshwater lake habitat preference varies between abundant bacterioplankton phyla

Despite their homogeneous appearance, aquatic systems harbour heterogeneous habitats resulting from nutrient gradients, suspended particulate matter and stratification. Recent reports suggest phylogenetically conserved habitat preferences among bacterioplankton, particularly for particle‐associated (PA) and free‐living (FL) habitats. Here, we show that independent of lake nutrient level and layer, PA and FL abundance‐weighted bacterial community composition (BCC) differed and that inter‐lake BCC varied more for PA than for FL fractions. In low‐nutrient lakes, BCC differences between PA and FL fractions were larger than those between lake layers. The reverse was true for high‐nutrient lakes. Nutrient level affected BCC more in hypolimnia than in epilimnia, likely due to hypolimnetic hypoxia in high‐nutrient lakes. In line with previous reports, we observed within‐phylum operational taxonomic unit (OTU) habitat preference conservation, although not for all phyla, including the phylum with the highest average relative abundance across all habitats (Bacteroidetes). Consistent phylum‐level habitat preferences may indicate that the functional traits that underpin ecological adaptation of freshwater bacteria to lake habitats can be phylogenetically conserved, although the levels of conservation are phylum dependent. Resolving taxa preferences for freshwater habitats sets the stage for identification of traits that underpin habitat specialization and associated functional traits that influence differences in biogeochemical cycling across freshwater lake habitats.     

The use of winter refuges by roach tagged with miniature radio transmitters

Adult roach (Rutilus rutilus) (13–25 cm) from two shallow lakes, one turbid and one with high water transparency, were tagged with surgically implanted miniature radio transmitters and their habitat use was monitored. Repeated tracking series over the course of the year provided information on seasonal changes in behaviour of roach in the two lakes. The results showed that the roach generally used the lakes only as summer habitat, whereas they moved into specific sheltered, restricted areas during winter. This behaviour was most pronounced in the lake with clear water, where roach gathered in large numbers in the stream in- and outlets as well as in other sheltered habitats where they remained rather passively during the 48 h tracking periods. In the turbid lake, many of the tagged roach, were even found in underground channels during the tracking in November. We see this behaviour as a response to low food abundance, decreased food demand and high predation risk, especially from birds. The tracking data from the clear lake showed that some tagged roach undertook swift habitat shifts from one refuge to the other. The traditional perception of the roach as a `lake fish' with an important structuring role in the lake ecosystem is undoubtedly true, but the present results suggest that roach are able to migrate between different habitat types and will do so if they can profit from this behaviour.     

MARINE‐DERIVED DINOFLAGELLATES IN ANTARCTIC SALINE LAKES: COMMUNITY COMPOSITION AND ANNUAL DYNAMICS1*[link]

The saline lakes of the Vestfold Hills in Antarctica offer a remarkable natural laboratory where the adaptation of planktonic protists to a range of evolving physiochemical conditions can be investigated. This study illustrates how an ancestral marine community has undergone radical simplification leaving a small number of well‐adapted species. Our objective was to investigate the species composition and annual dynamics of dinoflagellate communities in three saline Antarctic lakes. We observed that dinoflagellates occur year‐round despite extremely low PAR during the southern winter, which suggests significant mixotrophic or heterotrophic activity. Only a small number of dominant dinoflagellate species were found in each lake, in contrast to the species‐rich Southern Ocean from which the lake communities are believed to be derived. We verified that the lake species were representatives of the marine polar dinoflagellate community, and not freshwater species. Polarella glacialis Montresor, Procaccini et Stoecker, a bipolar marine species, was for the first time described in a lake habitat and was an important phototrophic component in the higher salinity lakes. In the brackish lakes, we found a new sibling species to the brackish‐water species Scrippsiella hangoei (J. Schiller) J. Larsen, previously observed only in the Baltic Sea.     

Behavioural strategy of large perch Perca fluviatilis varies between a mesotrophic and a hypereutrophic lake

Behaviour of large perch Perca fluviatilis was studied in two lakes differing in environmental state i.e. mesotrophic v. hypereutrophic. A total of 20 adult perch P. fluviatilis (29–42 cm total length) in each lake were tagged with radio‐transmitters, tracked and located eight times a day during six 24 h tracking periods over a year, enabling detection of differences in diel activity patterns and habitat use during summer and winter under two different environmental regimes. During summer, P. fluviatilis in the mesotrophic lake showed a distinct crepuscular activity pattern and a change from pelagic residency during daytime towards the littoral zone at night. In contrast, P. fluviatilis in the hypereutrophic lake were active during the entire diel cycle and were spread throughout the lake also during dark. During winter, crepuscular patterns of activity were seen in both lakes. Condition factor of large P. fluviatilis did not differ between the two lakes. Thus, it is suggested that P. fluviatilis in the hypereutrophic turbid lake adopted an alternative behaviour for successful foraging, being uniformly active throughout the diel cycle.     

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.