Dagmar Frisch

High dispersal capacity of a broad spectrum of aquatic invertebrates via waterbirds

Abstract.Speculation about the role of waterbirds in the dispersal of aquatic invertebrates pre-dates Darwin. However, there is a critical shortage of field studies quantifying such dispersal. We quantified the viability of aquatic invertebrates in the faeces of different waterfowl species collected in the field at different times during winter. Faeces were collected from four duck species (Northern Pintail Anas acuta, Mallard A. platyrhynchos, Shoveler A. clypeata, Eurasian Teal A. crecca) and Eurasian Coot Fulica atra in November 2004 and January 2005. We also collected soil samples from resting sites as an indicator of what may be transported on birds’ feet and plumage. Faecal and soil samples were incubated using two treatments (0.4 and 4.0 mS cm-1) to quantify the potential for dispersal between aquatic habitats of different salinities. We found that viable Nematoda, Rotifera, Copepoda, Ostracoda, Insecta (Tipulidae), and Daphnia and Moina cladocerans were transported internally by birds in the wild. We also found evidence that nematodes, rotifers, ostracods, copepods, tipulids, chironomids and hemipterans can be dispersed on birds’ feet and feathers. The overall incidence of hatching from all samples was higher in January (59.4%) than in November (11.5%). With the exception of bdelloid rotifers, we found no evidence that the potential for dispersal between two habitats would be impeded by salinity in the range tested. Our data suggest that the taxonomic range of dispersed invertebrates and the frequency of their dispersal via waterfowl has previously been underestimated.

Copepods come in first : rapid colonization of new temporary ponds

The sequence in which new colonists reach an empty habitat can be crucial for future development and species structure of communities. It is therefore important to assess species composition and abundance in the initial stages of habitat existence. In the present study we focussed on colonization of newly constructed temporary ponds in Donana, Southwest Spain, created by removing 30 or 60 cm of top soil. To confi rm that no egg-bank was present, we conducted hatching experiments using sediment cores from six new ponds and two reference sites. Hatching was not recorded in the sediment of the new ponds with the exception of two rotifer individuals. In contrast, in the reference sites a maximum of 103 individuals per sample hatched, including cyclopoids, cladocer- ans, ostracods and rotifers. In the fi eld, water samples were collected from seven ponds after 19 days of their fi rst hydroperiod. Cyclopoid copepods, mostly Metacyclops minutus, had arrived and dominated all sampled ponds. Other taxa were the monogonont rotifer Brachionus plicatilis in four ponds, and the cladoceran Moina brachiata in one pond. The abundance of zooplankton was negatively correlated with conductivity, suggesting that ponds with higher conductivity undergo delayed colonization. We suggest that fast dispersal and dominance of certain cyclo- poid copepods during early colonization is related to their ability to store sperm and fast individual development. Given that cyclopoid copepods can survive drought periods in the sediment of temporary ponds, fi rst and early colonization by cyclopoids is likely to have a profound effect on the propagule bank and future plankton communi- ties when the ponds refi ll.

Strong Spatial Influence on Colonization Rates in a Pioneer Zooplankton Metacommunity

The magnitude of community-wide dispersal is central to metacommunity models, yet dispersal is notoriously difficult to quantify in passive and cryptic dispersers such as many freshwater invertebrates. By overcoming the problem of quantifying dispersal rates, colonization rates into new habitats can provide a useful estimate of the magnitude of effective dispersal. Here we study the influence of spatial and local processes on colonization rates into new ponds that indicate differential dispersal limitation of major zooplankton taxa, with important implications for metacommunity dynamics. We identify regional and local factors that affect zooplankton colonization rates and spatial patterns in a large-scale experimental system. Our study differs from others in the unique setup of the experimental pond area by which we were able to test spatial and environmental variables at a large spatial scale. We quantified colonization rates separately for the Copepoda, Cladocera and Rotifera from samples collected over a period of 21 months in 48 newly constructed temporary ponds of 0.18–2.95 ha distributed in a restored wetland area of 2,700 ha in Doñana National Park, Southern Spain. Species richness upon initial sampling of new ponds was about one third of that in reference ponds, although the rate of detection of new species from thereon were not significantly different, probably owing to high turnover in the dynamic, temporary reference ponds. Environmental heterogeneity had no detectable effect on colonization rates in new ponds. In contrast, connectivity, space (based on latitude and longitude) and surface area were key determinants of colonization rates for copepods and cladocerans. This suggests dispersal limitation in cladocerans and copepods, but not in rotifers, possibly due to differences in propagule size and abundance.

Observing copepods through a genomic lens

BackgroundCopepods outnumber every other multicellular animal group. They are critical components of the worlds freshwater and marine ecosystems, sensitive indicators of local and global climate change, key ecosystem service providers, parasites and predators of economically important aquatic animals and potential vectors of waterborne disease. Copepods sustain the world fisheries that nourish and support human populations. Although genomic tools have transformed many areas of biological and biomedical research, their power to elucidate aspects of the biology, behavior and ecology of copepods has only recently begun to be exploited.DiscussionThe extraordinary biological and ecological diversity of the subclass Copepoda provides both unique advantages for addressing key problems in aquatic systems and formidable challenges for developing a focused genomics strategy. This article provides an overview of genomic studies of copepods and discusses strategies for using genomics tools to address key questions at levels extending from individuals to ecosystems. Genomics can, for instance, help to decipher patterns of genome evolution such as those that occur during transitions from free living to symbiotic and parasitic lifestyles and can assist in the identification of genetic mechanisms and accompanying physiological changes associated with adaptation to new or physiologically challenging environments. The adaptive significance of the diversity in genome size and unique mechanisms of genome reorganization during development could similarly be explored. Genome-wide and EST studies of parasitic copepods of salmon and large EST studies of selected free-living copepods have demonstrated the potential utility of modern genomics approaches for the study of copepods and have generated resources such as EST libraries, shotgun genome sequences, BAC libraries, genome maps and inbred lines that will be invaluable in assisting further efforts to provide genomics tools for copepods.SummaryGenomics research on copepods is needed to extend our exploration and characterization of their fundamental biological traits, so that we can better understand how copepods function and interact in diverse environments. Availability of large scale genomics resources will also open doors to a wide range of systems biology type studies that view the organism as the fundamental system in which to address key questions in ecology and evolution.

Phylogeny, molecular ecology and taxonomy of southern Iberian lineages of Triops mauritanicus (Crustacea: Notostraca)

We investigated the phylogeography of the main lineages in the tadpole shrimp Triops mauritanicus Ghigi in the south-western Iberian Peninsula, using mitochondrial 12S and 16S rDNA sequences. Our results indicate that a fourth, hitherto unknown main phylogenetic lineage occurs in Iberia, so that in total, the species is divided into six distinct clades, comprising T. m. mauritanicus, T. m. simplex Ghigi, and four as yet unnamed lineages that appear to be endemic to Iberia. Percentages of sequence divergence among the main clades in T. mauritanicus reach the range reported for recognized species in other notostracan lineages. A thorough morphological investigation also revealed that the differentiation among these lineages is higher than previously thought, and that populations of three of the main clades within T. mauritanicus can be reliably separated from each other and from the remaining lineages based on the morphology of adult males. The remaining clades also show a significant level of morphological differentiation, but include a certain proportion of populations for which the additional application of molecular methods is needed for a reliable determination. The geographic distributions of 12S haplotypes are indicative of frequent dispersal events and gene flow among populations belonging to the same main lineage, but give no evidence of recent migration events among different main lineages, suggesting that there is no gene flow among the latter. Our data thus suggest that the six main lineages within T. mauritanicus represent distinct species. We therefore describe the Iberian lineages as T. baeticus Korn n. sp., T. emeritensis Korn & Pérez-Bote n. sp., T. gadensis Korn & García-de-Lomas n. sp., and T. vicentinus Korn, Machado, Cristo & Cancela da Fonseca n. sp., and reinstate T. simplex Ghigi to full species status. Our data confirm the general, previously recognized pattern of a lower dispersal probability in gonochoric Triops taxa. However, we found evidence that passive dispersal in Triops may be further complicated by a strong habitat dependence of dispersal probability, mediated by prevailing dispersal vectors.

Differential transcriptomic responses of ancient and modern Daphnia genotypes to phosphorus supply

Little is known about the role of transcriptomic changes in driving phenotypic evolution in natural populations, particularly in response to anthropogenic environmental change. Previous analyses of Daphnia genotypes separated by centuries of evolution in a lake using methods in resurrection ecology revealed striking genetic and phenotypic shifts that were highly correlated with anthropogenic environmental change, specifically phosphorus (P)‐driven nutrient enrichment (i.e. eutrophication). Here, we compared the transcriptomes of two ancient (~700‐year‐old) and two modern (~10‐year‐old) genotypes in historic (low P) and contemporary (high P) environmental conditions using microarrays. We found considerable transcriptomic variation between ‘ancient’ and ‘modern’ genotypes in both treatments, with stressful (low P) conditions eliciting differential expression (DE) of a larger number of genes. Further, more genes were DE between ‘ancient’ and ‘modern’ genotypes than within these groups. Expression patterns of individual genes differed greatly among genotypes, suggesting that different transcriptomic responses can result in similar phenotypes. While this confounded patterns between ‘ancient’ and ‘modern’ genotypes at the gene level, patterns were discernible at the functional level: annotation of DE genes revealed particular enrichment of genes involved in metabolic pathways in response to P‐treatments. Analyses of gene families suggested significant DE in pathways already known to be important in dealing with P‐limitation in Daphnia as well as in other organisms. Such observations on genotypes of a single natural population, separated by hundreds of years of evolution in contrasting environmental conditions before and during anthropogenic environmental changes, highlight the important role of transcriptional mechanisms in the evolutionary responses of populations.

Long-term changes in metapopulation genetic structure: a quarter-century retrospective study on low-Arctic rock pool Daphnia.

Population genetic surveys approximately 25 years apart examined the distribution and abundance of asexual clones of the freshwater zooplankter Daphnia pulex complex in rock pools near Churchill, Manitoba, Canada. In 1984–1985, melanic members of this species complex were present in 131 rock pools at this site, but were only detected in 90 of these pools in 2007–2008. Allozymic surveys conducted during these two time periods revealed that 59 per cent of these populations showed unchanged clonal composition. Total clonal replacement occurred in 8 per cent of the populations, while the others (33%) included a mixture of ‘resident’ clones and new ‘colonists’. We discuss these changes in light of shifts in biotic and abiotic factors. We also discuss the use of rock pool habitats as ‘sentinel’ systems for examining long-term environmental changes in the ecological genetics of aquatic organisms in the Arctic.

Diversity of floodplain copepods (Crustacea) modified by flooding: species richness, diapause strategies and population genetics

We examined the impact of lateral and longitudinal hydrological connectiv- ity on copepod diversity (local species richness, diapause strategies and population genetics) in ponds of a North American floodplain. Results of a General Linear Model (GLM) which included three environmental pond variables showed that species rich- ness in these ponds was significantly influenced only by pond type (permanent ponds, temporary ponds with connections to other waterbodies during floods, isolated tempo- rary ponds). Local species richness in connected temporary ponds had significantly higher scores of local richness compared to isolated temporary ponds (means 7.4 and 2.7, respectively). Such enhancement of species diversity in connected ponds appears to be facilitated by increased dispersal during floods and also pertains to the diversity of diapause strategies in the ponds studied. Species with and without diapause were present in connected temporary ponds, but absent from isolated temporary ponds, which contained only species capable of diapausing during the dry season. To explore the significance of hydrological connectivity of longitudinal dispersal between up- stream and downstream sites, we used RAPD markers in populations of a calanoid co- pepod. The increase of average heterozygosity H (0.301 to 0.477) in an upstream- downstream direction and generally low genetic distances (Neis D < 0.05) suggested maintenance of gene flow by flood-mediated dispersal between connected ponds. Lower heterozygosity (0.289) of the population in a downstream pond protected by le- vees suggested the interruption of gene flow when dispersal by flooding is blocked.

The invasion history of the exotic freshwater zooplankter Daphnia lumholtzi (Cladocera, Crustacea) in North America: a genetic analysis.

Daphnia lumholtzi is a planktonic crustacean native to subtropical regions in Africa, Asia and Australia. Since its invasion to the southern USA in ~1990 it has spread across North America as far north as the Laurentian Great Lakes. We assessed invasion history using microsatellite makers and to explore the influence of mean annual temperature on the genetic structure along a latitudinal gradient in North America. Genotypic data were obtained from 9 microsatellite markers for 178 individuals from 13 populations (eight populations introduced to North America and five populations in the native range). Pairwise Fst values as well as Bayesian clustering showed a strong subdivision between native and introduced populations. Bayesian clustering identified multiple genetic clusters in recently invaded locations, suggestive of multiple invasions from various sources, including Asia and Africa. Using variation partitioning, we determined the amount of variation for genetic clusters of populations in the invaded range due to mean annual air temperature and the year of first detection. The results point to a primary introduction into the southern range of North America, with a subsequent northward expansion, and multiple introductions possibly from both the native range and by secondary spread from previously-invaded locations. Separate analysis of genetic clusters within the invaded range suggests additional effects of temperature conditions on geographic genetic structure, possibly as a consequence of D. lumholtzi’s tropical origin.

Paleogenetic records of Daphnia pulicaria in two North American lakes reveal the impact of cultural eutrophication.

Understanding the evolutionary consequences of the green revolution, particularly in wild populations, is an important frontier in contemporary biology. Because human impacts have occurred at varying magnitudes or time periods depending on the study ecosystem, evolutionary histories may vary considerably among populations. Paleogenetics in conjunction with paleolimnology enable us to associate microevolutionary dynamics with detailed information on environmental change. We used this approach to reconstruct changes in the temporal population genetic structure of the keystone zooplankton grazer, Daphnia pulicaria, using dormant eggs extracted from sediments in two Minnesota lakes (South Center, Hill). The extent of agriculture and human population density in the catchment of these lakes has differed markedly since European settlement in the late 19th century and is reflected in their environmental histories reconstructed here. The reconstructed environments of these two lakes differed strongly in terms of environmental stability and their associated patterns of Daphnia population structure. We detected long periods of stability in population structure and environmental conditions in South Center Lake that were followed by a dramatic temporal shift in population genetic structure after the onset of European settlement and industrialized agriculture in its watershed. In particular, we noted a 24.3-fold increase in phosphorus (P) flux between pre-European and modern sediment P accumulation rates (AR) in this lake. In contrast, no such shifts were detected in Hill Lake, where the watershed was not as impacted by European settlement and rates of change were less directional with a much smaller increase in sediment P AR (2.3-fold). We identify direct and indirect effects of eutrophication proxies on genetic structure in these lake populations and demonstrate the power of using this approach in understanding the consequences of anthropogenic environmental change on natural populations throughout historic time periods.