Marina Manca

Cladocera: predators and prey

Among the freshwater zooplankton community, Cladocera represent one of the most common elements of pelagic populations. Being almost exclusively filter feeders and algae users and, at the same time, the favourite prey of invertebrate and vertebrate predators, Cladocera represent the most important group in the plankton community of lakes as regards energy transfer along the food chain. Because of their short generation times and their high reproductive efficiency, predation by invertebrates, usually, has only a limited role in controlling their density. However, at high densities, invertebrate predators can provide an effective control of Cladocera populations. The intensive research on selective predation by vertebrates has demonstrated that this activity can be responsible, together with competitive interactions, for the dominance of different groups in the planktonic community: large Cladocera dominate when predation is low, Rotifera and small Crustacea dominate at high predation levels and high nannoplanktonic densities. These evidences on the role of vertebrate predation in structuring aquatic environments has greatly contributed to our better understanding of aquatic ecosystem functioning. In particular, it seems that the removal of large filter-feeding herbivorous Cladocera by zooplanktivorous fish can lead to worsening environmental conditions in eutrophicating lakes. In this respect, Cladocera appear to be the key group among zooplanktonic organisms, and their interactions the key factors in aquatic food chain management.

Reticulate evolution of the Daphnia pulex complex as revealed by nuclear markers

The study of species complexes is of particular interest to understand how evolutionary young species maintain genomic integrity. The Daphnia pulex complex has been intensively studied as it includes species that dominate freshwater environments in the Northern hemisphere and as it is the sole North American complex that shows transitions to obligate parthenogenesis. Past studies using mitochondrial markers have revealed the presence of 10 distinct lineages in the complex. This study is the first to examine genetic relationships among seven species of the complex at nuclear markers (nine microsatellite loci and one protein‐coding gene). Clones belonging to the seven species of the Daphnia pulex complex were characterized at the mitochondrial NADH dehydrogenase (ND5) gene and at the Lactate dehydrogenase (LDH) locus. K‐means, principal coordinate analyses and phylogenetic network analyses on the microsatellite data all separated European D. pulicaria, D. tenebrosa, North American D. pulex, D. pulicaria and their hybrids into distinct clusters. The hybrid cluster was composed of diploid and polyploid hybrids with D. pulex mitochondria and some clones with D. pulicaria mitochondria. By contrast, the phylogeny of the D. pulex complex using Rab4 was not well resolved but still showed clusters consisting mostly of D. pulex alleles and others of D. pulicaria alleles. Incomplete lineage sorting and hybridization may obscure genetic relationships at this locus. This study shows that hybridization and introgression have played an important role in the evolution of this complex.

Phytoplankton and Zooplankton Associations in a Set of Alpine High Altitude Lakes: Geographic Distribution and Ecology

Species composition and interactions, biomass dominance, geographic distribution and driving variables were investigated for two key elements of the pelagic food web of Alpine lakes, the phytoplankton and the zooplankton, based on a single sampling campaign during summer 2000. Altogether, 70 lakes were surveyed, 49 of which located in three different lake districts of the west and eastern Italian Alps and 21 in the central Austrian Alps (within the uppermost Danube catchment). In addition to the analysis of environmental variables affecting distribution and species structure of the two planktonic compartments, a brief review of the main research lines and hypotheses adopted in the past for the study of phytoplankton and zooplankton in high Alpine lakes is given. The lakes, investigated partly within the European project EMERGE (EVK1-CT-1999-00032) and partly within a regional project in the eastern Alps, comprise a wide range of morphological, chemical and trophic conditions. The phytoplankton communities were found to be diverse and mostly dominated by flagellates (chrysophytes, cryptophytes and dinoflagellates), and only to a lesser extent by non-motile green algae, desmids and centric diatoms. The zooplankton communities were mainly dominated by Alpine cladocerans and copepod species, while rotifers were abundant within one group of Italian lakes (sampled in early summer). The multivariate statistical analyses (CCA) showed that catchment features (i.e. percentage of vegetation cover and geochemical composition) and nitrate concentration are essential drivers for the phytoplankton, whereas for zooplankton also trophic status of the lakes and phytoplankton structure are important. The combined variance analysis of the lake clusters outlined by the multivariate analyses on phytoplankton and zooplankton data, respectively, allowed the identification of four principal lake types (three located on siliceous and one on carbonaceous bedrock), each one characterised by a certain combination of habitat features, which in their turn influence trophic state, and phytoplankton and zooplankton species composition and functionality.

Zooplankton (Cladocera) species turnover and long-term decline of Daphnia in two high mountain lakes in the Austrian Alps

We investigated long-term succession of sedimentary cladoceran assemblages in two morphologically divergent mountain lakes by utilizing sediment traps and previously available sediment data. We aimed to detect lake-specific changes in pelagic communities potentially attributable to climate warming under the presumption that lakes and biotic communities may respond individually to the same regional climatic forcing. Both lakes showed a similar community turnover, as Daphnia was first replaced by Chydorus cf. sphaericus and during the twentieth century by the latest colonizer Bosmina. The community succession was similar among the lakes and consistent with the regional temperature increase, although the timing of community shift, the magnitude of change, and taxa in question differed. Decline of Daphnia mismatched with historical fish stockings, but the eventual extirpation of Daphnia in one of the lakes corresponded to the start of fish introductions. We propose that the observed shifts were mainly governed by increasing temperatures and its limnoecological consequences (e.g., deeper mixing). We suggest that Bosmina distribution may be extending to lakes at higher altitudes as a response to climate warming, and that it may replace Daphnia as the key component of pelagic alpine food webs by coping in interspecific resource competition under changed limnological regimes.

Notes on Cladocera and Copepoda from high altitude lakes in the Mount Everest Region (Nepal)

The paper deals with Cladocera and Copepoda species from eight high altitude lakes in the Khumbu area. In all lakes, an endemic known diaptomid of the palearctic genus Arctodiaptomus is found. The dark and large Daphnia tibetana occurs in five of the lakes sampled, characterized by the persistence of the spine in adults (parthenogenetic and ephippial females, males). Apparently, this is an unknown feature of this species. The literature on the diagnostic traits of the different morphs described is reviewed. In addition, a transparent and smaller-sized Daphnia species occurs in two lakes. This is a D. longispina characterized by the absence in adults of the carapace spine (var. aspina Weretschagin, 1911). The presence of these two species is discussed in relation to water transparency, colour, and vertical distribution. Two hypotheses on the evolution of cuticular pigmentation in Daphnia are examined. In addition to these mostly dominant species, a macrothricid also typical of high altitude lakes in the Alps was found (Macrothrix hirsuticornis) together with two cosmopolitan Chydoridae.

The decline of Daphnia hyalina galeata in Lago Maggiore: a comparison of the population dynamics before and after oligotrophication

Abstract: We reconstruct the impact of different environmental cues on Daphnia seasonal dynamics by means of an analysis of demographic and reproductive parameters, as well as of the size structure of the population. Data from 1996, indicative of the most recent productivity of the lake, with those from 1973, when the lake was meso-eutrophic, allow a discussion of the relevance of the observed changes for the structure of the pelagic food web of Lago Maggiore. Daphnia summer dynamics is mainly controlled by Bythotrephes longimanus. This is quite a different situation from that of the past, when Leptodora kindtii was the prevalent predatory cladocera. The size structure of Daphnia population revealed a depletion of small ovigerous females with increases in the predator. Our data allow a discussion of the idea, recently proposed, of Bythotrephes longimanus as a visual sit- and -wait predator.

Reconstructing long-term changes in Daphnia's body size from subfossil remains in sediments of a small lake in the Himalayas

A combined analysis of modern zooplankton and fossil Cladocera assemblages from a Himalayan lake, Lake 40, revealed that the endemic Daphnia tibetana disappeared in the late-1980s, after persisting as the only Daphnia species for almost 3000 years. Daphnia head shields, which are rarely recovered from the sediments, were the most abundant Daphnia remains in the lake. The remains were of the Ctenodaphnia type; the smaller ones had a large central hole. Head shields of the same type were also found in zooplankton water samples, rich in exuviae as well as of intact specimens of D. tibetana, from a nearby lake. Small individuals had a distinct nuchal organ in the dorsal region of the head. We therefore postulated that the head shields with the hole were from young (newborn or individuals in the two first moults). Up to now, the nuchal organ has been described in laboratory populations of D. (Ctenodaphnia) magna, where it disappears quite early in life. It probably functions as an osmoregulatory organ, essential for the survivorship of late embryos and early juveniles. On the other hand, as far as we know, head shields with a hole have never been recorded in plankton or in sediments. In view of the fact that head shields were representative of Daphnia abundance, we used them to reconstruct changes in density and body size during ca. 3000 years. In fishless mountain lakes, mean Daphnia body size tends to increase toward the end of the productive season. The number of moults and maximum body size depend mainly on the duration of the ice-free period, and on summer temperatures. In cold years, when the productive season is short, the number of moults will be low, and the range of Daphnia body size will be narrow. We used Daphnia body size and abundance estimates, in addition to an analysis of changes in the Cladocera assemblage, to reconstruct past environmental conditions.

Evidence for short-lived oscillations in the biological records from the sediments of Lago Albano (Central Italy) spanning the period ca. 28 to 17 k yr BP

We report the results of analyses of pigments (derived from algae and photosynthetic bacteria), diatoms and invertebrate fossil remains (ostracods, cladocerans, chironomids) in two late Pleistocene sediment cores from Lago Albano, a crater lake in Central Italy. The record contains evidence for oscillations in lake biota throughout the period ca. 28 to 17 k yr BP. The earliest of these are contained in the basal 3.5 m of light olive-gray and yellowish-gray spotted muds sampled in core PALB 94-1E from 70 m water depth. The later oscillations are best represented in the more extended sediment sequence recovered from a second core site, PALB 94-6B, in 30 m water depth. The sediments at site 1E, containing the earlier oscillations (ca. 28-24 k yr BP), predate any sedimentation at the shallower site, from which we infer an initially low lake level rising to permit sediment accumulation at site 6B from ca. 24 k yr onwards. At site 6B, massive silts rich in moss remains are interbedded with laminated silts and carbonates. These sediments span the period ca. 24 to 17 k yr and are interpreted as representing, respectively, times of shallow water alternating with higher lake stands, when the lake was stratified and bottom water was stagnant. A range of mutually independent chronological constraints on the frequency and duration of the oscillations recorded in the lake biota indicate that they were aperiodic and occurred on millennial to century timescales. We interpret them as responses to climate forcing through its impact on lake levels and changing aquatic productivity. The time span they occupy, their frequency and their duration suggest that at least some of these changes may parallel both the Dansgaard-Oeschger events recorded in Greenland Ice Cores and the contemporary oscillations in North Atlantic circulation documented in marine sediment cores.

High-resolution paleolimnology opens new management perspectives for lakes adaptation to climate warming

Varved lake sediments provide opportunities for high-resolution paleolimnological investigations that may extend monitoring surveys in order to target priority management actions under climate warming. This paper provides the synthesis of an international research program relying on >150 years-long, varved records for three managed perialpine lakes in Europe (Lakes Geneva, Annecy and Bourget). The dynamics of the dominant, local human pressures, as well as the ecological responses in the pelagic, benthic and littoral habitats were reconstructed using classical and newly developed paleo-proxies. Statistical modelling achieved the hierarchization of the drivers of their ecological trajectories. All three lakes underwent different levels of eutrophication in the first half of the XXth century, followed by re-oligotrophication. Climate warming came along with a 2°C increase in air temperature over the last century, to which lakes were unequally thermally vulnerable. Unsurprisingly, phosphorous concentration has been the dominant ecological driver over the last century. Yet, other human-influenced, local environmental drivers (fisheries management practices, river regulations) have also significantly inflected ecological trajectories. Climate change has been impacting all habitats at rates that, in some cases, exceeded those of local factors. The amplitude and ecological responses to similar climate change varied between lakes, but, at least for pelagic habitats, rather depended on the intensity of local human pressures than on the thermal effect of climate change. Deep habitats yet showed higher sensitivity to climate change but substantial influence of river flows. As a consequence, adapted local management strategies, fully integrating nutrient inputs, fisheries management and hydrological regulations, may enable mitigating the deleterious consequences of ongoing climate change on these ecosystems.

A late glacial and holocene record of biological and environmental changes from the crater Lake Albano, Central Italy: An interdisciplinary european project (PALICLAS)

This paper reports the results of biological analyses (pigments, diatoms, chrysophyte cysts, cladocerans, chironomids and ostracods) of a ca. 14 m-long sediment core recovered from Lake Albano (Central Italy) in the course of the EU-funded project PALICLAS (PALaeoenvironmental analysis of Italian Crater Lake and Adriatic Sediments).A reconstruction of the environmental evolution and ecosystem response of Lake Albano during the last ca. 30 kyr was possible. Additional information on lake level oscillation is obtained from benthic and planktonic palaeocommunities. Several oscillations in the productivity and the level of the lake were detected in the oldest sediment layers (from ca. 30 kyr BP to ca. 17 kyr BP), followed by a long (ca. 5 kyr BP) period of low productivity in which cold, holomictic conditions prevailed. A period of high biological activity and, probably, meromictic conditions during the early-mid Holocene was detected. A clear impact of human activities in the catchment was found at ca. 4 kyr BP in the form of increased erosion, associated with a decline in the abundance of biological remains. Further signs of human impact on the lake ecosystem are recorded during the Roman period. Although large-scale environmental changes (e.g. regional climate changes) caused many of the observed biological changes, human activities were important during the mid-late Holocene.