Zooplankton Diversity and Pelagic Food Webs: Investigating Present and Past with Different Techniques

A special issue of Water (ISSN 2073-4441). This special issue belongs to the section "Biodiversity and Functionality of Aquatic Ecosystems".

Deadline for manuscript submissions: closed (23 December 2019) | Viewed by 36613

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Special Issue Editors

CNR IRSA, Verbania, Italy
Interests: ecology of freshwater zooplankton and analysis of zooplankton resting stages in lacustrine sediment, ecotoxicology, stable isotope analyses, and aquatic food webs
Special Issues, Collections and Topics in MDPI journals
Department of Human Sciences and Innovation for the Territory (DiSUIT), University of Insubria, Via Valleggio 11, 22100 Como, Italy
Interests: ecology; ecotoxicology
Special Issues, Collections and Topics in MDPI journals
Department of Earth and Environmental Sciences, University of Milano-Bicocca, Piazza Della Scienza 1, I-20126 Milano, Italy
Interests: ecology of lakes; planktonic food webs; freshwater chemistry; long-term limnological studies; anthropogenic and climate impacts on pelagic communities
Special Issues, Collections and Topics in MDPI journals

Special Issue Information

Dear Colleagues,

Zooplankton are of key importance in the structure and functioning of aquatic food webs. They contribute to a large part of the functional and structural biodiversity of predator and prey plankton communities. Promptly responding to long-term and seasonal changes in the physical and chemical environment, they are sensitive indicators of patterns and mechanisms of impact drivers, both natural and human-induced. In this volume, we aim at presenting evidence for both long-term and seasonal changes in zooplankton community structure and dynamics, investigating different approaches, from population dynamics to advanced molecular techniques and reconstructing past communities from subfossil remains in lake sediments.

Dr. Roberta Piscia
Prof. Dr. Roberta Bettinetti
Dr. Barbara Leoni
Dr. Marina Manca
Guest Editors

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Keywords

  • zooplankton
  • ecosystem functioning
  • aquatic food webs
  • population dynamics
  • functional biodiversity
  • C &N stable isotope analyses
  • persistent pollutants

Published Papers (12 papers)

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Research

15 pages, 4625 KiB  
Article
Changes in Planktivory and Herbivory Regimes in a Shallow South American Lake (Lake Blanca Chica, Argentina) Over the Last 250 Years
by David Carrozzo, Simona Musazzi, Andrea Lami, Francisco E. Córdoba and María de los Ángeles González Sagrario
Water 2020, 12(2), 597; https://doi.org/10.3390/w12020597 - 22 Feb 2020
Cited by 3 | Viewed by 2844
Abstract
Shallow lakes are vulnerable ecosystems impacted by human activities and climate change. The Cladocera occupy a central role in food webs and are an excellent paleoecological indicator of food web structure and trophic status. We conducted a paleolimnological study in Lake Blanca Chica [...] Read more.
Shallow lakes are vulnerable ecosystems impacted by human activities and climate change. The Cladocera occupy a central role in food webs and are an excellent paleoecological indicator of food web structure and trophic status. We conducted a paleolimnological study in Lake Blanca Chica (Argentina) to detect changes on the planktivory and herbivory regimes over the last 250 years. Generalized additive models were fitted to the time series of fish predation indicators (ephippial abundance and size, mucrone size, fish scales, and the planktivory index) and pheophorbide a concentration. The cladoceran assemblage changed from littoral-benthic to pelagic species dominance and zooplankton switched from large-bodied (Daphnia) to small-bodied grazers (Bosmina) ca. 1900 due to increased predation. The shift in planktivory regime (ca. 1920–1930), indicated by fish scales and the planktivory index, as well as herbivory (ca. 1920–1950), was triggered by eutrophication. Changes in planktivory affected the size structure of Bosmina, reducing its body size. This study describes the baseline for the lake as well as the profound changes in the composition and size structure of the zooplankton community due to increased predation and the shift in the planktivory regime. These findings will provide a reference status for future management strategies of this ecosystem. Full article
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16 pages, 2023 KiB  
Article
Evaluation of the Egg Bank of Two Small Himalayan Lakes
by Roberta Piscia, Sara Bovio, Marina Manca, Andrea Lami and Piero Guilizzoni
Water 2020, 12(2), 491; https://doi.org/10.3390/w12020491 - 11 Feb 2020
Cited by 2 | Viewed by 2476
Abstract
High mountain lakes are biodiversity treasures. They host endemic taxa, adapted to live in extreme environments. Among adaptations, production of diapausing eggs allows for overcoming the cold season. These diapausing eggs can rest in the sediments, providing a biotic reservoir known as an [...] Read more.
High mountain lakes are biodiversity treasures. They host endemic taxa, adapted to live in extreme environments. Among adaptations, production of diapausing eggs allows for overcoming the cold season. These diapausing eggs can rest in the sediments, providing a biotic reservoir known as an egg bank. Here, we estimated changes in abundance of the egg bank in two lakes in the Khumbu Region of the Himalayas, during the last ca. 1100 and 500 years, respectively, by analyzing two sediment cores. We tested viability of the diapausing eggs extracted from different layers of the sediment cores under laboratory conditions. We found that only diapausing eggs of the Monogont rotifer Hexarthra bulgarica nepalensis were able to hatch, thus suggesting that a permanent egg bank is lacking for the other taxa of the lakes, not least for the two Daphnia species described from these sites. Our results confirm previous studies suggesting that in high mountain lakes, the production of diapausing is mainly devoted to seasonal recruitment, therefore leading to a nonpermanent egg bank. The different ability of different taxa to leave viable diapausing eggs in the sediments of high mountain lakes therefore poses serious constraints to capability of buffering risk of biodiversity loss in these extremely fragile environments. Full article
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25 pages, 1794 KiB  
Article
Species Richness and Taxonomic Distinctness of Zooplankton in Ponds and Small Lakes from Albania and North Macedonia: The Role of Bioclimatic Factors
by Giorgio Mancinelli, Sotir Mali and Genuario Belmonte
Water 2019, 11(11), 2384; https://doi.org/10.3390/w11112384 - 14 Nov 2019
Cited by 8 | Viewed by 3447
Abstract
Resolving the contribution to biodiversity patterns of regional-scale environmental drivers is, to date, essential in the implementation of effective conservation strategies. Here, we assessed the species richness S and taxonomic distinctness Δ+ (used a proxy of phylogenetic diversity) of crustacean zooplankton assemblages from [...] Read more.
Resolving the contribution to biodiversity patterns of regional-scale environmental drivers is, to date, essential in the implementation of effective conservation strategies. Here, we assessed the species richness S and taxonomic distinctness Δ+ (used a proxy of phylogenetic diversity) of crustacean zooplankton assemblages from 40 ponds and small lakes located in Albania and North Macedonia and tested whether they could be predicted by waterbodies’ landscape characteristics (area, perimeter, and altitude), together with local bioclimatic conditions that were derived from Wordclim and MODIS databases. The results showed that a minimum adequate model, including the positive effects of non-arboreal vegetation cover and temperature seasonality, together with the negative influence of the mean temperature of the wettest quarter, effectively predicted assemblages’ variation in species richness. In contrast, taxonomic distinctness did not predictably respond to landscape or bioclimatic factors. Noticeably, waterbodies’ area showed a generally low prediction power for both S and Δ+. Additionally, an in-depth analysis of assemblages’ species composition indicated the occurrence of two distinct groups of waterbodies characterized by different species and different precipitation and temperature regimes. Our findings indicated that the classical species-area relationship hypothesis is inadequate in explaining the diversity of crustacean zooplankton assemblages characterizing the waterbodies under analysis. In contrast, local bioclimatic factors might affect the species richness and composition, but not their phylogenetic diversity, the latter likely to be influenced by long-term adaptation mechanisms. Full article
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21 pages, 6374 KiB  
Article
Dynamics of Mesozooplankton Assemblage in Relation to Environmental Factors in the Maryland Coastal Bays
by Efeturi U. Oghenekaro and Paulinus Chigbu
Water 2019, 11(10), 2133; https://doi.org/10.3390/w11102133 - 14 Oct 2019
Cited by 4 | Viewed by 2802
Abstract
The mesozooplankton composition and dynamics in coastal lagoons of Maryland, mid-Atlantic region, USA have received little scientific attention despite the fact that the lagoons have undergone changes in water quality in the past two decades. We compared mesozooplankton abundance and community structure among [...] Read more.
The mesozooplankton composition and dynamics in coastal lagoons of Maryland, mid-Atlantic region, USA have received little scientific attention despite the fact that the lagoons have undergone changes in water quality in the past two decades. We compared mesozooplankton abundance and community structure among sites and seasons, and between 2012, a year of higher than average salinity (33.4), and 2013 with lower than average salinity (26.6). It was observed that the composition, diversity, and abundance of mesozooplankton in 2012 differed from those of 2013. Barnacle nauplii were abundant in 2012 contributing 31% of the non-copepod mesozooplankton abundance, whereas hydromedusae were more dominant in 2013 and contributed up to 83% of non-copepod zooplankton abundance. Gastropod veliger larvae were more abundant in 2013 than in 2012 while larvae of bivalves, polychaetes, and decapods, in addition to cladocerans and ostracods had higher abundances in 2012. The abundance and diversity of mesozooplankton were explained by variations in environmental factors particularly salinity, and by the abundance of predators such as bay anchovy (Anchoa mitchelli). Diversity was higher in spring and summer 2012 (dry year) than in 2013 (wet year). The reduction of salinity in fall 2012, due to high freshwater discharge associated with Hurricane Sandy, was accompanied by a decrease in mesozooplankton diversity. Spatially, diversity was higher at sites with high salinity near the Ocean City Inlet than at sites near the mouth of tributaries with lower salinity, higher nutrient levels and higher phytoplankton biomass. Perhaps, the relatively low salinity and high temperature in 2013 resulted in an increase in the abundance of hydromedusae, which through predation contributed to the reduction in the abundance of bivalve larvae and other taxa. Full article
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13 pages, 4717 KiB  
Article
Cladoceran (Crustacea) Niches, Sex, and Sun Bathing—A Long-Term Record of Tundra Lake (Lapland) Functioning and Paleo-Optics
by Liisa Nevalainen, E. Henriikka Kivilä, Marttiina V. Rantala and Tomi P. Luoto
Water 2019, 11(10), 2008; https://doi.org/10.3390/w11102008 - 27 Sep 2019
Cited by 3 | Viewed by 3300
Abstract
Under fundamental ecosystem changes in high latitude lakes, a functional paleolimnological approach may increase holistic understanding of lake responses and resilience to climate warming. A ~2000-year sediment record from Lake Loažžejávri in the tundra of northern Finnish Lapland was examined for fossil Cladocera [...] Read more.
Under fundamental ecosystem changes in high latitude lakes, a functional paleolimnological approach may increase holistic understanding of lake responses and resilience to climate warming. A ~2000-year sediment record from Lake Loažžejávri in the tundra of northern Finnish Lapland was examined for fossil Cladocera assemblages to examine long-term environmental controls on aquatic communities. In addition, cladoceran functional attributes, including functional diversity (FD), UV absorbance (ABSUV) of Alona carapaces, and sexual reproduction (ephippia) in Bosmina and Chydoridae were analyzed. Cladoceran communities responded to a major change in benthic habitat quality, reflected as elevated (increasingly benthic) sediment organic matter δ13C signal since the 17th century. FD fluctuations showed association with climate oscillation, FD being generally higher during warm climate periods. These ecological changes were likely attributable to diversification of littoral-benthic consumer habitat space. ABSUV, irrespective of increases during the Little Ice Age (LIA) due to higher UV transparency of lake water, was lower under increasing autochthony (benthic production) suggesting establishment of physical UV refugia by the benthic vegetative substrata. Bosmina ephippia exhibited a decreasing trend associated with increasing benthic production, indicating favorable environmental regime, and, together with chydorid ephippia, transient increases during the climate cooling of the LIA driven by shorter open-water season. Full article
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14 pages, 2263 KiB  
Article
Stable Isotope Analysis and Persistent Organic Pollutants in Crustacean Zooplankton: The Role of Size and Seasonality
by Roberta Piscia, Michela Mazzoni, Roberta Bettinetti, Rossana Caroni, Davide Cicala and Marina Marcella Manca
Water 2019, 11(7), 1490; https://doi.org/10.3390/w11071490 - 18 Jul 2019
Cited by 6 | Viewed by 2887
Abstract
Zooplankton is crucial for the transfer of matter, energy, and pollutants through aquatic food webs. Primary and secondary consumers contribute to the abundance and standing stock biomass, which both vary seasonally. By means of taxa- and size-specific carbon and nitrogen stable isotope analysis, [...] Read more.
Zooplankton is crucial for the transfer of matter, energy, and pollutants through aquatic food webs. Primary and secondary consumers contribute to the abundance and standing stock biomass, which both vary seasonally. By means of taxa- and size-specific carbon and nitrogen stable isotope analysis, the path of pollutants through zooplankton is traced and seasonal changes are addressed, in an effort to understand pollutant dynamics in the pelagic food web. We analyzed zooplankton plurennial changes in concentration of polychlorinated biphenyls (PCBs) and dichlorodiphenyltrichloroethane and its relatives (DDTs) and in taxa-specific δ15N signatures in two size fractions, ≥450 µm and ≥850 µm, representative of the major part of zooplankton standing stock biomass and of the fraction to which fish predation is mainly directed, respectively. Our work is aimed at verifying: (1) A link between nitrogen isotopic signatures and pollutant concentrations; (2) the predominance of size versus seasonality for concentration of pollutants; and (3) the contribution of secondary versus primary consumers to carbon and nitrogen isotopic signatures. We found a prevalence of seasonality versus size in pollutant concentrations and isotopic signatures. The taxa-specific δ15N results correlated to pollutant concentrations, by means of taxa contribution to standing stock biomass and δ15N isotopic signatures. This is a step forward to understanding the taxa-specific role in pollutant transfer to planktivores and of zooplankton enrichment in PCBs and DDTs. Full article
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18 pages, 5103 KiB  
Article
Dynamics of Calanus Copepodite Structure during Little Auks’ Breeding Seasons in Two Different Svalbard Locations
by Kaja Balazy, Emilia Trudnowska and Katarzyna Błachowiak-Samołyk
Water 2019, 11(7), 1405; https://doi.org/10.3390/w11071405 - 09 Jul 2019
Cited by 12 | Viewed by 2949
Abstract
Populations dynamics of key zooplankton species in the European Arctic, Calanus finmarchicus and Calanus glacialis (hereafter defined as Calanus) may be sensitive to climate changes, which in turn is of great importance for higher trophic levels. The aim of this study was [...] Read more.
Populations dynamics of key zooplankton species in the European Arctic, Calanus finmarchicus and Calanus glacialis (hereafter defined as Calanus) may be sensitive to climate changes, which in turn is of great importance for higher trophic levels. The aim of this study was to investigate the complete copepodite structure and dynamics of Calanus populations in terms of body size, phenology and their relative role in the zooplankton community over time in different hydrographic conditions (two fjords on the West Spitsbergen Shelf, cold Hornsund vs. warm Kongsfjorden), from the perspective of their planktivorous predator, the little auk. High-resolution zooplankton measurements (taken by nets and a laser optical plankton counter) were adapted to the timing of bird’s breeding in the 2015 and 2016 summer seasons, and to their maximal diving depth (≤50 m). In Hornsund, the share of the Calanus in zooplankton community was greater and the copepodite structure was progressively older over time, matching the little auks timing. The importance of Calanus was much lower in Kongsfjorden, as represented mainly by younger copepodites, presumably due to the Atlantic water advections, thus making this area a less favourable feeding ground. Our results highlight the need for further studies on the match/mismatch between Calanus and little auks, because the observed trend of altered age structure towards a domination of young copepodites and the body size reduction of Calanus associated with higher seawater temperatures may result in insufficient food availability for these seabirds in the future. Full article
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11 pages, 2687 KiB  
Article
Zooplankton Size as a Factor Determining the Food Selectivity of Roach (Rutilus Rutilus) in Water Basin Outlets
by Robert Czerniawski and Tomasz Krepski
Water 2019, 11(6), 1281; https://doi.org/10.3390/w11061281 - 19 Jun 2019
Cited by 4 | Viewed by 2562
Abstract
Fish occurring in the outlets of water basins reduce the abundance of zooplankton. The study was performed at the outlet sections of the lake and waste stabilization pond of a sewage treatment plant. The aim of the study was to determine which zooplankton [...] Read more.
Fish occurring in the outlets of water basins reduce the abundance of zooplankton. The study was performed at the outlet sections of the lake and waste stabilization pond of a sewage treatment plant. The aim of the study was to determine which zooplankton is chosen more often by the roach (Rutilus rutilus), those drifting from the waste stabilization pond or from the lake. The zooplankton from the pond was dominated by Daphnia pulex while zooplankton from the lake was dominated by small planktonic rotifers. We observed that the larger the plankter-victim’s size, the faster the reduction of its number. The fish were more likely to feed on zooplankton drifting from the waste stabilization pond than from the lake. It was influenced by D. pulex individuals, attractive for fish due to their largest body size among the analyzed zooplankton. The significance of riverine zooplankton in the downstream food web may render this data even more important. Full article
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23 pages, 8508 KiB  
Article
The Copepod Acartia tonsa Dana in a Microtidal Mediterranean Lagoon: History of a Successful Invasion
by Elisa Camatti, Marco Pansera and Alessandro Bergamasco
Water 2019, 11(6), 1200; https://doi.org/10.3390/w11061200 - 08 Jun 2019
Cited by 12 | Viewed by 3827
Abstract
The Lagoon of Venice has been recognized as a hot spot for the introduction of nonindigenous species. Several anthropogenic factors as well as environmental stressors concurred to make this ecosystem ideal for invasion. Given the zooplankton ecological relevance related to the role in [...] Read more.
The Lagoon of Venice has been recognized as a hot spot for the introduction of nonindigenous species. Several anthropogenic factors as well as environmental stressors concurred to make this ecosystem ideal for invasion. Given the zooplankton ecological relevance related to the role in the marine trophic network, changes in the community have implications for environmental management and ecosystem services. This work aims to depict the relevant steps of the history of invasion of the copepod Acartia tonsa in the Venice lagoon, providing a recent picture of its distribution, mainly compared to congeneric residents. In this work, four datasets of mesozooplankton were examined. The four datasets covered a period from 1975 to 2017 and were used to investigate temporal trends as well as the changes in coexistence patterns among the Acartia species before and after A. tonsa settlement. Spatial distribution of A. tonsa was found to be significantly associated with temperature, phytoplankton, particulate organic carbon (POC), chlorophyll a, and counter gradient of salinity, confirming that A. tonsa is an opportunistic tolerant species. As for previously dominant species, Paracartia latisetosa almost disappeared, and Acartia margalefi was not completely excluded. In 2014–2017, A. tonsa was found to be the dominant Acartia species in the lagoon. Full article
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15 pages, 3144 KiB  
Article
Effects of Asian Dust and Phosphorus Input on Abundance and Trophic Structure of Protists in the Southern Yellow Sea
by Xi Chen, Guang-Xing Liu, Xiao Huang, Hong-Ju Chen, Chao Zhang and Yang-Guo Zhao
Water 2019, 11(6), 1188; https://doi.org/10.3390/w11061188 - 07 Jun 2019
Cited by 1 | Viewed by 2381
Abstract
To reveal the effects of Asian dust and phosphorus (P) input on the structure and function of micro-food web in the Yellow Sea, an experiment was conducted onboard the southern Yellow Sea where P was deficient. The response of the abundance and trophic [...] Read more.
To reveal the effects of Asian dust and phosphorus (P) input on the structure and function of micro-food web in the Yellow Sea, an experiment was conducted onboard the southern Yellow Sea where P was deficient. The response of the abundance and trophic structure of planktonic protists to different concentrations of dust and P were studied. The results showed that the sand-dust deposition presented variable effects on different sizes of protists during incubation periods. At the initial stage of incubation with dust, the amount of all sizes of autotrophic protists, especially 10–20 µm, were improved; on the contrary, the heterotrophic and mixotrophic protists were inhibited. At the late period, the increase of autotrophic protists was restricted, while the 2–5 µm heterotrophic and mixotrophic protists obviously increased. Similarly, adding P demonstrated the obviously positive effect on the 10–20 µm autotrophic protists at the initial period, and then the growth was restricted at the late period. These results were consistent with that of sand-dust deposition. Hence, it could be presumed that the positive effect of sand-dust deposition on autotrophic protists in the southern Yellow Sea was achieved by the release of P from dust. P in the early stage of sand-dust deposition promotes the growth of large-size autotrophic protists, which may accelerate red tides in eutrophic ocean. The stimulation of small-size heterotrophic protists at the late period of sand-dust deposition contributed to the material cycle and food transmission in the ocean. Therefore, the effects of sand-dust deposition on the abundance and trophic structure of different sizes of planktonic protists could change the structure of micro-food web in the southern Yellow Sea and further affected the ecological function of planktonic protists. Full article
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16 pages, 2524 KiB  
Article
Long-Term Changes in the Zooplankton Community of Lake Maggiore in Response to Multiple Stressors: A Functional Principal Components Analysis
by Andrea Arfè, Piero Quatto, Antonella Zambon, Hugh J. MacIsaac and Marina Manca
Water 2019, 11(5), 962; https://doi.org/10.3390/w11050962 - 08 May 2019
Cited by 9 | Viewed by 3041
Abstract
We describe the long-term (1981–2008) dynamics of several physico-chemical and biological variables and how their changes may have influenced zooplankton structure in Lake Maggiore (Italy). Data was available for the 1981–1992 and 1995–2008 periods. Standardized time-series for temperature and total phosphorus (TP), chlorophyll-a, [...] Read more.
We describe the long-term (1981–2008) dynamics of several physico-chemical and biological variables and how their changes may have influenced zooplankton structure in Lake Maggiore (Italy). Data was available for the 1981–1992 and 1995–2008 periods. Standardized time-series for temperature and total phosphorus (TP), chlorophyll-a, phytoplankton density (cel m−3), and cell size (µm3), as well as zooplankton structure (Copepoda, Cladocera, and Rotifera density, ind m−3) were smoothed using penalized B-splines and analyzed using Functional Principal Components (FPCs) to assess their dominant modes of variation. The first four FPCs explained 55% of 1981–1992 and 65% of 1995–2008 overall variation. Results showed that temperature fluctuated during the study period, particularly during 1988–1992 with a general tendency to increase. TP showed a declining trend with some reversions in the pattern observed in the years 1992, 1999, and 2000. Phytoplankton estimators and chlorophyll-a concentration showed a variable trend along the study period. Zooplankton groups also had a variable trend along the study period with a general increase in density of large carnivorous (mainly Bythotrephes longimanus) and a decrease of large herbivorous (mainly Daphnia), and a similar increase in the ratio of raptorial to microphagous rotifers. Our results suggest that the lake experienced a strong trophic change associated with oligotrophication, followed by pronounced climate-induced changes during the latter period. TP concentration was strongly associated with changes in abundance of some zooplankton taxa. Full article
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16 pages, 3160 KiB  
Article
Response of Cladocera Fauna to Heavy Metal Pollution, Based on Sediments from Subsidence Ponds Downstream of a Mine Discharge (S. Poland)
by Agnieszka Pociecha, Agata Z. Wojtal, Ewa Szarek-Gwiazda, Anna Cieplok, Dariusz Ciszewski and Andrzej Kownacki
Water 2019, 11(4), 810; https://doi.org/10.3390/w11040810 - 18 Apr 2019
Cited by 6 | Viewed by 2871
Abstract
Mining is recognized to deeply influence invertebrate assemblages in aquatic systems, but different invertebrates respond in different ways to mining cessation. Here, we document the response of the cladoceran assemblage of the Chechło river, S. Poland (southern Poland) to the cessation of Pb-Zn [...] Read more.
Mining is recognized to deeply influence invertebrate assemblages in aquatic systems, but different invertebrates respond in different ways to mining cessation. Here, we document the response of the cladoceran assemblage of the Chechło river, S. Poland (southern Poland) to the cessation of Pb-Zn ore mining. The aquatic system includes the river and associated subsidence ponds in the valley. Some ponds were contaminated during the period of mining, which ceased in 2009, while one of the ponds only appeared after mining had stopped. We used Cladocera to reveal how the cessation of mine water discharge reflected on the structure and density of organisms. A total of 20 Cladocera taxa were identified in the sediment of subsidence ponds. Their density ranged from 0 to 109 ind./1 cm3. The concentrations of Zn, Cd, Cu and Pb were much higher in sediments of the ponds formed during peak mining than in the ponds formed after the closure of the mine. Statistical analysis (CCA) showed that Alonella nana, Alona affinis, Alona sp. and Pleuroxus sp. strongly correlated with pond age and did not tolerate high concentrations of heavy metals (Cu and Cd). This analysis indicated that the rate of water exchange by the river flow and the presence of aquatic plants, affect species composition more than pond age itself. Full article
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