On the plurality of Chydorus sphaericus (O. F. Müller) (Cladocera,Chydoridae), and designation of a neotype from Sjaelsø, Denmark

Two readily distinguishable species of Chydorus sphaericus sens. lat. occur in Salmon Lake, Montana, differing from each other in size and shape of rostrum, headpore configuration, frequency of doubling of teeth on the postabdomen, pattern of reticulation of the shell, and morphology of the mature male, especially the postabdomen, postabdominal claws, copulatory hooks, and rostrum. In any such comparisons, individual specimens are not sufficient; populations are needed to sort out morphological characters associated with instar and sex and character's that change with increasing body size. Comparison of population structure and stage of reproduction can be accomplished by a size-frequency distribution and by specific morphological characters that enable the three male instars to be recognized individually.One species, thought at first to be Chydorus herrmanni, is very close to Chydorus sphaericus sens. str. from Denmark. The other taxon with a short, generally blunt labrum, is described as the new species Chydorus brevilabris. Because of morphological diversity among the entities currently listed as Chydorus sphaericus from around the World, it is certain that a complex of species is represented. To facilitate the eventual resolution of this problem, the population in Sjaelsø, Sjaelland, Denmark has arbitrarily been selected as the standard of comparison, and from this population a neotype and several allotypes have been designated. Cursory examination of various available populations suggests that C. sphaericus sens. str. and C. brevilabris represent a major dichotomy in the sphaericus complex, each branch consisting of an unknown number of closely-related species. Cautions are voiced against assuming that Chydorus sphaericus is a single cosmopolitan and ubiquitous species.This study has been aided by grant GB-29725 for the work in Denmark in 1971–72 and by my present grant DEB 76-20237, both from the National Science Foundation. I am indebted to D. F. Brakke for the samples from Salmon Lake and neighboring localities in Montana and information concerning these localities, and to Brenda J. Hann for dissections and drawings of trunk limbs and help in their interpretation. I am also indebted to Geoffrey Fryer, whose innocent inquiry precipitated this study.     

A contribution to the taxonomy ofC. sphaericus sens. lat. (Cladocera,Chydoridae)

Two distinct morphotypes ofChydorus sphaericus are recognised in Ireland. A reference collection of scanning electron micrographs of male and femaleC. sphaericus sens. str. has been compiled. This collection is used to compare the varying morphological features of Danish and Irish material to determine their taxonomic significance. However, no taxonomic decisions can be made about the Irish morphotypes until mature males and ephippial females become available. Comparison of population structures and stages of reproduction are accomplished by size-frequency distribution. This study illustrates the ever increasing value of scanning electron microscopy (SEM) in chydorid taxonomy, where characters such as shell-reticulation features may easily be misinterpreted using only light microscopy.     

Sex in a cyclical parthenogen: mating behaviour of Chydorus sphaericus (Crustacea; Branchiopoda; Anomopoda)

1. We describe the interactions during mating in Chydorus sphaericus, a cyclical parthenogenetic anomopod. Mating behaviour is more complex than previously assumed, with evidence for a diffusible chemical to which males react at the onset of mating, for reproductive isolation, and for postcopulatory mate guarding. 2. During mating, the male and female form a ‘mating cross’ that may be maintained for several hours, while copulation itself typically lasts less than a minute. Furthermore, males invariably attach to the right valve of females. Copulation involves intromission of the postabdomen between the valves, so that the gonopores approach the left ovarium. 3. This behaviour is reflected in the morphology of both sexes: males have a specialised anterior valve margin, postabdomen, first limb and rostrum, under selective pressure for successful mate guarding and copulation, while gamogenetic females have asymmetric ovaries, and a species‐specific setulation of the valves. Males of the structurally related Chydorus ovalis react to the presence of C. sphaericus, but fail to dock to females, suggesting a lock‐antilock element in the reproductive isolation of both species. 4. The morphological and ethological adaptations in C. sphaericus suggest that there is a strong selective pressure on mating behaviour in this cyclical parthenogen and specifically towards the formation of the ‘mating cross’.     

Responses to injured conspecifics and heterospecifics by water fleas,Chydorus sphaericus

Cladocerans have been shown to exhibit a variety of responses to chemical stimuli associated with predation risk, including those from predators and injured conspecifics. In this study, we tested the hypothesis that individuals of Chydorus sphaericus would respond with avoidance behavior to chemical cues from both injured conspecifics and injured heterospecifics. Avoidance behavior was seen in response to cues from C. sphaericus and Daphnia magna. Responses to cues from Daphnia pulex were intermediate between those of the other cues and the water control. We propose that a concentration effect may be responsible for the mixed responses to heterospecific cues and speculate that responses to a broad range of cues may help explain the expansive range of C. sphaericus.     

The response of three chydorid species to temperature, pH and food

The responses of three chydorid species, Chydorus sphaericus(O.F. Müller), Alona affinis (Leydig) and Alonopsis elongata (Sars) to temperature, pH and food type were examined. Egg development time of all species decreased with increasing temperature, although the degree of change was different for each species. C. sphaericus had the fastest development time at all temperatures, and A. elongata the slowest. pH also affected the egg development time of each species differently. A. elongata failed to reproduce at low and medium pH, the egg development time of C. sphaericus was fastest at high pH while that of A. affinis was fastest at low pH. Food type was found to have significant effects on the population growth of individual species. C. sphaericus populations grew equally well in all three food types provided, and grew more than the other two species when fed on an algae culture, and filtered pond water. A. affinisand A. elongata populations grew best when fed on a detritus food source. The responses of each species to the different variables tested are discussed in relation to field observations of their distribution and abundance.     

Spatial and temporal variation of benthic Cladocera (Crustacea) studied with activity traps in Lake Myvatn, Iceland

Benthic Cladocera were studied with a modified type of an activity trap in Lake Myvatn in 1990–1992. After feasibility experiments, the operation time and the distance of the traps from the benthic substrate were adjusted in order to minimize the effects of diurnal variation and the trapping of planktonic organisms. The trap catches of Eurycercus lamellatus were positively correlated with their abundance at the bottom as estimated by grab sampling (r = 0.910, P < 0.001). The usefulness of the activity trap was demonstrated by: (1) a lake-wide survey of the benthic Cladocera; (2) a study of the seasonal variation in the size distribution, abundance and sex ratio of E. lamellatus; and (3) a study of the seasonal succession of Chydorus sphaericus and Alona affinis. The variation of benthic Cladocera among 21 trap sites distributed on a 1–2-km scale across the lake exceeded the within-site variation. The sampling sites could be divided into five main groups based on cluster analysis. Eurycercus lamellatus was the most common species in the mat of filamentous green algae (Cladophorales). Alonella nana dominated the area of spring water inflow in the north basin and Macrothrix hirsuticornis the area of spring water inflow in the southeastern part of the lake. In other parts of the lake either Chydorus sphaericus or Alona quadrangularis tended to dominate. The size distribution and sex ratio of E. lamellatus was followed at two sites through one summer. In early summer most individuals were females less than 1.45 mm long. Around mid-summer they had grown to 0.69–3.1 mm. By the end of August the size distribution had become bimodal, with a large number of small males and a smaller number of much larger females. The seasonal succession in the abundance of E. lamellatus, A. affinis and C. sphaericus was followed at four sites over two seasons. With some exceptions the abundance of a species followed a similar seasonal trajectory on the different stations in any one year. There was, however, a marked difference between the two years (1991 and 1992), probably related to different temperatures.     

Chydorus arcticus n.sp., a new cladoceran crustacean (Chydoridae: Chydorinae) from the North Atlantic arctic and Subarctic areas

Chydorus arcticus n.sp. (Cladocera: Chydoridae: Chydorinae) is described, figured, and differentiated from the closely relatedC. sphaericus (O.F. Müller, 1785). The known distribution of the species is given, and some aspects of speciation of arctic crustaceans are pointed out.     

A Seasonal Quantitative Study of the Littoral Cladocera and Copepoda in a Bog Pond and an Acid Marsh in Newfoundland

A quantitative study was made of the littoral microcrustaceans in a bog pond and an acid marsh located on the Avalon Peninsula, Newfoundland, from May, 1972 to May, 1973. A total of 17 species of Cladocera and 7 species of Copepoda were found in La Manche Marsh, while 21 species of Cladocera and 5 species of Copepoda were collected in Round Pond. The maximum standing stocks were similar in the two localities. The dominant forms in the summer and fall in La Manche Marsh were Cladocera: Acantholeberis curvirostris, Biapertura intermedia, Chydorus sphaericus, and Ilyocryptus spinifer. In the winter the copepod, Macrocyclops albidus, was dominant, while Cyclops varicans rubellus was the most abundant species in the spring. Round Pond also was dominated by Cladocera in the summer and fall; namely, Acroperus alonoides, Alona rustica, Alonella excisa, Chydorus sphaericus, and Sida crystallina. The copepods, Eucyclops agilis and Macrocyclops albidus, were dominant in the winter and early spring.     

Cryptic species within the Chydorus sphaericus species complex (Crustacea: Cladocera) revealed by molecular markers and sexual stage morphology

The cosmopolitanism paradigm in the biogeography of freshwater invertebrates is currently being replaced by non-cosmopolitanism or continental endemism. Benthic water fleas (Cladocera) from the family Chydoridae were the first group of freshwater invertebrates for which non-cosmopolitanism and cryptic diversity was substantiated by morphological studies. Yet, little is known about genetic differentiation and evolutionary history of chydorid species complexes. Here we present the first analysis of the genetic versus morphological differentiation in a benthic cladoceran species complex—Chydorus sphaericus s. str. using sequence variation in a nuclear (ribosomal internal transcribed spacer 2, ITS-2) and a mitochondrial (cytochrome c oxidase subunit I, COI) genes in 50 Holarctic localities. We tested for continental endemism and cryptic diversity predicted by previous morphological studies. We found evidence for the presence of at least seven putative regional species in the Holarctic, at least three of them being distributed beyond a single continent. While the molecular and sexual stage characters showed general concordance on species lineages, parthenogenetic female characters lacked resolution or were unassociated with molecular lineages. We conclude that cryptic regional lineages of benthic cladocerans are apparent and that the sexual stages represent the most informative morphological source of species characters for this environmental indicator group.     

Occurrence of Cladocera (Crustacea) in subterranean waters in Yugoslavia

Contrary to some localities in France and Spain, Cladocera occur in hypogean waters in Yugoslavia very sporadically. Beside the stygobitic species, Alona hercegovinae from caves and Alona smirnovi from interstitial waters, particularly some other Chydoridae seem to be suited for hypogean life. Chydorus sphaericus, reputed as one of the most euryoecious cladocerans, occurs also the deepest in interstitial waters as well as in caves. Other species, found in hypogean waters are Simocephalus vetulus, Ilyocryptus sordidus, Eurycercus lamellatus, Chydorus ovalis, Leydigia leydigi, Acroperus harpae and Biapertura affnis.     

Particle selection over a broad size range by crustacean zooplankton

1. Preferences for particles over a broad size range were assessed in four crustacean zooplankton species under field conditions. The particles employed were six spherical algae with diameters of 2, 6, 13, 19, 38 and 61 μm. 2. Diaphanosoma brachyurum and Daphnia cucullata preferred the two smallest particles, Chydorus sphaericus the three smallest and Eudiaptomus gracilis the 19 μm particle. 3. These size preferences agree well with earlier determinations based on seston particles in the same lake, indicating consistent size preferences by these species over a broad size range.     

The freshwater fauna of Macquarie Island, including a redescription of the endemic water-flea Daphnia gelida (Brady) (Anomopoda: Crustacea)

The freshwater invertebrate fauna (excluding the Protozoa) of Macquarie Island is collated. This includes two Platyhelminthes, two Gastrotricha, three Tardigrada, 41 rotifera, at least eight Nematoda, nine Annelida, and 21 Arthropoda. The latter comprises six species of Anomopoda, two Copepoda, two Ostracoda, an Isopoda, five Acarina, and at least five species of Insecta with aquatic or semi-aquatic larvae. The freshwater Anomopoda (Cladocera) of Macquarie Island are re-evaluated, six species are now recognized and the largest, identified as Daphnia gelida (Brady), is redescribed. The records of both Alona weinecki Studer and Pleuroxus macquariensis Frey are confirmed, while that of Macrothrix hirsuticornis Norman and Brady is accepted with some reservations. Alona quadrangularis (O.F. Müller) is re-instated and records of Chydorus sphaericus O.F. Müller ascribed to C. patagonicus Ekman.     

Resource-niche complementarity and autotrophic compensation determines ecosystem-level responses to increased cladoceran species richness

This study examines the relationship between cladoceran species richness and ecosystem functioning. I conducted an experiment in which four cladocerans, Daphnia. magna, D. longispina, D. pulex and Chydorus sphaericus, were cultured in microcosms using different species combinations and levels of species richness. The results demonstrate that even within this closely related group of organisms the effects on ecosystem-level variables, such as total algae and zooplankton biomass, per capita productivity, and nutrient concentrations, as well as phytoplankton community structure, were highly variable between different combinations of these species. Since only four species where involved in this study, species-specific effects dominated the general relationship between species richness and ecosystem functioning. Particular combinations of species resulted in effects that indicated more efficient grazing. These effects, which were most pronounced in combinations including both D. magna and C. sphaericus, were manifested as an indirect effect as the prey community shifted towards grazing-resistant species. As a result, the productivity of the prey community decreased, because phytoplankton species with lower per capita productivity became more dominant. I suggest that the primary mechanism that caused this significant effect was complementarity in prey-size use of D. magna and C. sphaericus. In terms of prey-size range, D. pulex and D. longispina were redundant when D. magna was present and were quickly out-competed by the latter despite higher per capita filtering efficiency. The results show that different mechanisms are important for different combinations of species. Furthermore, the ability of the prey community to respond to changes of consumer species composition is an important factor in experiments in which consumer species richness is experimentally manipulated. Received: 2 November 1998 / Accepted: 8 September 1999     

The impact of three-spined sticklebacks on the Zooplankton of a previously fish-free pool

SUMMARY.

• 1 In the Camargue (southern France), most invertebrates inhabiting temporary shallow oligohaline marshes and pools are large-bodied, conspicuous and defenceless taxa that are restricted to fish-free environments. Management practices, by creating drainage canals, strongly affect the invertebrate community by allowing colonization by fish.
• 2 Sticklebacks colonizing the study pool were sampled daily and Zooplankton sampling was carried out at weekly intervals. From early October until the pool dried in June, 1365 sticklebacks entered and only 139 stickleback's fry and seven adults left the pool. Zooplankton abundance was high during the whole study period with a mean density of 1360 ind. 100cm^?2. Larger forms (Daphnia magna, Simocephalus vetulus, Arctodiaptomus wierzejskii) quickly disappeared after the first wave of sticklebacks entered the pool in November. Subsequently, smaller species (Diacyclops bicuspidatus odessanus, Chydorus sphaericus and ostracods) replaced the larger ones until the pool dried.
• 3 Food selectivity of sticklebacks and daily Zooplankton consumption were investigated under experimental conditions within the pool in spring. After introduction of sticklebacks, densities of Zooplankton species dropped quickly (D. b. odessanus, C. sphaericus) with the exception of Ostracoda. Estimates of daily Zooplankton consumption by adult sticklebacks varied between 2.8 and 10.7% of fish body weight. Examination of gut contents showed that fry mainly ingested Chydorus sphaericus.
• 4 Given that the stickleback is an annual species in the Camargue, and that few fry are able to leave the pool due to the early drying out, the colonization of such a former fish-free pool is not beneficial for the stickleback population. However, a more serious effect of such colonization is the progressive local extinction of the most typical Zooplankton species inhabiting these formerly fish-free environments.

     

Zooplankton (Cladocera, Ostracoda), Chironomidae and other benthic faunal remains in sediment cores from nine North African wetland lakes: the CASSARINA Project

Palaeolimnological studies were carried out on sediment cores from nine North African wetland lakes. The lakes represented a variety of habitats ranging from freshwater to mixohaline conditions and with and without direct connections to the sea. Sediment cores were examined for records of recent environmental change during the 20th century period. Faunal remains analysed in the sediments included those of Cladocera, Ostracoda, Chironomidae, Mollusca and, at the sites with marine connections, Foraminifera. Cyprideis torosa littoralis was the most common ostracod, occurring in sediments from the seven sites linked with the sea and also in brackish water Sidi Bou Rhaba. In acid Megene Chitane ostracods were scarce, being represented by a single species (Cypria ophtalmica). Candona neglecta completely disappeared at two sites (Sidi Bou Rhaba and Bokka) during the 20th century. Of the Cladocera, Chydorus sphaericus was common in the moderately mixohaline sites and but Heterocypris salina occurred only where marine salinities were occasionally achieved (in Zerga, Ichkeul and Korba). Microinvertebrate assemblages in the Nile Delta lake cores and to a less extent in Zerga showed a clear response indicative of more fresher conditions occurring during the latter part of the 20th century. However the freshening trend began prior to construction of the Aswan High Dam (mid 1960s).With the exception of the most saline site (Korba), microinvertebrate communiuties at all the CASSARINA sites have experienced major disturbances during the 20th century. Changes in freshwater availability associated with increased human usage of water resources is perceived as a major factor regulating the abundance and occurrence of aquatic microinvertebrate species at these sites     

The taxonomy and biogeography of the Cladocera

For a variety of reasons, including the analysis of a number of taxa having the same names on different continents, we have concluded overwhelmingly that the chydorid Cladocera are not cosmopolitan in distribution but instead are restricted to smaller regions by their specific ecological requirements for habitat type and also by long-term events in earth history. Recent study ofChydorus faviformis and species resembling it indicates there has been no effective exchange of genetic material between North America and South America, nor between Australia and Asia, nor even between China, Malaysia, and India in southern Asia. Moreover, the patterns of distribution are even narrower than this, as in North America, for example, taxa having the same names in the southern states as in the northern states are differentiated at the species level in some instances, possibly in most. Southern species push northward along the Atlantic Coast for varying distances, one species having reached Nova Scotia and Newfoundland probably during the warm interval in mid-Postglacial time. Thus, when species are studied closely to define their morphological limits, cosmopolitanism disappears, and patterns of distribution emerge that are very similar to those of other animals and plants. The ‘species’ that have been claimed to be cosmopolitan are being shown to be groups or complexes of morphologically similar species instead, each member species of which has a much more restricted distribution than the group or complex as a whole. To explain how the different continents can have such similar lists of ‘species’ without intercontinental dispersal of resting eggs occurring almost continuously, we are suggesting plate tectonics and the drifting of continents, either apart or together.     

Headshields of ephippial Chydorus piger Sars (Cladocera, Chydoridae) females from northern Finnish Lapland: a long period of gamogenesis?

Numerous headshields of ephippial Chydorus piger (currently called Paralona pigra) females were found from the Holocene sediments of Aitajärvi Lake, northern Finnish Lapland (69° 08′ N, 27° 14′ E). Headshields of ephippial females of the genus Chydorus s.l. can be easily identified since their posterior margin is elongated and pointed. According to analysis of fossil cladoceran remains, Chydorus piger lived in the lake throughout the Holocene but was never very common (less than 10% of preserved cladoceran remains). Headshields of ephippial females were found in most samples and they comprised ca 10–30% of all Chydorus piger headshields. No similar phenomenon was met in Chydorus cf. sphaericus which is common up to the most severe arctic conditions. Aitajärvi is situated north of the modern distribution area of Chydorus piger. I suggest that this species reacted to the stress of harsh climatic conditions outside of its distribution area by being gamogenetic during most of the ice-free period. The ephippial headshields could be valuable indicators of severe climatic conditions in pre-Holocene sedimentary environments.     

Long-term change of the littoral Cladocera in the Tatra Mountain lakes through a major acidification event

In our study, we focused on littoral Cladocera living and feeding in shallow shore parts of 46 mountain lakes in the Tatra Mountains (Slovakia and Poland). The studied lakes underwent a major acidification event in the 1980s and are now in the process of recovery. Lakes were divided into three categories based on their sensitivity to acidification: 5 extremely sensitive (ES), 11 acid sensitive (AS), and 30 non-sensitive (NS) lakes. In our study, we included historical data from the literature, and data from sediment core and littoral samples, which together represent the evolution of the littoral communities from a pre-industrial period up to the present. In total, 11 littoral species were found belonging to three cladoceran families. Most of the species were members of the family Chydoridae: Alona affinis, A. quadrangularis, A. rectangula, A. guttata, Acroperus harpae, Alonella excisa, A. nana, Chydorus sphaericus, and Eurycercus lamellatus. One species belonged to each family Daphniidae (Ceriodaphnia quadrangula) and Polyphemidae (Polyphemus pediculus). The most numerous littoral taxa were Alona affinis, Acroperus harpae, and Chydorus sphaericus. All species reacted to decreased pH levels during peak acidification in the 1980s by disappearing from most of the lakes of all categories; the only persisting species was Chydorus sphaericus. Most species returned to the lakes when pH started to increase in the 1990s, although their return was noticeably slower in AS lakes. Alona quadrangularis decreased its distribution range over the studied period; Polyphemus pediculus was mostly detected in the 1910s only. The number of species was highest in all lake categories when dwarf pine was present in the lake catchment. On the whole, the littoral community was richest in NS lakes.     

Relocation of Chydorus barroisi and related species (Cladocera,Chydoridae) to a new genus and description of two new species

Five taxa already in the literature are here removed from Chydorus to their own genus Ephemeroporus, and two new species — E. acanthodes and E. archboldi — are described, with E. acanthodes being designated the type species of the genus. These taxa, plus at least nine undescribed species and others undoubtedly waiting to be sorted out, constitute a tightly circumscribed group of species morphologically. The first two species described — E. barroisi and E. poppei — are nomina dubia for the present, as no specimens exist from the original collections, nor are any available from the type localities or reasonably close thereto. E. hybridus from Brazil has been characterized in greater detail through the availability of specimens from the type series, which has enabled one of the species in the E. hybridus group from North America to be judged conspecific with reasonable certainty. E. tridentatus, from Brazil, has been restored as a valid species, and the highly distinctive E. phintonicus from Sardinia and Algeria constitutes the seventh species in the genus. Chydorus nitidulus and Chydorus tilhoi, which have been suggested to be members of the barroisi complex, are not. What are presently called E. barroisi and E. hybridus, except for E. hybridus, sens. str., each consists of a cluster of species sharing the same number of teeth on the labrum and shell. Because of their wide, distribution, abundance, and frequency of occurrence, especially in South Asia, the species in the E. barroisi group will be especially meaningful to sort out.     

Cladocera response to Late Glacial to Early Holocene climate change in a South Carpathian mountain lake

This study explores changes in cladoceran composition in a high mountain lake of the Retezat (Lake Brazi), the South Carpathian Mountains of Romania, during the Late Glacial–Early Holocene (14,500–11,600 cal. yr. bp) transition using a paleolimnological approach. The lake had a species poor cladoceran community throughout this period. Daphnia longispina, Chydorus sphaericus and Alona affinis were the most common, showing marked fluctuations in their relative abundances through time. Distinct faunal response to warming at the Younger Dryas (YD)/Preboreal transition was recorded by increasing fossil densities and distinct community composition change: Alona affinis became dominant while numbers of Chydorus sphaericus dramatically decreased. In the Early Holocene, the productivity of Lake Brazi seem to have increased as reflected by higher numbers of Cladocera due to appearance of new species (Alona rectangula, A. quadrangularis and A. guttata) which are common in productive waters. Significant negative correlation was found between average dorsal length of daphnid ephippia and the NGRIP ^δ18O isotope values. Given the absence of fish predation, changes in Daphnia ephippia size were taken to indicate climatic change: larger ephippium size inferred cold conditions during the Late Glacial, while smaller size reflected climate warming during the Early Holocene. We conclude that Cladocera fossils are good indicators of climatic change that happened during the transition from the Late Glacial to the Holocene. We found that climatic conditions can be tracked either by size distribution of Daphnia ephippia (larger ephippium size under colder climate) and/or by community change of cladocerans.