Models of ontogenetic allometry in cladoceran morphology studies

Ontogenetic variation of morphological traits in cladocerans is usually analysed by a simple allometry function. We found that common cyclomorphic and inducible traits in Bosmina and Daphnia sometimes deviate from simple allometry, however. Consequently, simple allometric analysis may at times be inadequate since important aspects of the ontogenetic development may not be revealed. We suggest that other allometric models should be used more extensively in ontogenetic studies of cladocerans morphology. Especially, complex allometric functions may help analyse the effects of size-selective predation on morphological defence traits when predators prefer prey of intermediate size. Guest editor: Piet Spaak Cladocera: Proceedings of the 7th International Symposium on Cladocera     

Transitions in functional morphology from “large branchiopods” to Cladocera: Video and confocal microscopic studies of Cyclestheria hislopi (Cyclestherida) and Sida crystallina (Cladocera: Ctenopoda)

Great diversity is found in morphology and functionality of arthropod appendages, both along the body axis of individual animals and between different life-cycle stages. Despite many branchiopod crustaceans being well known for displaying a relatively simple arrangement of many serially post-maxillary appendages (trunk limbs), this taxon also shows an often unappreciated large variation in appendage morphology. Diplostracan branchiopods exhibit generally a division of labor into locomotory antennae and feeding/filtratory post-maxillary appendages (trunk limbs). We here study the functionality and morphology of the swimming antennae and feeding appendages in clam shrimps and cladocerans and analyze the findings in an evolutionary context (e.g., possible progenetic origin of Cladocera). We focus on Cyclestheria hislopi (Cyclestherida), sister species to Cladocera and exhibiting many “large” branchiopod characters (e.g., many serially similar appendages), and Sida crystallina (Cladocera, Ctenopoda), which likely exhibits plesiomorphic cladoceran traits (e.g., six pairs of serially similar appendages). We combine (semi-)high-speed recordings of behavior with confocal laser scanning microscopy analyses of musculature to infer functionality and homologies of locomotory and filtratory appendages in the two groups. Our morphological study shows that the musculature in all trunk limbs (irrespective of limb size) of both C. hislopi and S. crystallina comprises overall similar muscle groups in largely corresponding arrangements. Some differences between C. hislopi and S. crystallina, such as fewer trunk limbs and antennal segments in the latter, may reflect a progenetic origin of Cladocera. Other differences seem related to the appearance of a specialized type of swimming and feeding in Cladocera, where the anterior locomotory system (antennae) and the posterior feeding system (trunk limbs) have become fully separated functionally from each other. This separation is likely one explanation for the omnipresence of cladocerans, which have conquered both freshwater and marine free water masses and a number of other habitats.     

浙江两类饮用水源地浮游动物种类组成及其类群相关性分析

于2009年7月至2010年4月,在浙江省选取了10个水库和10个河网饮用水源地,进行4个季节的浮游动物采样调查。记录浮游动物101种(轮虫60,枝角类24和桡足类17种)。水库和河网轮虫的优势种分别为角突臂尾轮虫(Brachionus angularis)和针簇多肢轮虫(Polyarthratrigla),枝角类分别为颈沟基合溞(Bosminopsis deitersi)和长额象鼻溞(Bosmina longirostris),桡足类均为温剑水蚤(Thermocyclops)。水库中有一些能指示良好生态或水质的指示性种类。水库枝角类与桡足类群落之间周年在丰度和生物量上均有极显著的正相关关系(P<0.01),轮虫与桡足类类群之间的相关性最弱;河网枝角类与桡足类在夏、秋、冬季有显著的正相关性(P<0.05)。浮游动物类群之间的相关性程度,以水库的秋季与河网的夏季为最高。轮虫与浮游甲壳动物的种类丰富度随着饮用水源水体透明度的变化呈现相反的变化趋势。     

Laboratory studies on the life cycle of Leydigia acanthocercoides Fisher (1854) (Cladocera: Chydoridae)

The biology of Leydigia acanthocercoides has been studied under laboratory conditions with reference to longevity, instar duration, growth, fecundity and embryonic development at a temperature range of 28–30 °C. It has three preadult and thirteen adult instars. Under the given laboratory conditions this species produces 20 eggs during a life span of 23 days. The number of eggs produced is uniformly constant in all adult instars. The growth rate seems to be exponential in the early phase of the life cycle as in other Cladocera. The general pattern of embryonic development of L. acanthocercoides is similar to those of other tropical cladocerans though differences in the duration of total developmental period have been recorded.Part of Ph.D. thesis.     

Larval and post-larval development of the branchiopod clam shrimp Cyclestheria hislopi (Baird, 1859) (Crustacea, Branchiopoda, Conchostraca, Spinicaudata)

The larval and post-larval development of Cyclestheria hislopi is examined by SEM. There are at least nine stages (excluding the adult) – six larval and three post-larval stages. The first four stages are passed within the egg-membrane. The larval and the post-larval phase are separated by a profound change in morphology that takes place between stages VI and VII. The larva shifts from a dorso-ventrally flattened 'larval' appearance up to stage VI to a laterally flattened, more 'adult' appearance from stage VII. New morphological data have been revealed by this study, including (1) a large and globular larval dorsal organ; (2) the carapace starts its development from the segments of the first and second maxillae; (3) the anterior ramus of the second antenna in adult Cyclestheria hislopi is the endopod, and the posterior ramus the exopod. Direct development of the brood in Cyclestheria hislopi – unique among conchostracans – is compared with that of the Cladocera. If Cyclestheria is the sister group to the Cladocera, as favoured in this work, the classical neoteny theory of the Cladocera must be reconsidered, as there is no particular similarity between any adults of the Cladocera and any of the larval stages of Cyclestheria . It is suggested that Cyclestheria displays the type of development present in a cladoceran ancestor. A comparison between Cyclestheria and the Upper Cambrian 'Orsten' fossil Rehbachiella kinnekullensis reveals a remarkable similarity in the endite morphology of the trunk limbs.     

Patterns in the Distribution of Cladocerans (Crustacea: Branchiopoda) in Lakes Across a North–south Transect in Alaska, USA

The remains of cladocerans were examined from the surface sediments of 51 freshwater sites along a north–south transect spanning Alaska. We identified 27 cladoceran taxa from the sediments, consisting primarily of littoral chydorid species. Variations in cladoceran assemblages were related to measured physical and chemical variables using multivariate techniques. Redundancy analysis (RDA) indicated that lake depth, total phosphorus (TP), and altitude all had a significant influence in determining the composition of cladoceran assemblages. Cladoceran communities in tundra and forest-tundra lakes, which were relatively shallow and nutrient-poor, had relatively low abundances of pelagic Cladocera, and were primarily composed of several littoral chydorid species. Among pelagic cladoceran species, there was a distinct shift in dominance from the Bosminidae in lakes in the southern boreal forest region to Daphniidae in lakes in the northern boreal forest. Daphnia dominated lakes had significantly higher total phosphorus, specific conductivity, and calcium concentrations than lakes dominated by Eubosmina. Overall, the relative importance of physical and chemical factors in structuring cladocerans is similar to other previously studied regions, and suggests the Cladocera may be useful as ecological and paleoenvironmental indicators in this region.     

Predation pressure induced by seasonal fishing moratorium changes the dynamics of subtropical Cladocera populations

Seasonal fishing moratorium with selective fish harvesting is a common means of increasing fish production in Asia. Such manipulation might be expected to lead to a seasonal variation in abundance and composition of fish and predation pressure on zooplankton, and consequently a seasonal change in the Cladocera populations. In this study, we examined the seasonal dynamics of cladocerans in a subtropical plateau lake in southwestern China, Lake Erh, in which a fishing moratorium was implemented in spring and early summer. Cladocerans showed a bimodal seasonal distribution linked to variations in predation pressure induced by the seasonal fishing moratorium. When predation was low in winter because of cool water temperatures and decreasing fish density, Daphnia galeata appeared in the water column and increased towards spring. The cladoceran peaked in the middle of the fishing moratorium, coinciding with a clear water phase. The feeding rate increased with temperature in the late moratorium, allowing planktivorous fish to strongly suppress large daphnids. Small cladocerans benefited from the reduced competition with large daphnids by increasing their numbers rapidly, which resulted in a second peak of cladocerans in the fishing season.     

On the ontogeny of Leptodora kindtii (Crustacea,Branchiopoda, Cladocera), with notes on the phylogeny of the Cladocera

Leptodora kindtii, a large predaceous cladoceran, is among the most deviant species of the Cladocera. Therefore, its phylogenetic position has traditionally proven difficult to determine. Its many peculiar features include, among others, long, stenopodous, forwardly directed trunk limbs, a posteriorly placed dorsal brood pouch, a tri-lobed lower lip, and a long, segmented abdomen. This study describes the ontogeny of L. kindtii (Haplopoda), including general body proportions, appendages, the carapace, and other external structures in an attempt to facilitate the comparison of its aberrant morphology to that of other branchiopods. In general, the early embryos are similar to the early embryos of other cladoceran taxa with respect to body shape and size and position and orientation of the early limb buds. Many of the unusual features of L. kindtii appear late in ontogeny. The carapace appears at an early stage as a pair of dorsolateral swellings in a position corresponding to the gap between the mandibles and the first pair of trunk limbs; it later becomes posteriorly transposed by a gradual fusion of its more anterior parts to the dorsal side of the thorax. The tri-lobed "lower lip," under the labrum of the late embryo and the adult, develops as a fusion of the first maxillae (lateral lobes) to an elevated sternal region behind the mouth (median lobe). The stenopodous, segmented trunk limbs in the adult develop from embryonic, elongate, subdivided limb buds, similar to those seen in early stages of other branchiopods. Two conflicting possibilities for the phylogeny of the Cladocera, involving two different positions of L. kindtii (Haplopoda), are discussed. Several characters support a sister-group relationship between the Haplopoda and Onychopoda. However, some characters support the Anomopoda and Onychopoda as sister groups, leaving the Haplopoda outside this clade. In contrast to recent suggestions, we prefer to retain the term "Cladocera" in its original sense as comprising the Haplopoda, Ctenopoda, Anomopoda, and Onychopoda.     

Holoplankton of ship ballast water in the Port of Vladivostok

Ballast water samples were taken from three ships that arrived at the Port of Vladivostok (Peter the Great Bay, Sea of Japan) from ports of Japan (Sea of Japan and Pacific Ocean) and China (Yellow Sea and Yangtze River). The holoplankton in samples was presented by seven taxonomic groups, among which copepods (subclass Copepoda, 33 species) and cladocerans (subclass Cladocera, 5 species) dominated. In the samples, eight nonindigenous copepod species were revealed. The information contained in this paper may be of importance for the assessment of the risk of subsequent invasion of new species and the development of techniques for monitoring the qualitative and quantitative structures of ballast water.     

Cladocera and Anostraca from the Interior Plateau of British Columbia,Canada, as paleolimnological indicators of salinity and lake level

Cladoceran and anostracan species assemblages were identified from the surface sediments of 33 closed–basin lakes from the southern Interior Plateau of B.C. in order to explore their effectiveness as quantitative indicators of lakewater salinity and ionic composition. These lakes were chosen to maximize the range of lakewater salinity concentrations (freshwater through hypersaline) as well as brine composition (sulphate and carbonate dominated systems). The distribution of the anostracans and cladocerans were strongly correlated with lakewater salinity, ionic composition and lake depth. Based on these strong relationships significant predictive models were developed, using weighted-averaging techniques, to infer lakewater salinity based on the species composition of anostracans and cladocerans in surface sediments. Furthermore, models were developed to infer lake depth that are superior to previously used techniques based on the ratio of planktonic/littoral Cladocera. Given that the species composition of anostracans and cladocerans can be used to infer changes in salinity and lake level, and that their remains can be identified from sedimentary profiles, there is considerable potential in using their assemblages as paleolimnological indicators of past climatic conditions. This revised version was published online in July 2006 with corrections to the Cover Date.     

Interrelations of rotifers with predatory and herbivorous Cladocera: a review of Russian works

Publications in Russian on the influence of epibiontic rotifers on cladocerans as well as on predator-prey interrelations between cladocerans and rotifers are reviewed. Proales daphnicola and Brachionus rubens are common epibionts of Cladocera. The biology of these species is described including the choice of host, feeding, reproduction and impact on the host. Representatives of the families Daphniidae and Moinidae are most readily colonized. At high densities of epibiontic rotifers, a high percentage of young Cladocera die. Predators consuming cladocerans (mainly Bosminidae) belong to the family Asplanchnidae. Rotifers are consumed by the predatory cladocerans Leptodora and Polyphemus. The results of functional response experiments by the author with P. pediculus feeding on Synchaeta pectinata, Asplanchna priodonta, colonial and solitary Conochilus unicornis, and Platyias patulus are given. In the range of prey densities of 100–1600 1^–1 a functional response was found in all the rotifers except P. patulus. Colonies of C. unicornis were not consumed. The highest level of feeding rate saturation was observed in Synchaeta pectinata. Various adaptations in prey morphology prevented effective predation: coloniality, large size of the prey and hard lorica with spines.     

The role of solid waste materials as habitats for macroinvertebrates in a lowland dam reservoir

Eight hypereutrophic phytoplankton dominated ponds from the Brussels Capital Region (Belgium) were biomanipulated (emptied with fish removal) to restore their ecological quality and reduce the risk of cyanobacterial bloom formation. Continuous monitoring of the ponds before and after the biomanipulation allowed the effects of the management intervention on different compartments of pond ecosystems (phytoplankton, zooplankton, submerged vegetation and nutrients) to be assessed. Fish removal resulted in a drastic reduction in phytoplankton biomass and a shift to the clear-water state in seven out of eight biomanipulated ponds. The reduction in phytoplankton biomass was associated with a marked increase in density and size of large cladocerans in six ponds and a restoration of submerged macrophytes in five ponds. The phytoplankton biomass in the ponds with extensive stands of submerged macrophytes was less affected by planktivorous fish recolonisation of some of the ponds later in the summer. The two non-vegetated ponds as well as one pond with sparse submerged vegetation showed a marked increase in phytoplankton biomass associated with the appearance of fish. Phytoplankton biomass increase coincided with the decrease in large Cladocera density and size. One pond lacking submerged macrophytes could maintain very low phytoplankton biomass owing to large Cladocera grazing alone. The results of this study confirmed the importance of large zooplankton grazing and revegetation with submerged macrophytes for the maintenance of the clear-water state and restoration success in hypereutrophic ponds. They also showed that large Cladocera size is more important than their number for efficient phytoplankton control and when cladocerans are large enough, they can considerably restrain phytoplankton growth, including bloom-forming cyanobacteria, even when submerged vegetation is not restored. The positive result of fish removal in seven out of eight biomanipulated ponds clearly indicated that such management intervention can be used, at least, for the short-term restoration of ecological water quality and prevention of noxious cyanobacterial bloom formation. The negative result of biomanipulation in one pond seems to be related to the pollution by sewage water. Guest editors: B. Oertli, R. Cereghino, A. Hull & R. Miracle Pond Conservation: From Science to Practice. 3rd Conference of the European Pond Conservation Network, Valencia, Spain, 14–16 May 2008     

Zooplankton community changes in Lake Kinneret (Israel) during 1969–1985

Long-term records (1969–1985) of zooplankton density in Lake Kinneret indicated significant reduction of biomass (Copepods, Cladocera, Rotifera) and production (Copepods, Cladocera). Nauplius and adult copepod densities decreased but those of copepodites did not change. Mesocyclops was suppressed more than the smaller Thermocyclops and males of both genera became more abundant relative to the larger females. Ratios of small/large Cladocera densities became higher. Numbers of total cladocerans were stable, and therefore reduction of Cladocera grazing capacity is assumed. The abundant Keratella spp. were reduced. It is likely that intensification of fish visual-attack-predation pressure shifted the size-class structure towards smaller adult copepods and cladocerans. Reduction of Keratella spp. and copepod nauplii was probably affected by increasing pressure of fish filter-feeders. Data on fish food consumption, feeding behaviour and fisheries management suggested their direct impact on long-term changes of zooplankton in Lake Kinneret.     

Composition,size, and biomass of zooplankton in large productive Florida lakes

Crustacean zooplankton data were compiled from long-term observational studies at seven large shallow Florida lakes, to determine whether there are general characteristics in regard to species composition, body size, and biomass. In particular, we examined whether patterns in body size and species richness fit empirical models developed by Stanley Dodson. The lakes included range in size from 125 to 1730 km² and encompass mesotrophic to hyper-eutrophic conditions. We found that zooplankton biomass was strongly dominated by one species of calanoid copepod—Arctodiaptomus dorsalis. Large daphnids were absent, and Cladocera assemblages were dominated by small taxa such as Ceriodaphnia, Chydorus, and Eubosmina. The total number of species of pelagic cladocerans (8–12) was consistent with Dodson’s predictions based on lake area. The average size of crustacean zooplankton in Florida lakes is small in comparison with temperate communities. A. dorsalis is the smallest calanoid copepod in North America, and the mean length of Cladocera (0.6 mm) is consistent with Dodson’s results that size decreases from temperate to tropical zones. Total biomass of crustacean zooplankton was very low, ratios of zooplankton to phytoplankton biomass (0.01–0.1) are among the lowest reported in the literature, and the zooplankton displayed short-lasting early spring peaks in biomass. Cladocera were almost entirely absent in spring and summer. Factors known to occur in Florida lakes, which appear to explain these characteristics of biomass, include intense fish predation and high summer water temperature.     

Regulation of blue-green algal buoyancy and bloom formation by light,inorganic nitrogen,C02, and trophic level interactions

In highly eutrophic ponds, buoyancy of the gas-vacuolate blue-green alga Anabaenopsis Elenkinii (Miller) was regulated by complex interactions between chemical and physical parameters, as well as by biological interactions between various trophic levels. Algal buoyancy and surface bloom formation were enhanced markedly by decreased light intensity, and to a lesser extent by decreased CO₂ availability and increased availability of inorganic nitrogen. In the absence of dense populations of large-bodied Cladocera, early season blooms of diatoms and green algae reduced light availability in the ponds thus creating conditions favorable for increased buoyancy and bloom formation by A. Elenkinii. The appearance of blue-green algal blooms could be prevented by a reduced density of planktivorous fish, which allowed development of dense cladoceran populations. The cladocerans limited the growth of precursory blooms of diatoms and green algae, and given the resulting clear-water conditions, buoyancy of A. Elenkinii was reduced, and blue-green algal blooms never appeared.     

Influence of Chaoborus density on the effects of an insecticide on zooplankton communities in ponds

Zooplankton communities with a high or low density of Chaoborus larvae were established in outdoor concrete ponds, to which a carbamate insecticide, carbaryl, was applied at 0.1 or 0.5 mg l⁻¹. The lower concentration of the chemical was harmful only to Cladocera. The higher concentration damaged Chaoborus, Copepoda, and some rotifer species, as well as Cladocera. In the ponds with a low density of Chaoborus, chemical application altered the cladoceran community from dominance by Daphnia to that by Bosmina and Moina. In the ponds with a high density of Chaoborus, Chaoborus excluded cladocerans from the zooplankton community presumably by predation, and supported the dominance of rotifers. Cladocera did not recover after application of the chemical, even when Chaoborus was eliminated by the higher concentration of chemical. The relatively rapid recovery of Chaoborus seemed to interrupt the recovery of Cladocera.     

Changes in histone acetylation are associated with differences in accessibility of V(H) gene segments to V-DJ recombination during B-cell ontogeny and development

Although V(D)J recombination is thought to be regulated by changes in the accessibility of chromatin to the recombinase machinery, the mechanisms responsible for establishing "open" chromatin are poorly understood. We performed a detailed study of the acetylation status of histones associated with 11 V(H) gene segments, their flanking regions, and various intergenic elements during B-cell development and ontogeny, when V(D)J recombination is highly regulated. Histone H4 shows higher and more-regulated acetylation than does histone H3 in the V(H) locus. In adult pro-B cells, V(H) gene segments are acetylated prior to V(D)J rearrangement, with higher acetylation associated with J(H)-distal V(H) gene segments. While large regions of the V(H) locus have similar patterns of histone acetylation, acetylation is narrowly confined to the gene segments, their flanking promoters, and recombinase signal sequence elements. Thus, histone acetylation in the V(H) locus is both locally and globally regulated. Increased histone acetylation accompanies preferential recombination of J(H)-proximal V(H) gene segments in early B-cell ontogeny, and decreased histone acetylation accompanies inhibition of V-DJ recombination in a transgenic model of immunoglobulin heavy-chain allelic exclusion. Thus, changes in histone acetylation appear to be important for both promotion and inhibition of V-DJ rearrangement during B-cell ontogeny and development.     

The behavioural basis of prey selection by underyearling bream (Abramis brama (L.)) and roach (Rutilus rutilus (L.))

SUMMARY. A clear difference in the ability lo escape from fish predators exists between members of the Cladocera and Copepoda. The results of our laboratory studies have shown that underyearling roach and bream both found copepods more difficult to capture than cladocerans. However, bream were far more efficient than roach at catching the more elusive copepod prey. The basis for this difference was the greater strike ability of bream, most likely related to its more protrusible mouth. In their natural environment the two species of fish exhibited food resource partitioning with planktonic Cladocera predominating in roach guts and copepods and non-planktonic Cladocera composing the vast majority of the gut contents of bream. Differences in diet are partly due to their contrasting attack abilities. Additional variation may arise because of subtle differences in the timing and location of foraging.     

The evolutionary importance of behavioural development: the ontogeny and phylogeny of bird song

It has been proposed that ontogeny may be an important constraint on the evolution of morphological traits. In this paper, ontogeny is discussed as a possible constraint on behavioural evolution. A literature review of the development of song in oscine birds (Passeriformes; Passeres) shows that song development follows von Baer's law; that is, that development proceeds from early, generally distributed stages to later, specialized stages. Song ontogeny is found to parallel phylogeny in a lineage of sparrows (Emberizidae; Emberizinae). Possible cases of paedomorphosis (slowing down or truncation of development) in bird song are discussed, specifically in the family Mimidae and in the reed warbler, Acrocephalus palustris (Muscicapidae; Sylviinae). Finally, the implications of developmental constraints for the study of the adaptive significance of behaviour are discussed.     

Pelagic cladocera in upper volga water reservoirs

The results of cladoceran crustaceans studies in the pelagial of the Ivankovo and Uglich water reservoirs were generalized. In the period of 1973–1995, both waterbodies were similar in terms of Cladocera species composition and the dominating complex. The list of species composition of pelagic cladocerans has increased since the 1950s. The highest abundance of cladocerans was observed in the Ivankovo water reservoir. Daphnia cucullata G. Sars has been stably dominant in zooplankton summary biomass, while Chydorus sphaericus (O.F. Müller) and Bosmina longirostris (O.F. Müller) prevailed in terms of abundance. Changes in the pelagic cladoceran complex composition is evidence of the waterbodies’ eutrophication. The share of Cladocera in the zooplankton’s total summer biomass was 68 and 53% in the Ivankovski and Shoshinski stretches of the Ivankovo water reservoir, correspondingly, and 60% in the Uglich water reservoir (on average for 1970s–1990s).