The Rotifera of Malaysia and Singapore with remarks on some species

Summary An intensive and extensive sampling of all types of Malaysian and Singaporean freshwaters for zooplankton gave 165 species, sub-species and varieties of Rotifera, mainly belonging to the Monogononta. Bdelloida were rarely identified. The present records bring Malaysia and Singapore in line with other well studied tropical countries as far as Rotifera records are concerned. 7 new genera and over 50 species, subspecies, varieties and forms have been added to the Malaysian and Singaporean Rotifera records and interesting and marked morphological and size variation in species from previous records, have been noted.The richest (in species) fauna of Rotifera occurs in rice fields and ponds which have a seasonal succession of dry to wet conditions and vegetation which shelters the rotifers.The Malaysian and Singaporean Rotifera fauna is typically tropical with a dominance of the genus Brachionus over temperate genera like Keratella. Also very rare or missing are typical temperate genera like Ploesoma, Notholca and Synchaeta. The dominance of Brachionus in Malaysia is less evident than expected because only a very small proportion of samples were taken in limnetic situations where Brachionus clearly dominates in the tropics.     

Zooplankton Distribution in Tropical Reservoirs,South China

The zooplankton of 18 reservoirs of South China was investigated in 2000. 61 Rotifera species, 23 Cladoceras and 14 Copepodas were identified. The most frequent Rotifera genera were Keratella, Brachionus, Trichocerca, Diurella, Ascomorpha, Polyarthra, Ploesoma, Asplanchna, Pompholyx and Conochilus. Bosmina longirostris, Bosminopsis deitersi, Diaphanosoma birgei, D. brachyurum and Moina micrura were typical of Cladocera in the reservoirs. Phyllodiaptomus tunguidus, Neodiaptomus schmackeri and Mesocyclops leuckarti were the most frequent Copepoda and M. leuckarti dominated Copepoda in most reservoirs. High zooplankton species richness with low abundance was characteristic of the throughflowing reservoir, whereas low species richness with low abundance was found in the reservoir with the longest retention time. Relative high abundance and medium species diversity were the distinction of intermediate retention time reservoirs.     

The Species and Size Composition of Tropical Freshwater Zooplankton with Special Reference to the Oriental Region (South East Asia)

A contrast between temperate and tropical Cladocera, Copepoda and Rotifera zooplankton fauna shows a markedly different size composition and species diversity. There are fewer species, and these are generally smaller in the tropics than in either the North or South Temperate Zones. Larger species of the genera Daphnia and Simocephalus are relatively rare in the tropics, while members of Eurycerus and Saycia are absent. Members of the families Holopedidae, Leptodoridae and Polyphemidae increase in abundance toward the temperate regions. Two small cyclopoid copepods are common in tropical zooplankton, while many of the common larger species of other regions are absent or very rare. The chief rotifer components are Keratella tropica and species of Brachionus.     

The ciliate epibiont Epistylis pygmaeum: selection for zooplankton hosts, reproduction and effect on two rotifers

1. A clonal culture of the peritrich Epistylis pygmaeum was used for all observations and experiments. Motile cells preferentially attached to the eggs of three species of Brachionus but also attached to the body of adult B. angularis. Zooids on the transitory egg substratum developed only short stalks, while those on the body often developed long stalks and branched colonies. Selection for the eggs positions the ciliate near the cloaca, and thus high concentrations of fine particulate material excreted by the host. Settlement on eggs occurred equally well in the light and dark, and on moving and stationary eggs. 2. Motile Epistylis cells attached to a wide variety of rotifer and crustacean zooplankton, but exhibited some pronounced selectivity. They readily settled on the eggs of other rotifers (Epiphanes, Polyarthra), on the carapace of several cladocerans (Ceriodaphnia, Daphnia, Diaphanosoma), and on the egg sacs of a copepod (Tropocyclops). They settled less readily on the bodies of the rotifers Asplanchna and Synchaeta, and rarely or never settled on the rotifer Keratella, the cladocerans Bosmina and Scapholeberis, and the body of the copepod. 3. Epistylis populations initiated with a single zooid on Brachionus increased exponentially and often contained several hundred attached zooids and motile cells after 3 days at 20 °C. Observations of a culture initiated from a single telotroch provided new information about peritrich life cycles: (1) motile cells reproduced themselves at a rapid rate (λ = 4.26 day^?1); (2) telotrochs produced or transformed into swimming zooids and vice versa. Functions of the two types of motile cells remain to be clarified. Telotrochs likely are specialised for finding and attaching to hosts. Swimming zooids can feed and reproduce, producing both their own cell type and telotrochs. Together, they should enhance dispersal and population growth, especially when hosts are rare. 4. Life‐table experiments with two species of Brachionus showed that colonisation by Epistylis had no effect on adult survival but significantly decreased fecundity, by 29% in both cases. Zooids attached to eggs could be a weight burden, increase drag, and possibly inhibit egg development. Those on the body of B. angularis also could interfere with coronal cilia, inhibiting feeding and further slowing locomotion. The ability of E. pygmaeum to select and then interfere with its hosts indicates that this epibiont has the potential to influence the species structure of zooplankton communities.     

Energy utilization during swimming and cost of locomotion in larval and juvenile fish

Oxygen consumption and ammonia excretion rates increased in an accelerated manner in larvae and juveniles of whitefish (Coregonus sp.) as a function of swimming speed. The three-dimensional patterns of fish metabolic rates (expressed as energy consumed or nitrogen excreted) versus body weight and swimming speed show that the total standard metabolic rate (i. e. at extrapolated zero swimming speed) increased during early life of whitefish, followed by the expected decrease. This phenomenon might be due to the profound changes in oxidative and glycolytic enzyme activities during fish “metamorphosis”. Standard metabolic rate of ammonia excretion, as a principal product of protein catabolism in fish, decreased by one order of magnitude in early coregonid ontogenesis. This means that protein utilization as an energy source decreases as far as standard metabolism is concerned, but increases with swimming speed. This trend is opposite that in adult fish, where protein utilization in the overall energy supply is diminished at increasing swimming speed. The cost of locomotion offish larvae and juveniles demonstrates that the energy expenditure increases logarithmically with decreasing fish size but at an accelerated rate as compared to adult fish. This contradicts earlier estimates of lower cost of swimming in fish larvae than cost of paddle-propulsion swimming in small invertebrates or cost of flying in insects.     

The cost of incline locomotion in ghost crabs (<Emphasis Type="Italic">Ocypode quadrata</Emphasis>) of different sizes

It is well established that the metabolic cost of horizontal locomotion decreases as a regular function of animal body mass, regardless of body form and phylogeny. How body size affects the cost of incline exercise remains much less clear. Studies on vertebrates have led to the hypotheses that the cost of vertical work is independent of body mass and that the added cost of locomoting on inclines is lower for small animals. Studies on vertebrates and a few invertebrates provide evidence both for and against these hypotheses. To gain further insight into the cost of incline exercise, we measured oxygen consumption of small (2.33 ± 0.07 g) and large (46.66 ± 5.33 g) ghost crabs (Ocypode quadrata) locomoting horizontally and up a 20° incline. The slope of the oxygen consumption versus speed relationship (= minimum cost of transport) was not significantly different for small crabs exercising horizontally and on an incline. However, the intercept for incline exercise was significantly higher, indicating that small crabs used more energy during incline exercise than during horizontal exercise. Incline had no effect on the slope or intercept of the oxygen consumption versus speed relationship for large crabs. Our results suggest that the cost of incline locomotion may be large for small animals and that the cost is not independent of body size. Our results add to the growing body of research indicating that body mass is but one factor that determines the cost of incline locomotion and efficiency of vertical work.     

Induction of different defences by two enemies in the rotifer Keratella tropica: response priority and sensitivity to enemy density

1. Keratella tropica has qualitatively distinct spine‐development responses to kairomones released by the predatory rotifer Asplanchna and the cladoceran interference competitor Daphnia. Asplanchna induces a fourfold lengthening of the right posterior spine (to c.100 μm), a shortening or loss of the left posterior spine and a lengthening of two pairs of anterior spines. Daphnia induces a moderate elongation of both right and left posterior spines. This study tests three hypotheses regarding the sensitivity of these responses to enemy density, and the response priority when both enemies are present. 2. First, since K. tropica and Brachionus calyciflorus have similarly pronounced and effective spine‐development responses to Asplanchna, with no appreciable demographic cost, they should be similarly sensitive to Asplanchna density. This was the case. Both showed an exponential response to increasing Asplanchna density, well described by an asymptotic exponential regression model, and exhibited 50% maximal spine development at statistically similar Asplanchna densities–2.5 and 1.7 μg dry weight L^?1 (1.8 and 1.2 individuals L^?1), respectively. Strong selection for these Asplanchna‐induced responses clearly has led to a coupling of exuberant and effective morphological defence with an unrivalled sensitivity to predator density. 3. Second, since K. tropica’s response to Daphnia is much less pronounced and effective than its response to Asplanchna, it should be less sensitive to Daphnia density. This hypothesis was supported. Spine development increased linearly with increasing Daphnia density and was 50% maximal at 454 μg dry weight Daphnia L^?1, a biomass density 180 times greater than that inducing a comparable response to Asplanchna. 4. Third, since K. tropica’s response to Daphnia does not reduce Asplanchna predation, K. tropica should respond to Asplanchna when both enemies are present at densities sufficient to induce spine development. This was the case. The presence of Daphnia neither reduced nor increased the length of the right posterior spine; it only limited the extent to which the left spine was reduced or lost.     

Zooplankton Associations in the Middle Niger-Sokoto Basin (Nigeria: West Africa)

The species richness of the crustacean zooplankton (Cladocera and Copepoda) and Rotifera in the Middle Niger-Sokoto Basin is characterised from 351 samples collected from 109 localities. 26(4) Cladocera, 18(8) Copepoda and 38(8) Rotifera are recorded. Figures in brackets indicate first records of species for Nigeria. Zooplancton associations of the Middle Niger are compared with one of its major tributaries, River Sokoto, and seven of its minor tributaries using similarity indices of Sørensen and Pearsons Product Moment Cluster Coefficients. The Rivers Sokoto and Shagari consistently had the highest similarity of zooplankton associations, for all combinations of rivers studied with r = 0.54 (Cladocera), r = 0.91 (Copepoda) and r = 0.54 (Rotifera). Zooplankton associations of the Middle Niger are weakly correlated with all tributaries studied (r < 0.1). Species composition and hydrology of the River Sokoto Basin are also compared with previous studies in Nigeria. A Principal Component Analysis of species abundance accounted for 54 % of the total variance. It again confirmed the similarity of the branches Sokoto and Shagari. Two groups of species assemblages were identified. The occurrence of Asplanchna sp., Testudinella patina and Microcyclops varicans in one group appears related to microhabitat selection. The other group made up of Brachionus calyciflorus calyciflorus, Macrothrix spinosa, Lecane luna, Thermocyclops decipiens and Metacyclops minutus is suspected to be linked to physico-chemistry. Further studies on the hydrology and physico-chemistry are required for definitive identification of both factors.     

Sources contribution for benthic invertebrates: an inter-lake comparison in a flood plain system

Dwarfism in males is a common phenomenon in planktonic monogonont rotifers. Considering the small body size of males, some studies suggested that the development time of male rotifers is faster than that of females. However, empirical studies associated with such a speculation are scarce. In this study, we used two common rotifer species, Brachionus calyciflorus and Brachionus plicatilis (each with two geographical populations), to test the hypothesis that the development time of dwarf males is significantly shorter than that of females. Results showed that male eggs (embryos) were deposited much faster than female eggs (embryos) in both Brachionus species. However, the embryonic development time of these small male eggs (embryos) was longer than that of large female eggs (embryos). As a result, males needed significantly longer total development time than females. The total development time of males was 2–3 h longer than that of females in both Brachionus species. Male-producing and female-producing females in the two Brachionus species did not show any difference in development time. Because sexual reproduction begins at high population densities in both species, postponement of development in males will be advantageous by decreasing mating costs in male rotifers.     

Influence of cyanobacterial diet on Asplanchna predation risk in Brachionus calyciflorus

Asplanchna sylvestrii does not discriminate between groups of Brachionus calyciflorus fed either the cyanobacterium Anabaena flos-aquae or a control diet of Euglena gracilis. We based our analysis on the observed probabilities of attack, capture and ingestion during encounters between predator and prey. While A. sylvestrii was very sensitive to brachionid size, we found no significant affects of prey diet on predatory behavior. Thus, cyanobacterial diet did not influence the short-term predation risk of B. calyciflorus exposed to an effective predator. On the other hand, matched cohorts of A. sylvestrii fed B. calyciflorus cultured on the cyanobacterium reproduced more slowly than those fed the same prey cultured on the control food. With prolonged sympatry, therefore, the long-term risk of Asplanchna predation may be reduced for Brachionus by the latter's consumption of cyanobacteria.     

Maternal age and spine development in the rotifer Brachionus calyciflorus: increase of spine length with birth orders1

1. Maternal effects have long been known to influence phenotypic plasticity in rotifers. Females in Brachionus calyciflorus and several other species produce long‐spined offspring when the predatory rotifer Asplanchna is present; B. calyciflorus also develops short spines when food concentrations are low. These spines protect against predation and decrease food threshold concentrations. 2. Some strains of B. calyciflorus develop long spines even in the absence of Asplanchna and other environmental stimuli. We demonstrate in this study that spine length in such cases is dependent on the age of the mother. 3. In strains from Florida and Georgia, offspring spine length increased significantly with birth order, sometimes to lengths formerly observed only in the presence of Asplanchna. Significant variation in this trait was found among and within clones of a strain. Offspring body size also increased with maternal age. This is the first time maternal age has been shown to affect rotifer morphology. 4. These birth‐order effects may have important ecological implications and explain phenotypic plasticity and polymorphism in body size and spine length in populations when predators are absent and food concentrations are high. They may be a bet‐hedging mechanism to assure adaptation to rapid changes in predation pressure or food conditions.     

Resolving Shifting Patterns of Muscle Energy Use in Swimming Fish

Muscle metabolism dominates the energy costs of locomotion. Although in vivo measures of muscle strain, activity and force can indicate mechanical function, similar muscle-level measures of energy use are challenging to obtain. Without this information locomotor systems are essentially a black box in terms of the distribution of metabolic energy. Although in situ measurements of muscle metabolism are not practical in multiple muscles, the rate of blood flow to skeletal muscle tissue can be used as a proxy for aerobic metabolism, allowing the cost of particular muscle functions to be estimated. Axial, undulatory swimming is one of the most common modes of vertebrate locomotion. In fish, segmented myotomal muscles are the primary power source, driving undulations of the body axis that transfer momentum to the water. Multiple fins and the associated fin muscles also contribute to thrust production, and stabilization and control of the swimming trajectory. We have used blood flow tracers in swimming rainbow trout (Oncorhynchus mykiss) to estimate the regional distribution of energy use across the myotomal and fin muscle groups to reveal the functional distribution of metabolic energy use within a swimming animal for the first time. Energy use by the myotomal muscle increased with speed to meet thrust requirements, particularly in posterior myotomes where muscle power outputs are greatest. At low speeds, there was high fin muscle energy use, consistent with active stability control. As speed increased, and fins were adducted, overall fin muscle energy use declined, except in the caudal fin muscles where active fin stiffening is required to maintain power transfer to the wake. The present data were obtained under steady-state conditions which rarely apply in natural, physical environments. This approach also has potential to reveal the mechanical factors that underlie changes in locomotor cost associated with movement through unsteady flow regimes.     

Energetics and food requirements of the female snake Phillodryas chamissonis during the breeding season

Summary 1.The energetic performance and food requirements of female Phillodryas chamissonis was determined during its breeding season at different ambient temperatures. 2. If the long tailed snake is active for 24 weeks, 25 to 32 30-g rodents are required for reproduction and maintenance. This is slightly more than one rodent per week. 3. The energy cost of reproduction (38 to 48% of total metabolism) is close to the values reported for fish (35% of metabolizable energy). 4. Due to their low metabolic rates, the theoretical predation pressure (on a prey biomass basis) exerted by the snakes should be 2 to 2.5% of that expected for endothermic predators of similar body size.     

Zoogeography of the Southeast Asian Rotifera

The distribution and taxonomic composition of Rotifera in Southeast Asia is reviewed. For some countries, records are poor: Brunei, Cambodia and Laos are almost terra incognitafor rotifers (<10 taxa recorded), while the Thai rotifer fauna is the best documented (ca. 310 taxa on record). However, analysis of the available data is impeded by fuzzy taxonomy and the questionable reliability of many records. Most studies focus on the pelagic or littoral of freshwater habitats. Other habitats are largely ignored. Similarly, few studies deal comprehensively with illoricate Monogononta, sessile Flosculariacea and Collothecacea and, especially, Bdelloidea. The genera Lecane, Brachionus and Trichocerca are the best represented, with littoral taxa predominant. Fisheries-related studies dealing with highly productive pelagic environments tend to over report the contribution of Brachionus. Most taxa are thermophilic character, exemplified by the dominance of tropic-centred Lecaneand Brachionus. Some cold-water taxa have been recorded, but the relative climatological homogeneity of the region and low number of studies on high-altitude environments prevent the discrimination of clear latitudinal or altitudinal variation in the distribution of rotifers within Southeast Asia. The majority of Southeast Asian rotifers are widely distributed, including true cosmopolites and thermophilic taxa. There are several local or Oriental endemic Rotifera, mostly Lecane. The American Brachionus havanaensis and Keratella americana appear to have been introduced to the region. The taxonomy of some Rotifera described from the region is commented upon; Brachionus murphyiSudzuki is recognised as senior synonym of B. niwati Sanoamuang et al. (syn. nov.). Some cases of geographical and/or ecological vicariant species-pairs are suggested. The Southeast Asian rotifer fauna contains a sizeable fraction of taxa occurring in the tropical regions of the Old World, most of which also occur in tropical Australia or the Austro-Malayan region. A tropical Australasian faunal component is present, but consists of few taxa only. Hence, affinities between the rotifer fauna of the Ethiopian, Oriental and tropical Australian and Austro-Malayan regions are supported, rather than an affinity between the Indo-Asian or Indo-Malaysian and tropical Australian fauna.     

Locomotor performance and cost of transport in the northern flying squirrelGlaucomys sabrinus

We assess locomotor performance by northern flying squirrelsGlaucomys sabrinus Shaw, 1801 and test the hypothesis that gliding locomotion is energetically cheaper than quadrupedal locomotion. We measured 168 glides by 82 northern flying squirrels in Alaska. Mean glide distances varied from 12.46 m to 14.39 m, with a maximum observed glide distance of 65 m. Mean glide angles varied from 41.31° to 36.31°, and mean air speed ranged from 6.26 m/s to 8.11 m/s. There were no differences in the performance of male and female flying squirrels. We used models of transport cost to provide an initial assessment of the hypothesis that gliding locomotion is energetically less expensive than quadrupedal locomotion. For glides of average length, cost of gliding was less than cost of quadrupedal locomotion except when the animals climbed to the launch point very slowly or ran quickly. Thus the hypothesis that gliding is less expensive than quadrupedal locomotion is supported.     

A routine method of DNA-extraction from extremely small metazoans,e.g. single rotifer specimens for RAPD-PCR analyses

A simple and efficient method has been developed for isolating genomic DNA from single specimens of extremely small metazoan animals, e.g. rotifers (Rotifera: Monogononta). The amount and quality of the extracted DNA allows the Random Amplified Polymorphic DNA Polymerase Chain Reaction (RAPD-PCR) to be employed with up to three random primers per specimen. Reproducibility of RAPD-fingerprints has been checked and verified. Exemplarily, RAPD patterns of three Brachionus species (B. angularis, B. calyciflorus, B. urceolaris) amplified with the primer OPA 07 are demonstrated.     

Assessing metabolic constraints on the maximum body size of actinopterygians: locomotion energetics of Leedsichthys problematicus (Actinopterygii,Pachycormiformes)

Maximum sizes attained by living actinopterygians are much smaller than those reached by chondrichthyans. Several factors, including the high metabolic requirements of bony fishes, have been proposed as possible body‐size constraints but no empirical approaches exist. Remarkably, fossil evidence has rarely been considered despite some extinct actinopterygians reaching sizes comparable to those of the largest living sharks. Here, we have assessed the locomotion energetics of Leedsichthys problematicus, an extinct gigantic suspension‐feeder and the largest actinopterygian ever known, shedding light on the metabolic limits of body size in actinopterygians and the possible underlying factors that drove the gigantism in pachycormiforms. Phylogenetic generalized least squares analyses and power performance curves established in living fishes were used to infer the metabolic budget and locomotion cost of L. problematicus in a wide range of scenarios. Our approach predicts that specimens weighing up to 44.9 tonnes would have been energetically viable and suggests that similar body sizes could also be possible among living taxa, discarding metabolic factors as likely body size constraints in actinopterygians. Other aspects, such as the high degree of endoskeletal ossification, oviparity, indirect development or the establishment of other large suspension‐feeders, could have hindered the evolution of gigantism among post‐Mesozoic ray‐finned fish groups. From this perspective, the evolution of anatomical innovations that allowed the transition towards a suspension‐feeding lifestyle in medium‐sized pachycormiforms and the emergence of ecological opportunity during the Mesozoic are proposed as the most likely factors for promoting the acquisition of gigantism in this successful lineage of actinopterygians.     

Prey selection by Asplanchna girodi (Rotifera): the importance of prey defence mechanisms

1. Predator (laboratory-grown Asplanchna girodi and prey (several field-collected non-carnivorous rotifer species) interactions were studied in the laboratory as a function of both predator and prey densities. The clearance rates of A, girodi decreased with increasing prey density. Predator density had no effect on the feeding of A. girodi. 2. Asplanchna girodi selectively fed on Keratella cochlearis. Its clearance rates for K. cochlearis were much higher than those for Polyarthra and Brachionus. Short-spined forms (spine length less than 25 μm) showed a significantly higher susceptibility to predation than either the non-spined or the long-spined forms. 3. Large Asplanchna individuals fed selectively on reproductive females of K. cochlearis, thereby reducing the fecundity of their prey.     

Pygmy locomotion

The hypothesis that Pygmies may differ from Caucasians in some aspects of the mechanics of locomotion was tested. A total of 13 Pygmies and 7 Caucasians were asked to walk and run on a treadmill at 4–12 km · h^–1. Simultaneous metabolic measurements and three-dimensional motion analysis were performed allowing the energy expenditure and the mechanical external and internal work to be calculated. In Pygmies the metabolic energy cost was higher during walking at all speeds (P < 0.05), but tended to be lower during running (NS). The stride frequency and the internal mechanical work were higher for Pygmies at all walking (P < 0.05) and running (NS) speeds although the external mechanical work was similar. The total mechanical work for Pygmies was higher during walking (P < 0.05), but not during running and the efficiency of locomotion was similar in all subjects and speeds. The higher cost of walking in Pygmies is consistent with the allometric prediction for smaller subjects. The major determinants of the higher cost of walking was the difference in stride frequency (+9.45, SD 0.44% for Pygmies), which affected the mechanical internal work. This explains the observed higher total mechanical work of walking in Pygmies, even when the external component was the same. Most of the differences between Pygmies and Caucasians, observed during walking, tended to disappear when the speed was normalized as the Fronde number. However, this was not the case for running. Thus, whereas the tested hypothesis must be rejected for walking, the data from running, do indeed suggest that Pygmies may differ in some aspects of the mechanics of locomotion.