Mating behavior,spawning, parental care,and embryonic development of some marine pseudocerotid flatworms (Platyhelminthes: Rhabditophora: Polycladida) in Singapore

The reproductive biology of tropical marine cotylean polyclads is presently poorly known. Reproductive aspects of 16 polyclad species from the family Pseudocerotidae in the genera Acanthozoon, Nymphozoon, Phrikoceros, Pseudobiceros, Pseudoceros, Thysanozoon, and Tytthosoceros from Singapore were documented for the first time. Penis fencing was observed to be just a mating ritual and not necessary for insemination, not always aggressive, and could also result in eventual reciprocal insemination. Results showed that all species underwent similar embryonic developmental stages and hatched as Müller's larvae. Only Pseudoceros concinnus and Pseudoceros laingensis, with mean developmental times of >20 days and mean brood sizes of <1,000 eggs, displayed long‐term parental care. Polyclads producing larger brood sizes had shorter developmental times and only covered their egg masses for about one day. Phrikoceros baibaiye and Pseudobiceros spp. produced egg capsules with pointed opercula, whereas all other species possessed smooth, rounded opercula. All genera hatched with brownish‐orange larvae, except for Pseudoceros spp., which hatched with reddish‐purple larvae regardless of the initial embryo color (either grayish‐yellow or reddish‐purple). These could potentially complement current taxonomic characters in distinguishing polyclad genera and species.     

Seston capture by Hydropsyche siltalai and the accuracy of capture efficiency estimates

1. Suspension feeding by caseless caddisfly larvae (Trichoptera) constitutes a major pathway for energy flow, and strongly influences productivity, in streams and rivers. 2. Consideration of the impact of these animals on lotic ecosystems has been strongly influenced by a single study investigating the efficiency of particle capture of nets built by one species of hydropsychid caddisfly. 3. Using water sampling techniques at appropriate spatial scales, and taking greater consideration of local hydrodynamics than previously, we examined the size‐frequency distribution of particles captured by the nets of Hydropsyche siltalai. Our results confirm that capture nets are selective in terms of particle size, and in addition suggest that this selectivity is for particles likely to provide the most energy. 4. By incorporating estimates of flow diversion around the nets of caseless caddisfly larvae, we show that capture efficiency (CE) is considerably higher than previously estimated, and conclude that more consideration of local hydrodynamics is needed to evaluate the efficiency of particle capture. 5. We use our results to postulate a mechanistic explanation for a recent example of interspecific facilitation, whereby a reduction of near‐bed velocities seen in single species monocultures leads to increased capture rates and local depletion of seston within the region of reduced velocity.     

Determinative development in the polyclad turbellarian,Hoploplana inquilina

Summary

Blastomere deletion experiments at the two- and four-cell stages were carried out on the embryo of the polyclad turbellarian Hoploplana inquilina to further examine the relationship between spiral cleavage and early embryonic determination in primitive spiralians. Deletion of one cell at the two-cell stage resulted in “half” larvae that were abnormal in body shape, lobe development, and behavior. Deletion of one cell at the four-cell stage produced less abnormal “three-quarter” larvae which were still underdeveloped in one of the quadrants. A 3:1 ratio of one-eyed to two-eyed larvae implies that deletion of any one of three blastomeres results in loss of an eye, with two constituting the eye lineage and the third controlling the development of two eyes. The results demonstrate that the polyclad embryo is determined early in development, though significant cell interactions occur during cleavage, and suggest that determinative development and quartet spiral cleavage are always associated and probably represent a primitive, strongly conserved evolutionary condition.     

Dislodgement effect of natural semiochemicals released by disturbed triatomines: a possible alternative monitoring tool

The quick detection of domestic and peridomestic triatomines in their environments becomes difficult without the use of dislodgement substances that flush them out from their shelters. At present, tetramethrin 0.2% is being widely used in control programs. Although it is an efficient dislodging agent, its toxicity might affect the health of captured triatomines, of other insects and, to a lesser extent, of other animals, including humans. Here, we tested if semiochemicals released by disturbed adults of Triatoma infestans and/or Rhodnius prolixus can make larvae of the same species exit from their refuges. In a walking olfactometer we found that: 1) larvae of T. infestans were repelled by the odors released by disturbed adults of their own species and of R. prolixus, 2) larvae of R. prolixus did not change their behavior in the presence of odors released by adults of both species, and 3) activity levels were not modulated by these odors in any of both species. Besides, in pseudo‐natural conditions we found an increased flushing‐out activity of larvae of T. infestans when their shelters were sprayed with isobutyric acid or 3‐pentanol, and of larvae of R. prolixus when sprayed with 3‐methyl1butanol. We succeeded in this work to dislodge larvae of triatomines from artificial shelters using natural volatile compounds, allowing the capture of live bugs for further investigations (e.g., xenodiagnosis or genetic studies) and favoring ecological aspects (e.g., minimizing environmental insecticide‐contamination and non‐targeted mortality).     

Variation in food availability influences prey‐capture method in antlion larvae

Abstract 1. Larvae of a Myrmecaelurus sp. are unique among antlions because they have two prey‐capture methods; they either ambush prey at the surface, or dig pit traps that prey fall in to. It was hypothesised that larvae will use the capture method that maximises their net rate of energy gain, which will be influenced by food availability (encounter rate) and by past energy inputs (body condition). 2. Costs were estimated by measuring resting and activity metabolic rates and determining the duration of pit maintenance at various encounter rates with ants that served as prey. Benefits were estimated from the energy gained per ant captured at different encounter rates. 3. Net energy gained was higher with a pit than without one, and was influenced more by the differences in prey capture rate between the two capture methods, and less by the differences in energy costs associated with each method. The proportion of larvae that constructed pits was higher when they were in intermediate body condition than when in good or in poor body condition. 4. Thus, the use of one capture method or the other depends on a combination of the influences of past net energy gain and the antlion’s most recent change in encounter rate with prey. Ambushing without a pit may serve as a default when physiological constraints limit the larvae’s ability to invest in pit construction and maintenance, or when larvae are sated, and saving the energy of pit construction and maintenance is worthwhile.     

The effect of age at metamorphosis on the transition from larval to adult suspension‐feeding of the slipper limpet Crepidula fornicata

Slipper limpets use different ciliary feeding mechanisms as larvae and adults. Veliger larvae of Crepidula fornicata developed part of the adult feeding apparatus, including ctenidial filaments, neck lobe, and radula, before metamorphosis, but ctenidial feeding did not begin until well after loss of the larval feeding apparatus (velum) at metamorphosis. Earlier initiation of ctenidial feeding by individuals that were older larvae when metamorphosis occurred suggests continued development toward ctenidial feeding during delay of metamorphosis. Early juveniles produced a ciliary current through the mantle cavity and moved the radula in a grasping action before they began to capture algal cells on mucous strands or form a food cord. Either early juveniles could not yet form mucous strands or they delayed their production until development of other necessary structures. The neck canal for transporting food from ctenidium to mouth cannot develop before velar loss. In their first feeding, juveniles fed much like the adults except that the neck canal was less developed and the path of the food cord toward the mouth sometimes varied. As suspension feeders, calyptraeids lack the elaborations of foregut that complicate transition to juvenile feeding for many caenogastropods, but a path for the food cord must develop after velar loss. Why individuals can initiate ctenidial feeding sooner when they are older at metamorphosis is not yet known. The juveniles became sedentary soon after metamorphosis and were not observed to feed by scraping the substratum with the radula, in contrast to the first feeding by juveniles of another calyptraeid species, observed by Montiel et al. ( 2005 ).     

Male‐biased dispersal and the potential impact of human‐induced habitat modifications on the Neotropical bat Trachops cirrhosus

Gene flow, maintained through natal dispersal and subsequent mating events, is one of the most important processes in both ecology and population genetics. Among mammalian populations, gene flow is strongly affected by a variety of factors, including the species’ ability to disperse, and the composition of the environment which can limit dispersal. Information on dispersal patterns is thus crucial both for conservation management and for understanding the social system of a species. We used 16 polymorphic nuclear microsatellite loci in addition to mitochondrial DNA sequences (1.61 kbp) to analyse the population structure and the sex‐specific pattern of natal dispersal in the frog‐eating fringe‐lipped bat, Trachops cirrhosus, in Central Panama. Our study revealed that—unlike most of the few other investigated Neotropical bats—gene flow in this species is mostly male‐mediated. Nevertheless, distinct genetic clusters occur in both sexes. In particular, the presence of genetic differentiation in the dataset only consisting of the dispersing sex (males) indicates that gene flow is impeded within our study area. Our data are in line with the Panama Canal in connection with the widening of the Río Chagres during the canal construction acting as a recent barrier to gene flow. The sensitivity of T. cirrhosus to human‐induced habitat modifications is further indicated by an extremely low capture success in highly fragmented areas. Taken together, our genetic and capture data provide evidence for this species to be classified as less mobile and thus vulnerable to habitat change, information that is important for conservation management.     

Experimental evidence for the origins of determinative development in the polyclad turbellarians

Cell-deletion experiments were carried out on the embryo of the polyclad turbellarian Hoploplana inquilina to examine further the nature of development in primitive spiralians. The polyclads are of particular interest because they provide a link between the regulative development of acoels and the determinative development of annelids and molluscs. Single blastomeres were deleted at the two- and four-cell stages by puncture through the eggshell membrane with tungsten needles. All deletions resulted in abnormal larvae with consistent characteristics representing half or three-quarter Müller's larvae. The number of larval eyes was a particularly useful character in revealing mosaicism. This study establishes the polyclad embryo as determinative, but with important cell interactions also occurring during early development, and provides evidence that mosaicism became associated with spiral cleavage in the quartet form during the evolution of the Turbellaria.     

Benthic ctenophores (Platyctenida: Coeloplanidae) in South Florida: predator–prey interactions

The primary goal of this study was to demonstrate, from field observations and laboratory experiments, some key trophic roles of benthic ctenophores as predators and prey in subtropical communities. We examined individuals of two benthic platyctenid species: Coeloplana waltoni, a minute epibiont on octocorals in exposed, open‐water settings; and Vallicula multiformis, an associate of calm‐water biofouling communities and floating Sargassum spp. Laboratory observations of individuals of both ctenophore species revealed frequent capture and ingestion of diverse zooplankton taxa, especially crustaceans. Laboratory predation trials demonstrated the capture of dolphinfish (Coryphaena hippurus) eggs and larvae by both ctenophore species. Dolphinfish eggs and larvae larger than individuals of C. waltoni were captured but not ingested during 2‐h trial periods. These prey items were sometimes purloined and ingested by polyps of the ctenophore's octocoral host. Ingestion of dolphinfish eggs and larvae by individuals of C. waltoni was observed, however, after longer periods of exposure to prey. In predation trials, dolphinfish eggs and larvae were both captured and ingested by larger individuals of the ctenophore species V. multiformis. Field and laboratory observations revealed diverse invertebrate and fish taxa that prey on both ctenophore species. In the laboratory, the mean daily per capita consumption of individuals of C. waltoni by a pomacanthid fish ranged 0.5–2.8 individuals, and ranged 2.6–3.6 individuals for predation by an ovulid mollusc. Field population densities of these predators ranged 0.1–0.7 individuals per m² for the pomacanthid, and 0.2–1.1 individuals per m² for the mollusc. Laboratory feeding observations demonstrated frequent consumption of individuals of V. multiformis by a sea anemone, and by three species of brachyuran crabs. Field observations revealed eight fishes that probably feed incidentally on individuals of V. multiformis. These findings add to the limited knowledge base of predator–prey dynamics in both C. waltoni and V. multiformis.     

The effects of potential larval supply, settlement and post-settlement processes on the distribution of two species of filter-feeding caddisflies

1. Hydropsychid caddisfly larvae are often abundant in fast flow habitats and on highly textured substrata. Such patterns are usually inferred to result from habitat preference, but the roles of larval supply to the benthos and post‐settlement events are rarely examined for stream populations. We describe a manipulative field experiment that examined larval supply, habitat preference and post‐settlement events simultaneously for two co‐occurring species of hydropsychids. 2. Ten artificial channels were constructed in a natural riffle to create fast and slow flow habitats and within each channel we placed six artificial substrata whose surfaces had been modified to create three different texture treatments. We measured discharge through channels and monitored hydropsychid colonisation of substrata every 3–4 days. Half of the substrata had all hydropsychids and nets removed every 3–4 days (short‐term colonisation) whereas the remaining half were counted but nets and hydropsychids left attached (long‐term colonisation). Short‐term counts of nets and individuals were summed to allow comparison with long‐term colonisation substrata. 3. Smicrophylax sp. AV2 larvae were more abundant in fast flow channels than slow flow channels, but these differences were proportional to discharge through each of the channels, suggesting that supply of settlers can explain this pattern. Smicrophylax larvae were least abundant on smooth substrata, which suggests that this species selects habitats based on surface texture. Alternatively, Asmicridea sp. AV1 larvae were only found in fast flow channels and this most likely reflects an active habitat choice by this species, but there was no significant difference between different texture treatments. 4. There were no differences between summed, short‐ and long‐term counts of recruits for either species, but there were more nets than larvae, especially in slow flow channels, by the end of the experiment, suggesting that larval mortality or re‐dispersal after settlement could be considerable. 5. Our results indicate that supply, habitat selection at settlement and post‐settlement processes all contributed variously to the distribution of hydropsychid caddisfly larvae, but that each species was affected differentially by these factors. Larval supply and post‐settlement processes are rarely examined by stream researchers and our results demonstrate these factors deserve much more consideration. Calculating accurate larval supply rates to sites is challenging, but we suggest that such detailed information is necessary if we are to sort out what sets limits to distributions and the underlying population structure of stream invertebrate populations.     

Particle capture in ciliary filter‐feeding gymnolaemate and phylactolaemate bryozoans – a comparative study

Riisgård, H.U., Okamura, B. and Funch, P. 2009. Particle capture in ciliary filter‐feeding gymnolaemate and phylactolaemate bryozoans – a comparative study. —Acta Zoologica (Stockholm) 91 : 416–425. We studied particle capture using video‐microscopy in two gymnolaemates, the marine cheilostome Electra pilosa and the freshwater ctenostome Paludicella articulata, and three phylactolaemates, Fredericella sultana with a circular funnel‐shaped lophophore, and Cristatella mucedo and Lophophus crystallinus, both with a horseshoe‐shaped lophophore. The video‐microscope observations along with studies of lophophore morphology and ultrastructure indicated that phylactolaemate and gymnolaemate bryozoans with a diversity of lophophore shapes rely on the same basic structures and mechanisms for particle capture. Our study also demonstrates that essential features of the particle capture process resemble one another in bryozoans, brachiopods and phoronids.     

Myogenesis in two polyclad platyhelminths with indirect development,Pseudoceros canadensis and Stylostomum sanjuania

SUMMARY Myogenesis of two representatives of Platyhelminthes, Stylostomum sanjuania and Pseudoceros canadensis, was followed from egg deposition until well‐differentiated free‐swimming larval stages, using F‐actin staining and confocal laserscanning microscopy. Zonulae adhaerentes are the only structures to stain before 50% of development between egg deposition and hatching in S. sanjuania, and before 67% of development in P. canadenis. Subsequently, irregular fibers appear in the embryo, followed by a helicoid muscle close to the apical pole. Three longitudinal muscle pairs form, of which the dorsal pair remains more pronounced than the others. Gradually, new muscles form by branching or from double‐stranded muscle zones adjacent to existing muscles. This results in an elaborate muscular bodywall that consists of a single helicoid muscle as well as multiple circular and longitudinal muscles. Diverse retractor muscles insert at the sphincter muscles around the stomodeum. The overall arrangement and formation mode of the larval musculature appears very similar in both species, although only P. canadensis has a primary circular muscle posterior to the helicoid muscle. Muscle formation in the apical region of the embryo precedes that at the abapical pole and the primary longitudinal muscles form slightly later than the primary circular muscles. Myogenesis and larval myoanatomy appears highly conserved among polyclad flatworms, but differs significantly from that of other trochozoan clades. Our data suggest that the larval muscular ground pattern of polyclad larvae comprises a bodywall consisting of a helicoid muscle, circular and longitudinal muscles, several retractor muscles, and sphincter muscles around the stomodeum.     

The mussel filter–pump – present understanding,with a re‐examination of gill preparations

Filter feeding in mussels is a secondary adaptation where the gills have become W‐shaped and greatly enlarged, acting as the mussel filter–pump. Water pumping and particle capture in the blue mussel, Mytilus edulis, have been studied over many years. Here, we give a short status of the present understanding of ciliary structure and function of the mussel filter–pump, supplemented with new photo‐microscope and scanning electron microscopy (SEM) pictures of gill preparations. Pumping rate (filtration) and pressure to maintain flow have been extensively studied so the power delivered by the mussel pump to the water flow is known (1.1% of total respiratory power), but the actual cost based on gill respiration is much higher (19%), implying that the cost of maintaining of the large gill pump is considerable and that only relatively little energy can be saved by stopping or reducing the activity of the water‐pumping cilia so that continuous feeding with a ‘minimal scaled’ pump is cheaper than discontinuous feeding with a correspondingly larger pump. According to the present view, the pump proper is the beating lateral cilia (lc) on the gill filaments and particle capture is accomplished by the action of laterofrontal cirri (lfc) transferring particles from the main water current to the frontal gill filament currents driven by frontal cilia (fc). Unexplained aspects include retention efficiency according to particle size and the role of pro‐laterofrontal cilia (p‐lfc) placed between the lfc and fc. The structure of cilia and the mode of ciliary beating have been re‐examined in this study by new high‐resolution light and scanning electron microscopy of isolated gill preparations exposed to serotonin (5‐HT) stimulation which can activate the lc and lfc at low concentrations (10^?6 M), but removes the lfc from the interfilament canals at higher concentrations (10^?5 M).     

Versatile ciliary behaviour in capture of particles by the bryozoan cyphonautes larva

Cyphonautes larvae of a bryozoan, Membranipora membranacea, used several ciliary mechanisms to capture algal cells upstream from the lateral band of cilia that produces a feeding current. (1) Lateral cilia changed beat and a backcurrent occurred at the time and place that particles were retained. (2) Algal cells were sieved and held stationary at the upstream (frontal) side of a row of laterofrontal cilia that were not beating. (3) Localized extension of cilia toward the inhalant chamber, coincident with particle captures, indicated that laterofrontal cilia flick toward the inhalant chamber. These flicks may aid transport of captured particles toward the mouth. Thus my earlier report that larvae only sieve, in contrast to the adults (which have an active ciliary response) was in error. The similar ciliary bands in adult and larval bryozoans and in other lophophorates (brachiopods, and phoronids) suggest that these animals share a core repertoire of ciliary behaviours in the capture and concentration of suspended food particles.     

Ciliary sieving and active ciliary response in capture of particles by suspension-feeding brachiopod larvae

Larvae of a brachiopod, Glottidia pyramidata, used at least two ciliary mechanisms to capture algal cells upstream from the lateral band of cilia that produces a feeding/swimming current. (1) Filtration: the larvae retained algal cells on the upstream (frontal) side of a sieve composed of a row of stationary laterofrontal cilia. Movement of the laterofrontal cilia could not be observed during capture or rejection of particles, but the laterofrontal cilia can bend toward the beating lateral cilia, a possible mechanism for releasing rejected particles from the ciliary sieve. (2) Localized changes of ciliary beat: the larvae may also concentrate particles by a local change in beat of lateral cilia in response to particles. The evidence is that the beat of lateral cilia changed coincident with captures of algal cells and that captured particles moved on paths consistent with a current redirected toward the frontal side of the tentacle by an induced local reversal of the lateral cilia. The change of beat of lateral cilia could have been an arrest rather than a reversal of ciliary beat, however. The similar ciliary bands in adult and larval lophophorates (brachiopods, phoronids, and bryozoans) suggest that these animals share a range of ciliary behaviours. The divergent accounts of ciliary feeding of lophophorates could be mostly the result of different authors observing different aspects of ciliary feeding.     

ICK is essential for cell type‐specific ciliogenesis and the regulation of ciliary transport

Cilia and flagella are formed and maintained by intraflagellar transport (IFT) and play important roles in sensing and moving across species. At the distal tip of the cilia/flagella, IFT complexes turn around to switch from anterograde to retrograde transport; however, the underlying regulatory mechanism is unclear. Here, we identified ICK localization at the tip of cilia as a regulator of ciliary transport. In ICK‐deficient mice, we found ciliary defects in neuronal progenitor cells with Hedgehog signal defects. ICK‐deficient cells formed cilia with mislocalized Hedgehog signaling components. Loss of ICK caused the accumulation of IFT‐A, IFT‐B, and BBSome components at the ciliary tips. In contrast, overexpression of ICK induced the strong accumulation of IFT‐B, but not IFT‐A or BBSome components at ciliary tips. In addition, ICK directly phosphorylated Kif3a, while inhibition of this Kif3a phosphorylation affected ciliary formation. Our results suggest that ICK is a Kif3a kinase and essential for proper ciliogenesis in development by regulating ciliary transport at the tip of cilia.     

Opposed bands of cilia in the nonfeeding larvae of the serpulid annelid Salmacina tribranchiata

The nonfeeding planktonic larvae of marine invertebrates typically lack larval feeding structures. One puzzling exception to this generalization is the annelid clade Sabellidae, in which nonfeeding larvae possess ciliary bands (specifically, food groove and metatroch) that, to the best of our knowledge, have no function other than in feeding. Nishi and Yamasu (1992b, Bulletin of the College of Sciences, University of the Ryukyus, 54 , 107–121) published a scanning electron micrograph showing that nonfeeding larvae of the serpulid annelid Salmacina dysteri also possess food groove and metatrochal cilia. Here I demonstrate that nonfeeding larvae of Salmacina tribranchiata also bear ciliary bands identifiable as food groove and metatroch by position. High‐speed video of ciliary beat patterns shows that, together with the prototrochal cilia, these bands function in an opposed band system. The presence of feeding structures in nonfeeding annelid larvae is thus more widely distributed than previously recognized. The presence of feeding structures may make evolutionary transitions to planktotrophy more likely, and may underlie an inferred origin of larval feeding in the common ancestor of one of the two major clades of serpulid annelids, Serpulinae.     

Embryonic and larval development and fatty‐acid profile of Epinephelus marginatus spawned in captivity: tools applied to captive rearing

Development, ontogeny of the digestive system and the fatty‐acid (FA) profile, were analysed during development of Epinephelus marginatus. Larvae were analysed 7 and 17 days post fertilization (dpf) to evaluate fatty‐acid profile and morphological variables, respectively. Epinephelus marginatus larvae have relatively slow development of digestive structures, but were able to capture, ingest and digest prey by 5 dpf. Eggs were composed of high percentages of polyunsaturated fatty acids (PUFA) in phospholipids. The percentage of n3 PUFAs was higher than n6, especially docosahexaenoic acid (DHA), which exhibited higher levels compared with other marine species during the first 3 days of development, both in terms of phospholipids and triglycerides. The larvae present a high content of docosahexaenoic acid–eicosapentaenoic acid (DHA–EPA) and, during this phase, live food of small size was required (copepods or SS‐strain Brachionus rotundiformes), enriched with DHA–EPA. These results may guide future studies on the contribution of FAs required during this stage of the life cycle of E. marginatus, to advance knowledge of the use of these FAs throughout ontogeny and contribute to the culture of this species commercial production or restocking.     

The presence and location of small cardioactive‐like peptides in larvae of Crassostrea virginica

We examined the presence and location of small cardioactive peptide (SCP)‐like neuropeptides in both the central and peripheral nervous systems of D‐hinge, newly eyed, and pediveliger larvae of Crassostrea virginica. Results indicate that SCP‐like substances are present early in development (D‐hinge larvae), and that presence in the central and peripheral nervous systems increases as the larva develops toward metamorphic competence (pediveliger larvae). In addition, in newly eyed and pediveliger larvae, SCP‐like labeling is found in a varying number of neurons within all central ganglia, with the possible exception of the accessory ganglia. Varicose labeling of axons is also documented within the ganglia, commissures, and connectives, as well as in peripheral nerves. Peripheral tissues innervated by axons exhibiting SCP‐like immunoreactivity include the velum, foot, esophagus, mantle, and various musculatures. As indicated by the location of SCP‐like labeling in various organs and tissues, it is likely that these neuropeptides modulate muscle contraction or ciliary beating in molluscan larvae such as those of C. virginica.