Increased parasite abundance associated with reproductive maturity of the clam Anodonta piscinalis.

Several studies on vertebrates have demonstrated that reproductive activities may increase the parasite load, but this has not been shown in invertebrate hosts. We studied abundance of a potentially harmful gill parasite, the ergasilid copepod Paraergasilus rylovi, from the freshwater bivalve host Anodonta piscinalis in relation to reproductive maturity of the host in the field. Prevalence of this previously unstudied parasite varied from 90 to 100%, and the mean parasite abundance from 16.3 to 28.8 among 3 study populations. Abundance of P. rylovi increased with host size. In the maturating age groups (3-5 yr) the length-adjusted mean parasite abundance among mature, reproducing female clams that brooded glochidia larvae was 2 times higher than in nonreproducing females, the observed pattern being consistent among the 3 study lakes. Alternative, mutually nonexclusive explanations may be found for the result. For example, changes in clam behavior or filtration activity accompanying maturation can increase host exposure to parasites, or reproduction may decrease energy available to host immunologic defense. However, the present result indicates that maturation, and reproduction, is associated with increased parasite abundance in A. piscinalis, an invertebrate host.     

Perkinsosis in the Manila clam Ruditapes philippinarum affects responses to the harmful-alga, Prorocentrum minimum

The dinoflagellate Prorocentrum minimum is increasingly recognized as a harmful algal bloom (HAB) species that affects filter-feeding shellfish. An experiment was done to investigate possible interactions between parasitic diseases and exposure to P. minimum in Manila clams, Ruditapes philippinarum. Manila clams, with variable levels of infection with Perkinsus olseni, were exposed for three or six days to the benign phytoplankton species Chaetoceros neogracile or a mixed diet of C. neogracile and P. minimum. After three or six days of exposure, clams were assessed individually for condition index, parasite status, and plasma and hemocyte parameters (morphological and functional) using flow-cytometry. Histological evaluation was also performed on individual clams to assess prevalence and intensity of parasitic infection, as well as other pathological conditions.Prorocentrum minimum caused several changes in Manila clams, especially after six days of exposure, such as decreased hemocyte phagocytosis and size and clam condition index. Pathological conditions observed in Manila clams exposed to P. minimum were hemocyte infiltration in the intestine and gonad follicles, myopathy, and necrosis of the intestine epithelial cells. The parasite P. olseni alone had no significant effect on Manila clams, nor did it modulate the hemocyte variables in clams exposed to P. minimum; however, the parasite did affect the pathological status of Manila clams exposed to the P. minimum culture, by causing atrophy and degeneration of residual ova in the gonadal follicles and hyaline degeneration of the muscle fibers, indicating synergistic effects of both stressors on the host over a short period of time. Additionally, an in vitro experiment also demonstrated detrimental effects of P. minimum and exudates upon P. olseni cells, thus suggesting HAB antagonistic suppression of transmission and proliferation of the parasite in the natural environment over a longer period of time. The results of this experiment demonstrate the complexity of interactions between host, parasite, and HAB.     

The effects of water depth and density on the growth of a unionid clam*

SUMMARY. 1. Unionid clams from Narrow Lake, Alberta, were collected to quantify the natural variation in growth, to assess the natural variation in abundance, age and size distribution, and growth with water depth in the lake, and to conduct in situ experiments to directly test the effects of water depth (temperature) and clam abundance on clam growth. 2. The unionid clam, Anodonta grandis simpsoniana, showed wide variation in length at a given age. There were no significant differences in growth between clams collected at 1,3, 5, and 7m depths in the lake despite marked differences in water temperature. The wide variation in clam biomass within each depth zone may have masked possible effects of water depth. 3. The effect of water depth and variation in clam density on clam growth was tested directly by stocking clams into small enclosures at densities equivalent to 50, 150, 250, 350 and 450g m-2 (live weight) at each of 1, 3, 5 and 7 m depths in Narrow Lake (each depth and abundance treatment in triplicate). A uniform sandy substrate was used in all enclosures to eliminate any possible effect of substrate type on growth. 4. Mortality was negligible (0.9%) during the experiment. Clam density had no significant effect on clam growth which suggests that clam growth was not food limited in the lake. 5. Clams reared at 7 m grew more slowly than clams reared at 1, 3 and 5 m. Clams reared at 5 m grew more slowly than clams reared at 1 and 3m. Growth of clams reared at 1 and 3m did not differ. These differences in growth were strongly correlated with the measured differences in water temperature between depths. 6. Migration between depths probably accounts for the lack of a depth effect on clams growing in the natural habitat.     

Environmental distribution and persistence of Quahog Parasite Unknown (QPX)

Quahog Parasite Unknown (QPX) is the cause of mass mortality events of hard clams Mercenaria mercenaria from Virginia, USA, to New Brunswick, Canada. Aquaculture areas in Massachusetts, USA, have been particularly hard hit. The parasite has been shown to be a directly infective organism, but it is unclear whether it could exist or persist outside of its clam host. We used molecular methods to examine water, sediment, seaweeds, seagrass and various invertebrates for the presence of QPX. Sites in Virginia and Massachusetts were selected based upon the incidence of QPX-induced clam die-offs, and they were monitored seasonally. QPX was detectable in almost all of our different sample types from Massachusetts, indicating that the parasite was widely distributed in the environment. Significantly more samples from Massachusetts were positive than from Virginia, and there was a seasonal pattern to the types of samples positive from Massachusetts. The data suggest that, although it may be difficult to completely eradicate QPX from the environment, it may be possible to keep the incidence of disease under control through good plot husbandry and the removal of infected and dying clams.     

Aspects of the ecology and natural history of Paraergasilus rylovi (Copepoda, Ergasilidae) parasitic in unionids of Finland

The distribution of Paraergasilus rylovi in 17 populations of unionids was investigated. In 1 unionid population, the parasite was studied regarding host age, size, sex, and the reproductive period (occurrence of egg sacs). Results from pooled material from the years 1987--1989 and 1996 (southern Finland, 11 populations) indicated that Anodonta piscinalis (n = 1,359) is the main host (total mean prevalence 71% and intensity +/-SE of infection 16.4+/-0.6). Pseudanodonta complanata (n = 106) was infected occasionally (3% and 1.3+/-0.3), whereas Unio pictorum (n = 108) and U. tumidus (n = 17) were not infected. Results from 17 A. piscinalis populations showed that P. rylovi occurs in southern Finland but not in northern Finland. In A. piscinalis, the mean intensity of infection was higher in lake populations than in river populations. Both host age and length had a negative relationship with the intensity of P. rylovi infection. Host sex did not affect the intensity of infection. Egg sacs of P. rylovi were found from June to August. There was a tendency for higher intensities of infection in autumn. Infection by the digenean Rhipidocotyle fennica had no effect on the intensity of P. rylovi infection.     

Modulatory effects of hard clam (Mercenaria mercenaria) tissue extracts on the in vitro growth of its pathogen QPX

Quahog parasite unknown (QPX) is a fatal protistan parasite affecting cultured and wild hard clams Mercenaria mercenaria along the northeastern coasts of the USA and maritime Canada. Field investigations and laboratory transmission studies revealed some variations in the susceptibility of different hard clam stocks to QPX infection. In this study, we used in vitro QPX cultures to investigate the effect of plasma and tissue extracts from two different clam stocks on parasite survival and growth. Results demonstrated the presence of factors in clams that significantly modulate QPX growth. Extracts from gills and mantle tissues as well as plasma inhibited in vitro QPX growth, whereas foot and adductor muscle extracts enhanced parasite growth. Investigations of anti-QPX activities in plasma from two clam stocks displaying different susceptibility toward QPX disease in vivo demonstrated higher inhibition of QPX growth by plasma from New York (resistant) clams compared to Florida (susceptible) clams. Some clams appeared to be deficient in inhibitory factors, suggesting that such animals may become more easily infected by the parasite.     

Dynamics of biochemical components, lipid classes and energy values on gonadal development of R. philippinarum associated with the temperature and ingestion rate

This study evaluates the effect of temperature, coupled with ingestion rate, on the dynamics of biochemical components and lipid classes in R. philippinarum. The data are discussed with regard to sexual development and energy balance. Experimental protocol developed in the present study used two groups of the clam R. philippinarum: L (temperatures of 14 degrees C and 18 degrees C) and H (temperatures of 18 degrees C and 22 degrees C). The intra-group ingestion level was similar, although the ingestion level of the clams in the group H was 2.4 times higher than group L. We observed that R. philippinarum conditioned at 18 degrees C (18L) shows higher protein content, furthermore an important loss of organic weight was observed after 48 days. In such a situation, the clams use their own reserves (carbohydrates and glycogen) for sexual development while in situations without food stress (positive energy balance) and low temperature (14 degrees C) an accumulation of reserves is produced. Strikingly dissimilar behaviour in biochemical composition was observed for the 18H and 22H treatments, both with a positive energy balance. Despite similar protein content, the highest levels of carbohydrates were observed at the lower temperature (18 degrees C). Glycogen was also higher for the 18 degrees C treatment, although the differences were significant only in the males. Although the total lipids in R. philippinarum showed no significant differences in any treatment, they became apparent and related to sex when considering the individual lipid classes. There was no variation in lipid classes in the males between the 14L and 22H treatments despite the large disparity in the degree of sexual development. However, in the females significant differences in lipid classes (phospholipids, triglycerides) were observed. The results of this study show that a positive energy balance permits R. philippinarum gonadal development and accumulation of reserves both in low and high temperature conditions. In low temperature situations, gonadal development is slower and the energy reserves are accumulated in the form of carbohydrates. When the clams are conditioned at high temperatures, gonadal development is fast and complete, carbohydrates are consumed and lipids are accumulated.     

Effects of high temperatures on functional responses of haemocytes in the clam Chamelea gallina

The effects of high temperatures on the clam, Chamelea gallina, generally recognised as a low tolerant bivalve species, were studied by evaluating some functional responses of the haemocytes. The animals were kept for 7days at 20, 25 and 30 degrees C and total haemocyte count (THC), phagocytosis, lysozyme activity (in both haemocyte lysate and cell-free haemolymph), activity and expression of the antioxidant enzyme superoxide dismutase (SOD) (in both haemocyte lysate and cell-free haemolymph) were chosen as biomarkers of exposure to high temperatures. The survival-in-air test was also performed. During the experiment, the clams showed differing burrowing behaviour: the animals kept at 20 and 25 degrees C burrowed completely, whereas at 30 degrees C the clams progressively emerged from the sediment and then remained on the surface. The highest temperature significantly increased THC, whereas it decreased the phagocytic activity of haemocytes. The haemocyte size frequency distribution in clams kept at 30 degrees C showed that the cell population of about 8-10microm was markedly reduced compared to clams kept at 20 and 25 degrees C. In clams maintained at 25 degrees C, lysozyme activity was significantly increased in haemocyte lysate, whereas it was markedly decreased in cell-free haemolymph. Total SOD activity significantly decreased in haemocytes from clams held at 30 degrees C whereas it increased in cell-free haemolymph from clams held at 25 degrees C and 30 degrees C. A significant decrease in haemocyte Mn-SOD and Cu/Zn-SOD activities was found with increasing temperature. In cell-free haemolymph, the highest Mn-SOD activity was recorded at 30 degrees C, whereas the Cu/Zn-SOD activity showed no significant changes in clams maintained at different temperatures. SOD isoform expression exhibited different patterns in haemocyte lysate and cell-free haemolymph. The resistance to air exposure of clams kept at 30 degrees C was shown to decrease significantly, LT(50) values fell from 6days in clams kept at 20 degrees C and 25 degrees C to 4days in those kept at 30 degrees C.     

The effect of benthic prey abundance and size on red knot (Calidris canutus) distribution at an alternative migratory stopover site on the US Atlantic Coast

A population decline of the western Atlantic red knot (Calidris canutus rufa) has been linked to food limitation during the spring migratory stopover in Delaware Bay, USA. The stopover ecology at potential alternative sites has received little attention. We studied factors affecting red knot habitat selection and flock size at a coastal stopover site in Virginia in 2006–2007. The most common potential prey items were coquina clams (Donax variabilis) and crustaceans. Red knot foraging sites had more clams and crustaceans than unused sites in 2006. Prey abundance increased during the 2007 stopover period and remained high after the red knot peak. Red knot flock size in 2007 increased with mean clam shell length, and probability of flock presence decreased with increasing distance from night use locations. Our results suggest that red knots preferred coquina clams and that these clams were not depleted during the stopover period in 2007. Thus prey abundance did not appear to be a population-limiting factor at this coastal stopover site in Virginia in that year. Protection of coastal sites outside of Delaware Bay, many of which have been altered by human development, would likely benefit red knot population recovery, as they can apparently provide abundant food resources during at least some years.     

Occurrence of Perkinsus olseni (Protozoa: Apicomplexa) and other parasites in the venerid commercial clam Pitar rostrata from Uruguay, southwestern Atlantic coast

A study was conducted into the health status of natural populations of the venerid clam Pitar rostrata from Uruguay. Perkinsus sp. was detected in 22% of the clams. Severe hemocytic infiltration was detected in the tissues parasitized by this protozoan parasite. The sequencing of the ITS-5.8S gene cluster of the parasite confirmed that it belonged to the Perkinsus olseni species. Rickettsia or Chlamidia-like organisms were also found, with a prevalence of 11%, although without apparent host reaction; an unidentified species of Coccidia was found in the nephridia of 78% of the clams, with the intensity of infection ranging from moderate to high. A gregarine, Nematopsis-like organism was observed mainly in the epithelial cells of the intestine, without host response and with a prevalence of 56%. Of the metazoan parasites, trematodes were found in 11% of the individuals analyzed.     

Effects of temperature on hard clam (Mercenaria mercenaria) immunity and QPX (Quahog Parasite Unknown) disease development: II. Defense parameters

Quahog Parasite Unknown (QPX) is a protistan parasite affecting hard clams Mercenaria mercenaria along the Northeastern coast of the United States. The geographic distribution and occurrence of disease epizootics suggests a primary role of temperature in disease development. This study was designed to investigate the effect of temperature on constitutive and QPX-induced defense factors in M. mercenaria. Control and QPX-challenged (both experimentally and naturally) clams were maintained at 13, 21 and 27 °C for 4 months. Control and experimentally-infected clams originated from a southern broodstock (Florida, no prior reports of disease outbreak) while naturally-infected clams originated from a northern broodstock (Massachusetts, enzootic area). Standard and QPX-specific cellular and humoral defense parameters were assessed after 2 and 4 months. Measured parameters included total and differential hemocyte counts, reactive oxygen species production, phagocytic activity of hemocytes, lysozyme concentration in plasma, anti-QPX activity in plasma and resistance of hemocytes to cytotoxic QPX extracellular products. Results demonstrated a strong influence of temperature on constitutive clam defense factors with significant modulation of cellular and humoral parameters of control clams maintained at 13 °C compared to 21 and 27 °C. Similarly, clam response to QPX challenge was also affected by temperature. Challenged clams exhibited no difference from controls at 27 °C whereas different responses were observed at 21 °C and 13 °C compared to controls. Despite differences in infection mode (experimentally or naturally infected) and clam origin (northern and southern broodstocks), similarities were observed at 13 °C and 21 °C between QPX infected clams from Florida and Massachusetts. Clam response to temperature and to QPX exhibited interesting relationship with QPX disease development highlighting major influence of temperature on disease development.     

Epibiosis and parasitism of coquina clam Donax spp.: Location,location, location!

Cryptic species of coquina clams Donax fossor and D. variabilis carry the hydroid epibiont Lovenella gracilis and are infected with metacercariae of the monorchiid parasites Lasiotocus trachinoti and L. choanura. The associations among this host–epibiont–parasite system were investigated. Fifty clams were collected at low tide over 3 days in June 2020 in South Carolina from each of three groups: clams with no hydroid from the upper intertidal zone and clams with and without hydroids from the swash zone. Clams were measured, identified using a newly developed PCR-RFLP, and examined for infection by metacercariae. Parasites were identified based on cercarial morphology and on metacercariae habitat in the clams. D. fossor was most often found in the swash zone and D. variabilis in the upper intertidal zone. The hydroid was most often associated with D. fossor, which was more infected by both digeneans than D. variabilis. Mean abundance of metacercariae of L. choanura was higher than that of L. trachinoti in both clams and increased over time for both parasites, because higher infection was correlated with larger clams. Greater time spent in the water by individuals of D. fossor appears to best explain these results, with the presence of the hydroids also being associated with higher infection by metacercariae in this coquina. Integration of D. variabilis in both digenean life cycles appears to lead to a positive outcome for the parasites as prevalence and abundance of infection were high; however, because D. variabilis is most frequent in the upper intertidal, more emersed, zone, it is likely deleterious to the epibiont to establish on this clam.     

Mass mortality of the invasive bivalve <Emphasis Type="Italic">Corbicula fluminea</Emphasis> induced by a severe low-water event and associated low water temperatures

The Asian clam Corbicula fluminea, originating from Southeast Asia, was first recorded in Lake Constance in 2003 and developed local mass occurrences afterwards. Effects of harsh winter conditions in 2005/2006 associated with a strongly decreasing water level were studied at three different depths at and below the mean low water level (MLL, MLL ?1 m and MLL ?3 m). Low winter temperatures produced a massive die-off of the C. fluminea population. The mortality of the clams was size class and depth dependent. At the mean low water level (MLL), all clams died because of lying dry. However, at MLL ?1 m and at MLL ?3 m, mortality was a consequence of water temperatures around 2°C for nearly 3 months. At MLL ?3 m, clams >5 mm died later than young clams <5 mm and later than clams of all sizes at MLL ?1 m. But in late spring even the clams >5 mm at MLL ?3 m were dead and only about 1% of the overall population of C. fluminea survived the winter conditions until spring 2006.Lethal effects of low water temperatures on C. fluminea, which may become effective only after a time lag, were corroborated in an outdoor mesocosm experiment with constant water level and without predation.     

Gymnophallid digenean Parvatrema duboisi uses Manila clam as the first and second intermediate host

To identify the metacercariae of a gymnophallid trematode in the Manila clam Ruditapes philippinarum from the Ariake Sea, experimental infection and molecular analysis were conducted. Based on the morphology of adult worms obtained from experimentally infected mice, the parasite was identified as Parvatrema duboisi. Comparison of internal transcribed spacer (ITS) region of ribosomal DNA (rDNA) sequences for metacercariae and sporocysts from Manila clams and adult worms collected from wild wigeon Anas penelope showed minor differences ranging from 0 to 0.8%. These data strongly suggest that in the Ariake Sea, the parasite has a lifecycle using the Manila clam as the first and second intermediate hosts and wigeon as the definitive host.     

Behavioural interactions between ecosystem engineers control community species richness

Behavioural interactions between ecosystem engineers may strongly influence community structure. We tested whether an invasive ecosystem engineer, the alga Caulerpa taxifolia , indirectly facilitated community diversity by modifying the behaviour of a native ecosystem engineer, the clam Anadara trapezia , in southeastern Australia . In this study, clams in Caulerpa -invaded sediments partially unburied themselves, extending >30% of their shell surface above the sediment, providing rare, hard substrata for colonization. Consequently, clams in Caulerpa had significantly higher diversity and abundance of epibiota compared with clams in unvegetated sediments. To isolate the role of clam burial depth from direct habitat influences or differential predation by habitat, we manipulated clam burial depth, predator exposure and habitat ( Caulerpa or unvegetated) in an orthogonal experiment. Burial depth overwhelmingly influenced epibiont species richness and abundance, resulting in a behaviourally mediated facilitation cascade. That Caulerpa controls epibiont communities by altering Anadara burial depths illustrates that even subtle behavioural responses of one ecosystem engineer to another can drive extensive community-wide facilitation.     

Clams (Corbicula fluminea) as bioindicators of fecal contamination with Cryptosporidium and Giardia spp. in freshwater ecosystems in California

This study evaluated clams as bioindicators of fecal protozoan contamination using three approaches: (i) clam tissue spiking experiments to compare several detection techniques; (ii) clam tank exposure experiments to evaluate clams that had filtered Cryptosporidium oocysts from inoculated water under a range of simulated environmental conditions; (iii) sentinel clam outplanting to assess the distribution and magnitude of fecal contamination in three riverine systems in California. Our spiking and tank experiments showed that direct fluorescent antibody (DFA), immunomagnetic separation (IMS) in combination with DFA, and PCR techniques could be used to detect Cryptosporidium in clam tissues. The most analytically sensitive technique was IMS concentration with DFA detection of oocysts in clam digestive gland tissues, which detected 10 oocysts spiked into a clam digestive gland 83% of the time. In the tank experiment, oocyst dose and clam collection time were significant predictors for detecting Cryptosporidium parvum oocysts in clams. In the wild clam study, Cryptosporidium and Giardia were detected in clams from all three study regions by IMS-DFA analysis of clam digestive glands, with significant variation by sampling year and season. The presence of C. parvum DNA in clams from riverine ecosystems was confirmed with PCR and DNA sequence analysis.     

Effects of coral transplantation and giant clam restocking on the structure of fish communities on degraded patch reefs

Active restoration is being practiced to supplement conservation activities for the purpose of reversing the trend of reef degradation. In the last decade, the feasibility of different restoration approaches such as coral transplantation and restocking of other marine biota has been the focus of research and relatively few have examined experimentally its effects on the resultant communities. In this study, coral transplantation and giant clam restocking were applied on 25 degraded patch reefs (~ 25 m²) inside a marine sanctuary in Pangasinan, northwestern Philippines to examine their effects on the community structure of reef fishes. Five interventions or treatments were employed: 1) “coral” consisted of transplantation of a combination of Acropora spp. and Pocillopora spp. on concrete blocks; 2) “clam” consisted of restocking of Tridacna gigas; 3) “clam+coral” consisted of restocking of T. gigas with Acropora spp. transplanted on their shells; 4) “shell” consisted of deployment of T. gigas shells; and 5) “control” consisted of no intervention. Fish communities on the patch reefs were monitored monthly for 3 months before the intervention and were monitored further for 11 months after the intervention, including 1 recruitment season. After the intervention, the coral cover and the “other biota” category increased in the coral and clam+coral treatments, due to the transplanted corals and deployed giant clams. Consequently, the complexity of the substrate was enhanced. A month after the intervention, a rapid increase in the abundance and species richness of reef fishes on the coral, clam+coral and clam treatments was observed compared to the shell and control treatments. A change in species composition of reef fish assemblage was also apparent in the coral and clam+coral treatments relative to the clam, shell and control, especially 4 months after the intervention. The present experiment demonstrates the feasibility of improving the condition of degraded patch reefs, which can subsequently enhance the fish community. Results also show the importance of the underlying substratum and the abundance of live corals and clams to reef fishes.     

A coccidian infection of the kidney of the native littleneck clam, Protothaca staminea

During the summer of 1980, numerous exposed native littleneck clams (Protothaca staminea) were observed in Sequim Bay, Washington. The kidneys of these clams were found to be parasitized by a coccidian. Heavily parasitized kidney was characterized by massive epithelial destruction and an intense inflammatory response. In addition, large numbers of “brown cells” were present throughout the tissues of infected clams.Infected kidneys harbored several stages of the life history of the parasite. The present coccidian differs markedly in several respects from other coccidians in bivalves. First, the entire life history apparently occurs in the kidney of the clam, in contrast to the presumed involvement of an alternative host in other known coccidian parasites of bivalves. Second, a thick-walled spore is produced in addition to a thin-walled oocyst. The apparent presence of two types of dissemination stages is unknown in the Coccidia. Finally, the sporocysts are tetrazoic as opposed to dizoic or multiple sporocysts in previously described species. This parasite, therefore, appears to represent an undescribed genus and species of Coccidia.     

Movement and aggregation in the fluted giant clam (Tridacna squamosa L.)

Aquaculture has been the traditional focus of tridacnid giant clam research whereas their ecology and behaviour have received much less attention. This study was based on the observation that juvenile fluted giant clams (Tridacna squamosa), when evenly distributed in a tank, will move and aggregate over time. We observed movement in clams ranging from 10 to 313 mm in shell length and ‘climbing’ up the sides of tanks was noted for clams with lengths between 10 and 22 mm. Locomotion also occurred after byssal attachment to the substrate; there was a highly significant association between type of movement (i.e. translation, rotation and no movement) and presence of attachment. Tests for phototaxis were negative. Aggregation was examined by placing clams in regular patterns on grids. After three days in the aquarium and 24 h in the field, their positions were analysed to obtain a statistical parameter for ‘clumpiness’. This was found to be greater in the live clam runs than both random walk and random distribution simulations, suggesting that clams were attracted to conspecifics. The latter was tested by recording clam movement with respect to five types of fixed ‘targets’ (i.e. live clam, fouled clam shell, foul-free clam shell, random inanimate object and none). The test clams moved, non-significantly, towards live clam targets and displayed higher mobility compared to tests with other target objects; a negative correlation between mobility and clam length was also observed. A choice experiment using bidirectional water inflow with clam effluent as one source resulted in clams moving toward the effluent, offering the first direct support for positive chemotaxis among conspecifics in Bivalvia. Together, our results indicate the presence of chemical signalling among clams, leading to movement toward one another and clumping. Aggregation could serve several ecological functions, such as defence against predation, physical stabilisation and facilitation of reproduction. With worldwide decline in natural giant clam densities, the opportunity for conspecific clumping is reduced, and local stocks could be facing increased vulnerability to Allee affects.     

Biology of Microplitis kewleyi (Hymenoptera: Braconidae): A parasite of Agrotis ipsilon (Lepidoptera: Noctuidae) larvae

Microplitis kewleyi Muesebeck is a gregarious internal parasite of larvae of the black cutworm Agrotis ipsilon (Hufnagel). Studies of the biology of the parasite revealed that there was an inverse relationship between host instar and parasite preference. Duration of development from egg to pupa ranged from 18 days at 27°C to 68.7 days at 16°C. Development from egg to pupa took 13.5–21.6 days when fourth and first instar host larvae, respectively, were parasitized. A larger number of parasites emerged from hosts parasitized in the fourth instar (22.4) than the first instar (11.5). Parasite pupation occurred when the host was in the fifth/sixth instar, depending on the instar parasitized. Thirty‐nine per cent of host larvae exposed as first instars to parasites died before parasite emergence. This decreased to 0% for host larvae exposed as fourth instars. The sex ratio was 1:1.2 (M:F). Thirty‐seven per cent of hosts exposed diurnally were stung, compared to 24% exposed nocturnally. Mean daily progeny was highest (12) on the first day, decreasing to zero after 20 days. Percent host parasitism was also highest on the first day (35%) decreasing to nearly 0% after 18 days. There appear to be three parasite larval instars. Host larvae often remained alive after parasite emergence.