A reconsideration of the origins of the 'typical' Cretaceous inoceramid calcitic hinge plate in the light of new ultrastructural observations from some Jurassic 'inoceramids'

Abstract:  Marked differences occur between the hinge plates of Jurassic and Upper Cretaceous British 'inoceramid' species. Albian and Upper Cretaceous inoceramids had the poiriform calcite hinge plate with closely spaced resiliifers that is widely associated with this extinct bivalve family. The whole structure was dominated by compressible ligament. Jurassic species, in contrast, had an ultrastructurally less familiar aragonite hinge plate with widely spaced resiliifers and a ligament reminiscent of Isognomon with equally spaced elastic and compressional components. The wider taxonomic relationships of the Inoceramidae still remain uncertain, although the majority of the evidence links the family to the Pterioida. This study, however, suggests that the ancestral inoceramid did not necessarily have a calcitic hinge plate, but rather that a Jurassic 'inoceramid' species could have been developing such a structure via an aragonite precursor. Observations on British Kimmeridgian 'inoceramid' specimens indicate that calcite was being incorporated into the aragonite hinge plate by these times. Structures in the ligamentat of Jurassic species suggest that the fundamental change in mineralogy was initiated by valve architectural modification at the byssal notch, linked to inoceramid epifaunalism. Worldwide poor preservation of Lower Cretaceous inoceramids makes it very difficult to further prove this hypothesis via intermediate evolutionary stages leading to the fully poiriform calcite hinge plate observed in Albian and Upper Cretaceous species of Inoceramidae.     

Spatial and temporal patterns of parthenogenesis and parasitism in the freshwater snail Melanoides tuberculata

The Red Queen hypothesis predicts that sex should be more common in populations heavily infested with parasites, than in those without. This hypothesis was investigated in the aquatic snail Melanoides tuberculata, in which both sexual and parthenogenetic individuals exist in natural populations, and some populations are heavily infested by trematodes. The presence of fertile males and the higher genetic diversity of bisexual populations are indicative of sexual reproduction. We compared sites in 1990, 1999, and 2001, and we looked for a positive correlation between male and parasite frequencies. Male frequency was not correlated with the frequency of individuals infected by trematodes. This lack of correlation was reconfirmed in a retrospective power analysis. In a period of 9 years, male frequencies decreased but infection levels increased. These results do not support the Red Queen hypothesis. In samples with high male frequency the number of embryos was low, perhaps indicating that males may have a negative effect on embryo numbers. This effect of males on fitness could perhaps suggest that the cost of sex is fewer embryos. The reduction in embryo numbers may also represent a trade-off between mating and egg production costs.     

A quantitative estimation of the role of bithyniidae snails (gastropoda: Prosobranchia) in the ecosystems of the southern part of Western Siberia (Russia)

This study describes the species diversity, abundance and biomass of gastropods in the ecosystems of the southern part of Western Siberia (Karasuk River and Krotovo Lake, Karasukskii district, Novosibirsk Oblast). Eighteen species belonging to seven families were recorded, Lymnaeidae, Planorbidae, Bulinidae, Bithynhdae, Physidae, Succineidae, and Zonitidae. The percentage of Bithyniidae snails was assessed in comparison to other gastropods and in regard to the species diversity, abundance, and biomass. The role of Bithyniidae mollusks as a host for trematodes was assessed. The biodiversity of mollusks and trematodes was studied using the Shannon-Weaver index.     

Faster clonal turnover in high‐infection habitats provides evidence for parasite‐mediated selection

According to the Red Queen hypothesis for sex, parasite‐mediated selection against common clones counterbalances the reproductive advantage of asexual lineages, which would otherwise outcompete sexual conspecifics. Such selection on the clonal population is expected to lead to a faster clonal turnover in habitats where selection by parasites is stronger. We tested this prediction by comparing the genetic structure of clonal and sexual populations of freshwater snail Potamopyrgus antipodarum between years 2003 and 2007 in three depth‐specific habitats in Lake Alexandrina (South Island, New Zealand). These habitats differ in the risk of infection by castrating trematodes and in the relative proportion of sexual individuals. As predicted, we found that the clonal structure changed significantly in shallow and mid‐water habitats, where prevalence of infection was high, but not in the deep habitat, where parasite prevalence was low. Additionally, we found that both clonal diversity and evenness of the asexual population declined in the shallow habitat. In contrast, the genetic structure (based on F–statistics) of the coexisting sexual population did not change, which suggests that the change in the clonal structure cannot be related to genetic changes in the sexual population. Finally, the frequency of sexuals had no effect on the diversity of the sympatric clonal population. Taken together, our results show a more rapid clonal turnover in high‐infection habitats, which gives support for the Red Queen hypothesis for sex.     

Age-related characteristics of the infestation of populations of the littoral periwinkles Littorina obtusata and L. saxatilis by trematode parthenitae

Analysis of the age structure of parthenitae of trematodes of sympatric populations of L. obtusata and L. saxatilis was conducted in 1983 for 6 localities in Kandalaksha Bay of the White Sea. 10 species of trematodes were found. The main populational differences in the level of infection are caused by Microphallus piriformes. Species with secondary dispersion in the life cycle are characterized by the increase in the extent of infection in populations with age. In Microphallidae species of "pygmaeus" group the age structure of infection depends on the overage infection of populations: the increase in the parasitic press leads to the maximum turning towards the young part of the snail's population. The establishment of the age structure of infection is considered to be one of the regulation mechanisms in parasite--host relations on the populational level.     

Hemocytes from neonates of Biomphalaria glabrata are functionally less mature than those from adult snails

Gastropod molluscs, which serve as obligatory intermediate hosts for digenetic trematodes, possess an internal defense system (IDS) consisting of phagocytic hemocytes and plasma factors. This IDS is responsible for resistance to infection with larval trematodes, which are encapsulated and killed by hemocytes in incompatible snails. Like other physiological systems, the IDS probably undergoes maturation during early stages of life, and the relatively undeveloped state of the IDS in young snails has been hypothesized to be a factor in their increased susceptibility to infection with larval trematodes. In this study, hemocytes were examined in the BS-90 laboratory strain of Biomphalaria glabrata that is resistant to infection with Schistosoma mansoni as adults but susceptible to infection as neonates. Compared with hemocytes from adults, hemocytes from neonates had a smaller perimeter and lower intrinsic directional motility on glass microscope slides. Additionally, in vitro assays showed a lower association with fucoidan-linked polystyrene beads and less ability to produce superoxide anion in hemocytes from neonates compared to hemocytes from adults. These results support the hypothesis that the gastropod IDS undergoes maturation during growth. However, whether the observed differences between hemocytes of neonatal and adult BS-90 snails play a role in the susceptibility of the former and resistance of the latter to infection with S. mansoni is not known.     

Differences in the number of hemocytes in the snail host Biomphalaria tenagophila, resistant and susceptible to Schistosoma mansoni infection

The relationships between schistosomiasis and its intermediate host, mollusks of the genus Biomphalaria, have been a concern for decades. It is known that the vector mollusk shows different susceptibility against parasite infection, whose occurrence depends on the interaction between the forms of trematode larvae and the host defense cells. These cells are called amebocytes or hemocytes and are responsible for the recognition of foreign bodies and for phagocytosis and cytotoxic reactions. The defense cells mediate the modulation of the resistant and susceptible phenotypes of the mollusk. Two main types of hemocytes are found in the Biomphalaria hemolymph: the granulocytes and the hyalinocytes. We studied the variation in the number (kinetics) of hemocytes for 24 h after exposing the parasite to genetically selected and non-selected strains of Biomphalaria tenagophila, susceptible or not to infection by Schistosoma mansoni. The differences were analyzed referred to the variations in the number of hemocytes in mollusks susceptible or not to infection by S. mansoni. The hemolymph of the selected and non-selected snails was collected, and hemocytes were counted using a Neubauer chamber at six designated periods: 0 h (control, non-exposed individuals), 2 h, 6 h, 12 h, 18 h and, 24 h after parasite exposure. Samples of hemolymph of five selected mollusks and five non-selected mollusks were separately used at each counting time. There was a significant variation in the number of hemocytes between the strains, which indicates that defense cells have different behaviors in resistant and susceptible mollusks.     

The influence of the degree of infestation with trematode parthenites on the structure of penial glands in the mollusks Littorina saxatilis of different age

Histological and confocal microscopy studies of the structure of penial glands in the Littorina saxatilis males were carried out. The examined mollusks belong to two age groups and were at different stages of spontaneous infection with a trematode from the pygmaeus species group (Microphallus piriformes). Based on comparative analysis of microscopic sections of copulative organs in infested and non-infested mollusks, data on the modifications in histological structure ofpenial glands were obtained. From these data we can suggest that the infestation have an influence on all parts of the gland. Decrease of secret production and reduction of muscular capsule (down to its disappearance) were observed. We suppose that changes in the penial glands structure prevent their normal functioning. In the mollusks infested on reaching the sexual maturity, gradual reduction of distal part of reproductive system was observed, while in the mollusks infested before the sexual maturity, development of ancillary part of reproductive system was blocked.     

Population structure, parasitism, and survivorship of sexual and autodiploid parthenogenetic Campeloma limum

Two theories for the maintenance of sexual reproduction, the Red Queen hypothesis and mutation accumulation, suggest that the dispersal rates of sexuals and asexuals may determine the elimination or persistence of asexuals. Under higher dispersal rates of asexuals, asexuals may temporarily escape virulent parasites and reduce the effects of deleterious mutations. In the present study, I examine the population structure, parasite loads, and juvenile survivorship of Campeloma limum sexuals and autodiploid parthenogens from the southeastern U.S. Atlantic coastal plain. Using mtDNA sequence variation, it is shown that parthenogenetic haplotypes with limited sequence divergence are geographically widespread throughout this region and there is no significant population differentiation over a broad geographical scale. Sexual C. limum populations show significant mtDNA differentiation among and within river drainages and there is significant isolation by distance. These patterns are consistent with a recent origin and range expansion of parthenogens. Prevalence of infection by digenetic trematodes is significantly higher in autodiploid parthenogens, and the variance of prevalence is also higher in autodiploid parthenogens. I argue that the latter pattern indicates that unparasitized parthenogens have temporarily escaped these virulent parasites, but recolonization of these populations by trematodes results in high infection levels (> 40%), possibly due to reduced variation in resistance genes. I also examined whether the survivorship of juvenile sexuals and parthenogens varied under different stress levels. Sexual juveniles had twofold higher survivorship in all environments. Compared to polyploid parthenogens, autodiploid parthenogens may be less buffered against the effects of deleterious recessive alleles. I propose that the combined effects of higher parasitism and reduced juvenile survivorship of these autodiploid parthenogens accounts for the spatial distribution of sexual and parthenogenetic C. limum in the Atlantic coastal plain. Parthenogens may persist by higher dispersal rates into marginal habitats where there is a temporary escape from digenetic trematodes and competition with sexuals.     

Indirect effects of a parasite on a benthic community: an experiment with trematodes, snails and periphyton

1. Few studies have directly addressed the role played by parasites in the structure and function of ecosystems. Parasites influence the behaviour, reproduction and overall fitness of their hosts, but have been usually overlooked in community and ecosystem‐level studies. We investigated the effects of trematode parasites on snail–periphyton interactions. 2. Physa  acuta (Gastropoda: Pulmonata) snails infected with the trematode Posthodiplostomum minimum (often >30% of within‐shell biomass) grazed more rapidly than uninfected snails. Trematode effects on snail grazing indirectly affected the standing stock and community structure of periphyton. Populations of snails with 50% infected individuals reduced algal biomass by 20% more than populations with lesser (10% or 0%) infection rates. 3. The alga Cladophora glomerata dominated periphyton communities grazed by snail populations with 50% infection rates, whereas diatoms and blue–green algal taxa dominated when grazed by snail populations with lower infection rates. 4. Thus, trematodes indirectly affected periphyton communities by altering host snail behaviour, a trait‐mediated indirect effect. These results indicate that trematodes can indirectly influence benthic community structure beyond simple population fitness, with possible related effects on ecosystem function.     

Spatial and temporal variation of trematode infection in coexisting populations of intertidal gastropods Littorina saxatilis and L. obtusata in the White Sea

Trematode infection was studied in sympatric populations of the periwinkles Littorina saxatilis and L. obtusata in 2 regions of Kandalaksha Bay of the White Sea to assess host-parasite interactions at the population level. Twenty-seven spatially separated populations were each surveyed in 1984-1994; 2 heavily infected populations were investigated annually over a 16 yr period. Ten trematode species were found in the periwinkle populations. The closest association in spatial distribution and temporal dynamics was observed between 3 ecologically and morphologically similar trematodes of the 'pygmaeus' group: Microphallus piriformes, M. pygmaeus and M. pseudopygmaeus. For these 3 species, the prevalences were closely associated in the 2 host species when spatially separated sites from the 2 studied regions were considered, while in the 2 populations studied over the 16 yr period, a correlation was only observed between the infection levels of L. saxatilis and L. obtusata by either M. piriformes and immature microphallids. Likewise, within each host species, significant correlations were revealed between the prevalence of the different microphallids of the 'pygmaeus' groups. However, they were fewer and weaker when the long-term dynamics of infection in the 2 heavily infected populations were considered. Most other trematodes did not show significant association in prevalence either within or between the 2 host species on spatial and temporal scales.     

Encapsulation of the metacercariae of two species of trematodes in the skin and musculature of the stone cockscomb <Emphasis Type="Italic">Alectrias alectrolophus</Emphasis> (Pallas, 1814)

The structure of the coating surrounding the metacercaria of Cryptocotyle lingua in the skin and the metacercaria of Liliatrema skrjabini in the muscles of the stone cockscomb was studied. Metacercariae of these trematodes were surrounded by a cyst adjacent to the helminth and by a capsule. In both species, the cyst consisted of electron- dense homogeneous material, which was probably synthesized by the parasite. The capsules were formed from host tissues; in C. lingua, they mostly consisted of ordered layers of collagen fibers and in L. skrjabini they were formed from fibroblasts and, presumably, phagocytes. The differences in capsule structure of the studied metacercariae may be related to the physiological features of surrounding tissues or to the taxonomic attribution of the trematodes.     

The peculiarities of feeding of the American mink (<Emphasis Type="Italic">Neovison vison</Emphasis>) in the Southern Urals

The peculiarities of feeding of American mink in various waterbodies of the Southern Urals are described. The structure of the mink ration considerably depends on the type of water body. In all water bodies, fish, insects, and mammals were in the group of main forage; in lakes, mollusks or crayfish were added to this group. Birds and frogs in all water bodies were less important; reptiles (snakes) were accidental prey. It is revealed that the terrestrial mollusks played an important role in mink feeding. The differences in the structures of rations of the western and eastern populations of American mink were found.     

Long-term population dynamics of Littorina obtusata: the spatial structure and impact of trematodes

There are few examples of host population regulation by macroparasites in stable communities; however, strong impact of parasites on the host individual is obvious in many cases (for example increased mortality, a reduction in fecundity up to the complete castration). Associations between the host populations (periwinkle Littorina obtusata) and prevalence of trematodes was investigated using long-term data (1982–1997) of two L. obtusata populations in the White Sea, northwest Russia. We hypothesized that high prevalence of trematodes will reduce future host population density, and increase mortality. Using a general linear model, we found a significant negative correlation between host population density and the prevalence of the most abundant parasite, Microphallus piriformes in the previous year. We found no correlation between snail reproduction and the prevalence, but observed a significant reduction in middle-aged mollusk abundance which was associated with high prevalence. This indicates the importance of parasite-induced mortality for the dynamics of the host population. There was an association between trematode infection and L. obtusata populations that influence their distribution within littoral zone. The ‘source’ population, located in the lower section of the macrophyte zone, appears to be self-sustaining, controlling the whole population recruitment and dynamics.     

The distribution and incidence of larval trematodes in the freshwater fauna of the Wentloog level, South Wales

The Wentloog level, in South Wales, is a low-lying coastal plain dissected by a network of drainage channels. The latter contain a rich aquatic invertebrate fauna which harbour a wide range of larval digenetic trematodes. Few of the host species examined were not infected with larval trematodes. Miracidia together with their associated sporocysts, rediae and cercariae were more host specific than metacercariae but many of the miracidia which developed in pulmonate molluscs could do so in more than one genus. Metacercarial host specificity is less rigid, and usually species are only restricted to members of one phylum although several were found which could develop in two or more phyla. Most larval stages showed some host preference when they occurred in several host species and this was attributed to ecological or physiological factors or to mechanical barriers. Seasonal variation of infection rate occurs in all larval stages: infection with developing cercariae showed peaks during June and August, but metacercarial infection rates varied according to the host. In molluscs there may be two peaks of infection with metacercariae (as in the cercarial stages), but for many arthropods infections show a gradual increase through the year. The factors mainly responsible for these seasonal changes are considered to be the life-cycle of the intermediate hosts and the environmental temperature, the latter acting on the rate of development of the trematodes. Rainfall, and patterns of egg production of adult trematodes in the definitive hosts are also important.     

Feeding behavior,ration and size structure of a population of the predatory gastropod <Emphasis Type="Italic">Cryptonatica janthostoma</Emphasis> in Vostok Bay,Sea of Japan

The relationships between the shell height of the predatory gastropod Cryptonatica janthostoma and the shell length of its typical prey, the bivalve Ruditapes philippinarum, and the diameter of the borehole on the prey shell resulting from a successful attack of the predator were experimentally found and assessed statistically. The shell height of C. janthostoma calculated retrospectively from the borehole diameter using the obtained relationships was 17–52 mm. The prey of C. janthostoma are burrowing bivalves, whose populations are affected by the predator to a varying degree. In populations of medium-sized mollusks (R. philippinarum, Protothaca euglypta, P. jedoensis, and others), C. janthostoma feeds on mollusks larger than 7–10 mm; in species with a shell length greater than 100 mm (Callista brevisiphonata, Saxidomus purpuratus), it eats specimens of 10–58 mm. C. janthostoma apparently has no effect on populations of small-sized mollusks (Anisocorbula venusta) and mollusks with an active avoidance response (Clinocardium californiense).     

Parasites of Pleurobrachia pileus Muller, 1776 (Ctenophora), from Galway Bay, western Ireland

Specimens of Pleurobrachia pileus Müller from Galway Bay,western Ireland, were found to be parasitised by three trematodes,one isopod and one nematode. Among these, the trematode, Opechonabacillaris Molin and didymozoid larvae were the most abundant,infecting over 17% of the Pleurobrachia population. Peak infectionwas during the summer of each year when >40% of P. - pileuswere parasitised. Serious infection was found to be either accompaniedor followed by a sharp decline in the abundance of the ctenophore.A close, but negative, correlation was established between theabundance of P. pileus and the percentage of parasitic infection.The percentage of infection in larger P. pileus was higher thanin small specimens. No parasites were detected from P. pileus1 mm or less in length.     

Role of cell-cycle in regulating neuroepithelial cell shape during bending of the chick neural plate

Summary Neuroepithelial cells transform from spindle-shaped to wedge-shaped within the median and paired dorsolateral hinge points of the bending neural plate, but the mechanisms underlying these localized changes are unclear. This study was designed to evaluate further the hypothesis that localized wedging of neuroepithelial cells during bending involves basal cellular expansion resulting from alteration of the cell-cycle. Neurulating chick embryos were treated with tritiated thymidine, and transverse sections through the midbrain were examined autoradiographically. Parameters of the cell-cycle as well as nuclear position and size were assessed in the median hinge point, which contains predominantly wedge-shaped cells, and in adjacent lateral areas of the neural plate, which contain predominantly spindle-shaped cells. Both the DNA-synthetic phase and non-DNA synthetic portion of the cell-cycle were significantly longer in the median hinge point than in lateral neuroepithelial areas, some nuclei in both regions were located basally during these phases, and virtually all basal nuclei in the median hinge point were large. Additionally, the mitotic phase was significantly shorter in the median hinge point than in lateral areas. We present a model to explain how alteration of the cell-cycle in the median hinge point could generate wedging of cells in this region.     

The relationship between specialization and local abundance: the case of helminth parasites of birds

Positive relationships are commonly observed between the abundance of a species in a locality and the frequency of its occurrence among localities on a larger scale. This pattern may not hold for parasitic organisms when the average abundance of a parasite among its hosts is related to the number of host species in which it occurs, because of the additive investment in specific adaptations to counter host immune responses required for each host species in a parasites repertoire. For a rigorous test of the hypothesis that there is a trade-off between the number of host species that can be successfully exploited and the average abundance of parasites in those hosts, one needs to take into account the phylogenetic (or taxonomic) distances among the host species used by a parasite. Differences in immune responses are likely to increase with increasing phylogenetic distances. The trade-off hypothesis was tested in a comparative analysis of 393 species of trematodes, cestodes and nematodes parasitic in birds surveyed from the same geographical area, using an index of host specificity that measures the average taxonomic distances between a parasites known host species. After correcting for the influences of parasite phylogeny and other potential confounding variables, mean abundance was negatively correlated with the average taxonomic distance among host species for nematodes, and with the variance in taxonomic distances among hosts for cestodes. In the case of trematodes, these variables covaried positively. The trade-off between average infection success and how taxonomically distant a parasites host species are from each other was only found in two of the three groups of helminths investigated, possibly because of compensating features in trematodes, such as their ability to multiply asexually in intermediate hosts. These results provide empirical evidence consistent with the hypothesis that specialization allows greater local adaptation and therefore greater local population abundance, supporting key predictions regarding the evolution of ecological specialization.Electronic Supplementary Material Supplementary material is available for this article at     

Patterns of individual growth in the field slug <Emphasis Type="Italic">Deroceras agreste</Emphasis> (Gastropoda,Agriolimacidae)

The growth of individual Deroceras agreste slugs from the date of sampling to the end of life has been studied under laboratory conditions. Seven out of 37 mollusks (≈19%) have shown a finite growth pattern, with their body weight reaching a maximum and decreasing thereafter. The data on these mollusks are well approximated by the finite growth equation. Other mollusks have not reached the maximum weight. In many cases, periods of a sharp decrease in body weight with subsequent return to the initial growth curve (compensatory growth) have been observed. In three animals, the growth curve has changed after such a decrease. A comparison of growth parameters in different individuals with the aid of the Ford-Wallford equation has shown that differences between them lack statistical significance. In generally, the slug D. agreste appears to belong to animals with the finite growth pattern.