The fine structure of the germinal mass, brood cavity and birth canal of the rediae of the monoxenous hemiuroid digenean Bunocotyle progenetica Chabaud & Buttner, 1959.

Bunocotyle progenetica is a hemiuroid digenean whose sexual adults become fully developed and lay their eggs inside the rediae in the molluscan host. In this study, the fine structure of the germinal mass, brood cavity and birth canal in the B. progenetica rediae was examined using transmission electron and confocal microscopy. The large germinal mass attached to the body wall has a cellular composition typical for this organ. The characteristic traits of this germinal mass are weakly developed supporting tissue and the presence of deep lacunae opening into the brood cavity. These lacunae presumably participate in feeding the deeply lying embryos and facilitate their release into the brood cavity. The germinal mass is also characterized by intensive degeneration of cellular elements, which may represent a mechanism controlling the offspring number, limited in this species by the size of the redial brood cavity. The brood-cavity lining consists of flattened cells bearing lamellar projections and is connected anteriorly with the epithelium of the birth canal. The brood-cavity musculature, which is well developed in other hemiuroid digeneans, is significantly reduced in B. progenetica, most likely because their cystophorous cercariae remain inside the rediae, removing the need for muscle contractions pushing them through the brood cavity. The birth canal comprises three regions distinguished by the structure of the lining and muscle arrangement. The comparison of rediae of B. progenetica with parthenitae of other digeneans has shown that the organization of the redial reproductive apparatus in this species may have been influenced by life-cycle modification.

Patterns and processes influencing helminth parasites of Arctic coastal communities during climate change.

This review analyses the scarce available data on biodiversity and transmission of helminths in Arctic coastal ecosystems and the potential impact of climate changes on them. The focus is on the helminths of seabirds, dominant parasites in coastal ecosystems. Their fauna in the Arctic is depauperate because of the lack of suitable intermediate hosts and unfavourable conditions for species with free-living larvae. An increasing proportion of crustaceans in the diet of Arctic seabirds would result in a higher infection intensity of cestodes and acanthocephalans, and may also promote the infection of seabirds with non-specific helminths. In this way, the latter may find favourable conditions for colonization of new hosts. Climate changes may alter the composition of the helminth fauna, their infection levels in hosts and ways of transmission in coastal communities. Immigration of boreal invertebrates and fish into Arctic seas may allow the circulation of helminths using them as intermediate hosts. Changing migratory routes of animals would alter the distribution of their parasites, facilitating, in particular, their trans-Arctic transfer. Prolongation of the seasonal 'transmission window' may increase the parasitic load on host populations. Changes in Arctic marine food webs would have an overriding influence on the helminths' circulation. This process may be influenced by the predicted decreased of salinity in Arctic seas, increased storm activity, coastal erosion, ocean acidification, decline of Arctic ice, etc. Greater parasitological research efforts are needed to assess the influence of factors related to Arctic climate change on the transmission of helminths.

Genomic diversity of cercarial clones of Himasthla elongata (Trematoda, Echinostomatidae) determined with AFLP technique.

The aim of this study was to reveal genomic diversity formed during parthenogenetic reproduction of rediae of the trematode Himasthla elongata in its molluskan host Littorina littorea. We applied amplification fragment length polymorphism (AFLP) to determine the genomic diversity of individual cercariae within the clone, that is, the infrapopulation of parthenogenetic progeny in a single molluskan host. The level of genomic diversity of particular cercariae isolates from a single clone, detected with EcoR1/Mse1 AFLP reaction, was significantly lower than the variability of cercariae from different clones. The presence of intraclonal genomic diversity indicates a nonsexual shuffle of alleles during parthenogenesis in the rediae of H. elongata. The obtained polymorphic AFLP fragments were long enough to detect the sequences that may be responsible for clonal genomic variability. Based on this, AFLP can be recommended as a tool for the study of genetic mechanisms of this variability.

[PATTERNS IN CIRCULATION AND TRANSMISSION OF MARINE BIRD PARASITES IN HIGH ARCTIC: A CASE OF ACANTHOCEPHALAN POLYMORPHUS PHIPPSI (PALAEACANTHOCEPHALA, POLYMORPHIDAE)].

This study, based on the materials on parasitic infection of marine birds and invertebrates in Frantz Josef Land (FJL) collected in 1991-1993, focussed on the acanthocephalan Polymorphus phippsi. We identified this parasite, confirmed its species status and analysed its circulation and transmission patterns in high Arctic. The causes of its erroneous identification as P. minutus in several studies were also examined. In contrast to P. minutus, the transmission of P. phippsi is realized in marine coastal ecosystems. Its' main intermediate host in the Arctic is the amphipod Gammarus (Lagunogammarus) setosus, commonin coastal. areas of the shelf zone throughout the Arctic basin. P. phippsi population in FJL and the entire European Arctic is on the whole maintained by a single obligate final host, the common eider Somateria mollissima. Prevalence (P) of P. phippsi in this bird reached 100 %, with the maximal infection intensity (IImax) of 1188 and the mean abundance (MA) of 492.1. Other species of birds found to be infected with P. phippsi (Arctic turn, black guillemot, purple sandpiper and several gulls) are facultative and/or eliminative hosts. The most heavily infected birds were Arctic terns (P = 72.7%, IImax = 227, MA = = 47.1), which contained single mature acanthocephalans. For one of the FJL regions, infections flows of P. phippsi through various host categories were calculated. Involvement of birds unrelated to the common eider into the circulation of P. phippsi is facilitated by their feeding character in the Arctic. While coastal crustaceans are abundant, fish food is relatively scarce (polar cod, snailfishes), and so amphipods make up a considerable part of the diet of marine birds in FJL, if not most of it, as for instance in case of Arctic tern. This promotes an easy entry of the larvae of crustaceans-parasitizing helminthes (cestodes and acanthocephalans, including cystacanths P. phippsi) into non-specific hosts and opens broad colonization possibilities. Besides acanthocephalans, the phenomenon of non-specific parasitism has been shown for some cestodes circulating in the Arctic coastal ecosystems. Similar conditions for helminths transmission might have formed in marine coastal refugia during the glacial periods of late Pliocene-Pleistocene. According to the Arctic refugium hypothesis of Hoberg and Adams, this promoted parasitic colonization of phylogenetically distant hosts using similar foraging resources. Thus, present-day transmission patterns of helminthes in high Arctic can be, in a way, considered as a model allowing us to witness various stages of helminthes' speciation by host-switching.

[A case study of singular spectrum analysis application in parasitology: dynamics of prevalence of Cryptocotyle concavum and Bunocotyle progenetica trematode parthenitae in Hydrobia ventrosa snails at the White Sea].

In this study parasitological data were analyzed by different methods of revealing the structure of time series, namely auto-correlation analysis (ACA), Fourier spectrum analysis (SA) and singular spectrum analysis (SSA), and the results of these analysis were compared (SSA makes it possible to present non-stationary time series as a sum of independent components and to determine the contribution of each component into the dispersion of the initial series--Golyandina et al., 2001). This case study was based on the result of 10-year-long monitoring of changes in the prevalence of Cryptocotyle concavum and Bunocotyle progenetica trematode parthenitae in intertidal snails Hydrobia ventrosa at the White Sea (in total, 45 observations). ACA did not reveal any statistically significant oscillations in the analyzed series. The application of SSA and SA allowed us to reveal at least two quasi-periodical components. In addition, SSA made it possible to reveal a significant dome-shaped trend in the prevalence of B. progenetica parthenitae, which were described by SA as an oscillation with a period equal to the duration of the study, as well as to give proof that there was no trend in the changes of C. concavum parthenitae prevalence. The components (modes) extracted by the SSA described the changes in the prevalence better that the harmonics extracted by the SA. In particular, SSA modes (contrary to SA harmonics) reflected that the amplitude of oscillations of the B. progenetica prevalence increased as the prevalence grew. The sums of SSA modes correlated more with initial prevalence series that the sums of SA harmonics. A possible interpretation of the trends and modes extracted by the SSA in the light of the transmission features of the investigated trematode species in the study area was proposed.

In vitro encystment of Himasthla elongata cercariae (Digenea, Echinostomatidae) in the haemolymph of blue mussels Mytilus edulis as a tool for assessing cercarial infectivity and molluscan susceptibility.

Infectivity of Himasthla elongata cercariae to mussels, their second intermediate hosts, and resistance by these hosts to infection were assessed on the basis of the cercariae's ability to encyst in mussel haemolymph in vitro. A series of experimental in vivo infections of mussels with batches of cercariae, each batch released from a different single infected mollusc and referred to as a clone (due to their shared genotype), demonstrated that the results of the in vitro tests corresponded to the actual indices of infectivity/susceptibility of the parasites and their hosts. Most cercarial clones had high infectivity, with a few clones having very high or, at the other extreme, very low infectivity. A similar pattern was revealed in mussel resistance to cercarial infection. Most of the molluscs tested were moderately susceptible to cercarial infection, but at each extreme a small fraction (less than 10%) displayed very high or very low susceptibility. It was shown that there were no totally compatible or totally incompatible 'cercaria clone/mussel' combinations. Results obtained are compared with the data on intra-population variability using the characters parasite infectivity/host compatibility for trematode/mollusc-first intermediate host associations. Results are made relevant to actual infection levels in mussel settlements at the White Sea.

[Clonal variability in longevity of the cercariae of Himasthla elongata (Trematoda: Echinostomatidae)].

The study was carried out on Himasthla elongata cercariae shed by infected Littorina littorea snails. The infected periwinkles were collected from the settlement with the low prevalence of H. elongata. As shown earlier with the use of AFLP (Amplified Fragment Length Polymorphisms) method, rediae groups in all the infected periwinkles of this settlement arise from the infection of a mollusc with a single miracidium. Therefore, the cercariae shed by an infected mollusc have the same genotype or, in other words, represent a clone. The LT50 (the time during which 50% of cercariae perish in the experimental dish) were measured experimentally for cercariae Himasthla elongata belong to different clones. The investigated parameter demonstrated a high level of interclonal variability. Two groups of cercarial clones were identified: one of them was characterized by the high level of intraclonal variability in LT50 and the second, by the low one. It is assumed that the observed heterogeneity may be stipulated by different degrees of mitotic recombinations during formation of different cercarial clones.

Morpho-functional specialization of the branching sporocyst of Prosorhynchoides borealis Bartoli, Gibson & Bray, 2006 (Digenea, Bucephalidae).

Sporocysts of Prosorhynchoides borealis were obtained from the marine bivalves Abra prismatica and studied using transmission electron microscopy. The sporocyst body consists of a mass of branching and intertwining hollow tubules that ramify through the host's digestive gland and gonads. This study investigated the ultrastructure of the sporocyst branches which comprise alternate distended areas (brood chambers) with a relatively thin body wall, narrower portions with a thicker body wall (constricted areas) and terminal regions. Pronounced differences between these areas were revealed in the structure of their tegument and body cavity lining, as well as in the cellular composition of the subtegumental layers. Body wall composition in distended areas was consistent with the specialization for cercarial nurture in the brood chambers. The structure of the constrictions suggested a dual role of nutrient absorption and physical separation of adjacent brood chambers. Two types of terminal region were identified, one specialized for the investigation and penetration of host tissues and the other, in which the germinal cells are formed, for cercarial production. The overall structure of the sporocyst branches helps explain why this linear modular system, i.e. brood chambers and constrictions continuously growing into the host tissue, enables the sporocyst's long-term existence and can continuously produce cercariae in numbers comparable with those produced by rediae and/or daughter sporocyst infrapopulations in other digeneans. The origin of the nuclei in the outer tegumental layer of some branching bucephalid sporocysts is also discussed.

[Clonal variability in expression of geo- and photoorientation in cercariae of Himasthla elongata (Trematoda: Echinostomatidae)].

The study was carried out on Himasthla elongata, a digenean common in the coastal ecosystems of the northern European seas. This species utilises intertidal prosobranchs Littorina spp. as the first intermediate host, bivalves (in the White Sea, Mytilus edulis) as the second intermediate host and gulls as the final host. The periwinkles Littorina littorea infected with H. elongata rediae (parthenogenetic generations) were sampled in the intertidal of the White Sea (66 degrees 20' N, 33 degrees 38' E) and used as the source of cercariae. Periwinkles were collected from the settlement with the low prevalence of H. elongata. As shown earlier with the use of AFLP (Amplified Fragment Length Polymorphisms) method, rediae groups in all the infected periwinkles of this settlement arise from the infection of a mollusc with a single miracidium. Therefore, the cercariae shed by an infected mollusc have the same genotype or, in other words, represent a clone. Photo- and geoorientation of cercariae originating from different clones and aged 1 h and 6 h were analysed separately. It was shown that in general the larvae of each clone followed the behavioural pattern characteristics of the species (positive geoorientation and negative photoorientation). However, the degree of expression of this typical behaviour was different in different clones. An especially high variability was observed in the manifestation of geoorientation (in several clones, most larvae demonstrated negative geoorientation). Differences in the distribution of cercariae in the illumination gradient were almost equally associated with the interclonal variability and the age of the larvae. On the whole, as the age of cercariae increased, the positive geoorientation became more prominent, whereas the ratio of cercariae with the typical (negative) photoorientation decreased. Statistical analysis revealed significant differences between the cercarial clones both in the initial manifestation of geo- and photoorientation and in the changes in the character of these reactions with the larval age. Taking into account that each cercarial clone investigated had the same genotype, it seems very likely that the interclonal differences noted in this study are hereditary. Maintenance of a rather high level of genetic polymorphism by the character "expression of orientation reaction" in trematode cercariae may enhance the chances for successful transmission of these larvae. Such variability increases the scale of cercarial dispersion in space and promotes the successful infection of the hosts, whose behaviour is also subject to intra- and inter-population variability. Besides, cercariae whose behaviour deviates from the basic behaviour of the species may play the role of the population's potential for colonisation of new species of animal hosts.

[Effect of some pharmacological substances on the motility of the Cryptocotyle lingua cercaria (Heterophyidae)].

The effect of some biologically active substances (acetylcholine, serotonin, octopamine, sodium nitroprussid and FMRF-amide) on the motility of the Cryptocotyle lingua cercariae was studied. Solutions of FMRF-amide, octopamine, and sodium nitroprussid have no statistically significant influence on the motility of C. lingua. Acetylcholine and serotonin in solutions affected the motility through the prolongation of the active phase of swimming. Further research is required to elucidate the mechanisms underlying the cercarial motility.

[Microphallus kurilensis sp. nov., a new species of microphallids from the pygmaeus species group (Trematoda, Microphallidae) from the coastal areas of Okhotsk and Bering Seas].

The pygmaeus-species group is composed of close related species from the genus Microphallus in which metacercariae develop inside daughter sporocysts without encystment. Infection of periwinkles Littorina (Neritremna) spp. with intramolluscan stages of a new species of this group (Microphallus kurilensis sp. nov.) was recorded on the coasts of Sakhalin and Kuril islands, north of the Sea of Okhotsk and Chukchi Peninsula (the Bering Sea). Application of molecular methods allowed us to establish that M. kurilensis metacercariae are conspecific with one of the morphotypes of microphallid adults obtained from the intestine of the Pacific common eider (Somateria mollissima v-nigrum), which was shot in the north of the Sea of Okhotsk (Galaktionov, Olson, and Blasco-Costa, in press). The adults of the same morphotype were recorded in the Pacific common eider from the northwestern part of the Bering Sea (Chukchi Peninsula). In the course of experimental infection of the slaty-backed gull Larus schistisagus chicks with metacercariae of M. kurilensis, few microphallid adults were obtained. These adults were identical in their morphology with specimens of the microphallid morphotype from the Pacific common eider, which had been identified as M. kurilensis based on molecular data. Morphological description of metacercaria and adult of M. kurilensis and list of their differences from the same developmental stages of other species from pygmaeus-group are provided. It is concluded that M. kurilensis is transmitted in the host system including periwinkle Littorina (Neritrema) and seaducks (predominately, Pacific common eider). Most probably, distribution of M. kurilensis is not limited by the north Asiatic coast but expanded to the North American coast of the Pacific Ocean.

[Description of the maritae and determination of the species status of Microphallus pseudopygmaeus sp. nov. (Trematoda: Microphallidae)].

Microphallus pseudopygmaeus belongs to the "pygmaeus" microphallids, a group of closely related species with homotypic two-host life cycle. This cycle involves one intermediate host (littoral or sublittoral gastropods, mostly of the genus Littorina), and bird-invading metacercariae develop within daughter sporocysts. In spite of the fact that the name Microphallus pseudopygmaeus is widespread in special literature, the original description of this species (Galaktionov, 1980) was performed in contravention of the International Code of Zoological Nomenclature. This article is aimed to correct the above defect. The maritae of Microphallus pseudopygmaeus were obtained from the common eider ducklings which were infected experimentally with metacercariae of Microphallus sp. I Galaktionov, 1980. These metacercariae were extracted from the naturally infected mollusks Littorina saxatilis collected at the Barents Sea coast. The valid description of the species M. pseudopygmaeus, its comparison with other representatives of the "pygmaeus" microphallids and synonymy are provided. Detailed analysis of digeneans from marine and coastal birds preserved in the collections of the Zoological Institute RAS (St. Petersburg, Russia) and the Natural History Museum (London, UK) showed that in some cases M. pseudopygmaeus was misidentified as M. pygmaeus. It has been shown that M. pseudopygmaeus has an amphiboreal-arctic distribution and the spectrum of its first intermediate hosts is unequally wide for digeneans. It includes 15 species of north-Atlantic and north-Pacific mollusks belonging to different families and even orders of Prosobranchia. At the same time, the range of final hosts of M. pseudopygmaeus is limited by benthophagous marine ducks, first of all the common eider.

An ultrastructural study of the early cercarial development in Prosorhynchoides borealis (Digenea: Bucephalidae) with special reference to formation of the primitive epithelium.

Primitive epithelium and outer tegumental layer formation during early cercarial development was studied in Prosorhynchoides borealis using electron microscopy. It demonstrated that germinal cells freely floating in the sporocyst body cavity divide to give rise to naked cell aggregates. These early embryos are highly irregular in outline and are composed of blastomeres differing in size and structure. In embryos consisting of about 12-14 cells a few (possibly only two) superficial macromeres become concave and produce thin extensions which envelop the embryonic mass before fusing to form a syncytial primitive epithelium. This primitive epithelium forms syncytial connections with underlying embryonic cells. Primordial tegumental cells become apparent in late germinal balls below the primitive epithelium. These cells expand and fuse to give rise to an embryonic nucleated tegument. The embryonic tegument is connected to peripheral embryonic cells by thin cytoplasmic bridges until the basement lamina is formed. Subsequently, the primitive epithelium is shed by the embryos and the nuclei in the embryonic tegument undergo pyknotic degeneration. These results are analysed and compared with data from studies on other trematode species and it is concluded that the primitive epithelium is derived from the embryo in at least the majority of digeneans.

A description of the parthenogenetic metacercaria and cercaria of Cercaria falsicingulae I larva nov. (Digenea: Gymnophallidae) from the snails Falsicingula spp. (Gastropoda), with speculation on an unusual life-cycle.

Previously unknown gymnophallid parthenogenetic metacercariae (PM), referred to as Cercaria falsicingulae I larva nov., were found in the extrapallial cavities of the snails Falsicingula mundana (Yokoyama) and F. athera (Bartsch) on coasts of Sakhalin and the Kuril Islands. Unlike all other known PM, rather than producing metacercariae infective to the definitive host, their furcocercariae emerge into the environment. The developing cercariae and metacercariae of C. falsicingulae I are described and compared with other gymnophallid larvae from littoral molluscs in the region. Experimental evidence and analysis of metacercarial group composition in naturally infected molluscs indicate that some cercariae leave their molluscan hosts and penetrate other specimens of Falsicingula in which they develop into new cercariae-producing PM. Metacercariae with developing hermaphroditic reproductive organs were never observed in naturally infected molluscs. A probable life-cycle for C. falsicingulae I is presented in which cercarial/metacercarial production is switched seasonally in order to enable the infection of the definitive hosts by PM. Importantly, the 'cercaria - PM - cercaria' component of the life-cycle appears to be somewhat autonomous.

Infection patterns in white sea blue mussels Mytilus edulis of different age and size with metacercariae of Himasthla elongata (Echinostomatidae) and Cercaria parvicaudata (Renicolidae).

Infection of mussels Mytilus edulis L. by 2 trematode species was studied in a natural intertidal population in the Chupa inlet of the White Sea. The prevalence of metacercariae of Himasthla elongata (Mehlis, 1831) and Cercaria parvicaudata (Stunkard & Shaw, 1931) in mussels reached 100% in 3 to 4 yr old molluscs and remained at this level in older individuals. Infection intensity increased evenly with the age of the molluscan host, showing a tendency to decrease only in the oldest (9 yr old) mussels. These patterns of age dynamics of prevalence and infection intensity were associated with accumulation of trematode larvae in the course of the molluscs' lives. Ability of metacercariae to exist in mussels for long periods (at least 2.5 yr) was verified in the course of an experiment, during which infected molluscs were kept in a subtidal net cage. Decrease of infection intensity in the oldest individuals may reflect selective mortality of the most severely infected molluscs. Among mussels of the same age, higher infection intensity values occurred in larger individuals. This may be due to an enhanced pumping rate in large molluscs, which increases the probability of cercariae, free-living trematode larvae, infecting them via water currents.

One of the most complex life-cycles among trematodes: a description of Parvatrema margaritense (Ching, 1982) n. comb. (Gymnophallidae) possessing parthenogenetic metacercariae.

This study used light and electron microscopy to provide observations and morphometric details of the life-cycle of the gymnophallids (Trematoda, Digenea), Parvatrema margaritense (Ching, 1982) n. comb., the parthenogenetic metacercariae ('germinal sacs') of which were previously described by Ching (1982) as Cercaria margaritensis. The research was instigated by the discovery, on the Barents Sea coast, of a high prevalence of gymnophallid sporocysts and cercariae in the bivalve Turtonia minuta and an equivalent presence of distinctive gymnophallid metacercariae in the gastropod Margarites helicinus. Experiments and data obtained from naturally infected M. helicinus demonstrated that cercariae released from the bivalves invaded the gastropods to give rise to the metacercariae. Two generations (M1 and M2) of these parthenogenetic metacercariae were formed in the extrapallial cavities of their bivalve hosts and they, in turn, gave rise to a third generation (M3) which was shown to infect marine ducks such as the eider (Somateria mollissima). As only small numbers of cercariae are released from T. minuta, it was concluded that the inclusion of parthenogenetic metacercariae in the life-cycle is particularly significant. It allows each cercaria that infects M. helicinus to give rise to over 2000 invasive metacercariae. Evidence suggests that the parthenogenetic metacercariae are commensal rather that parasitic in the pallial cavities of their hosts. Implications of this for theories of early digenean evolution are discussed.

[A comparative analysis of the helminth fauna of kittiwake Rissa tridactyla (Linnaeus, 1758) and glaucous gull Larus hyperboreus Gunnerus, 1767 from different parts of the Barents Sea].

The article is based on the results of helminthological observations made on kittiwake Rissa tridactyla and glaucous gull Larus hyperboreus in 1991-2001 in different areas of the Barents Sea (Eastern Murman coast, Franz Josef Land, Novaya Zemlya, Spitzbergen). 18 helminth species (2 trematodes, 11 cestodes, 4 nematodes, and 2 acanthocephalans) were recorded in the kittiwakes and 19 (3 trematodes, 9 cestodes, 5 nematodes and 2 acanthocephalans) species were recorded in the glaucous gulls. Trematodes were absent in the birds collected at the Franz Josef Land and the northern island of Novaya Zemlya. 3 trematode species, namely Gymnophallus sp. (somateria?), Microphallus sp. 1 (M. pseudopygmaeus), and Cryptocotyle lingua were found in the glaucous gulls of western Spitzbergen. It was supposed that the life cycles of these parasites can be completed there. On the other hand, coastal ecosystems of Arctic archipelagoes turn out to be favourable for the transmission of some cestodes. This is closely connected with the regional traits in the marine bird diet, namely the increase of the amphipod (intermediate hosts of hymenolepidids and some dilepidids) and polar cod (supposed second intermediate host for some tetrabothriids) portion in Arctic. As a result, cestodes are the base of the helminth fauna of kittiwakes and glaucous gulls of the Barents Sea, by their species richness, prevalence and abundance. Nematodes and acanthocephalans were represented by a few species with low infection intensity. The main ecological factors affected the regional difference in the species richness and abundance of the helminths parasitising kittiwakes and glaucous gulls in the Barents Sea are proposed. Those are regional climatic features and regional traits in the behaviour and food priorities of birds, and also the distribution of the helminths intermediate hosts, invertebrates and fishes. The phenomenon of host specificity lowering with respect to the definitive host was recorded in some cestode species (Microsomacanthus diorchis, M. microsoma, and Arctotaenia tetrabothrioides) on the border of their distribution ranges, the coastal ecosystems of Arctic.

Evolutionary relationships within 'pygmaeus' group microphallids using genetic analysis and scanning electron microscopy.

There are four species of 'pygmaeus' microphallids, namely Microphallus pygmaeus, M. piriformes, M. pseudopygmaeus and M. triangulatus (Trematoda: Microphallidae) which are parasites of marine birds and their sporocysts give rise to transmissible metacercariae inside littoral gastropods (mostly littorines). Universally primed polymerase chain reaction (UP-PCR) showed no apparent pattern between genetic diversity of the metacercariae as estimated by genomic banding profiles and their geographic region or molluscan host species. At the same time UP-PCR product cross-hybridization showed that M. pseudopygmaeus and M. triangulatus are genetically very similar, indicating that these taxa represent one species complex. In contrast, M. pygmaeus and M. piriformes are genetically well separated from each other and also from the pseudopygmaeus-triangulatus complex. Scanning electron microscopy of ventral spines, and analyses of spine angles and the number of teeth per spine, showed that all species differed significantly from one another. It was concluded that M. piriformes represents the original western member of the 'pygmaeus' group. Microphallus pygmaeus probably diverged from M. piriformes as it progressively specialized for sea duck final hosts. Microphallus pseudopygmaeus and M. triangulatus diverged from each other and the piriformes-pygmaeus ancestral line relatively recently. Microphallus pseudopygmaeus specialized for adoption of a wide range of gastropod host species and M. triangulatus developed morpho-functional specialization associated with final host exploitation.

An ultrastructural study of excretory system development in the cercariae of Prosorhynchoides gracilescens (Rudolphi, 1819) and Prosorhynchus squamatus Odhner, 1905 (Digenea, Bucephalidae).

The ultrastructure of the developing excretory system of Prosorhynchoides gracilescens and Prosorhynchus squamatus cercariae is described. The development pattern was similar in both species. In early embryos the two main collecting tubes were composed of a layer of cells which were wrapped around the lumen. Later, the tubes fused and the excretory epithelium of the fusion zone and that of the lateral caudal ducts became a syncytium. The collecting tubes in the cercarial body retained their cellular organization. As the tails grew, additional excretory pores were formed in the tail stem where thickened portions of the caudal duct epithelium contacted the surface tegument. Following this, the distal portions of the lateral caudal ducts lost contact with the primary excretory pores and progressively degenerated. Excretory atrium development started with differentiation of secretory active cytons peripheral to the fusion zone. These cells gave rise to cytoplasmic extensions that penetrated the fusion zone wall to eventually form a continuous cytoplasmic layer. This layer eventually replaced some of the fusion zone excretory epithelium and became the lining of the excretory atrium. The anterior end of the fusion zone differentiated into an excretory bladder and a short posterior portion gave rise to the caudal vesicle.

An ultrastructural study of the cercarial excretory system in Bucephaloides gracilescens and Prosorhynchus squamatus.

The ultrastructure of the flame cells, capillaries, collecting tubes, excretory bladder, excretory atrium, caudal vesicle, lateral caudal ducts and excretory pores of cercariae of Bucephaloides gracilescens (Rudolphi, 1819) Hopkins, 1954 and Prosorhynchus squamatus Odhner, 1905 (Digenea: Bucephalidae) is described. Both species are essentially similar except for some details. The terminal parts of the protonephridia have all the structural features that are typical of trematodes. The collecting tubes in the cercarial body are composed of cells that are wrapped around the lumen. The main collecting tubes are joined to the excretory bladder syncytium by septate junctions. Features of P. squamatus excretory bladder epithelium indicate that it is involved in secretory activity, but this is not the case in B. gracilescens. In both species the luminal surface of the excretory bladder epithelium is increased by lamellae, and the basal plasma membrane forms invaginations. In the bladder syncytium of P. squamatus both apical lamellae and basal invaginations are more developed and mitochondria are also more numerous. The excretory atrium is lined by a syncytium with nucleated cytons located in the surrounding parenchyma. The atrium lining is not continuous with the body tegument and possesses specific secretory inclusions and a thick glycocalyx. Septate junctions connect the atrium syncytium to the excretory bladder epithelium at its anterior end and to the syncytial excretory epithelium lining the caudal vesicle and the lateral caudal ducts at its posterior. In the excretory pores the caudal duct syncytium is joined to the tegument by septate desmosomes.