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International audience; Despite considerable research effort, many aspects of the host-parasite relationships and parasite spatial distribution in invasive hosts remain poorly understood due to complex and context-dependent phenomena related to both the bioinvasions and the parasitism. Using macroecological patterns and theory is a useful approach to analyzing parasitological observations, but in practice parasite ecology and classical macroecology are disconnected. We propose a new framework that can use the conventional parasitology sampling data much more effectively. The innovative concept combines the data inferred from populations, infra- and component communities of parasites and the application of a macroecological approach in the analysis of complex and frequently hidden relationships in host-parasite systems. This comparative analysis draws on parasite data across regions and host species at different organizational (population vs. community) and hierarchical (infra vs. component community) levels of parasites. Our framework based on assessing and analysis of parasitological and ecological indexes, including descriptors of parasite species richness (individual and total), infection parameters, parasite aggregation (Taylor´s power law) and macroecological models (abundance-variance and abundance-occupancy relationships), can produce mechanistic explanations of the Enemy Release Hypothesis and unravel host-parasite relationships of an invasive host and its parasites. Moreover, abundance-variance and abundance-occupancy relationships, core-satellite species hypothesis, patterns on the aggregation and the frequency distribution of prevalence, infrapopulation size and individual parasite species richness provide useful tools to distinguish co-introduced and acquired parasites in communities of the invasive host based on quantitative descriptors. We hope that our framework becomes widely applied as it can potentially contribute to enhance future practice and research in biodiversity conservation and control of invasive species.

The major current challenge related to alien species is the absence of a comprehensive theoretical basis to explain the resistance from bioinvasions. One of the factors conferring resistance to invasion is the native species diversity of a recipient ecosystem. We aim to examine patterns of species diversity to understand the mechanisms underlying host-parasite relationships in alien species. The dataset represents 20 host-locality samples, including over 15,067,600 symbiotic organisms counted from 612 host individuals belonging to 7 gammarid species, two native and five invasive, sampled at 16 localities in the Baltic region of Poland. We asked whether diversity patterns depended on locality, habitat and host species, and which factors are deterministic in the variation of diversity. Our results suggest that both locality and host phylogeny are critical factors in determining the composition and abundance of symbiotic communities in gammarids. Out of these two factors, locality appears to be more determinant of the symbiotic community of gammarids than host species, whereas habitat condition was a stronger determinant of the symbiotic communities than geographic distance. The rich and uneven symbiotic communities of native gammarids keep the host-parasite system in equilibrium. Sharply dominated symbiotic species with a high load per host individual stabilize the host population. Simplified symbiotic communities of the invasive hosts showed a relatively even distribution of species with moderate loads per individual, with heavily infected gammarids being rare in the population. The survival rate of invasive gammarids is expected to be relatively high, facilitating their population growth and further spread. Complex communities of the native gammarids constructed from many symbiotic species may prevent host populations from undergoing explosive growth, while such mechanisms may be completely or partially reduced in invasive hosts.

Bipartite network analysis is a powerful tool to study the processes structuring interactions in ecological communities. In applying the method, it is assumed that the sampled interactions provide an accurate representation of the actual community. However, acquiring a representative sample may be difficult as not all species are equally abundant or easily identifiable. Two potential sampling issues can compromise the conclusions of bipartite network analyses: failure to capture the full range of interactions (sampling completeness) and use of a taxonomic level higher than species to evaluate the network (taxonomic resolution). We asked how commonly used descriptors of bipartite antagonistic communities (modularity, nestedness, connectance, and specialization [H2′]) are affected by reduced host sampling completeness, parasite taxonomic resolution, and their crossed effect, as they are likely to co-occur. We used a quantitative niche model to generate weighted bipartite networks that resembled natural host–parasite communities. The descriptors were more sensitive to uncertainty in parasite taxonomic resolution than to host sampling completeness. When only 10% of parasite taxonomic resolution was retained, modularity and specialization decreased by ~76% and ~12%, respectively, and nestedness and connectance increased by ~114% and ~345% respectively. The loss of taxonomic resolution led to a wide range of possible communities, which made it difficult to predict its effects on a given network. With regards to host sampling completeness, standardized nestedness, connectance, and specialization were robust, whereas modularity was sensitive (~30% decrease). The combination of both sampling issues had an additive effect on modularity. In communities with low effort for both sampling issues (50%–10% of sampling completeness and taxonomic resolution), estimators of modularity, and nestedness could not be distinguished from those of random assemblages. Thus, the categorical description of communities with low sampling effort (e.g., if a community is modular or not) should be done with caution. We recommend evaluating both sampling completeness and taxonomic certainty when conducting bipartite network analyses. Care should also be exercised when using nonrobust descriptors (the four descriptors for parasite taxonomic resolution; modularity for host sampling completeness) when sampling issues are likely to affect a dataset., Ecology, 104 (4), ISSN:0012-9658, ISSN:1939-9170

Introduced species can alter the dynamics and structure of a native community. Network analysis provides a tool to study host–parasite interactions that can help to predict the possible impact of biological invasions or other disturbances. In this study, we used weighted bipartite networks to assess differences in the interaction patterns between hosts and helminth parasites of native (Sea of Japan) and invasive (Black Sea and Sea of Azov) populations of Planiliza haematocheilus (Teleostei: Mugilidae). We employed three quantitative network descriptors, connectance, weighted nestedness and modularity, to gain insight into the structure of the host–parasite networks in the native and invaded areas. The role of parasite species in the networks was assessed using the betweenness centrality index. We analyzed networks encompassing the whole helminth community and subsets of species classified by their transmission strategy. The analyses were downscaled to host individual-level to consider intraspecific variation in parasite communities. We found significant differences between networks in the native and invaded areas. The latter presented a higher value of nestedness, which may indicate a co-occurrence between parasite species with many connections in the network and species with fewer interactions within the same individual-host. In addition, modularity was higher in the native area’s networks than those of the invaded area, with subgroups of host individuals that interact more frequently with certain parasite species than with others. Only the networks composed of actively transmitted parasites and ectoparasites did not show significant differences in modularity between the Sea of Azov and the Sea of Japan, which could be due to the introduction of a part of the native community into the invaded environment, with a lower diversity and abundance of species. We show that network analysis provides a valuable tool to illuminate the changes that occur in host–parasite interactions when an invasive species and its parasite community are introduced into a new area.

Parasites are often key players in biological invasions since they can mediate the impact of host invasions or can themselves become invasive species. However, the nature and extent of parasite‐mediated invasions are often difficult to delineate. Here, we used individual‐based, weighted bipartite networks to study the roles (degrees of interactions of individuals in a modular network according to their within‐ and among‐module connections) played by native and invasive host individuals to their parasite communities. We studied two phylogenetically and ecologically close fish species, Mugil cephalus s.l. and Planiliza haematocheilus (Teleostei: Mugilidae). Planiliza haematocheilus is native to the Sea of Japan and invasive in the Sea of Azov whereas, M. cephalus s.l. is native to both seas. Based on the common evolutionary history that drives native host–parasite networks, we hypothesised that 1) native networks have higher modularity than invaded ones; and 2) invasive hosts in the invaded area play a peripheral role to structure parasite communities. We analysed the whole parasite community and subsets based on transmission strategy and host specificity of the parasite species to establish whether modularity and host roles are related to these features in the native and invaded areas. All networks were found to be modular. However, modularity tended to be higher in networks of the native area rather than those of the invaded area. Host individuals of both fish species played similar roles in the native area, whereas invasive hosts played a peripheral role in the networks of the invaded area. We propose that long‐term monitoring of the roles of invasive hosts in parasite communities can be a useful proxy for estimating the maturity of the establishment of the invasive hosts in an ecosystem.

A new microsporidian infecting Gadus chalcogrammus Pallas, 1814 (Gadidae), is described based on morphological, ultrastructural, and molecular studies. This microsporidian parasite develops inside intramuscular spindle-shaped lesions measuring approximately 1-2 mm in width and 4-8 mm in length. Infected cells encapsulated by a host-produced wall containing a sponge-like acellular zone. Sporogony presumably proceeds via segmentation of sporogonial plasmodium, resulting in a variable number of spores. Sporogonial stages develop in sporophorous vesicles (SVs), abutting a moderately electron-dense thick walled coat of a homogeneous amorphous material. SVs space contains rare granular and tubular inclusions. Neighboring SVs often interconnected by bridges of the host cell cytoplasm that were limited by membrane comparable with SV coat. The elongate-ovoid spores, measuring 4.29 ± 0.38 × 2.51 ± 0.26 μm (N 104), possess a bipartite polaroplast and polar tube with 15-16 coils arranged in 2-3 layers. The angle of tilt of the polar tube coils is less than 30°. The sequence analysis of SSU rDNA coding region showed that the studied microsporidians differs from other fish muscle-infecting species at least in 17 bp (2.58%) and is closely related to Microsporidium cypselurus Yokoyama et al. (2002) infecting the flying fish from East China Sea. The parasite is provisionally positioned as Microsporidium theragrae sp. n.

1. Most species in ecological communities are rare whereas only a few are common. This distributional paradox has intrigued ecologists for decades but the interpretation of species abundance distributions remains elusive.2. We present Fuzzy Quantification of Common and Rare Species in Ecological Communities (FuzzyQ) as an R package. FuzzyQ shifts the focus from the prevailing species-categorization approach to develop a quantitative framework that seeks to place each species along a rare-commonness gradient. Given a community surveyed over a number of sites, quadrats, or any other convenient sampling unit, FuzzyQ uses a fuzzy clustering algorithm that estimates a probability for each species to be common or rare based on abundance-occupancy information. Such as probability can be interpreted as a commonness index ranging from 0 to 1. FuzzyQ also provides community-level metrics about the coherence of the allocation of species into the common and rare clusters that are informative of the nature of the community under study.3. The functionality of FuzzyQ is shown with two real datasets. We demonstrate how FuzzyQ can effectively be used to monitor and model spatio-temporal changes in species commonness, and assess the impact of species introductions on ecological communities. We also show that the approach works satisfactorily with a wide range of communities varying in species richness, dispersion and abundance currencies.4. FuzzyQ produces ecological indicators easy to measure and interpret that can give both clear, actionable insights into the nature of ecological communities and provides a powerful way to monitor environmental change on ecosystems. Comparison among communities is greatly facilitated by the fact that the method is relatively independent of the number of sites or sampling units considered. Thus, we consider FuzzyQ as a potentially valuable analytical tool in community ecology and conservation biology.

Neoechinorhynchus is known to be the most diverse acanthocephalan taxon with worldwide distribution; its species are characterized by uniformity of anatomical organization. The taxonomic status of Neoechinorhynchus agilis s. l. from grey mullets was recently reviewed that resulted in description of two new species, Neoechinorhynchus (Hebesoma) personatus and Neoechinorhynchus (Hebesoma) yamagutii. In the current study,18S rRNA partial gene sequences were obtained to molecularly characterize N. (N.) agilis, N. (H.) personatus and N. (H.) yamagutii from grey mullets, Chelonlabrosus and Mugil cephalus, in the North-East Atlantic, the Japan, Azov and Mediterranean Seas. The universal (F-566 and R-1200) and specific eukaryotic set primers were used to amplify specimens from each species of Neoechinorhynchus. It has been found that three species can be clearly recognized using universal primers, which allow to obtain sequences of 590 to 664 bp in length. The genetic divergences estimated between three species sequenced here were relatively high, at ranged between 2.08 and 6.57 %. Phylogenetic analysis demonstrates that the studied species of Neoechinorhynchus from grey mullet fish share common ancestor, despite their different geographic location, and are closely related. The terminal position of N. (N.) agilis and N. (H.) personatus on the evolution tree and the low genetic divergence found between them suggests the recent emergence of this group and that the colonization of the North-East Atlantic and the Mediterranean regions could represent a single event. Our phylogenetic analysis, which included several species of the subgenera Neoechinorhynchus and Hebesoma, showed that the latter is a polyphyletic taxon.

The biogeographic patterns of abundance and prevalence of helminths from Liza haematocheilus were studied across its native (Sea of Japan) and introduced (Sea of Azov) distribution ranges. Abundance-occupancy relationships (AORs) were tested for the core-satellite and enemy release (ERH) species hypotheses in eight and 14 host samples from the native and introduced host ranges, respectively. The AOR model fitted parasite data extremely well, irrespective of whether the host or the parasite species were native or invasive. Except for co-introduced monogeneans, species were less abundant and prevalent in the introduced host population than in the native one, which agrees well with the ERH. Two occupancy patterns were observed. A unimodal, right-skewed distribution of prevalence frequency was common for the acquired groups of helminth parasites in the introduced range, whereas a bimodal distribution was more common in the native range. Core species in the native range were monogeneans, adult and larval digeneans, whereas host-specific, co-introduced monogeneans were the only core species in the introduced range. Acquired grey-mullet specialists and host generalists infected only a small portion of the introduced host population with low mean abundance. These results indicate that strict host specificity, together with a direct life cycle, are the traits that enabled helminth species to entirely occupy the invasive host population. The AORs showed that parasite individuals tend to accumulate in a relatively small fraction of susceptible introduced hosts, probably as an adaptation to enhance mating opportunities, thereby providing a mechanistic explanation of the ERH. All this evidence suggests that co-introduced and acquired species use the introduced host population in very different ways. Therefore, we posit that the examination of AORs can be instrumental in understanding the role of co-introduced parasites in invasion theory.

[EN] To reach ethically and scientifically valid mean abundance values in parasitological and epidemiological studies this paper considers analytic and simulation approaches for sample size determination. The sample size estimation was carried out by applying mathematical formula with predetermined precision level and parameter of the negative binomial distribution estimated from the empirical data. A simulation approach to optimum sample size determination aimed at the estimation of true value of the mean abundance and its confidence interval (CI) was based on the Bag of Little Bootstraps (BLB). The abundance of two species of monogenean parasites Ligophorus cephali and L. mediterraneus from Mugil cephalus across the Azov-Black Seas localities were subjected to the analysis. The dispersion pattern of both helminth species could be characterized as a highly aggregated distribution with the variance being substantially larger than the mean abundance. The holistic approach applied here offers a wide range of appropriate methods in searching for the optimum sample size and the understanding about the expected precision level of the mean. Given the superior performance of the BLB relative to formulae with its few assumptions, the bootstrap procedure is the preferred method. Two important assessments were performed in the present study: i) based on CIs width a reasonable precision level for the mean abundance in parasitological surveys of Ligophorus spp. could be chosen between 0.8 and 0.5 with 1.6 and 1x mean of the CIs width, and ii) the sample size equal 80 or more host individuals allows accurate and precise estimation of mean abundance. Meanwhile for the host sample size in range between 25 and 40 individuals, the median estimates showed minimal bias but the sampling distribution skewed to the low values; a sample size of 10 host individuals yielded to unreliable estimates., SS and VS were supported by MEDEA project fellowships, Erasmus Mundus Action 2. CC-S was funded by project #MTM2014-52975-C2-1-R:" Inference in Structured Additive Regression (STAR) Models with Extensions to Multivariate Responses. Applications in Biomedicine", cofinanced by the Ministry of Economy and Competitiveness (SPAIN) and by the European Regional Development Fund (FEDER). This study is partially supported by Ministry of Education and Science of Ukraine, project #1/17.

One of the most intriguing questions in invasive biology is how an introduced species changes its population parameters in a new environment. Translocation of free-living species commonly results in co-introduction of their parasites. The current study focuses on the abundance pattern of the monogenean parasite Ligophorus llewellyni of the pacific so-iuy mullet, Planiliza haematocheila, across the native and introduced distribution ranges. We evaluated parasite release by the so-iuy mullet by comparing abundance patterns of L. llewellyni under effects of the host length, water temperature and month of the year in the Sea of Japan and the Sea of Azov. Generalised additive models applied to analysis of parasite abundance data showed that relationships between the mean number of L. llewellyni and the three tested independent variables were not linear. Our results suggest that the introduced host lost a large amount of parasite abundance due to the effect of warm climate in a new region, which is mediated by host defence mechanisms. The abundance of L. llewellyni rapidly rose in autumn, as fish activity and immune response decrease, reached the maximum in winter and began to fall in spring as a warm temperature facilitates the fish immune defence. The abundance of L. llewellyni showed an initial increase in response to fish growth and reached an asymptote. The response curves built for native and introduced regions reached an asymptote at different fish body lengths, reflecting the fish growth rate, which is higher in the introduced range of P. haematocheila. We found that the carried parasite species holds the same trend in relationships compared with its native area, between the mean number of monogeneans per host and independent variables increasing abundance with fish length, low temperature and cold months. Our results open new perspectives for future research on statistical modelling of parasite abundance across native and introduced distribution ranges in order to provide deeper insight into host-parasite interactions of invasive populations.

International audience; A number of hypotheses exist to explain aggregated distributions, but they have seldom been used to investigate differences in parasite spatial distribution between native and introduced hosts. We applied two aggregation models, the negative binomial distribution and Taylor's power law, to study the aggregation patterns of helminth populations from Liza haematocheilus across its native (Sea of Japan) and introduced (Sea of Azov) distribution ranges. In accordance with the enemy release hypothesis, we predicted that parasite populations in the introduced host range would be less aggregated than in the native host area, because aggregation is tightly constrained by abundance. Contrary to our expectation, aggregation of parasite populations was higher in the introduced host range. However, the analyses suggested that the effect of host introduction on parasite aggregation depends on whether parasite species, or higher level taxonomic groups, were acquired in or carried into the new area. The revealed similarity in the aggregation parameters of co-introduced monogeneans can be attributed to the repeatability and identity of the host-parasite systems. In contrast, the degree of aggregation differed markedly between regions for higher level taxa, which are represented by the native parasites in the Sea of Japan versus the acquired species in the Sea of Azov. We propose that the host species plays a crucial role in regulating infra-population sizes of acquired parasites due to the high rate of host-induced mortality. A large part of the introduced host population may remain uninfected due to their resistance to native naïve parasites. The core concept of our study is that the comparative analysis of aggregation patterns of parasites in communities and populations, and macroecological relationships, can provide a useful tool to reveal cryptic relationships in host-parasite systems of invasive hosts and their parasites.

International audience; The abundance and aggregation patterns of helminth communities of two grey mullet hosts, Liza haematocheilus and Mugil cephalus, were studied across 14 localities in Atlantic and Pacific marine areas. The analysis matched parasite communities of (i) L. haematocheilus across its native and introduced populations (Sea of Japan and Sea of Azov, respectively) and (ii) the introduced population of L. haematocheilus with native populations of M. cephalus (Mediterranean, Azov-Black and Japan Seas). The total mean abundance (TMA), as a feature of the infection level in helminth communities, and slope b of the Taylor’s power law, as a measure of parasite aggregation at the infra and component-community levels, were estimated and compared between host species and localities using ANOVA. The TMA of the whole helminth community in the introduced population of L. haematocheilus was over 15 times lower than that of the native population, but the difference was less pronounced for carried (monogeneans) than for acquired (adult and larval digeneans) parasite communities. Similar to the abundance pattern, the species distribution in communities from the invasive population of L. haematocheilus was less aggregated than from its native population for endoparasitic helminths, including adult and larval digeneans, while monogeneans showed a similar pattern of distribution in the compared populations of L. haematocheilus. The aggregation level of the whole helminth community, endoparasitic helminths, adult and larval digeneans was lower in the invasive host species in comparison with native ones as shown by differences in the slope b. An important theoretical implication from this study is that the pattern of parasite aggregation may explain the success of invasive species in ecosystems. Because the effects of parasites on host mortality are likely dose-dependent, the proportion of susceptible host individuals in invasive species is expected to be lower, as the helminth distribution in the invasive host was featured by a higher number of uninfected hosts and a shorter distribution tail when compared with native species.

2 Section of Hydrobiont Diseases, Pacifi c Research. Fisheries Center, Shevchenko str., 4, Vladivostok, 690950 Russia Taxonomic Status of Neoechinorhynchus agilis (Acanthocephala, Neoechinorhynchidae), with a De- scription of Two New Species of the Genus from the Atlantic and Pacifi c Mullets (Teleostei, Mugili- dae). Tkach, Ie. V., Sarabeev, V. L., Shvetsova, L. S. — Th e wide variability in morphological features, geographical and host ranges of mullet acanthocephalan parasite Neoechinorhynchus agilis (Rudolphi, 1819), raises the question of taxonomic status of this species. Rudolphi's type and Yamagutis voucher specimens, as well as our own material from the WW Pacifi c and NE Atlantic region were used herein to provide comparative morphological analysis. Th e study revealed three diff erent species of Neoechino- rhynchus, N. (N.) agilis and N. (H.) personatus Tkach, Sarabeev et Shvetsova, sp. n. in the Atlantic and N. (H.) yamagutii Tkach, Sarabeev et Shvetsova, sp. n. in the Pacifi c. Strong morphological and morpho- metric diff erences were found between three described herein species from diff erent hosts and regions. Th e dividing of N. agilis into three species, two of them are new, provides a basis for the further revision of host-geographical records of mullet acanthocephalan parasites. Таксономический статус Neoec hinorhynchus agilis (Acanthocephala, Neoechinorhynchidae), с описанием двух новых видов рода от кефалевых рыб (Teleostei, Mugilidae) Атлантики и Пацифики. Ткач Е. В., Сарабеев В. Л., Швецова Л. С. — Широкое географическое распространение, богатый список хозяев, а также значительная вариабельность метрических признаков аканто- цефалы Neoechinorhynchus agilis (Rudolphi, 1819) ставит вопрос о таксономическом статусе этого вида. Для проведения сравнительного морфологического анализа был использован типичный материал Рудольфи и ваучерные экземпляры Ямагути, а также собственный материал, собранный в западной части Тихого океана и северо-восточной Атлантике. В результате исследования было обнаружено три вида акантоцефал Neoechinorhynchus, в том числе два вида из Атлантики: N. (N.) agilis и N. (H.) personatus Tkach, Sarabeev et Shvetsova, sp. n., и один вид из Тихого океана, N. (H.) yamagutii Tkach, Sarabeev et Shvetsova, sp. n. Описанные в работе виды хорошо различаются как морфологически, так и метрически. Разделение N. agilis на три вида, два из которых описаны впервые, является основой для дальнейшего пересмотра находок этой акантоцефалы из разных регионов и от разных хозяев. Ключевые слова : Средиземное море, Азово-Черноморский бассейн, cеверо-восточная Атлан- ти ка, Chelon labrosus, Mugil cephalus.

The search for phylogenetic signal in morphological traits using geometric morphometrics represents a powerful approach to estimate the relative weights of convergence and shared evolutionary history in shaping organismal form. We assessed phylogenetic signal in the form of ventral and dorsal haptoral anchors of 14 species of Ligophorus occurring on grey mullets (Osteichthyes: Mugilidae) from the Mediterranean, the Black Sea and the Sea of Azov. The phylogenetic relationships among these species were mapped onto the morphospaces of shape and size of dorsal and ventral anchors and two different tests were applied to establish whether the spatial positions in the morphospace were dictated by chance. Overall significant phylogenetic signal was found in the data. Allometric effects on anchor shape were moderate or non-significant in the case of evolutionary allometry. Relatively phylogenetically distant species occurring on the same host differed markedly in anchor morphology indicating little influence of host species on anchor form. Our results suggest that common descent and shared evolutionary history play a major role in determining the shape and, to a lesser degree in the size of haptoral anchors in Ligophorus spp. The present approach allowed tracing paths of morphological evolution in anchor shape. Species with narrow anchors and long shafts were associated predominately with Liza saliens. This morphology was considered to be ancestral relative to anchors of species occurring on Liza haematocheila and M. cephalus possessing shorter shafts and longer roots. Evidence for phylogenetic signal was more compelling for the ventral anchors, than for the dorsal ones, which could reflect different functional roles in attachment to the gills. Although phylogeny and homoplasy may act differently in other monogeneans, the present study delivers a common framework to address effectively the relationships among morphology, phylogeny and other traits, such as host specificity or niche occupancy.

A new multivalvulid myxozoan parasite, Kudoa unicapsula n. sp., is described from the intestinal mesentery, intestine and pyloric caeca of the thin-lipped grey mullet Liza ramada (Risso 1826) and the golden grey mullet L. aurata (Risso, 1810) from the Mediterranean coastal waters of Spain. It is characterized by the presence of elongated, rice corn-like white cysts of 0.47-0.56 x 0.18-0.38 mm, filled with tetracapsulate, slightly asymmetric spores, rectangular in apical view and tear-shaped in lateral view with four polar capsules of considerably different size and slightly unequal spore valves with rounded edges, overlapping each other on the apex of the spore. One large polar capsule includes a polar filament coiled in two to three turns, and the other three polar capsules, which are very small, posses only a rudimental filament. Both light and electron microscopy data showed that this species differs from all previously described Kudoa spp. with unequal polar capsules. The molecular analysis based on 18S and 28S ribosomal deoxyribonucleic acid DNA sequence data of K. unicapsula n. sp. indicates a close relationship and thus phylogenetic clustering together with K. trifolia, a myxozoan from the same host and the same geographical location.

A new species of myxozoan, Kudoa trifolia sp. n., was found in various organs of the golden grey mullet, Liza aurata (Risso), and the thinlip mullet, L. ramada (Risso), from the western Mediterranean. Spores developed in subspherical plasmodia of 0.28-1 mm diameter within connective tissue, predominantly in the spleen, the outer wall of the gall bladder and the gut, the mesenteries and occasionally also in the gills. The spores of K. trifolia differ from the commonly known shape of Kudoa by considerable enlargement of one of the four valve cells, thus forming a 'spore body', which contains the major part of the binucleate sporoplasm. Scanning electron microscopy of the spores revealed the presence of grape-like appendages, which occur in bundles terminally on the valve cells. Phylogenetic analysis based on the 18S rDNA sequence of K. trifolia showed that this species is deeply embedded in the genus Kudoa despite its aberrant morphology and host tissue location. This suggests important amendments to the morphological diagnosis of the genus Kudoa.

A comparative morphological study of specimens of Ligophorus spp. from Mugil cephalus in western Mediterranean and the Black Sea localities has been carried out, indicating the presence of 2 distinct forms, i.e., Ligophorus chabaudi and Ligophorus cephali n. sp. A detailed description of the latter and an up-to-date redescription of L. chabaudi are provided. The existence of these 2 morphological species was additionally supported by principal component analysis based on 19 metric characters of 87 specimens arranged in samples defined by parasite species and geographical locality. The analysis indicated consistent differences between species but not between localities; this pattern was well supported by jackknife procedures. Linear discriminant analyses showed that the main metric differences between the 2 species were attributable to the lengths of the dorsal and ventral bars and to the uncinulus length; use of the first 2 variables was enough to allocate all specimens studied to either form. The erection of L. cephali n. sp. raises the number of sympatric Mediterranean species of Ligophorus on M. cephalus to 3. The known geographical ranges of these species are compared.

Three species of the bunocotyline genus Saturnius Manter, 1969 are described from the stomach lining of mugilid fishes of the Mediterranean and Black Seas. Two of the species are new: S. minutus n. sp. occurs in Mugil cephalus off the Mediterranean coast of Spain; and S. dimitrovi n. sp., a parasite of M. cephalus off the Bulgarian Black Sea coast and the Spanish Mediterranean coast, was originally described as S. papernai by Dimitrov et al. (1998). In addition, S. papernai Overstreet, 1977 is redescribed from M. cephalus off the Spanish Mediterranean coast and from Liza aurata and L. saliens off the Bulgarian Black Sea coast. The three species are distinguished morphometrically using univariate and multivariate analyses. These results were verified using Linear Discriminant Analysis which correctly allocated all specimens to their species designations based on morphology (i.e. 100% successful classification rate) and assigned almost all specimens to the correct population (locality). The following variables were selected for optimal separation between samples: the length of the forebody, ventral sucker and posterior testis, the length and width of the posteriormost pseudosegment, and the width of the muscular flange at ventral sucker level.

Quantitative complex analyses of parasite communities of invaders across different native and introduced populations are largely lacking. The present study provides a comparative analysis of species richness of helminth parasites in native and invasive populations of grey mullets. The local species richness differed between regions and host species, but did not differ when compared with invasive and native hosts. The size of parasite assemblages of endohelminths was higher in the Mediterranean and Azov–Black Seas, while monogeneans were the most diverse in the Sea of Japan. The helminth diversity was apparently higher in the introduced population of Liza haematocheilus than that in their native habitat, but this trend could not be confirmed when the size of geographic range and sampling efforts were controlled for. The parasite species richness at the infracommunity level of the invasive host population is significantly lower compared with that of the native host populations that lends support to the enemy release hypothesis. A distribution pattern of the infracommunity richness of acquired parasites by the invasive host can be characterized as aggregated and it is random in native host populations. Heterogeneity in the host susceptibility and vulnerability to acquired helminth species was assumed to be a reason of the aggregation of species numbers in the population of the invasive host.

Within ectoparasitic fish monogeneans, the genus Ligophorus contains a high number of species from which several were recently described. The precise determination of their taxonomic status requires robust diagnostic morphologic features that rely predominantly on a restricted set of sclerotized structures. In the present study, these morphological characters were used for the reconstruction of a phylogenetic tree, which was compared with a tree built from molecular data (28S and ITS1 DNA sequences). Thirty-eight morphological characters were used in 29 species of Ligophorus from the Atlantic and Pacific regions and 5 species within close genera of Dactylogyridae. The morphological and molecular phylogenetic trees are congruent and suggest that the genus Ligophorus is monophyletic, and that species parasitizing Liza spp. and Chelon labrosus occupy basal positions. The present study suggests that host switching is a common event in this host–parasite association, because about half of the species infecting the same host species are not close relatives. Following host switching, dispersal with vicariance is probably an important force shaping the present distribution and diversity of Ligophorus. The pattern of occurrence of Ligophorus spp. on Mugil cephalus supports that reproductive isolation and therefore parallel speciation are taking place among these parasitic organisms.

An important question in the study of phenotypic evolution is whether characters are independent of each other or behave and evolve as integrated modules. Morphological integration and modularity provide a powerful framework for the analysis of the evolution of morphological traits. We used geometric morphometrics and phylogenetically independent contrasts (PIC) to test four different modularity hypotheses in the haptoral anchors of 14 monogenean species of Ligophorus. Integration between the modular units identified was further evaluated with two-block partial least squares analysis. Roots and points represented two modules in the dorsal and ventral anchors, but modularity was not statistically supported when parasite phylogeny was accounted for, which may indicate convergent evolution related to host characteristics and gill morphology. In contrast, PIC revealed medial and lateral modules in ventral anchors only. Moreover, we found evidence for ventral and dorsal anchor pairs forming two modules, supporting the notion that they play different functional roles. Integration between all identified modules was strong. We conclude that there is modular structure in the anchors of Ligophorus spp., accounted by adaptive and phylogenetic factors acting at different levels, and ventral and dorsal anchors evolve as integrated modules with specific roles in attachment.

The taxonomic framework of Ligophorus, monogenean specialists of the gills of grey mullets (Mugilidae), is evaluated and its interspecific relationships are assessed for the first time using molecular data. The position of Ligophorus within the paraphyletic Ancyrocephalinae is re-assessed based on newly sequenced species. Furthermore, the relationship between morphometric and genetic interspecific similarities is evaluated. Partial 28S and complete ITS1 rDNA sequences from representatives of 14 of the 16 nominal species of Ligophorus from the Mediterranean, Black and Azov Seas were analysed together with published sequences of members of the Dactylogyridae. The phylogenetic analyses of the Dactylogyridae (i) confirmed the position of Ligophorus within the marine Ancyrocephalinae; (ii) revealed a sister relationship between Ergenstrema and Ligophorus, whose species are all exclusive parasites of grey mullets; and (iii) substantiated the affinities of Ergenstrema with the marine Ancyrocephalinae. The phylogenetic analysis restricted to Ligophorus confirmed the distinct status of the included species. The ITS1 region provided the highest divergence between species and phylograms with the strongest branch support. Both the 28S and ITS1 phylograms revealed two main clades. One included species from hosts with Mediterranean and NE Atlantic distribution and another was formed by species parasitising several Liza spp., including Lz. haematocheilus from the Northwestern Pacific, and Mugil cephalus, which suggests an origin outside the Mediterranean for the latter clade. The phylogenetic evidence presented herein indicated that a combination of host-switching and lineage duplication events accounted for the diversification of this genus in the Mediterranean basin. The agreement between molecular and morphological interspecific similarities observed in Ligophorus supports the validity of morphometric characters used for species identification.

Ligophorus mugilinus was originally described on the flathead mullet Mugil cephalus from the Gulf of Mexico. The species was later recorded in the Mediterranean Sea on the flathead mullet and in the Caribbean Sea on the white mullet Mugil curema. After examination of the type material and additional specimens of Ligophorus from the northwest Atlantic coast, the Caribbean Sea, and the Mediterranean and Black Sea, we observed morphological differences concerning haptoral and male copulatory structures between L. mugilinus from the northwest Atlantic and the forms from the Mediterranean and Black Seas. Accordingly, the latter specimens are described as a new species, Ligophorus mediterraneus n. sp. A redescription of L. mugilinus, including new morphometric data, is also given. The morphometric analysis of the Caribbean specimens suggested that their ascription to L. mugilinus is uncertain, but additional material is needed to clarify their specific identity. The results of this study, together with previous zoogeographical evidence, indicate that flathead mullets from different seas harbor different species complexes of Ligophorus. Given the coastal habits of the host, it is suggested that open oceans could act as geographical barriers favoring speciation within disjointed flathead mullet populations.

Ligophorus pilengas Sarabeev & Balbuena, 2004 and L. gussevi Miroshnichenko & Maltsev, 2004, both recently described as new species from the so-iuy mullet Mugil soiuy in the Sea of Azov and Black Sea, were compared. A morphometric analysis of both forms strongly suggests that they represent the same species. According to the ‘Principle of priority’, L. pilengas is proposed as a senior synonym of L. gussevi.

The monogenean Ligophorus chabaudi was originally described on the gills of the flathead mullet, Mugil cephalus, and was subsequently reported on the So-iuy mullet, Mugil soiuy. However, the morphology of sclerotized parts and multivariate statistical analyses suggest that the form from the So-iuy mullet represents a new species. This study provides a description of the new species Ligophorus pilengas n. sp. and provides additional morphological data concerning the morphology of the ventral bar that might be useful for the diagnosis of Ligophorus. Ligophorus pilengas n. sp. is the second species of Ligophorus reported on the So-iuy mullet. Zoogeographical records indicate that L. pilengas n. sp. was probably introduced to the Black Sea and the Sea of Azov from the western Pacific Ocean together with its host. Balbuena Diaz-Pines, Juan Antonio, J.A.Balbuena@uv.es

Dicrogaster contracta and D. perpusilla were originally described about 100 years ago from the thicklip grey mullet Chelon labrosus. Subsequent studies have reported D. contracta in several mullet species in the Mediterranean, Azov and Caspian Seas, but D. perpusilla has never been reported again. Since previous evidence suggests that the distinguishing features between these two species are uncertain, this study provides new and additional data on the variability of morphometric and meristic characters of D. contracta in order to assess their usefulness for differentiating this species from D. perpusilla. Body measurements, egg counts and shape statistics of three sets of specimens of D. contracta from different hosts and localities were compared with similar information from previous studies, including the original descriptions of D. contracta and D. perpusilla. The results showed consistent differences between the three sets in metric measurements, egg counts and shape statistics. In addition, the values of morphometric and meristic characters of D. perpusilla fell within the range of variation recorded for D. contracta. Neither conventional body measurements, eggs counts, nor shape statistics seemed useful to distinguish D. contracta from D. perpusilla, suggesting that they represent the same species. Accordingly, the valid designation of the species should be D. perpusilla because it is the type-species of the genus. This study proposes a neotype for D. perpusilla, since no type-specimens seem to be available, and provides additional morphologic data that might be useful for diagnosis of the genus Dicrogaster.