Parasite assemblages in volatile host stocks: inter- and intra-cohort variability restrict their value as biological tags for squid stock assessment.

The Argentine shortfin squid, Illex argentinus, inhabits in the southwest Atlantic; it is a semelparous species which grows rapidly along its 1 year lifespan. The identification of its stocks is critical for sustainable fishery exploitation. Parasites have been used as biological indicators in a lower number of studies dealing with squids, therefore a validation of this methodology is necessary. The intra- and inter-cohort variability of parasite assemblages in the summer-spawning stock of I. argentinus was analysed to assess their value as indicators of stock structure. Four squid samples from the continental shelf of central Patagonia, corresponding to 3 consecutive cohorts, were examined for metazoan parasites. Results evidenced heterogeneity in terms of parasite assemblage composition and structure, dominated by short-lived gastrointestinal parasites, with a strong influence of host size, but no effect of squid sex. These changes are related to their recent habitats and diets, which change with ontogeny and migrations, clouding any interpretation of patterns when samples spatially or temporally separated are compared. Many squid species share these characteristics; therefore, it is recommended that the use of parasites as biological tags should be restricted to simultaneous sampling, while size or age must be considered for deriving proper conclusions.

Ecology and effects of metazoan parasites of fish in transitional waters.

Given the abundance, heterogeneity and ubiquity of parasitic organisms, understanding how they influence biodiversity, evolution, health and ecosystem functionality is crucial, especially currently when anthropogenic pressures are altering host­parasite balances. This review describes the features, roles and impacts of metazoan parasites of fish occurring in transitional waters (TW). These aquatic ecosystems are highly productive and widespread around the globe and represent most favourable theatres for parasitism given the availability of hosts (invertebrates, fishes and birds) and an increased probability of parasite transmission, especially of those having complex life cycles. Fascinating examples of how parasitism can influence different hierarchical levels of biological systems, from host individuals and populations to entire aquatic communities, through effects on food webs come from this kind of ecosystem. Edible fish of commercial value found in TW can harbour some parasite species, significantly reducing host health, marketability and food safety, with possible economic and public health consequences. Many TW are historically exploited by humans as sources of relevant ecosystem services, including fisheries and aquaculture, and they are highly vulnerable ecosystems. Alteration of TW can be revealed through the study of parasite communities, contributing, as bioindicators, for assessing environmental changes, health and restoration. Fish parasites can provide much information about TW, but this potential appears to be not fully exploited. More studies are necessary to quantify the ecological, economic and medical impacts fish parasites can have on these important ecosystems.

Two new species of Gyrodactylus von Nordmann, 1832 parasitizing Cnesterodon decemmaculatus (Poeciliidae) from the southern limit of the family in the Neotropical region.

In this study, we followed an integrative taxonomy approach to describe two new species of Gyrodactylus von Nordmann, 1832, and to identify specimens of G. breviradix Vega, Razzolini, Arbetman, and Viozzi, 2019, all three collected from ten spotted live-bearer Cnesterodon decemmaculatus (Jenyns, 1842), an endemic and widespread poeciliid from the Pampean region, which is the southernmost occurring species of the Poeciliidae in the Americas. Gyrodactylids were first characterized morphologically and mophometrically, and when possible, sequences of the Internal Transcribed Spacers (ITS1-5.8S-ITS2) and the cytochrome oxidase II (COII) were used to delimit species. Gyrodactylus breviradix, Gyrodactylus marplatensis n. sp., and Gyrodactylus pampeanus n. sp. were found on the fins and body surface of C. decemmaculatus in La Tapera Creek, Mar del Plata, Buenos Aires province, Argentina. A phylogenetic analysis combining newly generated sequences of one of the new species, G. marplatensis n. sp., and of G. breviradix, along with those available in GenBank for a further 36 species of Gyrodactylus, revealed that G. marplatensis n. sp. is a sister taxon of Gyrodactylus decemmaculati Vega, Razzolini, Arbetman, and Viozzi, 2019. Genetic distances for the ITS and COII gene were estimated among Gyrodactylus spp. and further supported the validity of the new species. Overall, morphometric and molecular data coincided in delimiting the new taxa, thus demonstrating the value of integrative taxonomy for the erection of new species of Gyrodactylus and species identification.

Dendrapta nasicola n. sp. (Copepoda: Siphonostomatoida: Lernaeopodidae) a parasite from the olfactory sacs of Bathyraja scaphiops (Norman, 1937) in the South Western Atlantic.

During a parasitological survey of the olfactory sacs of 21 species of Rajiformes (Chondrichthyes) from the Southwestern Atlantic Ocean, copepods referable to Dendrapta Kabata (1964) (Siphonostomatoida: Lernaeopodidae) were found parasitizing the cuphead skate Bathyraja scaphiops (Rajiformes: Arhynchobatidae). Morphological analyses using both light and electron microscopy revealed that they belong to a new species. It can be easily distinguished from its congeners by the ratio between lengths of posterior process and trunk (1:0.8), the large to width ratio of trunk (1:0.7) and the armature of the antennule (1, 1, 5 + 1 aesthete). Dendrapta cameroni longiclavata is raised to full specific status, as Dendrapta longiclavata n. comb. Kabata & Gusev, 1966.

Novel polymorphic microsatellite loci in Anisakis pegreffii and A. simplex (s. s.) (Nematoda: Anisakidae): implications for species recognition and population genetic analysis.

The species of Anisakis constitute one of the most widespread groups of ascaridoid nematodes in the marine ecosystem. Three closely related taxa are recognised in the A. simplex (s. l.) complex, i.e. A. pegreffii, A. simplex (s. s.) and A. berlandi. They are distributed in populations of their intermediate/paratenic (fish and squids) and definitive (cetaceans) hosts. A panel of seven microsatellite loci (Anisl 05784, Anisl 08059, Anisl 00875, Anisl 07132, Anisl 00314, Anisl 10535 and Anisl 00185), were developed and validated on a total of N = 943 specimens of A. pegreffii and A. simplex (s. s.), collected in fish and cetacean hosts from allopatric areas within the range of distribution of these parasite species. In addition, the locus Anisl 7, previously detected in those Anisakis spp., was investigated. The parasites were first identified by sequence analysis of the EF1 α-1 nDNA. The panel of the microsatellites loci here developed have allowed to: (i) detect diagnostic microsatellite loci between the two species; (ii) identify specimens of the two species A. pegreffii, A. simplex (s. s.) in a multi-marker nuclear genotyping approach; (iii) discover two sex-linked loci in both Anisakis species and (iv) estimate levels of genetic differentiation at both the inter- and intra-specific level.

Diversity of Empruthotrema Johnston and Tiegs, 1992 parasitizing batoids (Chondrichthyes: Rajiformes and Myliobatiformes) from the Southwest Atlantic Ocean, with description of three new species.

During an extensive research project involving 519 specimens of batoids, including 13 species of Rajiformes and Myliobatiformes (Chondrichthyes) from the Argentine Sea, three new species of Empruthotrema were found and are described using morphologic characteristics and two molecular markers: LSU rDNA and COI mtDNA. The new species can be distinguished from their congeners by the number and distribution of the marginal loculi, the length and morphology of male copulatory organ, and the presence of eyespots. Additionally, multivariate analysis identified the dimensions of the pharynx and ejaculatory bulb as diagnostic features. Host specificity and previous records of the genus in the region are discussed. This is the first description of new species in this genus for the Southwestern Atlantic Ocean, as well as for arhynchobatid hosts.

Distribution patterns of two species of Corynosoma (Acanthocephala: Polymorphidae) in fishes from Southwestern Atlantic.

Corynosoma australe and C. cetaceum are the most frequently reported acanthocephalans in fish from the Argentine Sea, particularly in central and northern areas. Their definitive hosts are otariids and odontocete cetaceans, respectively. The low specificity of these larvae, in combination with high infective capability and long survival periods in fish, make them potentially good biological markers for stocks and other biological features of their fish hosts. In order to determine the distribution patterns of these species and their determining factors, a large dataset composed by newly collected fish samples, published and unpublished data from previous studies by the authors in the region were analysed in relation to host and environmental variables. The complete dataset comprised a total of 5084 fish, belonging to 29 species distributed in 21 families and 9 orders. Host size and trophic habits arose as the main determinants of abundance for both species of Corynosoma, showing higher abundances on larger fish and on higher trophic levels, as it is usual for trophically transmitted parasites. Biogeographic province and depth (indirectly representing the temperature of water) were the main drivers of the spatial distribution, displaying a latitudinal pattern associated to the temperature clines created by the interaction of Malvinas and Brazil currents, determining a decrease in abundance southwards and towards the deeper areas. No patterns were found regarding the distribution of definitive hosts. The knowledge of these distribution patterns of Corynosoma spp. in fish at regional scale, as well as of their causes, provides useful information to design management and conservation policies thus contributing to maintain the full and sustainable productivity of fisheries.

Rhinoxenus (Dactylogyridae) parasitizing piranhas (Serrasalmidae) at its southernmost limit of distribution (Paraná River, Argentina), with the description of two new species.

Three piranha species, Serrasalmus maculatus, S. marginatus and Pygocentrus nattereri, living sympatrically in the lower Paraná River (Argentina) were examined searching for nasal monogeneans to know its diversity and distribution. Four species of monogeneans belonging to Rhinoxenus were found parasitizing the nasal cavities. Two new species are described, and new morphological data of 2 previously described species is provided. Rhinoxenus argentinensis n. sp. is characterized by having the male copulatory organ (MCO) as a coiled tube with a reel shaped-base; an elongated accessory piece articulated to base of MCO; a sinistral vagina with a sclerotized vestibule, and a sclerotized cap of the ventral anchor laterally modified forming a triangular expansion. Rhinoxenus paranaensis n. sp. is characterized by having a MCO as a coiled tube with a reel-shaped base; an accessory piece with an elongate proximal portion, a dilated distal portion with digitiform projections articulated to base of MCO; a sinistral vagina with 5-6 cuticular spine-shaped processes, and a sclerotized cap of the ventral anchor modified laterally forming a bilobate expansion. Additionally, multivariate discriminant analyses showed significant morphometric differences in the ventral anchors among Rhinoxenus species parasitizing 'piranhas '.

Parasites of the Brazilian flathead Percophis brasiliensis reflect West Atlantic biogeographic regions.

With the aim of evaluating the utility of marine parasites as indicators of zoogeographical regions in the South West Atlantic, we analyzed data on assemblages of long-lived larval parasites of 488 specimens of Percophis brasiliensis distributed in 11 samples from nine localities covering the entire distribution of the species in the Argentine biogeographical Province. Near half a million long-lived parasite individuals belonging to 17 species present in the whole sample displayed clear latitudinal patterns. Data for parasite assemblages at infracommunity and component community levels were analysed in relation to the geographical distance. Significant similarity decay of parasite assemblages over distance was observed, with those based on abundances and mean abundances showing departures from predicted values of regressions. These departures were represented by higher dissimilarities between samples coming from different zoogeographical regions than between those caught within the same region, independently of the distance separating them. Consequently, zoogeographical regions were identified in a distance-decay context. Multivariate analyses corroborated a close fit of similarity between assemblages to existing zoogeographical classifications. Regressions representing distance decay of similarity, and the identification of their outliers, can therefore shed light on the existence of discontinuities or uniformities in the geographic distribution of parasite assemblages and, in turn, in the zoogeography of their fish hosts.

Three-dimensional morphology of rigid structures as a tool for taxonomic studies of Dactylogyridae (Monogenea).

Dactylogyridae is overwhelmingly the most abundant and diverse taxon among monogeneans in continental waters of South America. Their small body size requires considerable sampling effort and training for collecting and identifying the worms from the gills, skin, nasal cavities, and other microhabitats. Indeed, diagnostic characteristics as sclerites and male copulatory complex are generally less than 100-µm long and are essential for taxonomic description and identification of species. Here, a combination of simple and routine methods for three-dimensional morphological studies on hard structures is proposed for dactylogirids: SDS treatment for clarification of specimens and enzymatic digestion with proteinase K for freeing sclerotized structures, followed by laser confocal microscopy. This method is applicable to fresh or fixed specimens and does not require staining or dehydration. Indeed, stable autofluorescence emission is detectable at 500-530 nm for bars, anchors, and male copulatory complex when excited by argon laser. Advantages of this protocol over previous methodologies for taking laser confocal images are discussed. Open access software for image processing was used for three-dimensional reconstruction of sclerotized structures generating models and full 360° rotation videos.

Morphological and molecular evidence for a new species of Pseudanisakis Layman & Borovkova, 1926 (Nematoda: Ascaridida), parasitizing Rajiformes in southern Southwest Atlantic waters.

Pseudanisakis argentinensis n. sp. is proposed to accommodate parasitic nematodes found in six skate species (Rajidae and Arhynchobatidae) examined from southern Southwest Atlantic waters. The new species differs from its congeners by the following combination of characters: a cupola on each lip, males with 8-12 pairs of precloacal genital papillae, a larger size for both males and females, a greater length-to-breadth ratio of the ventriculus and the presence of a small knob on the tip of the tail. Allometric growth was observed for several morphometric features; however, the slopes of the allometric relationships across host species exhibited non-significant differences and were considered as a strong evidence for conspecificity. Congruent results were obtained after the genetic characterization of the mitochondrial cytochrome c oxidase subunit 1 gene of worms obtained from different skate species, whose values of genetic divergence (1.3) lay within the range of intraspecific variation. Previous records of specimens referred to as Pseudanisakis tricupola in skates from South American waters are regarded as conspecific with P. argentinensis n. sp.

A new species of Hedruris Nitzsch, 1821 (Nematoda: Hedruridae) parasitic in the freshwater fish Oligosarcus jenynsii (Günther, 1864) (Characidae) from Argentina.

Nematodes belonging to genus Hedruris Nitzsch, 1821 (Nematoda: Hedruridae) were found in the stomach of the freshwater fish Oligosarcus jenynsii (Günther, 1864) (Characidae) from a shallow eutrophic lake in Argentina. Morphological comparisons with congeneric relatives showed that these nematodes belong to a new species, for which H. bifida n. sp. is proposed. Hedruris bifida n. sp. can be distinguished from its congeners by the following combination of characters: bifid deirids, absence of precloacal papillae and nine pairs of postcloacal papillae and non-mammillated eggs. The new species closely resembles Hedruris suttonae Brugni & Viozzi, 2010¸ the only known species from Argentina; however, the bifid deirids in the new species and the distribution pattern of perianal spines in females clearly distinguish both species. Hedruris bifida n. sp. represents the second nominal species of the genus in Argentina and the eighth species described in the Neotropical region.

Dactylogyrid monogeneans parasitising Cyphocharax voga (Hensel) (Teleostei: Curimatidae) from the Pampas region, Argentina: new and previously described species.

Most studies on dactylogyrid monogeneans in Argentina have been carried out during 1980s and 1990s. Many of these species have been later synonymised and other remain under a confusing taxonomic status, particularly those parasitising Cyphocharax voga (Hensel) (Teleostei: Curimatidae). In order to clarify the identity of dactylogyrids, new material was collected from fishes in Lake Chascomús, Buenos Aires Province, Argentina. A total of four species was found in the gills of C. voga. Two known species, Curvianchoratus singularis (Suriano, 1980) Suriano, 1986 and Palombitrema triangulum (Suriano, 1981) Suriano, 1997, are redescribed and their generic and specific status discussed, and two new species are described. Urocleidoides surianoae n. sp. can be distinguished from its congeners by having an anterior medial projection in the ventral bar and a laminar ligament connecting the base of the male copulatory organ and accessory piece. Annulotrematoides bonaerensis n. sp. differs from its congeners principally by having a ventral bar with an anterior medial projection. The diversity of dactylogyrids harboured by C. voga indicates the need of further studies in the Pampas region, which will provide interesting and valuable sources of evidence for future zoogeographical and evolutionary research on dactylogyrids in the Neotropics.

A new species of Dendromonocotyle Hargis, 1955 (Monogenea: Monocotylidae) from the skin of Zearaja chilensis (Guichenot) (Rajiformes: Rajidae) from the Argentine Sea.

Dendromonocotyle rajidicola n. sp. is described from the dorsal surface of the yellownose skate Zearaja chilensis (Guichenot) (Rajiformes) caught on the Argentine shelf. Dendromonocotyle rajidicola n. sp. can be distinguished from the other 17 species in the genus by the morphology of the distal portion of the male copulatory organ and by the unique morphology of the sclerotised proximal portion of the vagina. This is the first species of Dendromonocotyle to be described from a host in the Rajiformes and also the first record of this genus in the southwestern Atlantic Ocean.

Parasites of Urophycis brasiliensis (Gadiformes: Phycidae) as indicators of marine ecoregions in coastal areas of the South American Atlantic.

The potential value of parasites as ecosystem markers was tested by analyzing the metazoan assemblages of Urophycis brasiliensis caught in four locations distributed in three ecoregions of the Warm Temperate Southwestern Atlantic. A total of 5,001 metazoan parasites belonging to 33 species were found. The identified parasites varied across locations in terms of presence, prevalence, and abundance, and their multivariate analyses resulted in clear similarity patterns. No differences were observed between two locations of the same ecoregion, whereas an evident separation of samples was observed across ecoregions in support of the existing hypotheses regarding the ecoregional division of the southwestern Atlantic. We proposed that parasite assemblages, which are composed of several metazoan phyla, are potentially useful as ecosystem indicators. This suggestion is derived from the combined evidence of the evolutionary history and biogeography of multiple lineages, which is expected to be more efficient in capturing recurrent patterns in overall biodiversity than individual lineages. Furthermore, as many parasites have complex life cycles, their distribution patterns are dependent not only on environmental conditions but also on the distribution and population density of all hosts involved in their life cycles, adding further sources of distributional variability that act synergistically to define robust geographical patterns. The selection of long-lived parasites and their comparative analysis provided evidence supporting the existence of three different stocks in the four sampled areas. The best parasite tags were those with low specificity in fish hosts, constituting promising biological tags for the stock discrimination of other fish species in the region.

Merizocotyle euzeti sp. n. (Monogenea: Monocotylidae) from the nasal tissue of three deep sea skates (Rajidae) in the southwestern Atlantic Ocean.

A new species of Merizocotyle Cerfontaine, 1894 (Monogenea: Monocotylidae) is described from the nasal tissues of three deep sea rajid skates: the southern thorny skate, Amblyraja doellojuradoi (Pozzi), broadnose skate, Bathyraja brachyurops (Fowler), and yellownose skate, Zearaja chilensis (Guichenot), collected off Buenos Aires Province, Argentina, southwest Atlantic Ocean. Two additional species of sympatric rajid, the white-dotted skate, Bathyraja albomaculata (Norman), and the Patagonian skate, Bathyraja macloviana (Norman), were also examined but no merizocotylines were found. The taxonomy of the Merizocotylinae is not widely accepted and, as a result, the status of Thaumatocotyle and Mycteronastes, and their proposed synonymy with Merizocotyle are currently under discussion. The new species differs from its congeners by having a unique haptoral structure, 6 peripheral loculi that are asymmetrically arranged (one much smaller, indistinctly located in the left or right side of the haptor). The presence of the new species in three sympatric species of Rajidae belonging to distinct genera and subfamilies, as well as its absence in sympatric congenerics indicates the lack of phylogenetic host specificity. Host ecology and geographical distribution appear to be more important than host phylogeny in determining the distribution of this parasite across potential hosts in the region. This constitutes the first record of Merizocotyle in the southwestern Atlantic Ocean.

Genetic and morphological evidence reveals the existence of a new family, genus and species of Echinorhynchida (Acanthocephala).

Gymnorhadinorhynchus gen. n. is proposed to accommodate its type species, G. decapteri sp. n., a parasite of the marine fish Decapterus punctatus (Cuvier), caught from the coastal waters of Brazil. Gymnorhadinorhynchus decapteri sp. n. was morphologically most similar to species of two echinorhynchid families, the Rhadinorhynchidae and the Cavisomidae, particularly in the structure of the proboscis and the absence of somatic spines, respectively. This combination of morphological features made it difficult to assign our specimen to an extant family of the Acanthocephala. Therefore, in order to clarify the systematic placement of G. decapteri, a molecular phylogenetic analysis was performed based on the SSU and LSU rDNA and the mitochondrial cox1 gene sequences obtained for the new taxon and other 26 acanthocephalan species. The results of parsimony and maximum likelihood analyses, using individual, combined and concatenated sequence data, consistently indicate that the specimens do not belong to any known family of the Echinorhynchida. Rather, G. decapteri represents a distinct lineage that is closely related to the Transvenidae, but distantly related to both the Rhadinorhynchidae and the Cavisomidae. Gymnorhadinorhynchidae fam. n. is therefore erected. This newly described family can be distinguished from other families of Echinorhynchida by the combination of the following morphological characters: a proboscis cylindrical with 10 rows of 22-26 hooks, dorsoventral differences in proboscis hooks, basal hooks forming a ring and being abruptly larger than anterior hooks, absence of trunk spines and presence of four tubular cement glands. This combination, in addition to several molecular autapomorphies, justifies the erection of a new genus, Gymnorhadinorhynchus gen. n., in order to accommodate this new species.

Low similarity between parasite communities of ten sympatric carangid species.

Host phylogeny and ecological convergence are two factors thought to influence the structure of parasite communities. The aims of this study were to determine the diversity of metazoan parasites of 10 sympatric fish species of the family Carangidae from the southeastern Gulf of California, and to analyze their similarity at infracommunity and component community levels, in order to determine if the host species, particularly those congeneric with similar ecological characteristics, exhibit similar assemblages of parasites. In total, 874 fish specimens were examined and 40 parasite species were identified. The component community was composed by 21 parasite species in Caranx caninus, 20 in C. caballus, 11 in C. vinctus, five in Chloroscombrus orqueta, four in Carangoides otrynter, seven in Hemicaranx leucurus, eight in Selene brevoortii, 14 in S. peruviana, and 11 in Trachinotus rhodopus. The metazoan parasite communities of C. vinctus, Ch. orqueta, H. leucurus, and S. brevoortii are reported here for the first time. The parasite communities of the remaining six carangid species have been reported from regions other than the Gulf of California. All fish species differed significantly regarding the diversity of their parasite infracommunities. This possibly is due to different patterns of habitat use among fish species, and because of the differential host specificity among parasite taxa. Nonetheless, when the analysis was restricted to common parasite species, some fish showed similar parasite infracommunities, particularly congeners of the genus Selene as well as C. caballus and C. vinctus. The component communities of species of Selene were highly similar (>65%), but the three species of Caranx were not. This result supports the hypothesis that congeneric fish species with similar ecological filters harbor similar parasite communities. However, the difference observed between C. caninus and C. caballus suggests that these species, despite being evolutionary and ecologically related, have different physiological or immunological characteristics (compatibility filters) that may result in different parasite communities.

Pseudascarophis brasiliensis sp. nov. (Nematoda: Cystidicolidae) parasitic in the Bermuda chub Kyphosus sectatrix (Perciformes: Kyphosidae) from southeastern Brazil.

A new species of Pseudascarophis (Nematoda: Cystidicolidae) found in the stomach of Kyphosus sectatrix (Linnaeus) (Kyphosidae), off Rio de Janeiro, Brazil, is described. The new species can be differentiated from the other congeners by the presence of lateral alae, distinct but inconspicuous cephalic papillae at the anterior end, three pairs of precloacal and one pair of adcloacal papillae in males, egg morphology and morphometry of glandular oesophagus and spicules. Pseudascarophis tropica is transferred to Ascarophis as Ascarophis tropica (Solov'eva) comb. n. due to its ambiguous diagnosis.

Chondracanthid copepod parasites of dories (Zeiformes: Zeidae) with the description of a new species of Chondracanthus from waters off northern Argentina.

A new species of parasitic copepod, Chondracanthus hoi sp. n. (Copepoda: Chondracanthidae), is described based on specimens of both sexes collected from the buccal cavity and gill arches of the silvery john dory, Zenopsis conchifer (Lowe) (Zeiformes: Zeidae), from waters off northern Argentina (35-36 degrees S, 53-54 degrees W). Female of C. hoi differs from its congeners by the following combination of characters: presence of five pairs of trunk processes, antennule with four knobs tipped with small setae and absence of denticles on the terminal process of maxilla. Chondracanthids and zeiform fishes have been proposed as an example of co-speciation; this assumption is derived from a series of analyses based on incomplete records of both geographical distribution and host range of some parasite species, as well as misidentification offish hosts. These inconsistences observed during our bibliographical analyses are also discussed.