Emendation of the genus Neoheterobothrium and a proposal of a new genus Paraheterobothrium (Monogenea: Diclidophoridae) for five species of diclidophorids from Pleuronectiform fishes.

Neoheterobothrium chilense González, Oliva & Acuña, 2002 is redescribed based on newly collected specimens from the type host, Hippoglossina macrops and type locality (Coquimbo, Northern Chile). Type specimens of Neoheterobothrium hippoglossini Piasecki, Wierzbicka & Kempter, 2000 and Choricotyle exilis Crane, 1972 are also redescribed. These three diclidophorid monogeneans infecting pleuronectiform flatfishes have pre-ovarian seminal receptacle, ootype inside the semicircle of ovary and elongated pharynx, differentiated from the most similar Orbocotyle Euzet & Suriano, 1975 in that members of the latter genus possess post-ovarian seminal receptacle, post-ovarian ootype and cylindrical or pyriform pharynx. Paraheterobothrium n. g. is proposed for P. chilense n. comb. (type), P. hippoglossini n. comb., P. exilis n. comb., P. papillosum n. comb. and P. syacii n. comb. A redescription of Neoheterobothrium affine (Linton, 1898), the type species of the genus, based on voucher specimens confirmed differences from the new genus in the presence of lamellate plate in the anterior jaw of clamps, absence of seminal receptacle and ovoid pharynx. The genus Neoheterobothrium was emended and N. cynoscioni (MacCallum, 1917) and N. mcdonaldi Payne, 1987 were removed from the genus. Neoheterobothrium now comprises N. affine (type), N. insulare Oliva & Luque, 1995, N. hirame Ogawa, 1999, and N. paralichthyi Suriano & Labriola, 1999, all of them parasites of flatfishes of the genus Paralichthys. Molecular analyses with the internal transcribed spacer 1 distinguished the clade for Paraheterobothrium spp. (P. chilense n. comb. and P. papillosum n. comb.) and Neoheterobothrium spp. (N. hirame, N. affine and N. paralichthyi), supporting this taxonomic emendation.

Eurythenes atacamensis sp. nov. (Crustacea: Amphipoda) exhibits ontogenetic vertical stratification across abyssal and hadal depths in the Atacama Trench, eastern South Pacific Ocean.

Eurythenes S.I. Smith in Scudder, 1882 (Crustacea: Amphipoda) are prevalent scavengers of the benthopelagic community from bathyal to hadal depths. While a well-studied genus, molecular systematic studies have uncovered cryptic speciation and multiple undescribed lineages. Here, we apply an integrative taxonomic approach and describe the tenth species, Eurythenes atacamensis sp. nov., based on specimens from the 2018 Atacamex and RV Sonne SO261 Expeditions to the southern sector of the Peru-Chile Trench, the Atacama Trench (24-⁠21°S). Eurythenes atacamensis sp. nov. is a large species, max. observed length 83.2 mm, possesses diagnostic features, including a short gnathopod 1 palm and a chelate gnathopod 2 palm, and a distinct genetic lineage based on a 16S rRNA and COI phylogeny. This species is a dominant bait-attending fauna with an extensive bathymetric range, spanning from 4974 to 8081 m. The RV Sonne SO261 specimens were recovered along a 10-station transect from abyssal to hadal depths and further examined for demographic and bathymetric-related patterns. Ontogenetic vertical stratification was evident across the trench axis, with only juveniles present at abyssal depths (4974-6025 m). Total length-depth analysis revealed that the size of females was unrelated to depth, whereas juveniles followed a sigmoidal relationship with a step-up in size at depths >7200 m. Thus, these bathymetric trends suggest that juveniles and females employ differing ecological strategies in subduction trench environments. This study highlights that even dominant and ecologically important species are still being discovered within the abyssal and hadal environments. Continued systematic expeditions will lead to an improved understanding of the eco-evolutionary drivers of speciation in the world's largest ecosystem.

Testing for deterministic succession in metazoan parasite communities of marine fish.

Parasite communities are similar to free-living communities; decay of similarity over geographic distance, theory of island biogeography, species-area relationships and nestedness have been documented in both communities. Ecological succession has been studied in free-living communities but has rarely been examined in parasite communities. We use seriation with replication to test the hypothesis that succession of parasite community structure is deterministic, thus developing throughout consecutive changes along the fish ontogeny, via a seriated pattern. 12 306 marine fishes (95 species) were studied. In 40 species, a seriated pattern was detected; 25 had a tendency towards a seriated pattern, and for 31 species, succession was at random. Age-classes for each host species explained deterministic successional patterns for whole parasite communities and ectoparasites. Richness and number of age-classes explained this pattern for endoparasites. Seriated successional pattern was evident for parasite communities of long-lived marine fish, indicating that parasite communities follow sequential changes over time, like many free-living communities.

A review of the potential genes implicated in follicular atresia in teleost fish.

In recent years, numerous studies conducted on teleost fish have highlighted the contribution of transcriptomic studies in elucidating the physiological mechanisms underlying the molecular events of oogenesis and follicular atresia, enabling the identification of potential genes and molecular networks that participate in both the reproductive cycle and the process of follicular atresia. Atresia can affect the reproductive potential of females by reducing the healthy eggs that a female can spawn in both aquaculture and wild populations. The substantial diversity of reproductive strategies exhibited by teleost fish has contributed to the difficulty in identifying common genes between species, but a set of core genes has emerged as potential markers for atresia in relation to apoptosis/autophagy, lipid metabolism, oxidative metabolism and other physiological processes similar to those identified in other vertebrates, even mammals. We review the current status of the genes that have been identified in ovaries with atretic oocytes. Our primary goal is to review the current status regarding gene expression during gonadal development and follicular atresia. This information will enable us to understand the factors and expression patterns involved in the follicular atresia of teleost fish.

Calicotyle hydrolagi n. sp. (Monogenea: Monocotylidae) infecting the deep-sea Eastern Pacific black ghost shark Hydrolagus melanophasma from the Atacama Trench, with comments on host specificity of Calicotyle spp.

We describe Calicotyle hydrolagi n. sp. (Monogenea: Monocotylidae) infecting the cloaca of deep-water Eastern Pacific black ghost sharks, Hydrolagus melanophasma captured as bycatch at a local fishery for Patagonian toothfish Dissostichus eleginoides, (Nototheniidae) in the Atacama Trench using morphological and nucleotide (LSU rRNA and SSU rRNA) data. This new species is differentiated from its congeners by a number of characters, including the absence of a cecal diverticula, the size and shape of the male copulatory organ and the shape of the vagina, as well as by differences in molecular data (SSU rRNA and LSU rRNA). The suitability of some sclerotized structures such as the male copulatory organ (MCO) as a taxonomic character is discussed; specifically, we found that the relationship between MCO and total length exhibit different trends in members of Calicotyle isolated from sharks, skates and chimaeras. Additional efforts to obtain sample of Calicotyle species and further molecular studies based on ribosomal and mitochondrial genes are necessary to clarify the degree of host specificity in this genus. Additionally, this is the first report of a member of Calicotyle to be reported in the Southeastern Pacific Ocean.

Two new species of Acanthocotyle Monticelli, 1888 (Monogenea: Acanthocotylidae), parasites of two deep-sea skates (Elasmobranchii: Rajiformes) in the South-East Pacific.

BACKGROUND: Parasites of deep-sea fishes from the South-East Pacific (SPO) are poorly known. Of c.1030 species of fish found in this area, 100-150 inhabit the deep-sea (deeper than 200 m). Only six articles concerning metazoan parasites of fish from deep-waters of SOP are known, and nine monogenean species have been reported. Currently, ten species are known in Acanthocotyle Monticelli, 1888 (Monogenea) and when stated, all of them are found in shallow waters (10-100 m). Acanthocotyle gurgesiella Ñacari, Sepulveda, Escribano & Oliva, 2018 is the only known species parasitizing deep-sea skates (350-450 m) in the SPO. The aim of this study was the description of two new species of Acanthocotyle from two Rajiformes. METHODS: In September 2017, we examined specimens of two species of deep-sea skates (Rajiformes), Amblyraja frerichsi (Krefft) and Bathyraja peruana McEachran & Myyake, caught at c.1500 m depth off Tocopilla, northern Chile, as a by-catch of the Patagonian tooth fish Dissostichus eleginoides Smitt fishery. Specimens of Acanthocotyle were collected from the skin of the skates. Morphometric (including multivariate analysis of proportional measurements, standardized by total length), morphological and molecular analyses (LSU rRNA and cox1 genes) were performed in order to identify the collected specimens. RESULTS: The three approaches used in this study strongly suggest the presence of two new species in the genus Acanthocotyle: Acanthocotyle imo n. sp. and Acanthocotyle atacamensis n. sp. parasitizing the skin of the thickbody skate Amblyraja frerichsi and the Peruvian skate Bathyraja peruana, respectively. The main morphological differences from the closely related species Acanthocotyle verrilli Goto, 1899 include the number of radial rows of sclerites, the non-discrete vitelline follicles and the number of testes. CONCLUSIONS: The two species of monogeneans described here are the only recorded parasites from their respective host species in the SPO. Assessing host specificity for members of Acanthocotyle requires clarifying the systematics of Rajiformes.

Metazoan parasite communities of two deep-sea elasmobranchs: the southern lanternshark, Etmopterus granulosus, and the largenose catshark, Apristurus nasutus, in the Southeastern Pacific Ocean.

Two deep-sea shark species were obtained as by-catch of the local fishery of the Patagonian toothfish, Dissostichus eleginoides, at depths ranging from 1000 to 2200 m off central and northern Chile. A total of 19 parasite taxa were found in 133 specimens of the southern lanternshark, Etmopterus granulosus, (n = 120) and largenose catshark, Apristurus nasutus, (n = 13). Fourteen taxa (four Monogenea, one Digenea, four Cestoda, one Nematoda, two Copepoda, one Annelida and one Thecostraca) were found in E. granulosus, whereas five taxa (one Monogenea, two Cestoda and two Nematoda) were found in A. nasutus. Representatives of Cestoda showed higher values of prevalence and a greater intensity of infection; this pattern is consistent with reports for elasmobranchs, but the monogenean richness was higher than that previously reported for related deep-sea sharks. Regarding E. granulosus, a positive and significant correlation between host length and abundance was found for six (four ectoparasites, one mesoparasite, and one endoparasite) of the 14 taxa recorded, but prevalence was significantly correlated with host length only for the monogenean Asthenocotyle sp. Although the sample size for A. nasutus was limited, we compared richness, abundance, diversity and evenness at the infracommunity and component community levels. All of these variables were higher for E. granulosus, but at the infracommunity level, abundance was higher for A. nasutus. All the parasite taxa (except two) represent new host and geographical records.

Morphological and molecular characterisation of digenean parasites of the Galápagos sheephead Semicossyphus darwini (Jenyns) with the re-description of Labrifer secundus Manter, 1940 (Lepidapedidae) from the Humboldt Current Large Marine Ecosystem.

The Humboldt Current Large Marine Ecosystem (HCLME) is recognised as one of the largest and most productive marine ecosystems. The estimated number of teleost fish species for the Chilean coast influenced by the HCLME reaches c.1,030 but digeneans have been reported for 39 teleost species, i.e. less than 4% of the potential hosts. During a survey of parasites of the Galápagos sheephead wrasse Semicossyphus darwini (Jennyns) (Perciformes: Labridae) from off northern Chile, we obtained representatives of three digenean species, Helicometrina nimia Linton, 1910 (Opecoelidae), Proctoeces humboldti George-Nascimento & Quiroga, 1983 (Fellodistomidae) and Labrifer secundus Manter, 1940 (Lepidapedidae). Labrifer secundus, originally described from Semicossyphus pulcher (Ayres, 1854) off the coast of Baja California, Mexico is re-described. Helicometrina nimia is a common parasite from littoral fishes in the HCLME. Semicossyphus darwini is a new host for these three digenean species.

A new species of Proctoeces and reinstatement of Proctoeces humboldti George-Nascimento and Quiroga 1983 (Digenea: Fellodistomidae) based on molecular and morphological evidence.

The most studied digenean of marine organisms in Chile is by far Proctoeces humboldti, a parasite of the intestine of the clingfish Sicyases sanguineus and gonad of the keyhole limpet Fissurella spp. (progenetic metacercariae). The mussel Perumytilus purpuratus has been suggested as the first intermediate host for this digenean. In a study examining the parasites of S. sanguineus from central Chile, we found specimens of Proctoeces showing significant morphological differences with P. humboldti. To assist in the resolution of the taxonomic identification of these specimens, as well sporocysts obtained from the mussel P. purpuratus from central and northern Chile, phylogenetic studies using DNA sequences from the SSU rRNA, as well the LSU rRNA and Cox 1 gene were performed. Results showed that the clingfish S. sanguineus is a host for two species of Proctoeces (P. humboldti and P. syciases n. sp.) along the northern and central Chilean coast, without geographic separation; the mussel P. purpuratus is the first intermediate host for P. syciases n. sp. but not for P. humboldti in central and northern Chile. Fissurellids (Archaeogastropoda) along the Chilean coast harbor only progenetic stages of P. humboldti, but there is no evidence of progenesis for P. syciases. The reinstatement of Proctoeces humboldti is strongly suggested.

Molecular and Morphological Evidence Demonstrating Two Species of Helicometrina Linton 1910 (Digenea: Opecoelidae) in Northern Chile.

The opecoelid Helicometrina nimia Linton, 1910 has been reported from numerous marine fishes along the Pacific and Atlantic coasts of the Americas. Along the Chilean coast, H. nimia is found in fishes belonging to at least 9 families. This surprisingly low host specificity of H. nimia raises question about the correct identification of specimens assigned to this species. Here we evaluate whether H. nimia specimens isolated from sympatric fish species in northern Chile but with different diets and found in different habitats (water column and demersal) are the same species. Our results demonstrate that specimens from the shallow benthic fish Labrisomus philippii (Steindachner) do not correspond to H. nimia but instead belong to a new species of Helicometrina. This species is described and distinguished from H. nimia using morphological descriptions and 2 molecular markers (the cytochrome c oxidase subunit 1 gene and the V4 region of the SSU rRNA gene). The new species Helicometrina labrisomi (Digenea: Opecoelidae), is found in the intestine of L. philippii (Steindachner, 1866) (Pisces: Labrisomidae), a shallow benthic fish that inhabits the northern coast of Chile. We also studied the related Helicometrina nimia Linton, 1910 from the benthopelagic fishes Paralabrax humeralis (Valenciennes, 1828) and Acanthistius pictus (Tschudi, 1846) (Serranidae). The new species differs from H. nimia by a combination of characters that include ovary shape, number of uterine loops, and position of the genital pore. Our results indicate that morphological characteristics, such as body size, extent of the vitellarium, shape of the testes, and cirrus sac size and extent, traditionally used in the taxonomy of Helicometrina are highly variable. In contrast, meristic and morphological characteristics, such as a lobed ovary, the number of uterine loops, dimensions of the pharynx, and the opening of the genital pore, are highly constant.

Parapedocotyle prolatili gen. n. et sp. n., a representative of a new subfamily of the Diclidophoridae (Monogenea), a gill parasite of Prolatilus jugularis (Teleostei: Pinguipedidae) from Chile.

Parapedocotylinae, a new subfamily, is proposed to accommodate the gen. n. et sp. n. Parapedocotyle prolatili (Monogenea: Diclidophoridae), a gill parasite of the Pacific sandperch, Prolatilus jugularis (Valenciennes) (Pinguipedidae) from northern Chile (30 degrees 56'S; 71 degrees 20'W). Among the Diclidophoridae Cerfontaine, 1895, the species of the Pedocotylinae Yamaguti, 1963 are unique by bearing the first pair of clamps (most posteriorly) in a haptoral projection. Pedocotyle MacCallum, 1913, the only genus in the Pedocotylinae, is characterised by the first pair of clamps non-pedunculate, modified and non-functional, without accessory suckers, and clamps of pairs 2-4 being pedunculate and functional. In contrast, the first pair of clamps in Parapedocotyle is well developed and functional at the terminal end of a long haptoral appendix and having clamp pairs 2-4 pedunculated, modified and apparently not functional. Seminal receptacle is preovarian in Parapedocotyle in opposition to its postovarian position in Pedocotyle. These differences justified the erection of the new subfamily Parapedocotylinae. The new subfamily is also supported by genetic analyses (18S rDNA, 28S rDNA and cox1 sequences) demonstrating that the Pedocotylinae and Parapedocotylinae belong to different clades in the Diclidophoridae.

A new gonad-infecting species of Philometra (Nematoda: Philometridae) from the red cusk-eel Genypterus chilensis (Osteichthyes: Ophidiidae) off Chile.

A new nematode species, Philometra genypteri sp. n. (Philometridae), is described from male and female specimens found in the ovary of red cusk-eel, Genypterus chilensis (Guichenot) (Ophidiidae, Ophidiiformes), from the eastern South Pacific Ocean off the coast of northern Chile. Based on light and scanning electron microscopy examination, the new species differs from most other gonad-infecting Philometra spp. in the length of spicules (126-135 µm), a gubernaculum with dorsal lamella-like structures on its distal end, and the number and arrangement of genital papillae in the male. From a few congeneric, gonad-infecting species with unknown males, it can be distinguished by some morphological and biometrical features found in gravid female (absence of caudal projections, length of first-stage larvae or oesophagus, moderately developed anterior oesophageal inflation) and by the host type (fish family) and geographical distribution. P. genypteri is the first philometrid species reported from a fish belonging to the order Ophidiiformes and the second nominal species of Philometra recorded from marine fishes of the eastern South Pacific.

Regulatory factors and structure of a component population of the spionid Polydora bioccipitalis infesting the surf clam Mesodesma donacium.

Spionidae, particularly polydorids, are common polychaete parasites of edible mollusks around the world. However, our understanding of the regulatory factors and population structure of these parasites is scant. In this study involving Polydora bioccipitalis and the surf clam Mesodesma donacium we evaluated (1) the environmental correlates of the prevalence and mean intensity of the infestation, (2) the relationship between the number of egg capsules and juvenile and adult parasites and the time elapsed since infestation, and (3) the spatial patterns of juveniles and adults within the host. Environmental factors showed no significant correlations with prevalence and mean intensity, suggesting that these factors do not act directly as regulators. Rather, storm surges seemingly induced clam stranding, which in turn affected both the prevalence and intensity of the infestation. The numbers of juveniles and egg capsules in blisters were significantly related to the time since infestation, suggesting mechanisms of use and expansion of the space within the host. Juvenile worms showed an aggregated distribution that was probably related to the episodic nature of infestation events, whereas adults exhibited uniform distributions that probably reflect territorial behavior and reproductive strategies.

Molecular and experimental evidence refuse the life cycle of Proctoeces lintoni (Fellodistomidae) in Chile.

The mussel Perumitylus purpuratus has been described as the first intermediate host for the digenea Proctoeces lintoni (Fellodistomidae) in the Chilean coast. The study of more than 3000 specimens of P. purpuratus, taken off northern Chile revealed the absence of sporocysts. Experimental infection of mussels with eggs obtained from the known host for P. lintoni was unsuccessful. We analyze the V4 region of the SSU rRNA of living sporocysts and cercariae obtained from P. purpuratus from central Chile in order to confirm the proposed life cycle for this digenea. Our results demonstrated that sporocysts and cercariae obtained from P. purpuratus do not belong to P. lintoni but to an undescribed digenea from the Chilean coast.

Long-term parasitic association between the boring polychaete Polydora bioccipitalis and Mesodesma donacium.

Polydora and related genera are common pests for molluscs. Establishing differences between native species and recent invaders provides the basis for understanding the effect of parasites and has implications for resource management. P. biocipitalis has been reported as a recently introduced species to the Chilean-Peruvian coast, raising concerns about its threat to native bivalve species. In contrast, studies on the infestation of P. bioccipitalis on the surf clam Mesodesma donacium, one of the most important species for shellfisheries, suggest a long-term parasitic relationship. The present study analyses infested (i.e. blistered) fossil shells of M. donacium deposited during the Holocene and Middle Pleistocene epochs and critically reviews evidence supporting the hypothesis of the recent introduction of P. bioccipitalis to the Chilean-Peruvian coast. The blistering pattern seen on fossil and recent shells can be considered species-specific for the infestation of M. donacium by P. bioccipitalis. No evidence was actually found on vectors, introduction pathways or distribution range to support the status of P. bioccipitalis as an introduced species. On the contrary, our findings point to a long-term association, at least for several hundred thousand years, between M. donacium and P. bioccipitalis.

Local variability in metazoan parasites of the pelagic fish species, Engraulis ringens: implications for fish stock assessment using parasites as biological tags.

Parasites have been used successfully as biological tags in population studies, mainly in marine fishes, but also in marine mammals, crustaceans and molluscs. Almost all published information dealing with parasites as biological tags evaluates differences between localities. However, local variability in the component community has not been assessed. In this work, we examined whether local variation of the metazoan parasite fauna of Engraulis ringens, extracted from five independent samples from two nearby localities in northern Chile, can be a factor causing bias in stock identification. Our results show that local variability, as estimated by a single sample, may suffice to represent component community variability with no need for replicated data.

Metazoan parasite fauna as a biological tag for the habitat of the flounder Hippoglossina macrops from northern Chile, in a depth gradient.

Quantitative changes in the parasite communities of the flounder Hippoglossina macrops (Steindachner) were studied along a depth gradient. Samples were obtained from the waters off Coquimbo, Chile (29 degrees 18'S to 30 degrees 50'S), at depths ranging from 160 to 342 m. Samples were assigned to 3 depths, i.e., shallow water (less than 200 m), midwater (from 200 to 300 m), and deep water (more than 300 m). Twelve parasite species were recorded: Neoheterobothrium chilensis and Entobdella hippoglossi (Monogenea); Holobomolochus chilensis and Protochondria longicauda (Copepoda); Gliptonobdella sp. (Hirudinea); Nybelinia surmenicola, Scolex pleuronectis, and Neobothriocephalus adspinosus (Cestoda); Floridosentis sp. and Corynosoma australe (Acanthocephala); Anisakidae (Nematoda); and an unidentified hemiurid (Digenea). Univariate analyses showed that C. australe has its highest prevalence and mean intensity of infection in hosts from shallow waters. Floridosentis sp. showed significant differences along the depth gradient, with higher mean intensity and prevalence of infection in fish from midwater. Among the ectoparasites, only N. chilensis exhibited differences in mean intensity with depth, where intensity of infection increased with depth of host habitat. A canonical multivariate analysis demonstrated that parasite burdens can be a good predictor of the environment (shallow, mid-, or deep water) occupied by the flounders.

Neoheterobothrium chilensis n. sp. (Monogenea: Diclidophoridae), a parasite of the bigeye flounder Hippoglossina macrops (Paralichthyidae: Pleuronectiformes) from northern Chile.

Neoheterobothrium chilensis n. sp. (Monogenea: Diclidophoridae) from the gills of the bigeye flounder Hippoglossina macrops (Paralichthyidae: Pleuronectiformes) from northern Chile (30 degrees 11'S, 71 degrees 51'W) is described and compared with the known species in the genus. Diagnostic characters include the position of the seminal receptacle, the number of hooks in the male copulatory organ, the number of testes, and the relative size of the clamp peduncles.

Metazoan parasite fauna of the bigeye flounder, Hippoglossina macrops, from Northern Chile. Influence of host age and sex

The metazoan parasite fauna of Hippoglossina macrops (n = 123) from northern Chile (30°S) is quantitatively described for the first time, and the role of host age and sex was evaluated. Twelve parasite species were recovered, including 5 ectoparasites (2 Monogenea, 2 Copepoda and 1 Piscicolidae) and 7 endoparasites (1 Digenea, 3 Cestoda, 2 Acanthocephala, and 1 Nematoda). The copepod Holobomolochus chilensis, the monogenean Neoheterobothrium sp., the adult acanthocephalan Floridosentis sp. and the hirudinean, Gliptonobdella sp. are new geographical and host records. The most prevalent ectoparasitic species were the monogenean, Neoheterobothrium sp. and the copepod, H. chilensis. Among endoparasites, the acanthocephalans Floridosentis sp. and Corynosoma australe were most prevalent and abundant. Prevalence and mean intensity of infection for most parasitic species were not affected by host sex, however the prevalence of Floridosentis sp. was significantly greater in males. Intensity of infection was positively correlated with host age for Neoheterobothrium sp., and negatively correlated for Floridosentis sp. and H. chilensis. The helminth species richness of the host H. macrops was lower compared to related flatfishes from the Northern Hemisphere. The relationship of the helminth fauna of H. macrops, its feeding habits and ecological habitats are discussed