Co-migration fidelity at a stopover site increases over time in African-European migratory landbirds.

Migratory species are changing their timing of departure from wintering areas and arrival to breeding sites (i.e. migration phenology) in response to climate change to exploit maximum food availability at higher latitudes and improve their fitness. Despite the impact of changing migration phenology at population and community level, the extent to which individual and species-specific response affects associations among co-migrating species has been seldom explored. By applying temporal co-occurrence network models on 15 years of standardized bird ringing data at a spring stopover site, we show that African-European migratory landbirds tend to migrate in well-defined groups of species with high temporal overlap. Such 'co-migration fidelity' significantly increased over the years and was higher in long-distance (trans-Saharan) than in short-distance (North African) migrants. Our findings suggest non-random patterns of associations in co-migrating species, possibly related to the existence of regulatory mechanisms associated with changing climate conditions and different uses of stopover sites, ultimately influencing the global economy of migration of landbirds in the Palearctic-African migration system.

Parasites and prey of the nursehound shark Scyliorhinus stellaris (Linnaeus, 1758): Insights into hidden trophic web interactions in the Mediterranean Sea.

The metazoan parasite community and the stomach contents of the nursehound shark Scyliorhinus stellaris from the Gulf of Naples (central Mediterranean Sea) were studied. The nursehound shark harboured a poor parasite community composed of a species of gill monogenean (Hexabothrium appendiculatum) and three intestinal cestode taxa (Acanthobothrium coronatum and two unidentified species of the genera Yamaguticestus and Scyphophyllidium), all represented by adult stages. Hosts were mostly parasitized by individuals of A. coronatum, which was the most abundant species and contributed to almost 80% of the total number of parasites found. Conversely, other trophically transmitted parasites (i.e., Yamaguticestus sp. and Scyphophyllidium sp.) showed low prevalence and abundance. The parasite infracommunity was poor, showing low values of species richness, total mean abundance, and diversity indices. Overall, 52 prey items belonging to 13 taxa were identified in the stomach contents. Cephalopods were the most important prey items (represented by nine taxa) and the most diverse and abundant group. In the multivariate space provided by a principal component of mixed data (PCAmix), nursehound sharks distributed along two main axes, related to individual traits (first axis) and stomach contents (including empty ones, second axis). A logistic regression based on the first two axes of the PCAmix showed a significant influence of host individual traits and, to a lesser extent, of stomach contents, regarding the probability of being infected by A. coronatum. Alongside specific traits already associated with parasites transmission, our results highlight the importance of cephalopods in transferring cestode infections through trophic web interactions in the top-predator nursehound shark.

Drivers of parasite communities in three sympatric benthic sharks in the Gulf of Naples (central Mediterranean Sea).

Sharks play a key role in the functioning of marine ecosystems and maintenance of trophic web balance, including life cycles of parasites co-occurring in their habitats. We investigated the structure of parasite communities of three sympatric shark species (Etmopterus spinax, Galeus melastomus, and Scyliorhinus canicula) and explored both the influence of host features in shaping the communities and their role as biological indicators of environment stability in the Gulf of Naples (central Mediterranean Sea), a geographical area characterized by strong anthropic pressure. Parasites found were all trophic transmitted helminths with a complex life cycle, except Lernaeopoda galei, that is a ecto-parasite copepod. Communities were all similarly impoverished with 4-5 component species and low values of species richness and diversity. Higher abundance of cestode larvae of the genus Grillotia was found in G. melastomus, although their dominance in all host species suggests that the three sharks have a similar role as intermediate/paratenic hosts in local food webs. Similarly, high abundance of Grillotia larvae could also suggest the occurrence of high abundance of largest top predators in the area. Host morphological (fork length in S. canicula and G. melastomus and body condition index in G. melastomus) and physiological (sex and gonadosomatic and hepatosomatic indices in S. canicula) variables were differently correlated to parasite community structures depending by host species. Potential reasons for the present impoverished parasite communities are discussed.

A molecular and ecological study of Grillotia (Cestoda: Trypanorhyncha) larval infection in small to mid-sized benthonic sharks in the Gulf of Naples, Mediterranean Sea.

AIM: Trypanorhyncha cestodes comprise a wide range of heteroxenous parasites infecting elasmobranchs as definitive hosts. Limited data exist on the larval infection of these cestodes and the role of intermediate and paratenic hosts in the life cycle of these parasites. We investigated the factors that determine the occurrence and the level of infection of Grillotia plerocerci in the skeletal muscles of various benthonic sharks and analyzed the parasites through an integrative taxonomic approach. LOCATION: Mediterranean Sea. METHODS: Sharks obtained as bycatch of commercial trawling activities (i.e., Etmopterus spinax, Galeus melastomus, and Scyliorhinus canicula) were used in this study. Data from a limited number of Dalatias licha and Scyliorhinus stellaris were also included. Grillotia plerocerci were molecularly characterized using the partial 28S large subunit rDNA. Boosted regression trees were used to model the relationship between the abundance of infection with both morphological and physiological predictors in each host. RESULTS: Plerocerci of Grillotia were detected in all shark species except S. stellaris. Host species significantly differed in terms of parasite abundance, with the highest and lowest prevalence and abundance of infection detected in G. melastomus and E. spinax, respectively. The relative influence of the traits involved in explaining the parasite abundance was related to the host size in G. melastomus, while both morphology- and physiology-related traits explained the patterns observed in E. spinax and S. canicula. The 28S rDNA sequences shared an identity of ∼99.40% with a Grillotia species previously found in the Mediterranean Sea. At intraspecific level, two different genotypes were found. A first type was retrieved only from D. licha, whereas a second type was found in G. melastomus, E. spinax, and S. canicula. MAIN CONCLUSIONS: Present results suggest that the two genotypes could be involved in different consumer-resource systems and confirm most of the examined shark species as transport hosts of Grillotia species for unknown larger top predators.

Host biological factors and geographic locality influence predictors of parasite communities in sympatric sparid fishes off the southern Italian coast.

Host biological factors and habitat influence the faunal assemblages and biodiversity worldwide, including parasite communities of vertebrate and invertebrate hosts. The ecological relationship between hosts and parasites can be mediated by interaction of host's biological factors, as their physiological condition, diet and size, with the environmental components, somehow influencing the features of parasite infection in host populations. Here, we used boosted regression tree models to study the parasite communities of two sympatric sparid fishes, the salema Sarpa salpa and the white seabream Diplodus sargus, to investigate the role of specific host's traits in two contiguous coastal areas along the southern-western Tyrrhenian coast of Italy characterized by different degree of deterioration. Results showed that overall and across all parasite groups (ecto-, endo- and ecto- plus endo-parasites), sampling localities were the most important predictors of abundance, species richness, and diversity for salema. Moreover, seasonality was the main predictor of endo-parasite abundance, while size-related factors explained most of the variation in species richness and diversity. In the white seabream, size-related factors and reproductive cycle-related factors were the most important predictors for the overall parasite abundance and parasite richness, respectively. Our findings suggest that the parasite community of salema and white seabream responded differently to specific biological factors, highlighting how the environmental conditions under which they live may exert a strong influence on the parasite communities of each host fish.

A network approach to identify bioregions in the distribution of Mediterranean amphipods associated with Posidonia oceanica meadows.

Although amphipods are key components of the macro-fauna associated with Posidonia oceanica meadows, to date no studies focused on the structure and diversity of their assemblages across the whole Mediterranean Sea. Here, we applied a network approach based on modularity on a dataset mined from literature to identify biogeographic modules and to assess the biogeographic roles of associated localities. We also correlated the patterns evidenced with the biogeographic distribution of amphipod groups by means of a multivariate analysis. Modularity analysis highlighted four biogeographic modules bounded by the main Mediterranean biogeographic divides and evidenced a decrease in species diversity along a NW-SE gradient. Assemblages associated with Central-Western Mediterranean and, to a lesser extent, Tunisian modules showed the highest species richness and were identified as hubs, characterized by species with regional distributions that behave as source in a biogeographic context. The paleogeographic history of the host seagrass and the ecology of associated amphipods, both suggest the joint effect of species persistence and post-Last Glacial Maximum expansion in explaining the pattern of amphipod distribution in the Mediterranean Sea.

Genetic identification and distribution of the parasitic larvae of Anisakis pegreffii and Anisakis simplex (s. s.) in European hake Merluccius merluccius from the Tyrrhenian Sea and Spanish Atlantic coast: implications for food safety.

The consumption of the hake Merluccius merluccius is widespread in European countries, where this fish has a high commercial value. To date, different larval species of Anisakis have been identified as parasites in M. merluccius from European waters, Anisakis pegreffii and Anisakis simplex (s. s.) being the two most common. The aim of the study is to present data on the occurrence of Anisakis spp. larvae in the viscera and flesh of M. merluccius. Consequently, the distribution and infection rates of different species of Anisakis in different sites (viscera, and dorsal and ventral fillets) were investigated in hake caught in the central Tyrrhenian Sea (FAO 37.1.3) and the NE Atlantic Ocean (FAO 27 IXa). A sample of N=65 fish individuals (length>26 cm) was examined parasitologically from each fishing ground. The fillets were examined using the pepsin digestion method. A large number (1310) of Anisakis specimens were identified by multilocus allozyme electrophoresis (MAE) and mtDNA cox2 sequence analysis; among these, 814 larvae corresponded to A. simplex (s. s.) and 476 to A. pegreffii. They were found to infect both the flesh and the viscera. The two species co-infected the same individual fish (both in the viscera and in the flesh) from the FAO 27 area, whereas only A. pegreffii was found in hake from the Tyrrhenian Sea. The average parasite burden of A. pegreffii in hake from the Tyrrhenian Sea was significantly lower to that observed from hake off the Atlantic coast of Spain, both in prevalence and in abundance. In addition, whereas no significant difference in overall prevalence values was recorded between the two Anisakis species in the viscera of the FAO 27 sample, significant differences were found in the abundance levels observed between these species in the flesh, with A. simplex (s. s.) exhibiting significantly higher levels than that observed for A. pegreffii (p<0.001). Given that the pathogenic role in relation to man is known for these two species of Anisakis, both the flesh inspection and the infection rates of the different anisakid species assume particular importance in terms of assessing the risk they pose to humans.

Temporal stability of parasite distribution and genetic variability values of Contracaecum osculatum sp. D and C. osculatum sp. E (Nematoda: Anisakidae) from fish of the Ross Sea (Antarctica).

The Ross Sea, Eastern Antarctica, is considered a "pristine ecosystem" and a biodiversity "hotspot" scarcely impacted by humans. The sibling species Contracaecum osculatum sp. D and C. osculatum sp. E are anisakid parasites embedded in the natural Antarctic marine ecosystem. Aims of this study were to: identify the larvae of C. osculatum (s.l.) recovered in fish hosts during the XXVII Italian Expedition to Antarctica (2011-2012); perform a comparative analysis of the contemporary parasitic load and genetic variability estimates of C. osculatum sp. D and C. osculatum sp. E with respect to samples collected during the expedition of 1993-1994; to provide ecological data on these parasites. 200 fish specimens (Chionodraco hamatus, Trematomus bernacchii, Trematomus hansoni, Trematomus newnesi) were analysed for Contracaecum sp. larvae, identified at species level by allozyme diagnostic markers and sequences analysis of the mtDNA cox2 gene. Statistically significant differences were found between the occurrence of C. osculatum sp. D and C. osculatum sp. E in different fish species. C. osculatum sp. E was more prevalent in T. bernacchii; while, a higher percentage of C. osculatum sp. D occurred in Ch. hamatus and T. hansoni. The two species also showed differences in the host infection site: C. osculatum sp. D showed higher percentage of infection in the fish liver. High genetic variability values at both nuclear and mitochondrial level were found in the two species in both sampling periods. The parasitic infection levels by C. osculatum sp. D and sp. E and their estimates of genetic variability showed no statistically significant variation over a temporal scale (2012 versus 1994). This suggests that the low habitat disturbance of the Antarctic region permits the maintenance of stable ecosystem trophic webs, which contributes to the maintenance of a large populations of anisakid nematodes with high genetic variability.

Parasite communities of icefish (Chionodraco hamatus) in the Ross Sea (Antarctica): influence of the host sex on the helminth infracommunity structure.

Parasite communities of Chionodraco hamatus were investigated from Terra Nova Bay (Ross Sea, Antarctica) during host spawning time. Special attention was given to helminth infracommunities and effect of host sex on its structure. A total of 21 taxa including 5 ecto-parasites and 16 endo-parasites were identified. The number of ecto and endo-parasite species per individual host ranged from 1 to 3 and 3 to 10, respectively, while the mean numbers of parasite specimens per individual host were 4.7 and 1309.7, respectively. The rich abundance of infection suggests a rich concentration of helminth intermediate/paratenic hosts in the coastal waters of Terra Nova Bay. Chionodraco hamatus serves as a definitive host for 10 helminth taxa, while it acts as an intermediate/paratenic host for 6 helminth taxa. Larvae of 6 helminth taxa for which C. hamatus serves as intermediate/paratenic host represented 98.7% of all specimens found. Of these, the tetraphyllidean and diphyllobothridean cestodes and the nematode Contracaecum osculatum s.l. were the most prevalent and abundant. 'Larval' infracommunities had significantly higher species richness, total abundance and diversity than 'adult' infracommunities, suggesting the important role of C. hamatus in supporting the life cycles of those parasites in the study area as a paratenic/intermediate host. Significant differences in the pattern of helminth infracommunities of larval forms between male and female fish were found. These differences could be caused by physiological, and most probably by behavioral differences between sexes suggesting that sex is an important factor influencing parasite burden in C. hamatus during reproductive season.

Genetic and morphological approaches distinguish the three sibling species of the Anisakis simplex species complex, with a species designation as Anisakis berlandi n. sp. for A. simplex sp. C (Nematoda: Anisakidae).

Numerous specimens of the 3 sibling species of the Anisakis simplex species complex (A. pegreffii, A. simplex (senso stricto)), and A. simplex sp. C) recovered from cetacean species stranded within the known geographical ranges of these nematodes were studied morphologically and genetically. The genetic characterization was performed on diagnostic allozymes and sequences analysis of nuclear (internal transcribed spacer [ITS] of ribosomal [r]DNA) and mitochondrial (mitochondrial [mt]DNA cox2 and rrnS) genes. These markers showed (1) the occurrence of sympatry of the 2 sibling species A. pegreffii and A. simplex sp. C in the same individual host, the pilot whale, Globicephala melas Traill, from New Zealand waters; (2) the identification of specimens of A. pegreffii in the striped dolphin, Stenella coeruleoalba (Meyen), from the Mediterranean Sea; and (3) the presence of A. simplex (s.s.) in the pilot whale and the minke whale, Balaenoptera acutorostrata Lacépède, from the northeastern Atlantic waters. No F1 hybrids were detected among the 3 species using the nuclear markers. The phylogenetic inference, obtained by maximum parsimony (MP) analysis of separate nuclear (ITS rDNA region), combined mitochondrial (mtDNA cox2 and rrnS) sequences datasets, and by concatenated analysis obtained at both MP and Bayesian inference (BI) of the sequences datasets at the 3 studied genes, resulted in a similar topology. They were congruent in depicting the existence of the 3 species as distinct phylogenetic lineages, and the tree topologies support the finding that A. simplex (s.s.), A. pegreffii, and A. berlandi n. sp. (= A. simplex sp. C) represent a monophyletic group. The morphological and morphometric analyses revealed the presence of morphological features that differed among the 3 biological species. Morphological analysis using principal component analysis, and Procrustes analysis, combining morphological and genetic datasets, showed the specimens clustering into 3 well-defined groups. Nomenclatural designation and formal description are given for A. simplex species C: the name Anisakis berlandi n. sp. is proposed. Key morphological diagnostic traits are as follows between A. berlandi n. sp. and A. simplex (s.s.): ventriculus length, tail shape, tail length/total body length ratio, and left spicule length/total body length ratio; between A. berlandi n. sp. and A. pegreffii: ventriculus length and plectane 1 width/plectane 3 width ratio; and between A. simplex (s.s.) and A. pegreffii: ventriculus length, left and right spicule length/total body length ratios, and tail length/total body length ratio. Ecological data pertaining to the geographical ranges and host distribution of the 3 species are updated.

Detritus-based assemblage responses under salinity stress conditions in a disused aquatic artificial ecosystem.

BACKGROUND: Despite the plethora of approaches, the sensitivity of the methods to measure the relationship between the abundance and biomass curves in stressed detritus-based ecosystems still remain to be refined. In this work, we report the comparison between biomass and abundance in a set of detritus-based macrozoobenthic assemblages located in six sampling pools with different salinity in an artificial aquatic ecosystem (disused Tarquinia Saltworks), using two diversity/dominance approaches (Abundance/Biomass Comparisons, or ABC, and Whittaker plots). We also evaluated the contribution of abundances and biomasses diversity (Simpson index) and nestedness, which measures the order by which macroinvertebrates colonized the detrital resource. RESULTS: The outputs obtained by both ABC curves and Whittaker plots highlight two different thresholds in assemblage structure: between about 44 and 50 practical salinity unit (psu) and between 50 and 87 psu, respectively. The first threshold was due to a turnover in taxon composition between assemblages, the second threshold (evidenced by Whittaker plots) was due to a change in taxon richness (lower in pools with higher salinity: i.e. > 50 psu). Moreover, a normal-shaped pattern in diversity (Simpson index) emerged, suggestive of an intermediate disturbance effect. The nested pattern did not show significant differences when considering the density and biomass of the sampled taxa, providing similar threshold of salinity in the relative contribution of macrozoobenthos on nestedness. CONCLUSIONS: The use of detailed (ABC and Whittaker plots) and macroscopic (Simpson index and nestedness) approaches is proposed to identify thresholds in the structuring and functioning of detritus-based community of disused aquatic ecosystems: in particular, the inclusion of the parameter of biomass (scarcely utilized in community-based research) appears crucial. The responses of macrozoobenthic assemblages to the salinity stress conditions, in term of abundance and biomass, using a detritus food source (Phragmites australis leaves), may also highlight, by comparing macroscopic and detailed approaches, structuring and functioning patterns to consider for the management of disused artificial ecosystems.

Infaunal macrobenthic community dynamics in a manipulated hyperhaline ecosystem: a long-term study.

BACKGROUND: Understanding the responses of ecological communities to human-induced perturbations is crucial for establishing conservation goals. Ecological communities are dynamic entities undergoing fluctuations due to their intrinsic characteristics as well as anthropogenic pressures varying over time. In this respect, long-term studies, based on large spatial and temporal datasets, may provide useful information in understanding patterns and processes influencing the communities' structure. Theoretical evidence suggests that a role of biodiversity is acting as a compensatory buffer against environmental variability by decreasing the temporal variance in ecosystem functioning and by raising the level of community response to perturbations through the selection of better performing species. Therefore, the spatial and temporal changes in the specialization of the community components may be used as an effective tool to monitor the effects of natural and anthropogenic alterations of the environment in dynamic systems. We examined the temporal dynamics of macroinvertebrate community structure in the hyperhaline habitat of Tarquinia Saltworks (central Italy). We aimed at: (i) investigating the relationships between the level of community specialization and the alterations of the environment across fourteen years; (ii) comparing the ability of aggregate community parameters such as the average abundance vs. species specialization in describing patterns of community composition. RESULTS: We arranged the data in three sub-sets according to three periods, each characterized by different environmental conditions. The mean abundance of sampled macroinvertebrates showed a significant change (p < 0.01) only in the community inhabiting the saltwork basin closely connected to the sea, characterized by the highest environmental variation (i.e. the coefficient of variation, CV, of the aggregate environmental variability over the study period, CVrange = 0.010 - 0.2). Here we found marine species like Modiolus adriaticus (Lamarck, 1819), Neanthes irrorata (Malmgren, 1867), and Amphiglena mediterranea (Leydig, 1851), which inhabited the saltworks during the halt period but disappeared during the subsequent eutrophication phase. Conversely, species specialization showed a significant decrease for each sampled community in the presence of habitat degradation and a recovery after ecological restoration. The widest fluctuations of specialization were recorded for the community inhabiting the saltwork basin with the highest long-term environmental variability. CONCLUSIONS: Recent advances have shown how the increased temporal and spatial variability of species' abundance within the communities may be a signal of habitat disturbance, even in the absence of an apparent decline. Such approach could also be used as a sensitive monitoring tool, able to detect whether or not communities are subjected to increasing biotic homogenization. Also, the increased functional similarity triggered by habitat degradation may impact on species at higher trophic levels, such as the waterbirds wintering in the area or using it as a stopover during migration.

Parasitic infection by larval helminths in Antarctic fishes: pathological changes and impact on the host body condition index.

We examined pathological changes and relationship between body condition index (BCI) and parasitic infection in 5 species of fish, including 42 icefish Chionodraco hamatus (Channichtyidae), 2 dragonfish Cygnodraco mawsoni (Bathydraconidae), 30 emerald rock cod Trematomus bernacchii, 46 striped rock cod T. hansoni and 9 dusty rock cod T. newnesi (Nototheniidae) from the Ross Sea, Antarctica. All parasites were identified by a combination of morphology and mtDNA cytochrome-oxidase-2 sequence (mtDNA cox2) analysis, except Contracaecum osculatum s.l., for which only the latter was used. Five larval taxa were associated with pathological changes including 2 sibling species (D and E) of the C. osculatum species complex and 3 cestodes including plerocercoids of a diphyllobothridean, and 2 tetraphyllidean forms including cercoids with monolocular and bilocular bothridia. The most heavily infected hosts were C. hamatus and C. mawsoni, with C. hamatus most often infected by C. osculatum sp. D and sp. E and diphyllobothrideans, while C. mawsoni was most often infected with tetraphyllidean forms. Histologically, all fish showed varying severity of chronic inflammation associated with larval forms of helminths. Diphyllobothrideans and C. osculatum spp. were located in gastric muscularis or liver and were associated with necrosis and mild to marked fibrosis. Moderate multifocal rectal mucosal chronic inflammation was associated with attached tetraphyllidean scolices. C. hamatus showed a strong negative correlation between BCI and parasite burden.