High parasite diversity in the amphipod Gammarus lacustris in a subarctic lake.

Amphipods are often key species in aquatic food webs due to their functional roles in the ecosystem and as intermediate hosts for trophically transmitted parasites. Amphipods can also host many parasite species, yet few studies address the entire parasite community of a gammarid population, precluding a more dynamic understanding of the food web. We set out to identify and quantify the parasite community of Gammarus lacustris to understand the contributions of the amphipod and its parasites to the Takvatn food web. We identified seven parasite taxa: a direct life cycle gregarine, Rotundula sp., and larval stages of two digenean trematode genera, two cestodes, one nematode, and one acanthocephalan. The larval parasites use either birds or fishes as final hosts. Bird parasites predominated, with trematode Plagiorchis sp. having the highest prevalence (69%) and mean abundance (2.7). Fish parasites were also common, including trematodes Crepidostomum spp., nematode Cystidicola farionis, and cestode Cyathocephalus truncatus (prevalences 13, 6, and 3%, respectively). Five parasites depend entirely on G. lacustris to complete their life cycle. At least 11.4% of the overall parasite diversity in the lake was dependent on G. lacustris, and 16% of the helminth diversity required or used the amphipod in their life cycles. These dependencies reveal that in addition to being a key prey item in subarctic lakes, G. lacustris is also an important host for maintaining parasite diversity in such ecosystems.

Parasitic nematodes of marine fishes from Palmyra Atoll, East Indo-Pacific, including a new species of Spinitectus (Nematoda, Cystidicolidae).

Here, we present the results of a taxonomic survey of the nematodes parasitizing fishes from the lagoon flats of Palmyra Atoll, Eastern Indo-Pacific. We performed quantitative parasitological surveys of 653 individual fish from each of the 44 species using the intertidal sand flats that border the atoll's lagoon. We provide morphological descriptions, prevalence, and mean intensities of the recovered seven species of adult nematode (Pulchrascaris chiloscyllii, Capillariidae gen. sp., Cucullanus bourdini, Cucullanus oceaniensis, Pseudascarophis sp., Spinitectus (Paraspinitectus) palmyraensissp. nov., Philometra pellucida) and three larval stages (Pulchrascaris sp., Hysterothylacium sp., Cucullanus sp.). We recorded: Pulchrascaris chiloscyllii from Carcharhinus melanopterus; Capillariidae gen. sp. from Chaetodon lunula, Lutjanus fulvus, and Ellochelon vaigiensis; Cucullanus bourdini from Arothron hispidus; Cucullanus oceaniensis from Abudefduf sordidus; Pseudascarophis sp. from Chaetodon auriga, Chaetodon lunula, and Mulloidichthys flavolineatus; Spinitectus (Paraspinitectus) palmyraensissp. nov. from Albula glossodonta; Philometra pellucida from Arothron hispidus; and three larval forms, Pulchrascaris sp. from Acanthurus triostegus, Acanthurus xanthopterus, Rhinecanthus aculeatus, Platybelone argalus, Carangoides ferdau, Carangoides orthogrammus, Caranx ignobilis, Caranx melampygus, Caranx papuensis, Chaetodon auriga, Chanos chanos, Amblygobius phalaena, Asterropteryx semipunctata, Valencienea sexguttata, Kyphosus cinerascens, Lutjanus fulvus, Lutjanus monostigma, Ellochelon vaigiensis, Mulloidichthys flavolineatus, Upeneus taeniopterus, Gymnothorax pictus, Abudefduf septemfasciatus, Abudefduf sordidus, and Stegastes nigricans; Hysterothylacium sp. type MD from Acanthurus triostegus, Carangoides ferdau, Chaetodon lunula, Chanos chanos, Kyphosus cinerascens, Abudefduf sordidus, and Arothron hispidus; and Cucullanus sp. from Caranx ignobilis. Spinitectus (Paraspinitectus) palmyraensissp. nov. (Cystidicolidae) is described from the intestine of roundjaw bonefish Albula glossodonta. All the nematode species reported in this study represent new geographical records. We discuss how our survey findings compare to other areas of the Indo-Pacific, and the way the relatively numerical dominance of trophically transmitted larval stages likely reflect the intact food web of Palmyra Atoll, which includes a large biomass of large-bodied top predator sharks and ray-finned fishes.

Parasitic copepods (Crustacea, Hexanauplia) on fishes from the lagoon flats of Palmyra Atoll, Central Pacific.

We surveyed copepods parasitic on the fishes at Palmyra, a remote atoll in the Central Indo-Pacific faunal region. In total, we collected 849 individual fish, representing 44 species, from the intertidal lagoon flats at Palmyra and recovered 17 parasitic copepod species. The parasitic copepods were: Orbitacolaxwilliamsi on Mulloidichthysflavolineatus; Anuretesserratus on Acanthurusxanthopterus; Caligusconfusus on Carangoidesferdau, Carangoidesorthogrammus, Caranxignobilis, Caranxmelampygus, and Caranxpapuensis; Caliguskapuhili on Chaetodonauriga and Chaetodonlunula; Caliguslaticaudus on Rhinecanthusaculeatus, Pseudobalistesflavimarginatus, M.flavolineatus, Upeneustaeniopterus, Chrysipteraglauca, and Epinephalusmerra; Caligusmutabilis on Lutjanusfulvus and Lutjanusmonostigma; Caligusrandalli on C.ignobilis; Caligus sp. on L.fulvus; Caritusserratus on Chanoschanos; Lepeophtheiruslewisi on A.xanthopterus; Lepeophtheirusuluus on C.ignobilis; Dissonussimilis on Arothronhispidus; Nemesis sp. on Carcharhinusmelanopterus; Hatschekialongiabdominalis on A.hispidus; Hatschekiabicaudata on Chaetodonauriga and Chaetodonlunula; Kroyerialongicauda on C.melanopterus and Lernanthropus sp. on Kyphosuscinerascens. All copepod species reported here have been previously reported from the Indo-Pacific but represent new geographical records for Palmyra, demonstrating large-scale parasite dispersion strategies.

Monogenea of fishes from the lagoon flats of Palmyra Atoll in the Central Pacific.

A survey of the monogeneans of fishes from the lagoon flats of Palmyra Atoll detected 16 species already reported from the Indo-West Pacific faunal region. A total of 653 individual fish from 44 species were collected from the sand flats bordering the lagoon of the atoll. Eighteen species of fish were infected with monogeneans. The monogenean species recovered were: Benedenia hawaiiensis on Acanthurus xanthopterus, Chaetodon auriga, Chaetodon lunula, Mulloidichthys flavolineatus, Pseudobalistes flavimarginatus and Rhinecanthus aculeatus; Ancyrocephalus ornatus on Arothron hispidus; Euryhaliotrema annulocirrus on Chaetodon auriga and Chaetodon lunula; Euryhaliotrema chrysotaeniae on Lutjanus fulvus; Euryhaliotrema grandis on Chaetodon auriga and Chaetodon lunula; Haliotrema acanthuri on Acanthurus triostegus; Haliotrema aurigae on Chaetodon auriga and Chaetodon lunula; Haliotrema dempsteri on Acanthurus xanthopterus; Haliotrema minutospirale on Mulloidichthys flavolineatus; Haliotrematoides patellacirrus on Lutjanus monostigma; Neohaliotrema bombini on Abudefduf septemfasciatus and Abudefduf sordidus; Acleotrema girellae and Acleotrema parastromatei on Kyphosus cinerascens; Cemocotylella elongata on Caranx ignobilis, Caranx melampygus and Caranx papuensis; Metamicrocotyla macracantha on Crenimugil crenilabris; and Pseudopterinotrema albulae on Albula glossodonta. All these monogenean-host combinations represent new geographical records. The monogenean species composition of the Palmyra Atoll is similar to that of the Hawaiian Islands. However, the number of species recovered was lower compared with other localities within the Indo-West Pacific, perhaps due to the geographical isolation of Palmyra Atoll.

Molecular analyses reveal high species diversity of trematodes in a sub-Arctic lake.

To identify trematode diversity and life-cycles in the sub-Arctic Lake Takvatn, Norway, we characterised 120 trematode isolates from mollusc first intermediate hosts, metacercariae from second intermediate host fishes and invertebrates, and adults from fish and invertebrate definitive hosts, using molecular techniques. Phylogenies based on nuclear and/or mtDNA revealed high species richness (24 species or species-level genetic lineages) and uncovered trematode diversity (16 putative new species) from five families typical in lake ecosystems (Allocreadiidae, Diplostomidae, Plagiorchiidae, Schistosomatidae and Strigeidae). Sampling potential invertebrate hosts allowed matching of sequence data for different stages, thus achieving molecular elucidation of trematode life-cycles and exploration of host-parasite interactions. Phylogenetic analyses also helped identify three major mollusc intermediate hosts (Radix balthica, Pisidium casertanum and Sphaerium sp.) in the lake. Our findings increase the known trematode diversity at the sub-Arctic Lake Takvatn, showing that digenean diversity is high in this otherwise depauperate sub-Arctic freshwater ecosystem and indicating that sub-Arctic and Arctic ecosystems may be characterised by unique trematode assemblages.

Comparing mechanisms of host manipulation across host and parasite taxa.

Parasites affect host behavior in several ways. They can alter activity, microhabitats or both. For trophically transmitted parasites (the focus of our study), decreased activity might impair the ability of hosts to respond to final-host predators, and increased activity and altered microhabitat choice might increase contact rates between hosts and final-host predators. In an analysis of trophically transmitted parasites, more parasite groups altered activity than altered microhabitat choice. Parasites that infected vertebrates were more likely to impair the host's reaction to predators, whereas parasites that infected invertebrates were more likely to increase the host's contact with predators. The site of infection might affect how parasites manipulate their hosts. For instance, parasites in the central nervous system seem particularly suited to manipulating host behavior. Manipulative parasites commonly occupy the body cavity, muscles and central nervous systems of their hosts. Acanthocephalans in the data set differed from other taxa in that they occurred exclusively in the body cavity of invertebrates. In addition, they were more likely to alter microhabitat choice than activity. Parasites in the body cavity (across parasite types) were more likely to be associated with increased host contact with predators. Parasites can manipulate the host through energetic drain, but most parasites use more sophisticated means. For instance, parasites target four physiological systems that shape behavior in both invertebrates and vertebrates: neural, endocrine, neuromodulatory and immunomodulatory. The interconnections between these systems make it difficult to isolate specific mechanisms of host behavioral manipulation.

Reef fishes have higher parasite richness at unfished Palmyra Atoll compared to fished Kiritimati Island.

We compared parasite communities at two coral atolls in the Line Islands chain of the central Pacific (Kiritimati Island and Palmyra Atoll). Palmyra Atoll is relatively pristine while Kiritimati Island is heavily fished. At each island, we sampled five fish species for helminth and arthropod endoparasites: Chromis margaritifer, Plectroglyphidodon dickii, Paracirrhites arcatus, Acanthurus nigricans, and Lutjanus bohar. The surveys found monogeneans, digeneans, cestodes, nematodes, acanthocephalans, and copepods. Parasite richness was higher at Palmyra compared to Kiritimati for all five fish species. Fishes from Palmyra also tended to have more parasites species per host, higher parasite prevalence, and higher parasite abundance than did fishes from Kiritimati. The lower parasitism at Kiritimati may result from a simplified food web due to over fishing. Low biodiversity could impair parasite transmission by reducing the availability of hosts required by parasites with complex life cycles. Most notably, the lower abundances of larval shark tapeworms at Kiritimati presumably reflect the fact that fishing has greatly depleted sharks there in comparison to Palmyra.

Ecosystem energetic implications of parasite and free-living biomass in three estuaries.

Parasites can have strong impacts but are thought to contribute little biomass to ecosystems. We quantified the biomass of free-living and parasitic species in three estuaries on the Pacific coast of California and Baja California. Here we show that parasites have substantial biomass in these ecosystems. We found that parasite biomass exceeded that of top predators. The biomass of trematodes was particularly high, being comparable to that of the abundant birds, fishes, burrowing shrimps and polychaetes. Trophically transmitted parasites and parasitic castrators subsumed more biomass than did other parasitic functional groups. The extended phenotype biomass controlled by parasitic castrators sometimes exceeded that of their uninfected hosts. The annual production of free-swimming trematode transmission stages was greater than the combined biomass of all quantified parasites and was also greater than bird biomass. This biomass and productivity of parasites implies a profound role for infectious processes in these estuaries.

An efficient strategy to estimate intensity and prevalence: sampling metacercariae in fishes.

Accurate estimates of population-level parameters of parasites, such as prevalence and mean intensity, require large sample sizes. The processing of such samples becomes an overwhelming task when parasites are abundant, as with trematode metacercariae in fishes. In the present study, a subsampling method reduced processing time while maintaining an accurate estimation of metacercariae prevalence and intensity across 3 trematode species and 2 fish species. By double sampling, we generated regression models to predict total intensity from a combination of subsamples. The key to this approach lies in choosing the best strategy from a large number of potential subsampling routines. We selected the most efficient routine by weighing the costs and benefits of each. This approach, however, could not provide an estimate of parasite abundance when no parasites occurred in the initial subsample. To estimate prevalence accurately, our subsampling algorithm prescribed an additional sampling sequence using a new, optimal regression model. In addition, we optimized the technique to measure three parasite species infecting a single host simultaneously. This efficient subsampling procedure decreased the overall processing time per host by up to 91% while obtaining accurate (R2 > 0.8) estimates for both prevalence and intensity.