Changes in the parasite fauna of whiting Odontogadus merlangus L. with age and sex of host, season, and from different areas in the vicinity of the Isle of Man

Infestation of whiting with the helminths Derogenes varicus Müller, Hemiurus communis Odhner, Grillotia erinaceus Van Beneden (plerocercoids) and Contracaecum clavatum Rudolphi, and the copepod Lernaeocera branchialis L., increased with the age of the fish but infestation with the helminths Lecithaster gibbosus Rudolphi, Stephanostomum pristis Looss and Podocotyleatomon and the copepod Clavella uncinata Müller decreased as the fish aged. Infestation with the gill monogenean Diclidophora merlangi Kuhn remained at much the same level in fish 1–5+ years old. Derogenes, Hemiurus, Lecithaster, Cotracaecum and Stephanostomum showed seasonal fluctuations in the levels of their populations in whiting. In all cases the parasites reached a main peak in the summer and early autumn, a time when juvenile digeneans were most apparent.
Whiting from Morecambe Bay were less infested with Clavella than whiting taken from other areas. Experimental evidence suggests that the lower salinity of Morecambe Bay may be responsible; however, this could not account for the lower infestation of Morecambe Bay whiting by Diclidophora which appeared to be unaffected by a lowered salinity. Whiting from the east side of the Isle of Man were more infested with Lernaeocera but less infested with Diclidophora than fish from the west side of the island. The myxosporidian Myxobolus aeglefini Auerbach was externally visible in the sclera of whiting from the east side of the island but absent from the sclera of whiting from the west side of the island. It is, therefore, suggested that whiting to the east of the Isle of Man form a separate sub-stock from those to the west of the island.     

The effects of Lernaeocera branchialis on the Merlangius merlangus population in the Medway Estuary

Whiting entering estuarine waters during autumn are largely free from the copepod Lernaeocera branchialis (L.). Infection soon follows and, between 1973 and 1975, an average 40.4% whiting were infected. The parasite's effects on fish weight over this period are assessed and, in a separate study, its effects on the host's lipid biochemistry determined. Mature adults of L. branchialis caused a significant reduction in body weight. Liver lipid content was also lower in these individuals, by more than 50%. Phospholipid levels were lower but cholesterol readings were greater in infected fish. Fatty acid composition of liver lipids appeared unaffected.     

Copepod ectoparasites of Merlangius merlangus and Platichthys flesus

Seasonal changes in the levels of copepod ectoparasitic infections from Merlangius merlangus and Platichthys flesus were examined and related to the annual migrations of young fish into an estuary. Lernaeocera branchialis (L.) was the only parasite common to both fish species; its life-cycle stages infected the branchial chambers of 96·7% flounders ( P. flesus ) and 40·4% whiting ( M. merlangus ). Infection in the two fish species exhibited similar seasonal fluctuations and a temporal relationship between the parasite's occurrence on its two hosts was apparent. Whiting were also parasitized by Clavella uncinata (Milller); 26·9 % fish were affected. The copepods' distribution was highly localized with the majority restricted to the long gill rakers of the first gill arch. Infection was related to size of whiting up to 12·5 cm; there was little increase beyond this length. In addition to L. branchialis , flounder was also host to Acanthochondria depressa (T. Scott) and Lepeophtheirus pectoralis (Müller). A. depressa showed a preference for the ocular side of the body with its developing stages attached mainly to the second gill arch and the adults located in the posterior pocket of the branchial chamber. L. pectoralis was associated with the paired fins. Factors which could result in such localized distributions are discussed.     

The common dab as definitive host for the Pennellid copepods Lernaeocera branchialis and Haemobaphes cyclopterina

Seven (0.039%) and five (0·028%) of 18 163 common dab from the east coast of Scotland were definitive hosts to Lernaeocera branchialis and to Haemobaphes cyclopterina , respectively. Both represent new host records. Muscle necrosis and localized haemorrhage were associated with H. cyclopterina infestation and as described from other definitive hosts.     

Experimental studies on the development of Lernaeocera branchialis (Copepoda: Pennellidae): population processes from egg production to maturation on the flatfish host

Quantitative population dynamical information derived from laboratory- and field-based experiments is provided for the fish-parasitic copepod, Lernaeocera branchialis, infecting flounder (Platichthys flesus) and whiting (Merlangius merlangus). Adult, post-metamorphosis females from whiting can produce more than one set of egg-strings. The mean number of eggs in each egg-string pair was 1445. At 10 °C these eggs took about 12.7 d after extrusion before hatching of NI nauplii began. Hatching took up to 12 days to be completed with an exponentially declining pattern of output over this period. In the laboratory about 44% of the egg-string egg population successfully passed through the NI to NII nauplius moult and the NII to copepodid moult to produce infective copepodids, a process lasting about 2 d. The non-feeding copepodids had a maximum survival time at 10 °C of 18 d, with a time to 50% survival of 7.5 d. In laboratory infection experiments at 10 °C, copepodids infected flounder and passed through all their developmental stages to adulthood and copulation in a minimum of 25 d. Field experiments on the seabed off Lowestoft in June 1987 with a sea temperature of about 16 °C suggested that the developmental period in those conditions could be as short as 11 d. Previously uninfected flounder in the field experiments became naturally infected with copepodids at a mean rate of not less than 30 parasites per fish d^–1.     

Effect of concurrent infections of Lernaeocera Branchialis (Copepoda) and Trypanosoma murmanensis (Protozoa) on Atlantic cod, Gadus morhua

A study was initiated to ascertain the prevalence, effect and interaction of the adult stages of the parasitic copepod, Lernaeocera branchialis, on Atlantic cod concurrently infected with a hematozoan, Trypanosoma murmanensis, by comparing condition (K) factor, organ somatic indices, hematological values and lipid concentrations of the liver from infected and uninfected fish of comparable length. Prevalence of the copepod varied from 9 to 21% and was highest in fish examined on the southern coast of Newfoundland. Body condition and blood values were significantly lower in young cod infected with two parasites whereas only K-factor was altered in two size classes parasitized by one copepod. Although lipid concentrations were similar in three length groups of fish without and infected with L. branchialis, the pooled values were significantly greater among the infected group. Mortality occurred only in juvenile cod infected with the copepod and T. murmanensis concurrently. Cod which survived dual infections had lower K-factor and blood values than controls or fish harboring single, parasitic infections. It is concluded that an infection with an adult L. branchialis does not impair the health of fish in this area of the northwestern Atlantic Ocean, but the presence of T. murmanensis concurrently can potentiate its effect.     

Survey of the metazoan ectoparasites of the European flounder Platichthys flesus (Linnaeus, 1758) along the north-central Portuguese coast

A survey was undertaken to identify metazoan ectoparasite species on the European flounder, Platichthys flesus (Linnaeus, 1758), in 4 different locations off the north-central Portuguese coast. Parasites of 7 different taxa were found: Caligus diaphanus, Caligus sp., and Lepeophtheirus pectoralis (Copepoda: Caligidae); Acanthochondria cornuta (Copepoda: Chondracanthidae); Holobomolochus confusus (Copepoda: Bomolochidae); Nerocila orbignyi (Isopoda: Cymotholdae); and praniza larvae (Isopoda: Gnathiidae). Lernaeocera branchialis, a common European flounder parasite in the North and Baltic Seas, was not observed among the surveyed fish. Caligus diaphanus, Caligus sp., and Nerocila orbignyi are new host records. The high prevalence and intensity values recorded for L. pectoralis and A. cornuta suggest that both parasite species are common to the European flounder along the north-central Portuguese coast. In contrast, infection levels with respect to the other parasite taxa were, in most cases, comparatively lower, thereby indicating that they only occur occasionally among flounders in the surveyed area.     

Notes on helminth parasites of the whiting Odontogadus m. merlangus (L.) from the northern Irish Sea

In an examination of 1352 whiting from the Irish Sea 13 helminth species were discovered. One species, Scolex polymorphus Rudolphi, was a new species associated with whiting and 7 other species were new species for whiting from the Northern Irish Sea. The species of the digenean genus Podocotyle found in whiting is probably an ecotype of P. atomon. It was concluded that the bothridial structure of 5. polymorphus cannot be used for diagnostic purposes nor to determine early and late stages of the larvae. The number of Diclidophora merlangi Kuhn (Monogenea) from each monthly sample with eggs in the uterus is directly proportional to temperature, but egg production under experimental conditions was greatest at 6° C. Several theories are proposed to explain these apparently contradictory observations. The life-history of the digenean Stephanostomum pristis Looss, appears to be unusual in that the 2nd intermediate host may be planktonic. The nematode Contra caecum clavatum Rudolphi is found both as larvae and adults but larvae comprise the greater part of the worm population. The discovery of small stage II larvae in the stomach of whiting supports the theory that only one intermediate host is necessary for the development of Contmcaecum . There were significantly more female C. clavatum than male but a 1: 1 sex ratio is demonstrated by the acanthocephalan Echinorhynchus gadi Muller. Early and late forms of the plerocercoid of Grillotia erinaceus Van Beneden are identified. The early form occurred predominantly in the young whiting.     

Experimental transmission, development, and effects of a parasitic copepod, Lernaeocera branchialis, on Atlantic cod, Gadus morhua

Development and effects of adult Lernaeocera branchialis were studied following experimental transmission to Atlantic cod, Gadus morhua. Growth from the detection of pennella stages to mature adults was approximately 9-10 mo (September-June of the following year) at which time eggs were released and the adult parasites degenerated. Cod from all size groups were susceptible but the prevalence of infection was greater in small fish. Peak mortality, about 30%, occurred within 4 mo after infection and was greatest in young fish with multiple infections. Death was associated with emaciation, blood loss, open lesions, and probable occlusion of branchial blood vessels and/or ventral aorta. Infected fish, especially subadults that survived, consumed less food, gained less weight, had a lower conversion factor, and were substantially smaller than uninfected controls through a 32-wk period. Many of these fish displayed hyperactivity. Large cod that survived the infection and harboured 1 parasite were apparently unaffected but died when stressed. A previous infection conferred no protection against reinfection. Multiple infections delayed gonadal maturation and resulted in significantly lower gonadal somatic indices than in controls. Exposure of infected fish to crude oil fractions or to infection with a hemoprotozoan, Trypanosoma murmanensis, culminated in mortality, weight loss, or low organ somatic indices. It is estimated that considerable losses through mortality and weight gain in young fish occur each year in coastal areas from infections by L. branchialis, particularly in one area where it was estimated that 20% of the population was infected.     

Ultrastructural studies on the early cuticular metamorphosis of adult female Lernaeocera branchialis (L.) (Copepoda,Pennellidae)

After its final moult and fertilization an adult female of the marine fish-parasitic copepod, Lernaeocera branchialis, begins an extensive metamorphosis. This commences while the parasite is still on the flounder intermediate host and is completed once the female has established itself on the whiting final host. One early component of the metamorphosis is a considerable elongation of the parasite's abdominal region. S.e.m. and t.e.m. studies have revealed that part of the mechanism of the elongation consists of a straightening out of a highly folded abdominal cuticle. Before fertilization, the epicuticle and outer procuticular layers of this integument are thrown into a series of transverse, 4–6 µm deep pleats or folds with a density of 1–1.2 folds/µm of abdominal length. Straightening these folds can generate an approximately 6-fold length increase. The folds are already present beneath the female chalimus IV cuticle when the epidermis of this development stage starts to secrete the adult cuticle. Immediately before the final moult, the adult cuticle is super-folded with the whole cuticle displaying second-order folds, 8–10 µm deep.The capacity of Lernaeocera to engage in extensive cuticular modifications without recourse to a moult is compared with similar abilities shown by some insect species.     

Effect of sedimentological and hydrodynamical changes in the intertidal areas of the Oosterschelde estuary (SW Netherlands) on distribution,density and biomass of five common macrobenthic species: Spio martinensis (Mesnil), Hydrobia ulvae (Pennant), Arenicola marina (L.), Scoloplos armiger (Muller) and Bathyporeia sp.

In order to evaluate the impact of the construction of the storm-surge barrier and secondary dams on macrobenthos of the tidal flats in the Oosterschelde (SW Netherlands), changes in distribution, density and biomass of five common species (Spio martinensis, Hydrobia ulvae, Arenicola marina, Scoloplos armiger and Bathyporeia sp) were analysed.Data on macrobenthos were collected from 1979 to 1989 on five different tidal flats. Changes in sediment texture and hydrodynamic factors during the construction and after the completion of the coastal engineering project were taken into account.Three severe winters in a row caused more disturbance in the population of the main predator of S. armiger than did the hydrodynamical changes. A temporary prolongation of the emersion time (in 1986 and 1987) caused a temporary decrease in juvenile A. marina. But afterwards they still occupy the same nursery grounds. Increased wave action on the edges of the flats probably created new niches for Bathyporeia sp. and Spio martinensis, replacing other benthic species.It is not yet clear what has caused the decline of H. ulvae in many places in the Oosterschelde estuary. Parasitic infestation is one of the possibilities.     

Seasonal population dynamics of Paraquimpevia tenevvima (Linstow) (Nematoda) in the European eel Anguilla anguilla (Linnaeus) in three localities of southwest England

Seasonal population dynamics of Puraquzmperia tenerrima (Linstow) in the European eel Anguillu anguillu (Linnaeus) have been investigated in three localities in Devon, southwest England. The nematode is common and exhibits very similar seasonal patterns in changes of prevalence and abundance in the three localities. The population increases over winter through to late spring or early summer as the recruitment of the new generation occurs throughout this period. Reproduction of the nematode starts in spring. A sharp decrease of prevalence and abundance accompanies or follows reproduction, and infection levels are very low in autumn and early winter. The dispersion pattern of the nematode varies with the infection levels from over-dispersion in spring and early summer to almost random dispersion in autumn and winter. It is suggested that P. tenerrimu is a common specific parasite of eels, especially in small streams or rivers and that insect larvae or species of crustaceans might serve as intermediate hosts for the nematode.     

The population dynamics of the parasitic copepodeLernaeocera lusci (Bassett-Smith, 1896) on its definitive host

The mesoparasitic copepodLernaeocera lusci (Bassett-Smith, 1896) was recovered from first-year bib (Trisopterus luscus L.) in the Voordelta (Southern Bight of the North Sea) from May until December 1989. Analysis of the seasonal abundance and of the population structure showed that transmission of infective stages to bib mainly occurred from June to September. From September to December the overall prevalence fluctuated around 70%. Maximum parasite population size (47/10⁴m²) and the highest total egg number were recorded in September and October, respectively. It was found that total parasite mortality was significantly influenced by mortality of hosts carrying parasites. Natural mortality probably contributed a small percentage to total parasite mortality. Calculation of the temporal mean-variance regression equation revealed that the parasites were aggregated within the definitive host population.     

Is reduced body growth of cod exposed to the gill parasite Lernaeocera branchialis a cost of resistance?

Three hundred and sixty-nine cod Gadus morhua were individually marked and caged for 19 months. During this period, each cod was inspected several times for Lernaeocera branchialis . Growth in four groups of cod, identified by their infection history, were compared. During the caging, 79% of the cod remained uninfected, 8·5% were infected, but lost the parasite, 8% were infected with one parasite and 4·5% were infected with more than one parasite. The infected fish either harboured the parasite at caging or were infected during the study period. The highest rate of increase, both in body mass and in standard length ( L _S), was recorded in the group of male fish infected with one parasite throughout the experimental period. Conversely, those males free from infection showed significantly lower growth. The observed differences in growth could not be explained by changes in variables related to reproductive strategies. The alternative explanation for these results is that resistance to L. branchialis was associated with costs in terms of reduced growth of body mass and L _S.     

Infection of estuarine fish populations by Cryptocotyle lingua (Creplin)

Fish populations in the Medway Estuary become infected with Cryptocotyle lingua (Creplin) shortly after their inshore migrations. The present study examines infection levels between 1973 and 1975 in four fish species; whiting, Merlangius merlangus (L.), pouting, Trisopterus luscus (L.), plaice, Pleuronectes platessa L. and flounder, Platichthys flesus (L.). Frequency of infection was highest in flounder, followed by pouting, plaice and whiting. Unusually, parasite density was lowest in plaice even though gastropod molluscs (the parasite's first intermediate host) were most common in their diet. Seasonal levels of infection are discussed and prove useful indicators to the migratory movements of individual fish populations. In whiting and pouting, incidence of infection increased with length of host while no relationship between infection and age of host was observed in either flatfish. Metacercariae were not distributed randomly over the body surface; in gadoids they were concentrated above the lateral line while in pleuronectids they were most intense in the fins and gills.     

The fish populations in inshore waters of the West Coast of Scotland. The distribution, abundance and growth of the whiting (Merlangius merlangus L.)

The whiting ( Merlangius merlangus ) is an important commercial species from the West Coast of Scotland. This paper describes the distribution, abundance, age composition and growth of juvenile whiting which are found in the inshore areas and sea lochs. No spawning was observed in the areas and it is assumed there was an active migration of whiting from the open sea. They were first caught by the bottom trawl in July, reached maximum abundance in November and December and thereafter declined in numbers until few remain by early summer. An exception to this occurs in Loch Etive and possible explanations for the fact that whiting tend to remain for longer periods and grow at a slower rate in this loch are discussed.     

Population stability over a ten-year period in the short-lived fish Liparis liparis (L.)

A ten-year study of the population dynamics of the sea-snail, Liparis lipuris , in the Bristol Channel, England is reported. This small fish which only lives for 1 year undertakes a regular seasonal migration from birth in marine waters into the estuary from which it retreats during the winter to return to the spawning grounds by early spring. At Hinkley Point, Somerset (salinity 22–3O%), abundance has been shown to be negatively correlated with water temperature presumably because of the avoidance of warmer inshore waters during mild winters. When the temperature effect is removed from the data the peak winter abundance of the population was found to have a coefficient of variation of only 27%. For a population which only comprises of age group individuals, this shows a remarkable stability. No statistically significant relationship was found between the abundance of sea-snail and either their predators or prey. However, there were indications of a negative relationship with the abundance of whiting, Merlangius merlangus , which was the most abundant predator. We conclude that the population of a short-lived marine fish living within a physically variable marine environment can be constrained within tight limits. Further. sea-snail reproductive success has been independent of variation in local physical conditions.     

Lernaeocera branchialis: a potential pathogen to cod ranching

A study was carried out to determine the effect of Lernaeocera branchialis on Atlantic cod infected in the laboratory and in the field and also to ascertain its effect on cod-ranching. Sixty-four percent (308) of 481 cod acquired infections in the laboratory and 33% (159) of the infected fish died over a 4-yr period. About 74% of the deaths occurred within 4 mo of the infection. Monthly samples of cod collected adjacent to a cod-ranching operation showed an initial prevalence of 30% that subsequently decreased in the following 2 mo to 15%. Prevalence of the infection also decreased among the initial field sample of cod that were kept alive, from 30 to 17% during the same 2-mo period and to 9% after 8 mo and was associated with death caused by the parasite. Cod examined at intervals after infection showed evidence of reduced weight gain, lower liver somatic index, liver lipid, and blood values than controls. A field sample taken from the same area during the summer of the following year indicated a prevalence of 12%. This higher than usual prevalence (4-6%) was associated with retention of the intermediate host, Cyclopterus lumpus, that provided an additional source of infective stages. It is likely that increased parasitism could affect the success of the cod-ranching operation in view of the parasite's devastating effects on its hosts.     

The population biology of Camallanus lacustris (Zoega) in eels, Anguilla anguilla (Linnaeus), and their status as its host

The population biology of Camallanus lacustris (Zoega) and the status of the eel, Anguilla anguilla (Linnaeus), as its definitive host have been studied in a small Devon lake. No clear seasonal pattern in prevalence and abundance was observed, and recruitment of the new generation may occur in all seasons. However, reproduction exhibits some seasonality as adult nematodes were the commonest stage in the parasite population from early summer onwards and production of first-stage larvae appeared to occur primarily during this period. These larvae were viable, and shown experimentally to be infective to copepods. The occurrence of the nematode and its development to full maturity in eels, the absence of its typical definitive host ( Perca fluviatilis Linnaeus) from the lake and the absence of the nematode from other species of fish in the lake indicate that eels serve as the only, true definitive host of the nematode in this locality. It is suggested that account must be taken of this alternative host when the population of C. lacustris is investigated in the future.     

Sex, horizontal transmission, and multiple hosts prevent local adaptation of Crithidia bombi, a parasite of bumblebees (Bombus spp.)

Local adaptation within host-parasite systems can evolve by several non-exclusive drivers (e.g., host species-genetic adaptation; ecological conditions-ecological adaptation, and time-temporal adaptation). Social insects, especially bumblebees, with an annual colony life history not only provide an ideal system to test parasite transmission within and between different host colonies, but also parasite adaptation to specific host species and environments. Here, we study local adaptation in a multiple-host parasite characterized by high levels of horizontal transmission. Crithidia bombi occurs as a gut parasite in several bumblebee species. Parasites were sampled from five different host species in two subsequent years. Population genetic tools were used to test for the several types of adaptation. Although we found no evidence for local adaptation of the parasite toward host species, there was a slight temporal differentiation of the parasite populations, which might have resulted from severe bottlenecks during queen hibernation. Parasite populations were in Hardy-Weinberg equilibrium and showed no signs of linkage disequilibrium suggesting that sexual reproduction is an alternative strategy in this otherwise clonal parasite. Moreover, high levels of multiple infections were found, which might facilitate sexual genetic exchange. The detection of identical clones in different host species suggested that horizontal transmission occurs between host species and underpins the lack of host-specific adaptation.