Spatial distributions of two species of monogeneans on the gills of Siganus fuscescens (Houttuyn) and their seasonal dynamics in caged versus wild-caught hosts

Two monogeneans species, Polylabris mamaevi Ogawa & Egusa, 1980 and Tetrancistrum nebulosi Young, 1967, are sympatric on the gills of Siganus fuscescens (Houttugn, 1782) in the South China Sea. An investigation into their distribution on the host's gills based on monthly samples from both wild and caged populations revealed that these 2 species co-occurred on the same gill arches, but they were segregated. P. mamaevi attached to the distal half of the gill filaments, whereas T. nebulosi attached to the proximal half. This difference may partly be attributed to microhabitat specialization. Microhabitat distributions were intensity dependent for both species. As intensity increased, monogeneans were increasingly found on the second, third, and even fourth gill arches, and niche breadths increased. The changes in distribution and niche breadths at the host gill arch level in relation to differences in host size and sampling period were considered to represent a functional response to increased intensity. Differences in host resources, in either net-caged or wild populations, did not significantly influence the parasite distributions at the host gill arch level. Simultaneous infections of these 2 species were common, but there was no evidence of negative interspecific interactions.     

黄颡鱼鳃部寄生单殖吸虫和鳋类的空间分布特点

研究了黄颡鱼Pelteobagrus fulvidraco(Richardson)鳃部的寄生单殖吸虫(Monogeneans)和鳋类(Copepods)的空间分布特点.寄生于黄颡鱼鳃部的单殖吸虫和鳋类在两侧鳃上的感染强度及其差异表明,两种寄生虫对于鳃片的左右位置没有明显的选择性;在黄颡鱼四鳃间的分布存在极显著的差异(PP<0.01);在鳃片的各区均存在选择性;对鳃丝的各分段也均有极显著的选择性.       

Ecology of a diplozoon parasite on the gills of the African cyprinid Barbus neumayeri

In oxygen‐deficient waters, the difficulties of oxygen uptake in gill parasites and their fish hosts may influence host and parasite densities, site selection by the parasite, and effects of the parasite on host condition. This study quantified the prevalence and intensity of the gill monogenean Neodiplozoon polycotyleus in the African cyprinid fish Barbus neumayeri from an intermittent forest stream in western Uganda. Oxygen levels were low in the stream over the 12‐month study, averaging only 2.5 mg litre^?1 (monthly range = 1.2–4.3 mg litre^?1). However, parasite prevalence was high (47.2%), suggesting high tolerance to low oxygen in N. polycotyleus. The prevalence of parasites varied with host body size, with the highest frequency of occurrence in the middle size classes. Prevalence also varied over the year; seasonal peaks of rainfall coincided with a lower frequency of N. polycotyleus. The significantly nonrandom frequency distribution of parasites among hosts suggests regulation of parasite numbers. Of the hosts infected, 37.1% harboured one N. polycotyleus parasite, and 62.9% harboured two parasites. No fish were infected with more than two diplozoons. There was evidence for strong site specificity by N. polycotyleus within hosts; 77.7% of the parasites were located on the filaments of the second gill arch, which may relate to increased oxygen availability. In addition, only one of the 178 infected fish had more than one parasite on one side of the branchial basket. Although N. polycotyleus is undoubtedly parasitic, we found no evidence of a negative parasitic effect on the condition or reproductive status of B. neumayeri.     

Gill monogeneans of the chub mackerel, Scomber japonicus from Madeiran waters of the Atlantic Ocean, Portugal

Five species of monogeneans were recovered from the gill filaments of 181 chub mackerel, Scomber japonicus, from the Madeiran waters of the Atlantic Ocean, Portugal, during 2004/2005. The monogenean Pseudokuhnia minor showed the highest prevalence (98.68%) and a mean intensity of 28.23, followed by Kuhnia scombri (prevalence of 43.71% and mean intensity of 2.69) and K. scombercolias (prevalence of 39.1% and a mean intensity of 1.81). Kuhnia sprostonae and Grubea cochlear were rare, occurring in only one and five fish hosts respectively. No correlation between fish host length and mean intensity of infection with the three most abundant monogeneans was found. However, significant differences in prevalence and abundance were found in relation to date of sampling for P. minor, and all parasites were aggregated in their distribution.     

Evolutionary expansion of the Monogenea

The evolutionary expansion of the monogeneans has taken place in parallel with the diversification of the fish-like vertebrates. In this article the main trends in monogenean evolution are traced from a hypothetical skin-parasitic ancestor on early vertebrates. Special consideration is given to the following topics: early divergence between skin feeders and blood feeders; diversification and specialization of the haptor for attachment to skin; transfer from host to host, viviparity and the success of the gyrodactylids; predation on skin parasites and camouflage; colonization of the buccal and branchial cavities; diversification and specialization of the haptor for attachment to the gills; phoresy in gill parasites; the development of endoparasitism and the origin of the cestodes; the success of dactylogyroidean gill parasites; the uniqueness of the polyopisthocotyleans; ovoviviparity and the colonization of the tetrapods. Host specificity has been the guiding force of coevolution between monogeneans and their vertebrate hosts, but the establishment of monogeneans on unrelated hosts sharing the same environment (host-switching) may have been underestimated. Host-switching has provided significant opportunities for evolutionary change of direction and is probably responsible for the establishment of monogeneans on cephalopod molluscs, on the hippopotamus and possibly on chelonians. There are indications that host-switching may be more common in monogeneans that spread by direct transfer of adults/juveniles from host to host. A limitation on the further expansion of monogeneans is the need for water for the dispersal of the infective larva (oncomiracidium).     

Host impact of monogenean Lepidotrema bidyana infection and intensity estimates for onsite monitoring

We developed a rapid effective method for accurate estimation of intensity for the monogenean Lepidotrema bidyana, a gill parasite of silver perch Bidyanus bidyanus. This parasite requires monitoring because high-intensity infections reduce host growth and can lead to secondary bacterial and fungal infections. The most accurate method for counting L. bidyana was visual examination of fresh gills. There was a significant relationship between fish size and parasite intensity; however, there was no significant relationship between fish condition and parasite intensity. Parasite intensity estimates were generated by using the mean intensity of worms on the posterior hemibranch on the first left gill arch, compared to the total mean intensity of worms on all hemibranchs. Estimates were validated by predicting L. bidyana intensity from a random sample of silver perch obtained from aquaculture ponds. Parasite intensity estimates correlated strongly to real counts, and this method can be used to accurately predict parasite intensity on an individual host, and thus represents an improvement over previous methods.     

Infection dynamics of Cichlidogyrus tilapiae and C. sclerosus (Monogenea, Ancyrocephalinae) in Nile tilapia (Oreochromis niloticus L.) from Uganda

The infection dynamics of the gill monogeneans Cichlidogyrus tilapiae and C. sclerosus on Oreochromis niloticus with respect to habitat type (reservoir, stream, ponds and cages), host sex, size and seasons was determined between January and November 2008. During the study period, 45.2% of the 650 fish examined were infected with Cichlidogyrus spp. The infected hosts harboured an average of 8.6 ± 3.4 parasites/fish. Across habitat types, the proportion of infected fish was not statistically different. In contrast, the number of parasites recorded on infected fish from different habitat types differed significantly. The highest parasite number was recorded in reservoir-dwelling fish and lowest in stream-dwelling hosts. Concerning sex, more female O. niloticus were infected and harboured a high number of parasites than male and sexually undifferentiated fish. A weak negative relationship was found between rainfall and monthly parasite infections. However, a higher number of parasites and proportion of infected hosts were found during dry than in wet seasons, except in ponds. Results of this study show that differential exposure due to changes in fish behaviour associated with habitat modification and sex may account for the infection difference across the sampled sites. Meanwhile, rainfall and the associated hydrological events are important factors regulating monogenean infections in tropical aquatic environments. The continuous presence of Cichlidogyrus spp. in fish provides evidence of possible parasite outbreaks, indicating the application of biosecurity measures as crucial for the success of intensive fish farming.     

Mast cell responses to Ergasilus (Copepoda), a gill ectoparasite of sea bream

Immunocytochemical, light microscopy and ultrastructural studies were conducted on gill of sea bream, Sparus aurata L., naturally parasitized with the important parasitic copepod Ergasilus sp. to assess pathology and cellular responses. Thirty-seven S. aurata were examined from a fish farm; 26 (70%) were parasitized, with infection intensity ranging from 3 to 55 parasites per fish. Hosts were divided into two groups, lightly infected fish (<15 parasites per fish) and heavily infected fish (>15 parasites per fish). In histological sections, the copepod encircled gill lamellae with its second antennae, compressed the epithelium, provoked hyperplasia and hemorrhage, occluded arteries and often caused lamellar disruption. Fusion of the secondary lamellae due to epithelial hyperplasia was common in all infected fish; heavily infected fish showed more intense branchial inflammation. In both healthy and infected fish, mast cells (MCs) were free within the connective tissue inside and outside the blood vessels of the primary lamellae and made close contact with vascular endothelial cells, mucous cells and rodlet cells (RCs). MCs were irregular in shape with a cytoplasm filled by numerous electron-dense, membrane-bound granules. Immunostaining of primary and secondary gill filaments with an antibody against the antimicrobial peptide (AMP) piscidin 3 (anti-piscidin 3 antibody, anti-HAGR) revealed a subpopulation of MCs that were positive. These MCs were more abundant in gills of heavily infected fish than in either lightly infected or uninfected fish (ANOVA, P<0.05). Our report documents the response of gill to ectoparasite infection and provides further evidence that mast cells and their AMPs may play a role in responding to branchial ectoparasite infections.     

Lernanthropus kroyeri infections in farmed sea bass Dicentrarchus labrax: pathological features

Twenty sea bass Dicentrarchus labrax L. from a fish farm (floating cage) in Greece were examined for the presence of parasites. The gills of 7 (35%) fish were infected with adult female specimens of the parasitic copepod Lernanthropus kroyeri van Beneden, 1851, and the intensity of infection ranged from 1 to 24 parasites per host. The most infected portion of the gills appeared to be the primary lamellae. Erosion, desquamation and necrosis of the secondary lamellae were noticed near the site of copepod attachment; furthermore, the terminal claw of the second antennae lacerated tissue and vessels of infected gill. Parasitism by L. kroyeri affected the host's condition factor (mean +/- SE in uninfected vs parasitized; 1.88 +/- 0.04 vs 1.66 +/- 0.12; p < 0.05).     

红鳍原鱼白鳃部寄生单殖吸虫生态位分析

鱼类寄生单殖吸虫越来越多受到寄生虫学家和生态学家的关注 ,生态位分析也是生态学研究的一个重要方面 ,国外学者在这方面进行了较多的工作[5～ 8] 。近年来 ,我国单殖吸虫生态学的研究也吸引了不少科学家的兴趣[1～ 3 ,9] 。红鳍原鱼白 (Ｃｕｌｔｅｒｉｃｈｔｈｙｓｅｒｙｔｈｒｏｐｔｅｒｕｓ)是我国的一种自然经济鱼类 ,许多学者研究了不同水体红鳍原鱼白的寄生虫种类组成[4 ] 。作者现将红鳍原鱼白寄生单殖吸虫生态位分析作一报道 ,为研究寄生虫群落结构提供必要的背景参数。1　材料与方法1.1　材料来源洪湖系长江和汉江支流东荆河之间…     

Population and seasonal distribution of Procamallanus daccai in Eutropiichthys vacha in Kaptai Lake, Chittagong, Bangladesh

The population and seasonal distribution of Procamallanus daccai (Nematoda: Procamallanidae) in Eutropiichthys vacha (Siluridae) from the Kaptai Lake were studied for a period of 20 months during 1986-1987. A total of 208 fish were necropsied and 5652 worms were recovered, of which 2392 were from 87 infected male fish and 3260 were from 89 infected female fish. Prevalence values varied from 50% to 95%. The intensity and abundance of nematodes fluctuated over the study period and are statistically significant (F=14.87, P<0.0001), and showed a seasonal preference for the summer period (F=33.39, P<0.05). The mean intensity was the highest in the smallest size group at 66.3+/-12.7 which gradually decreased to 4+/-1.89 in the largest size group. No significant differences were found between parasites from male and female hosts, except in the summer. Male parasites were less abundant in the population. Female worms were grouped into immature, maturing and gravid and all these stages were present throughout the year. The highest occurrence of both immature and gravid P. daccai was found during the months of February to April each year. Peak recruitment seems to occur during this period due to the presence of high quantities of immature worms.     

Pathogenicity of the gill artery worm Philometra obturans (Nematoda) in northern pike (Esox lucius) in southwest Finland

The pathogenicity of female Philometra obturans (Prenant, 1886) was investigated. Fish samples were taken in May and June 2001 from Lake Littoistenj?rvi, in southwest Finland. The intensity of infection varied between 1 and 4 worms, these being found in the afferent artery of the 2 outermost gill arches. The prevalence was 33.9% (n = 59). Host condition and tissue reactions were investigated through dissection and analysis of histopathology. The parasites obstructed the gill arteries, and the arterial wall was stretched around intact worms. Hypertrophy of the arterial wall was observed, especially when only disintegrating remains of the parasites were present. Infected pike were in a poorer condition after catch, transportation, and holding in aquaria and thus less tolerant to stress than uninfected pike. This was correlated with the number of infected gill arteries (P < 0.05). The results show that infection with this nematode has a detrimental effect on the host when combined with environmental or physiological stress and may result in host mortality.     

Field preservation of monogenean parasites for molecular and morphological analyses

In the present work we examined the efficacy of three different chemical solutions (EtOH 70%, DMSO-NaCl solution, and Longmire buffer) in field preservation of fish gills to be subsequently screened for monogenean specimens destined to morphological and molecular analyses. Degree of difficulty in collecting monogeneans from gills, morphological state of parasites, integrity of their DNA and reliability of sequence reading were observed and qualitatively compared to those of gills and parasites stored in 5% formalin and 99% ethanol. Data were collected over a period of 2 months. Storage in Longmire buffer resulted in dissociation of gills and parasites, while both DMSO and 70% ethanol provided a fine physical and molecular preservation of gills and monogeneans, allowing rapid collection of parasites from lamellae, and easy extraction, amplification and sequencing of parasitic DNA.     

Involvement of host recognition by oncomiracidia and post-larval survivability in the host specificity of Heterobothrium okamotoi (Monogenea: Diclidophoridae)

Heterobothrium okamotoi, a monogenean gill parasite, shows high host specificity for tiger puffer, Takifugu rubripes. In the present study, in vivo and in vitro experimental infections were conducted using various fish species, including T. rubripes, to understand the mechanisms of specificity. In in vivo experiments, T. rubripes, grass puffer, Takifugu niphobles, olive flounder, Paralichthys olivaceus, and red sea bream, Pagrus major, were exposed to oncomiracidia of H. okamotoi labelled with a fluorescent dye, 5- (and -6) carboxyfluorescein diacetate succinimidyl ester, and the numbers of parasites on the gills and skin were recorded at intervals. Oncomiracidia were attached to gills and skin of all the experimental fish species immediately after exposure, and the infection intensity on T. rubripes was higher than that on T. niphobles and much higher than those on the other two species. After 2 days, the attached parasites remained on the gills of T. rubripes, but disappeared from the other hosts. During in vitro experiments, gill filaments excised from seven different fish species (four fish species used in the in vivo experiments and panther puffer, Takifugu pardalis, southern flounder, Paralichthys lethostigma and spotted halibut, Verasper variegates) were exposed to oncomiracidia and the attachment to each fish species and subsequent larval behaviour was observed. The percentage of post-larvae that attached to T. rubripes was slightly higher than those which attached to congeneric fish species and much higher than those of non-tetraodontid fish species. In vivo and in vitro experiments demonstrated that oncomiracidia of H. okamotoi have an affinity for their natural host, T. rubripes, and congeneric fish species. The disappearance of attached post-larvae from 'alien' hosts within 2 days during in vivo experiments suggested that host recognition by oncomiracidia and subsequent post-larval survivability are involved in the host specificity of H. okamotoi.     

Determinants of host-specificity in parasites of freshwater fishes.

Factors responsible for interspecific variability in host-specificity were investigated within 15 genera (including 176 species) of metazoan parasites found in Canadian freshwater fish. For each species in a genus, the parasite's number of known hosts was determined from published host-parasite records. The effects of the total number and mean size of potential hosts (i.e. all fish species belonging to the family or families that include a parasite's known hosts) on number of hosts of congeneric species were evaluated using multiple regressions. Since parasite species that have been recorded often tend to have greater numbers of known hosts than do seldom-recorded parasites, it was necessary to control for the confounding effect of study intensity. In all parasite genera, whether from highly specific taxa such as monogeneans or from less host-specific ones, there was a positive relationship between the number of potential hosts and the number of known hosts. However, no consistent relationships were observed between the mean size of potential hosts and number of known hosts. These results suggest that the availability of suitable host species may have been a key factor limiting the colonization of new hosts by fish parasites.     

Parasite fauna of introduced pumpkinseed fish Lepomis gibbosus: first British record of Onchocleidus dispar (Monogenea)

A survey of pumpkinseed Lepomis gibbosus from a stillwater fishery in southern England revealed infections by the ancyrocephalid monogenean Onchocleidus dispar at 100% prevalence. Host specificity of O. dispar to North American centrarchid fish suggests that it is a non-native parasite, introduced to Britain with L. gibbosus. Mean intensity of O. dispar was significantly higher in male (25.5 parasites) compared to female (15) and immature (7) hosts, but was not influenced by host standard length or sampling time. This sex-biased parasitism is likely to be related to both ecological and physiological factors, such as differential exposure to infective stages during nest building, or higher susceptibility to infection due to enhanced host stress levels. O. dispar significantly dominated the external surfaces and the outer gill arches. Analysis of the spatial distribution of the parasite between the 9 regions of gill surface determined that as density increased, higher numbers of parasites were found on the anterior and central gill regions compared to posterior (distal) regions. Apparent microhabitat selection is discussed in terms O. dispar life cycle, maximising oxygen availability, avoiding unstable areas and increasing intraspecific contact. The absence of O. dispar in the gills of roach, rudd and gudgeon sampled from the same fishery supports the assumption that this parasite is currently of little threat to native fish populations.     

Spatial distribution and coexistence of monogenean gill parasites inhabiting two damselfishes from Moorea island in French Polynesia

The ectoparasite fauna of two damselfishes, Stegastes nigricans and Dascyllus aruanus, from Moorea Island in French Polynesia was investigated. Gills of these damselfishes were infected with congeneric Monopisthocotylea Monogenea belonging to the genus Haliotrema. Stegastes nigricans were found to harbour a guild of three Haliotrema species whereas only one species inhabited D. aruanus. Microhabitat distribution, inter- and intraspecific competition and interspecific associations on the gill were studied. Observations on site preference revealed no spatial segregation between the three congeneric species inhabiting the gills of S. nigricans. Juvenile and adult monogeneans of that guild occurred on the same microhabitat. The dominant species Haliotrema sp. 1 did not expand on the microhabitat when the intensity of infection increased. Interspecific association tests revealed positive and negative associations. Haliotrema sp. 4 expanded its distribution on the gills of Dascyllus aruanus when the intensity of infection increased suggesting the likelihood of intraspecific competition. Juvenile and adult monogeneans of Haliotrema sp. 4 appeared to segregate as a result of intraspecific competition. This competition may exist to enhance resource availability when the gill habitat is limited. Overlaps between niche breadth and species microhabitat were revealed for monogenean species inhabiting S. nigricans. Interspecific competition did not appear to play an important role in the distribution of S. nigricans congeneric ectoparasites. Reinforcement of reproductive barriers may have led to the avoidance of hybridization.     

A possible evolutionary lag in the relationship between freshwater mussels and European bitterling

The coevolutionary dynamics between European bitterling Rhodeus amarus and freshwater unionid mussels, which the former parasitize by laying eggs on their gills, were tested. In a series of experiments fish preferences and mussel responses were compared in parasites and hosts of recent (Europe) and ancient (Asia) sympatry. Rhodeus amarus readily oviposited on the gills of all mussel species tested. Fish that laid their eggs on the gills of Asian Anodonta woodiana, however, suffered a dramatic reduction in reproductive success compared to fish that oviposited on the gills of European mussels: Unio pictorum , Unio tumidus , Anodonta anatina and Anodonta cygnea . This difference was the result of egg ejection behaviour by mussels rather than the unsuitability of the internal gill environment for European bitterling embryo development. The ejection response of mussels with a long sympatry with European bitterling was considerably more pronounced than that of mussels with a substantially shorter sympatry. The data support a coevolutionary arms race between bitterling and mussels and point to an evolutionary lag in the relationship between R. amarus and its European mussel hosts.     

The niche of Salsuginus thalkeni, a gill parasite of Fundulus zebrinus

Distribution of a monogenean parasitic helminth Salsuginus thalkeni on the gills of the fish Fundulus zebrinus is described by calculation of mean positions and niche breadths on the linear spatial resource gradients gill filament length, gill arch length and arch number. All distributions are given for parasites in the presence and absence of various combinations of potential competitors, namely the 6 other parasite species that occupy the same host species. Filament niche breadth was narrowest in the absence of potential competitors; breadth on arch was widest in the presence of potential competitors. Breadth on both resources was correlated positively with mean number of parasites per individual. Arch breadth exhibited cyclic seasonal changes, being lowest in early to mid-summer. Mean position exhibited no repeated pattern of variation on either resource. The results are considered consistent with predictions about the niche structures of species in unsaturated noninteractive specialist communities.     

Parasitic community of Fransciscodoras marmoratus (Reinhardt, 1874) (Pisces: Siluriformes, Doradidae) from the upper S?o Francisco River, Brazil.

One hundred and thirteen specimens of Franciscodoras marmoratus (Reinhardt, 1874) were collected in the upper S?o Francisco River (18 degrees 12' 32" S, 45 degrees 15' 41" W, state of Minas Gerais) between September, 1999 and January, 2004 to investigate their parasite fauna. From this total, 45 (39.8%) were afflicted by at least one parasite species. The parasitic richness consisted of six species represented by Hirudinea (n = 20), Monogenoidea (n = 25), Eucestoda (n = 55), Nematoda (n = 1, n = 2) and Acanthocephala (n = 41) found in the dry and wet periods making a total of 144 specimens. Proteocephalus renaudi Chambrier & Vaucher, 1994 was the only species with prevalence higher than 10% and a typical aggregate distribution pattern. The prevalence, intensity and abundance of P. renaudi were not influenced by the total length or sex of the hosts or by the collection periods. The relative condition factor indicated that the health of the P. renaudi hosts was not significantly affected in relation to fish not infected by parasites. The fish stocked in tanks before necropsy were opportunistically infested by Lernaea cyprinacea Yashuv, 1959. The various parasites found indicate that F. marmoratus is omnivorous and a potential definitive host. The parasite species, except for Acanthocephala, have expanded their known geographic distribution to the S?o Francisco River Basin. The parasite community was considered isolationist because of the low endoparasite diversity, infrapopulations with low intensity, lack of evidence of parasite interactions and sparse signs of parasite aggression against their hosts.