The genus Argulus (Crustacea: Branchiura) in Africa: identification keys

Two keys to the African species of Argulus have been constructed, the first of which is applicable to either sex and the second is based on the male accessory copulatory structures.     

The occurrence of Argulus (Crustacea: Branchiura) in Scotland

Records of the distribution of the genus Argulus in Scotland are given. The occurrence of A. coregoni Thorell in the River Clyde is confirmed.     

Laying, development and hatching of eggs of the fish ectoparasite Argulus japonicus (Crustacea: Branchiura)

Egg laying of the fish-louse Argulus japonicus was observed and examined experimentally. The effect of temperature on development time and hatching yielded an inverse exponential function. Hatching started after 61–10 days in a temperature span of 15–35 °C. Eggs are laid in strings on hard substrata and covered by a gelatinous material. Females lay between 1–9 strings, 5–226 eggs per string, arranged in 1–6 rows. Four embryonic developmental stages were recognized and the mean hatching efficiency was 50% in the optimal temperature range of 20–30 °C. Hatching efficiency was not related to either the number of eggs in a string or the total number of eggs laid by any particular female. Argulus japonicus displays continuous egg-laying activity with a possibility of an overwintering mechanism which suggests a seasonality of a sort.     

Seasonal and vertical patterns of egg-laying by the freshwater fish louse Argulus foliaceus (Crustacea: Branchiura)

Argulus foliaceus is a damaging fish ectoparasite for which new control measures are being developed based on egg-removal. In an attempt to develop further understanding of seasonal and vertical egg-laying patterns in this parasite, egg-laying activity was monitored over the period 14 April to 17 November 2003 in 2 rainbow trout Oncorhynchus mykiss fisheries in Northern Ireland, UK. At Site 1, egg-laying was continuous from 21 April to 17 November, when water temperature was above 8 to 10 degrees C. At Site 2, egg-laying was continuous from 4 June to 29 October. In the early months of the season, egg-laying was recorded mainly within the top 1 m of the water column; however, a significant shift to deep water egg-laying was recorded between 7 July and 17 November at Site 1 and between 20 August and 29 October at Site 2. Egg clutches were preferentially laid at depths of up to 8.5 m during this time (Site 2), a feature of egg-laying hitherto unappreciated. Temperature and dissolved oxygen did not differ significantly among depths, but there was an increase in water clarity over time. However, the precise environmental triggers for deep water egg-laying are still unclear. These new insights into the reproductive behaviour of this species will be useful in developing control methods based on egg-removal.     

Comments on the morphology of the pre-oral spine in Argulus (Crustacea: Branchiura)

A histological study of the pre-oral spine and associated structures in Argulus japonicus reveals that two exceptionally long and slender muscles casuse retraction of the spine into the spinal sheath. Originating from the body wall mid-dorsally the two muscles proceed through the nerve ring and the blood-filled lumen of the spinal sheath before being inserted on a bolster of tissue constituting the base of the spine. Movement of the spinal sheath is affected by two muscles originating from a common apodeme in the dorsal body wall, just posterior to the nauplius eye, and inserted on the lateral walls of the base of the spinal sheath. Two ducts, each leading from a glandular complex in the lateral parts of the carapace, proceed rostrally through the haemocoelic spaces. After passing the tritocerebrum laterally they accompany the retractor muscles of the spine on their course to the base of the spine. Passing through the bolster of tissue at the base of the spine the two ducts run all the way up to the tip of the spine where they open separately. The glandula praeboscoidalis is not associated with the pre-oral spine since its duct runs posteriorly in the wall of the proboscis. The labial spines are associated with giant glandular cells located beneath the optic tracts. © 1992 Wiley-Liss, Inc.     

Structure of the adult ovary and oogenesis in Argulus japonicus thiele (Crustacea: Branchiura)

The adult ovary of the branchiuran Argulus japonicus is a single, median, long sac-like organ located in the thorax above the alimentary canal. A long germarium, including oogonia, very early previtellogenic oocytes, and young somatic cells (interstitial cells), is embedded in the dorsal ovarian wall along the median longitudinal line of the ovary. The ovarian wall, consisting of a layer of the ovarian epithelial cells, is folded repeatedly and distinctively in the lateral and ventral portions of the ovary. Growing oocytes, previtellogenic and vitellogenic, occur on the outer surface of the ovarian wall, not in the ovarian lumen. The smaller oocytes are located nearer to the germarium and the larger ones on the more ventral surface of the ovary. These structural features of the branchiuran ovary are compared with those of other crustaceans and the pentastomids to consider their phylogenetic implications. J. Morphol. 231:29–39, 1997. © 1997 Wiley-Liss, Inc.     

Argulus papuensis n. sp., a new fish louse (Crustacea: Branchiura) from Papua New Guinea

A new species of Argulus (Crustacea: Branchiura) is described from a sleeper, Bunaka herwerdeni (Weber), in Papua New Guinea. It is characterised by the shape of the respiratory areas, the number and shape of supporting sclerites in the suckers of the first maxilla, as well as the accessory copulatory structures on the third and fourth legs of the male.     

Growth and development of Argulus coregoni (Crustacea: Branchiura) on salmonid and cyprinid hosts

The obligate fish ectoparasite Argulus coregoni is strictly specific to salmonids and is very rarely found on other fish species. The ability of the parasite to grow and complete its life cycle on a cyprinid host, Rutilus rutilus, was compared with that on a typical salmonid host, Oncorhynchus mykiss. Rearing experiments were run for 42 d with newly hatched metanauplii in flow-through tanks. Body length and sex of the parasites were recorded every 5 d. Growth rates on O. mykiss exceeded those on R. rutilus from the age of 2 wk, at which time the parasites reached a length of about 3.5 mm. Males grew faster than females at the beginning of the experiment up to a length of 2.5 to 3.0 mm; thereafter, a faster growth rate was observed in females. In another experiment, association of parasites with the hosts was monitored and residence time defined as the period between attachment and first detachment from the host. Longer residence time was observed on O. mykiss than on R. rutilus; female parasites stayed on both fish species longer than did males. Faster growth of parasites could be associated with longer uninterrupted periods of attachment to hosts, since frequent detachment means higher energy losses and less time available for feeding. Despite its slower growth on R. rutilus, A. coregoni matured and laid egg clutches, but took 5 d longer than on O. mykiss. The potential of A. coregoni to complete its life cycle on cyprinids could have important ecological consequences, creating an infection reservoir when the main salmonid hosts are rare or temporarily missing.     

Ectoparasitic Argulus coregoni (Crustacea: Branchiura) hedge their bets – studies on egg hatching dynamics

Unpredictability in the temporal availability of susceptible hosts is likely to act as a selection pressure affecting the life history strategies of parasites. In highly variable environments the future of the lineage can be secured by spreading the risk, for example, by producing descendants that differ in their timing of emergence. Counter to this, in predictable environments a single "best-adapted" phenotype is expected. We asked whether ectoparasitic Argulus coregoni egg hatching pattern can be explained as a genetically canalized individual trait; an instance of phenotypic plasticity or bet-hedging. We collected egg clutches laid by individual A. coregoni females in early and late reproductive period of the lice population and randomized the clutches within 3 treatments. Intra- and inter-clutch variability in the hatching dynamics of A. coregoni eggs was monitored and the reproductive potential assessed. On average A. coregoni females laid 317 (SD±176.6) eggs. We found that the plasticity in the hatching dynamics among A. coregoni eggs was remarkable. Noticeable peaks in hatching were followed each of the repeated artificial "winter treatments" in 1°C. Repeated 2 weeks cold treatments induced relatively bigger hatching peaks than 2 days cold treatments compared to controls at room temperature. However, in all treatments, egg clutches hatched through an extended period of 7 months on average and the total hatching percentages were similar. We found that intra-clutch variability in hatching among eggs laid by single A. coregoni females was greater than inter-clutch variability. Our data support the predictions of the adaptive bet-hedging strategy in relation to egg-hatching dynamics. Response to cooling and bet-hedging may be adaptive for such species like A. coregoni since by synchronizing the life-cycle with the seasonal environment will assist transmission and parasite fitness.     

Survivorship and fecundity of Argulus bengalensis (Crustacea; Branchiura) under laboratory conditions

The Life table features and the fecundity schedule of the fish louse Argulus bengalensis (Branchiura: Argulidae) were estimated under laboratory conditions using Cirrhinus mrigala as the model host. The cohort-based Life table revealed that survival of A. bengalensis declines as a function of age. Life expectancy was observed to be 34?days for a newly emerged adult. The mean number of eggs produced per egg strip was 86 (±8.89 S.E.). The sex ratio 2:1 was biased towards males. The net reproductive rate (R₀) varied between 50 and 487 while the cohort generation time (T_c) ranged between 24 and 26?days. The intrinsic rate of increase (r) was 0.14–0.23, while the finite rate of increase (λ) remained between 1.16 and 1.28. The Life table features and the fecundity schedule of A. bengalensis are comparable with congenerics of temperate regions. Early reproduction (between 21 and 23?days) and an extended postreproductive phase (47?days) of females are characteristic of A. bengalensis and a reflection for survival under uncertain conditions and a parasitic mode of life.     

Experimental evidence for a hierarchy of mate- and host-induced cues in a fish ectoparasite, Argulus coregoni (Crustacea: Branchiura)

Argulus coregoni is an ectoparasite primarily infesting freshwater salmonids. Sexually reproducing parasites such as A. coregoni are confronted with a dilemma between finding a mate and the costs involved in doing so; if mating partners are unavailable on a host, by leaving to search for a mate on a new host, the parasite is exposed to risks such as predation and energy loss. The utilization of chemical cues could enhance the probability of finding a host and/or a suitable mating partner and thus decrease the level of costs associated with detachment from the host. In this study we constructed a Y-maze arena to determine if adult A. coregoni respond to mate- and host-related chemical cues. We also tested the directional response towards light, since it has been suggested that photic cues are the most important cues for juvenile A. coregoni locating a host. Our results showed that both sexes were attracted to light and fish odour. Free-swimming A. coregoni males responded to chemical cues produced by adult females but not vice versa. The hierarchy of these stimuli was analyzed by pitting the cues against one another in the Y-maze, showing that light was the most salient stimulus for both male and female parasites. Moreover, male parasites were more strongly attracted towards light and fish odour than female odour. In another experiment in a semi-natural environment, we examined whether the ability of A. coregoni males to detect female odour influences their host choice. Free-swimming males did not preferentially infest fish infected with female parasites over parasite-free fish. We suggest that a hierarchy of stimulus responses exists, whereby free-swimming parasites first respond to host-related signals and most dominantly to visual cues. However, cues connected to mate finding may become a priority for late adult stages and/or once the parasite has attached to the host.     

A new species of the genus Argulus (Crustacea: Branchiura) parasitic on Atractosteus tropicus (Pisces: Lepisosteidae) from Tabasco,Mexico

A new species of Argulus is described from the garfish Atractosteus tropicus in the Tabasco wetlands, Mexico. Comparisons are made with what appear to be its closest relatives. This species is characterised by the presence of two notches in the inner margin of the large respiratory area, seven to nine supporting sclerites in the suckers, and the characteristic shape of the sexual modifications of the second, third and fourth legs in the male.     

Spatial distribution and hatching of overwintered eggs of a fish ectoparasite, Argulus coregoni (Crustacea: Branchiura).

The habitat distribution of overwintered eggs, which were found to be the only source of spring recruitment of Argulus coregoni Thorell, was studied at a commercial fish farm in Central Finland. The frequency of occurrence of egg clutches in the deep parts of the canals and ponds was 50 to 80% and the percentage cover of the surface of stones with egg clutches was 1.7 to 6.4%, while in the shallow parts these values were 8 to 27% and 0.1 to 0.3%, respectively. A greater proportion of empty egg-shells was observed in shallow water in the mid-May, suggesting an earlier hatching there stimulated by the increased temperature and higher illumination. Under laboratory conditions, only elevated UV illumination, but not diurnally fluctuating temperature, significantly accelerated hatching. Normally overwintered eggs produced a pronounced peak of hatched larvae at the end of May and hatching continued at a much slower rate throughout the summer. Eggs that overwintered twice, first normally and then for a second time buried under sediments, were exposed to the same laboratory conditions simultaneously with normally overwintered eggs, but their hatching was delayed until August. The hatching rate was low, but markedly increased in December.     

The male reproductive system and mechanism of sperm transfer in Argulus japonicus (Crustacea: Branchiura)

The histomorphology of the male reproductive system and surface morphology of the “peg-and-socket” in Argulus japonicus are described from serial sagittal and transverse sections and scanning electron micrographs. The prostate complex consists of a glandular part, a reservoir for storing the secretion, and an efferent duct opening into the ejaculatory duct. The openings of both the vas deferens and the prostate duct into the ejaculatory duct are guarded by sphincters. The ejaculatory ducts, which are lined by tall columnar epithelial cells, do not open into the cuticle-lined genital atrium but are blind-ending tubes. This observation and results obtained from observing live specimens, as well as the fact that no spermatophores are formed, suggest that semen could leave the ejaculatory duct only after puncturing of its walls. It is suggested that sperm transfer is accomplished in the following manner: during copulation contraction of the muscular walls of the vas deferens and prostate duct causes semen to be pumped into the ejaculatory duct, which is then closed off by sphincters and a high internal pressure is developed. When a spermathecal spine penetrates the walls of the ejaculatory duct, semen flows from the ejaculatory duct into the spermathecal vesicle due to the higher pressure in the ejaculatory duct. This mechanism is analogous to the sucking up of fluid with a hypodermic syringe. © 1993 Wiley-Liss, Inc.     

The ultrastructure of the digestive cells of Argulus japonicus, Thiele 1900 (Crustacea: Branchiura)

The ultrastructure of the cells of the digestive system of Argulus japonicus is described with the use of transmission electron microscopy. Specimens of Argulus japonicus were collected from the Vaal Dam in South Africa and fixed in Todd's fixative. The samples were post fixed in osmium tetroxide and embedded in resin. The anterior midgut is composed mostly of R cells while the enteral diverticula are composed mainly of R cells in the proximal diverticules and of F cells in the distal diverticula. The posterior midgut is composed of very large papilliform B cells and of R cells. The R cells in the anterior midgut probably absorb nutrients including lipids. The F cells are filled mostly with rough endoplasmic reticulum, suggesting enzyme synthesis, while the B cells portrayed endocytotic vesicles, indicating intracellular digestion of predigested food. The R cells of the posterior midgut are less active than cells present in the anterior midgut. E cells and peritrophic membrane were not observed.     

Description of the previously unknown male of Argulus kosus Avenant-Oldewage, 1994 (Crustacea: Branchiura)

Argulus kosus Avenant-Oldewage, 1994 was described from two females collected in Kosi Bay, South Africa. Individuals of both sexes have now been found associated with freshwater, estuarine and marine fishes in Lake St Lucia on the coast of northern KwaZulu/Natal. The male is described, and A. smalei Avenant-Oldewage & Oldewage, 1995 is synonymised with A. kosus.