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Many entomopathogens regulate the development of their insect hosts. However, the influence of mermithid nematodes on the development of their host remains unclear. In the current study, we provide insights into how Ovomermis sinensis parasitism affects the development of Helicoverpa armigera. We observed that O. sinensis arrests host development, as evidenced by the reduced body size and failure of Helicoverpa armigera to pupate. Moreover, midgut replacement of the host was significantly blocked by parasitism. Furthermore, juvenile hormone (JHIII) titers of the host were dramatically elevated by parasitism, but JH esterase (JHE) activities were strongly inhibited. By contrast, steroid hormone (20-hydroxyecdysone, 20E) titers of the host were significantly depressed by parasitism on days 4-6. The expression profiles of hormone-related genes in the host also showed similar patterns with the hormone titer. For this reason, rescue experiments were performed by injecting 20E and JHIII into developmentally arrested hosts. Notably, the midgut replacement of the host was rescued by the injection of 20E, whereas JHIII injection resulted in negative effects. Altogether, O. sinensis arrests H. armigera midgut replacement by reducing 20E and maintaining JH, thereby causing developmental arrests. Our study is the first report of the possible mechanism of mermithid nematodes in regulating insect development., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier Inc. All rights reserved.)

Mermithids (phylum Nematoda) and hairworms (phylum Nematomorpha) somehow drive their arthropod hosts into water, which is essential for the worms' survival after egression. The mechanisms behind this behavioural change have been investigated in hairworms, but not in mermithids. Establishing a similar mechanistic basis for host behavioural change between these two distantly related parasitic groups would provide strong convergent evidence for adaptive manipulation and insight into how these parasites modify and/or create behaviour. Here, we search for this convergence, and also contrast changes in physiology between hosts infected with immature and mature mermithids to provide the first ontogenetic evidence for adaptive manipulation by disentangling host response and pathology from the parasite's apparent manipulative effects. We used SWATH-mass spectrometry on brains of Forficula auricularia (earwig) and Bellorchestia quoyana (sandhopper), infected with the mermithids Mermis nigrescens and Thaumamermis zealandica , respectively, at both immature and mature stages of infection, to quantify proteomic changes resulting from mermithid infection. Across both hosts (and hairworm-infected hosts, from earlier studies), the general function of dysregulated proteins was conserved. Proteins involved in energy generation/mobilization were dysregulated, corroborating reports of erratic/hyperactive behaviour in infected hosts. Dysregulated proteins involved in axon/dendrite and synapse modulation were also common to all hosts, suggesting neuronal manipulation is involved in inducing positive hydrotaxis. Furthermore, downregulation of CamKII and associated proteins suggest manipulation of memory also contributes to the behavioural shift.

We examined manipulation of mosquito behavior by the parasitic mermithid nematode, Strelkovimermis spiculatus. This nematode species typically infects early instar host larvae and emerges after parasitic development to kill last-instar larvae. Parasitized adults, however, have occasionally been reported from field collections. We obtained low rates (1.7-11.5%) of parasitized adults in laboratory exposures only when Culex pipiens pipiens fourth-instar larvae nearing pupation were exposed to infective nematodes. This did not allow an adequate interval for parasitic development in immature host stages. Parasitized adult females in a multiple-choice assay were three times more likely to seek water than a blood source (63.1 vs. 20.5%), whereas uninfected females were twice as likely to seek blood than water (64%3.9 vs. 32.6%). This altered host behavior benefits the parasite by providing the only mechanism for dispersal and colonization of new host habitats while concurrently avoiding risks from the defensive behaviors associated with blood-feeding. Behavioral alternation in Cx. p. pipiens larval hosts was also examined using larvae infected as second instars to allow for a normal duration of parasitic development. As larvae neared pupation and parasite emergence, parasitized larvae became more spatially aggregated than unparasitized larvae. This altered host behavior benefits the parasite by providing a corresponding increase in post-parasite aggregation, which facilitates formation of large mating clusters and concomitantly reproductive success. Parasites derive fitness gains by overriding host autonomy, whereas hosts have zero fitness once parasitism is established, suggesting a coevolutionary response is inoperative and that the behavioral modifications may be adaptive., (Copyright © 2019 Elsevier Inc. All rights reserved.)

Certain species of parasites have the apparent ability to alter the behaviour of their host in order to facilitate the completion of their own life cycle. While documented in hairworms (phylum Nematomorpha), the ability for mermithid parasites (from the sister phylum Nematoda) to force hosts to enter water remains more enigmatic. Here, we present the first experimental evidence in a laboratory setting that an insect which normally never enters open water (the European earwig Forficula auricularia) will readily enter the water when infected with a mermithid nematode (Mermis nigrescens). Only adult mermithids appear capable of inducing this polarising shift in behaviour, with mermithid length being a very strong predictor of whether their host enters water. However, mermithid length was only weakly associated with how long it took an earwig to enter water following the beginning of a trial. Considering the evidence presented here and its alignment with a proteomic investigation on the same host-parasite system, this study provides strong evidence for adaptive behavioural manipulation and a foundational system for further behavioural and mechanistic exploration.

The hemocoel of 26 of 30 (86%) eastern grass shrimps, Palaemon paludosus (Gibbes, 1850) (Decapoda: Palaemonidae), captured during June 2017 from several freshwater lakes near Leesburg and Lake Kissimmee, Florida, was infected by juveniles of a nematode species (Mermithidae sp.). Some infected eastern grass shrimps were preserved whole for histopathology, whereas others were dissected to excise parasitic juvenile nematodes, and still others were kept alive in glass aquaria such that post-parasitic (emerged) juvenile nematodes could be opportunistically observed alive and ultimately preserved. Parasitic and post-parasitic juvenile nematodes had cuticle cross-fibers, cephalic papillae, cup-shaped amphids, a horn-shaped vagina, a trophosome, and a caudal appendage, which collectively diagnosed them as Mermithidae sp. They differed from those of nematomorphs (Nematomorpha) by lacking 4 giant cells anteriorly, tegumental bristles, scale or plate-like areoles, a bifurcate or trifurcate posterior end, and an anus. A phylogenetic analysis of the small subunit rDNA (18S) that included all of the available mermithid sequences corresponding to morphologically diagnosed specimens recovered Mermithidae sp. within the clade of mermithids and sister to Ovomermis sinensis Chen, Jian, and Ren, 1991 . This is the first record of a mermithid infection in a decapod and first report of a mermithid infection in an aquatic crustacean from North America (another mermithid infects a terrestrial isopod there). The high prevalence of infection and the multiple geographic localities harboring infected eastern grass shrimps indicated that these infections were not spurious. Because no other decapod is confirmed as a mermithid host, we suspect that these specimens likely represent a new species with a life cycle worth studying, since none for a mermithid involving a decapod nor a crustacean has been elucidated to date. We also provide a table of all mermithid and nematomorph infections in crustaceans.

Background: Mermithid nematodes, such as Ovomermis sinensis, display a broad host range including some lepidopteran pests. Infective juveniles penetrate their host through the cuticle, complete their growth within the hemocoel and eventually kill the host upon their emergence. Hence, mermithid nematodes are considered potential biological control agents of insect pests. Our previous data indicate that the infection rate of O. sinensis on cotton bollworm (Helicoverpa armigera) is low, which may be largely due to the strong immune system of the host. However, current knowledge on the interactions of mermithid nematodes with their hosts and the mechanisms employed by hosts to defend themselves against mermithid nematodes is limited., Results: Here, we investigated the response of H. armigera to O. sinensis infection. Parasitism by O. sinensis caused a sharp decline in the survival rate of H. armigera. The hemocytic phagocytosis ability, antibacterial activity, and phenoloxidase (PO) activity in plasma of H. armigera increased at 1 d post parasitism (dpp) but decreased at 3 dpp. Further, we investigated gene expression in the fat body of parasitized and non-parasitized H. armigera larvae at 1, 3, and 5 dpp using a digital gene expression system. In total, 41, 60 and 68 immune-related differentially expressed genes were identified at 1, 3, and 5 dpp, respectively. These genes encoded pattern recognition receptors (PRRs), antimicrobial peptides (AMPs), serine proteases (SPs), SP inhibitors, mucins and other immune-related proteins. The expression of most PRRs, AMPs, SPs, and mucins was upregulated in the fat body of larvae at 1 dpp, downregulated at 3 dpp, and then again upregulated at 5 dpp by O. sinensis. The increased expression of SP inhibitors may contribute to the inhibited PO activity at 5 dpp., Conclusions: This study demonstrates that parasitism by O. sinensis modulates the immune reaction of the host H. armigera by altering the expression of immune-related genes. Our data provide a basis for future investigation of the molecular mechanisms employed by the mermithid nematode O. sinensis to modulate the immunity of the host H. armigera. These data will also likely facilitate the improvement of success in parasitism of H. armigera by O. sinensis.

Mermithidae is a family of endoparasitic nematodes known to cause intersexuality in arthropods. Intersexes of the genus Culicoides parasitized by mermithids have been the object of several studies aiming to describe their particular morphology. Culicoides intersexes are specimens with male genitalia and feminized sexually dimorphic structures, i.e. antennae, mouthparts and wings. To date, these specimens have only been described qualitatively and a quantitative approach supported by statistical analysis is lacking. Here we conduct morphometric analyses of sexually dimorphic structures in a sample of Culicoides circumscriptus that includes 34 intersexes with the aim of describing precisely the intersexual morphology. The morphology of antennae and the mouthparts was studied by multivariate statistical analysis of linear measures, and wing form by implementing geometric morphometrics techniques. While intersex wings proved to have a similar size to male wings, their shape was intermediate between males and females. However, when allometric shape variation was removed, the wing shape of intersexes was almost identical to that of females. The intersex antennae were morphometrically of the female type, especially when size variation was considered. In contrast, the measured mouthparts (the labrum and the third palpal segment) were halfway between males and females, even when body size was considered. Overall, the antennae and the wings showed a higher degree of feminization than the mouthparts. These findings indicate that the degree of feminization depends both on the morphological structure and on body size. Moreover, we propose that the feminization of the wings and antennae has an adaptive meaning for the parasite, which would favor female-like traits in order to access more easily its breeding sites, where the parasite has plenty of new hosts to infect. Female-like antennae would be beneficial to detect these sites, while having female-like wings would favor the host's capacity of dispersal to these sites., (Copyright © 2016 Elsevier Inc. All rights reserved.)

The impact of parasites on host population dynamics depends on local abundance of the parasites, which may vary considerably across spatial scales. In sand hopper populations, mermithid parasites have major impacts on host dynamics, which may vary among spatially separated populations due to the sand hopper's wide, patchy distribution. The present study compared the abundance and biomass of a mermithid parasite (Thaumamermis zealandica Poinar et al., 2002) in sand hoppers (Bellorchestia quoyana (Milne-Edwards)) both within and among disconnected beaches. In addition, several variables were measured and tested as potentially important predictors of the parasite abundance and biomass. It was found that geographic isolation may only be responsible for minor differences in parasite populations compared with other factors. Host size was identified as the most important predictor of mermithid parasite abundance, but epibiont abundance, kelp patch mass and host density were poor predictors of abundance. These factors were also poor predictors of parasite biomass in hosts. This study further supports the notion that studies aiming to elucidate population dynamics or patterns should sample thoroughly across both spatial and temporal scales.

Mating aggregations in the mosquito parasitic nematode, Strelkovimermis spiculatus, were investigated in the laboratory. Female postparasites, through their attraction of males and, remarkably, other females, drive the formation of mating clusters. Clusters may grow in size by merging with other individual or clusters. Female molting to the adult stage and reproductive success are enhanced in larger clusters. Male mating behavior is initiated when the female begins to molt to the adult stage by shedding dual juvenile cuticles posteriorly. Males coil their tail around the adult cuticle, migrating progressively along the female in intimate synchrony with the molting cuticle until the vulva is exposed and mating can occur. The first arriving male is assured of access to a virgin female, as his intermediate location between the vulva and subsequently arriving males blocks these competitors. Males deposit an adhesive gelatinous copulatory plug into and over the vulva before departing the female. Fecundity was greater in larger mating clusters, but this was a function of a greater rate of molting which is a prerequisite for mating. Males compete for virgin females by emerging and molting to the adult stage earlier than females. Mating aggregations have previously only been examined in snakes, but these studies have tended to be observational as snakes offer a challenging system for study. The relatively easy to culture and manipulate mermithid system may offer a model for experimental studies of male-male competition, protandry, copulatory plugs and female choice in mating clusters., (Copyright © 2014 Elsevier Inc. All rights reserved.)

The fauna of blackflies of North Armenia is represented by 12 species from the genus Simulium. The species composition in the Tavush Region is most diverse (10 species) versus that in the Shirak (n = 5) and Lori (n = 3) Provinces, which is due to a wide variety of Simuliidae breeding sites. Among the bioregulators of blackflies, there are microsporidia of 4 species, cabbageworms (Mermithidae), and caddisflies of the genus Hydropsyche. There is a preponderance of microsporidia among the blackfly bioregulators. The highest percentage (60%) of the larvae infected with microsporidia is noted in the second half of summer, which is related to their environmental conditions.

Strelkovimermis spiculatus (Poinar and Camino 1986) is a nematode parasite of mosquitoes isolated from the Neotropical region. We investigated the host range of this parasite in mosquito populations for a better understanding of the dynamics of temporary breeding sites. Five grassy-pool habitats filled by rainwater were sampled from the summer 2007 through the fall 2008. Eight mosquito species were collected: Anopheles albitarsis, Culex chidesteri, Culex dolosus, Culex maxi, Aedes albifasciatus, Psorophora ciliata, Psorophora cyanescens, and Psorophora albigenu. Six of these species were parasitized: Cx. chidesteri, Cx. dolosus, Cx. maxi, Ae. albifasciatus, Ps. ciliata, and Ps. cyanescens. The occurrence of this mermithid in natural mosquito populations was increased from the end of winter to the end of the spring. Prevalence ranged from 11% to 100%. High levels of infections were registered only in Ae. albifasciatus larvae, the most abundant mosquito species (95%), followed by Cx. dolosus (2.7%). Strelkovimermis spiculatus completed its development in all infected mosquito larvae. The presence of S. spiculatus in six natural mosquito populations increases the number of susceptible species to 24., (© 2013 The Society for Vector Ecology.)

The responses of eggs in diapause and the infective stage of the nematode, Strelkovimermis spiculatus, to larvae of its host, Culex pipiens pipiens, were investigated in the laboratory. The results indicated that the presence of the host induced the egg hatching. The hatching rate increased when larger numbers of host larvae were present. Second instar mosquito larvae induced significantly higher hatching rates than any other stages. These findings explain how S. spiculatus synchronizes its life cycle with its host life cycle and population dynamics to increase its fitness when the natural habitat is constantly covered by water. Direct exposure of the nematode eggs to host larvae resulted in consumption of as many as 20 eggs per host. The eggs consumed caused 0-70% host mortality depending on the number consumed, which indicated an infection path other than cuticle penetration although it may represent a rare situation in nature. The result of host cue assays showed that the combination of chemical cues and physical vibration induced the highest egg hatching, which may increase the chance of host availability after hatching. However, once hatched, the nematodes ignored vibrations and used only chemical cues for host location. These findings suggest that eggs hatch synchronously with the most susceptible mosquito stage and with peak mosquito larval density., (Copyright © 2012 Australian Society for Parasitology Inc. All rights reserved.)

We assessed the number of Strelkovimermis spiculatus preparasites obtained from a known initial number of nematode eggs and the effect of abiotic conditions (temperature, photoperiod, flooding-drying) on the number of emerged preparasites. Two egg groups were maintained: one continuously flooded, another with flooding-drying cycles (every 15, 30, 60 days). Each egg group was studied at 25°C and 14:10 (L:D) and 16°C and 12:12 (L:D). The flooded eggs contained a higher overall percentage of S. spiculatus preparasites compared to the wet-dry-cycle eggs. The conditions of continuous flooding at 16°C and 12:12 (L:D) produced the maximum percent of emerged J2s (30±15%). Preparasites were recorded by 7 (25°C) and 14 (16°C) days, suggesting this period as the minimum time for embryonic development. The preparasite-emergence time observed from the same flooded-egg batch (98 and 112 days at 25°C and 16°C, respectively) suggested a nonsynchronous hatching, possibly through nonuniform egg embryonation. The time of exposure to drought in the assays did not significantly affect the total average percentage of J2s obtained at 25°C and 14:10 (L:D), whereas at 16°C the number of emerged J2s diminished with a prolongation of the drying period. The oviposition period was also recorded only at 16°C and 12:12 (L:D): S. spiculatus eggs were detected at 12.6 days after postparasite emergence, and oviposition was complete at 51days under those conditions. We propose a flooding schedule to optimize the mass-rearing of S. spiculatus., (Copyright © 2012 Elsevier Inc. All rights reserved.)

Little is known about the mechanism by which mermithid nematodes avoid encapsulation responses of insect hosts. In this study, we investigated the influence of the mermithid nematode Ovomermis sinensis on host Helicoverpa armigera hemocyte number, encapsulation activity, spreading behavior and cytoskeleton. Parasitism by O. sinensis caused a significant increase in the total hemocyte counts (THC) and plasmatocyte numbers of H. armigera. However, in vivo encapsulation assays revealed that hemocyte encapsulation abilities of H. armigera were suppressed by O. sinensis. Moreover, parasitism by O. sinensis changed the spreading behavior and cytoskeletons of the host hemocytes. The results suggested that O. sinensis could actively suppress the hemocyte immune response of its host, possibly by destroying the host hemocyte cytoskeleton. This is the first report of a possible mechanism by which mermithid nematodes suppress encapsulation responses of insect hosts.

Parasitic nematodes of the family Mermithidae were found within four specimens of Auchenorrhyncha from two families (Cicadellidae and Delphacidae). This appears to be the the first extant example of mermithid parasitism of Auchenorrhyncha in Europe. The insect hosts were collected from agricultural grassland field margins at three locations in Ireland during a farmland biodiversity study in 2007.

Specimens of Isomermis wisconsinensis, Gastromermis viridis, Mesomermis flumenalis, and Mesomermis camdenensis have been found in 14 out of 28 studied sites. Their hosts were the following blackfly species: Cnephia dacotensis, Cnephia mutata, Prosimulium sp., Simulium decorum, Simulium tuberosum, Simulium venustum/verecundum complex, and Simulium vittatum. Superparasitism was observed in several simuliid larvae and, occasionally, with 2 different mermithid species. The prevalence of different mermithid species varies according to seasons: Mesomermis genus is more abundant during winter, whereas Isomermis and Gastromermis are found in higher number during summer. The study of environmental parameters related to mermithid presence or absence shows that only stream depth makes a difference.

A single eye is present in females of the nematode Mermis nigrescens. A pigment cup occupies the entire cross section near the anterior tip of the worm, and the curved cuticle at the tip becomes a cornea. The shading pigment is hemoglobin instead of melanin. The eye has been shown to provide a positive phototaxis utilizing a scanning mechanism; however, the eye's structure has not been sufficiently described. Here, we provide a reconstruction of the eye on the basis of light and electron microscopy of serial sections. Hemoglobin crystals are densely packed in the cytoplasm of expanded hypodermal cells, forming the cylindrical shadowing structure. The two putative photoreceptors are found laterally within the transparent conical center of this structure where they would be exposed to light from different anterior fields of view. Each consists of a multilamellar sensory process formed by one of the dendrites in each of the two amphidial sensory nerve bundles that pass through the center. Multilamellar processes are also found in the same location in immature adult females and fourth stage juvenile females, which lack the shadowing pigment and exhibit a weak negative phototaxis. The unique structure of the pigment cup eye is discussed in terms of optical function, phototaxis mechanism, eye nomenclature, and evolution.

A wide range of parasites are known to cause behavioral changes in their hosts and parasitized insects are especially amenable to the study of such changes. The majority of studies addressing parasite-induced behavioral alterations have focused on parasites with complex life cycles and the adaptive nature of such changes. Behavioral changes caused by parasitoids, single-host parasites that kill their host upon emergence, have been studied less and the adaptive nature of these changes is likely to be different than those in complex life cycles. I investigated behavioral alterations in Aedes aegypti mosquito larvae infected with parasitoid nematodes (family Mermithidae). I conducted several experiments in which I tested the following hypotheses: 1) Mermithid nematodes induce behavioral changes in mosquito larvae and the changes are density dependent. 2) Different species of mermithid nematodes induce similar changes in mosquito larvae behavior. 3) Behavioral alterations vary with mermithid developmental stage. 4) Mosquito larvae infected with mermithid nematodes behave similarly to uninfected food-deprived mosquito larvae. I found that 4th instar Ae. aegypti infected with Romanomermis culicivorax or Strelkovimermis spiculatus exhibited resting behaviors significantly more often than uninfected controls but that intensity of infection did not affect activity levels. In earlier instars, infected mosquito larvae were more active than uninfected control larvae in some behaviors associated with feeding. There was no significant difference between infected and uninfected food-deprived mosquitoes in nine of the ten behaviors observed. The decrease in activity of late instar Ae. aegypti larvae infected with mermithids may be a parasitoid adaptation that reduces the risk of predation and thus increases host and parasitoid survival. The increase in feeding activity in earlier instars as well as the similarity between uninfected food-deprived and infected Ae. aegypti behavior may indicate that these behaviors are adaptive for the parasitoid, increasing nutritional acquisition for successful parasitoid development.

The present report describes a mermithid nematode (Nematoda: Mermithidae) and a gordiid hairworm (Nematomorpha: Chordodidae) from Early Cretaceous Burmese amber dated at 100-110 million years. The mermithid, Cretacimermis protus sp. n., is emerging from a biting midge (Diptera: Ceratopogonidae) while the hairworm, Cretachordodes burmitis, gen. n., sp. n. had already emerged from its host. These rare specimens represent the first fossil mermithid parasite of a ceratopogonid midge and second oldest described nematode and the earliest known and only Mesozoic fossil of the phylum Nematomorpha. A list of previously described fossil mermithids is included.

Population interactions among mosquitoes in the Culex vishnui subgroup, which are vectors of Japanese Encephalitis, and their natural enemies were studied in Pondicherry, India. We tested the hypothesis that the breakdown of interactions between the larvae and their natural enemies due to drought followed by rain was responsible for the sudden increase in the vector population above the threshold for disease transmission during the heavy rainy period. We randomly sampled mosquito larvae and their predators in different breeding habitats and subjected the mean densities of prey, predator, and mosquito larvae infected with parasites/pathogens to covariate analysis to understand the interaction between prey and their natural enemies in relation to environmental factors. In rice fields, neither prey nor predator showed any positive correlation with temperature, RH, or the number of rainy days. However, the pathogen/parasite of mosquito immatures showed a positive correlation with RH. Among the mosquito predators, notonectids exhibited a significant positive correlation with Cx. vishnui larvae. The parasitic Romanomermis iyengari and pathogenic Coelomomyces anopheliscus also showed positive correlations with immatures. No parasites and pathogens of mosquito larvae were recorded in shallow water pools (SWP) or cement tanks (CT) during the study period. Important predators recorded in SWP were notonectids, damselfly nymphs, Diplonychus indicus, and hydrophilids. Dragonfly nymphs, gerrids, and tadpole shrimps were recorded in CT. In CT, prey and their predators were positively correlated with RH and rainy days. In SWP, there was a highly significant correlation between prey, predators and environmental factors. We conclude that rice fields are a stable ecosystem where regular interaction occurs between larvae and their natural enemies and a sudden increase in mosquito populations is uncommon. In transient habitats, no such stability is present and they become more important as breeding habitats in terms of seasonality and number. Shallow water pools should be seriously considered for the control of these vectors.

The following new recent species of mermithid (Nematoda) parasites of Neotropical ants are described: Meximermis ectatommi n.gen., n. sp. from workers of Ectatomma ruidum Roger in Southern Mexico, Agamomermis cephaloti n. sp. from a worker of Cephalotes minutus (Fab.) collected in Brazil, Agamomermis ecitoni n. sp. from a worker of Eciton burchellii Westwood collected in Venezuela and Agamomermis costaricensis n. sp. from a worker of Odontomachus hastatus (Fab.) collected in Costa Rica. In addition, a fossil mermithid parasite of a worker of Linepithema sp. from Dominican amber is described as Heydenius myrmecophila n. sp. These parasites represent the first descriptions of Neotropical ant mermithids and together with previous records listed here, show that ant infection by mermithids in the Neotropics is widespread and has occurred for at least 20-30 million years.

Studies of the interactions between two or more species of symbiotic microorganisms in a single host animal have lagged behind those involving interactions between hymenopteran or dipteran parasitoids. We screened the larvae of four species of black flies (Cnephia ornithophilia, Simulium furculatum, Simulium innoxium, and Simulium vandalicum) to determine if the colonization and prevalence of trichomycete fungi were influenced by ichthyosporeans, mermithid nematodes, microsporidia, or other species of trichomycetes. Differences in the layering of the peritrophic matrix of microsporidian-infected larvae, compared with that of uninfected larvae, were associated with greater levels of colonization by the midgut-inhabiting trichomycete Harpella melusinae. Colonization levels of different species of trichomycetes were significantly correlated with one another in only 2 of 11 cases. Aside from the consistent attachment of the trichomycete Simuliomyces microsporus to the ichthyosporean Paramoebidium chattoni, we found little evidence that prevalence and colonization levels of trichomycetes were influenced by other symbiotes, including other trichomycetes. This trend might reflect the physical separation of trichomycetes, which are gut specific, from other symbiotes. In addition, any combinations of symbiotes that were wholly incompatible would not have been observed in our field-collected material.

Hawaiian Tetragnatha spiders inhabiting small forest fragments on the Big Island of Hawaii are parasitized by mermithid nematodes. This is the first report of mermithid nematodes infecting spiders in Hawaii, and an initial attempt to characterize this host-parasite interaction. Because immature mermithids were not morphologically identifiable, a molecular identification was performed. A phylogenetic analysis based on 18S small ribosomal subunit nuclear gene sequences suggested that Hawaiian spider mermithids are more closely related to a mainland presumptive Aranimemis species that infects spiders, than to an insect-infecting mermithid collected on Oahu, HI, or to Mermis nigrescens, also a parasite of insects. Measured infection prevalence was low (ranging from 0 to 4%) but differed significantly among forest fragments. Infection prevalence was associated significantly with fragment area, but not with spider density nor spider species richness. Results suggest that mermithid populations are sensitive to habitat fragmentation, but that changes in infection prevalence do not appear to affect spider community structure.

Strelkovimermis spiculatus is a common parasite of culicid species in Argentina. Effect of temperature on longevity and infectivity of juvenile preparasites of S. spiculatus was determined at 4, 10, 20 and 27 degrees C. Three containers with 100 ml of dechlorinate water and 300 preparasites (12 hour-old), were placed for each day and temperature, during 40 days (total = 480 containers). Survived preparasites were counted on 12 containers per day (three for each temperature). When number of survived preparasites was determined, second instar larvae of Aedes aegypti were added to each container in a 10:1 ratio (preparasites:mosquito) to determine infectivity of daily survived preparasites. Longevity of preparasites decreased at higher temperatures. Maximum longevity of preparasites maintained at 4, 10, 20 and 27 degrees C were 35, 30, 25 and 27 days, respectively. Survivorship of preparasites, exposed to the same temperatures, varied from 57% to 100% at day two, from 21% to 77% at day five and from 9% to 33% at day ten. Infectivity of preparasites maintained at temperatures from 4 to 27 degrees C was always higher than 70%. Extended longevity with maintenance of the infectivity capacity of preparasites, are important attributes to consider S. spiculatus an effective mean of controlling a large number of culicid species between 4 and 27 degrees C.

Hemoglobins are best known as oxygen transport proteins. Here we describe a hemoglobin from the parasitic nematode Mermis nigrescens (Mn-GLB-E) that has an optical, light shadowing function. The protein accumulates to high concentration as intracellular crystals in the ocellus of mature phototactic adult females while also being expressed at low concentration in other tissues. It differs in sequence and expression pattern from Mn-GLB-B, a second Mermis globin. It retains the structure and oxygen-binding and light-absorbing properties typical of nematode hemoglobins. As such, recruitment to a shadowing role in the eye appears to have occurred by changes in expression without modification of biochemistry. Both globins are coded by genes interrupted by two introns at the conserved positions B12.2 and G7.0, which is in agreement with the 3exon/2intron pattern model of globin gene evolution.

Three experiments were conducted to investigate the possibility of maintaining the Echinomermella matsi-Strongylocentrotus droebachiensis system in the laboratory. The experiments were performed by injecting E. matsi larvae taken directly from gravid female nematodes into the mouths of sea urchins. In all experiments, this treatment resulted in a higher infection in treated animals than in unmanipulated controls. The successful establishment of larvae indicates that E. matsi has a monoxenous life cycle. The growth of larvae in experimentally infected hosts was slow, indicating that the generation time of the parasite is of the same magnitude as the life expectancy of the host, 1-2 yr. This slow growth rate suggests that considerable resources will be needed to maintain the system in the laboratory.

A 12-month study of Culicoides in Bulloch County, Georgia, U.S.A., revealed that twelve of 23,859 specimens (0.05%) of C.stellifer were intersexes as a result of being parasitized by mermithid nematodes. Of these, eleven had male genitalia and female type antennae and one had female genitalia with male type antennae.

Egg production by the mermithid nematode Strelkovimermis spiculatus, a parasite of mosquitoes, was examined over a period of 34 days. Oviposition did not occur in the absence of males. Egg production was best when males were continuously present (6.4 x 10(3) +/- 0.9 x 10(3) eggs/female). Fewer eggs were produced when males were removed after 7 days (2.8 x 10(3) +/- 0.2 x 10(3) eggs/female) and oviposition partially recovered after males were returned 11 days later (4.4 x 10(3) +/- 0.5 x 10(3) eggs/female). The nematodes deposited substantially more eggs in sand (6.4 x 10(3) +/- 0.9 x 10(3)/female) than in water (1.9 x 10(3) +/- 0.3 x 10(3)/(female).

Romanomermis yuanenesis (Mermithidae: Nematoda) was found in Henan, China (Song and Peng, 1987), which has a broad host range in Culicinae mosquito and has been used successfully in field test for control of culex tritaeniorhynchus, culex fatigans and Aedes albopictus in Sichuan, Yunnan, Guangxi and Henan Provinces. This study was attempted to determine the viability and infectivity of preparasitic larvae in various temperatures. The cultures containing R. yuanenesis eggs were flooded 2h with distilled water, filtered and blocked with 1% agarose. Put the filter paper into water, then the motile preparasites separated from the unhatched eggs and got through the agarose membrane into water. About 200ml water containing preparasites free from eggs were held at 26 degrees C-28 degrees C, 16 degrees C-18 degrees C and -2 degrees C to 2 degrees C for test. The motility or lack of motility was used as the criterion to distinguish the living and dead nematodes. The rate of infection of mosquitoes and the rate of parasitism of nematodes were used to show the infectivity of the preserved preparasites. The results showed that at -2 degrees C to 2 degrees C, more than 90% of preparasitic larvae of R. yuanenesis survived for 8 days and the rate of mosquito infection was 87.5% to 100%, but at 26 degrees C-28 degrees C and 16 degrees C-18 degrees C the survival times of 90% preparasites were only 24 hours and 48 hours respectively. It indicates the low temperature preservation may prolong the survival time and keep the infectivity of these preparasitic larvae.

Laboratory studies allowed determine that female Romanomermis iyengari Welch, 1964, had a reproductive potential of 1975, 29 eggs in an average time of 21,76 days in distilled water with pH of 4.5 and a temperature of 25 +/- 2 degrees C. It was found that females of this nematode began oviposition in an average time of 14.65 days from the moment of emersion of postparasites of the host larvae. In studying the embryonic development, it was observed that the time elapsed from oviposition to egg hatching and the emersion of preparasites was 290,74 hours. It was determined that the optimum time to begin the exploitation of cultures and obtain a greater yield of preparasites under laboratory conditions is the sixth week of storage.

The effect of temperature (20 degrees-35 degrees C) on different stages of Romanomermis iyengari was studied. In embryonic development, the single-cell stage eggs developed into mature eggs in 4.5-6.5 days at 25-35 degrees C but, required 9.5 days at 20 degrees C. Complete hatching occurred in 7 and 9 days after egg-laying at 35 and 30 degrees C, respectively. At 25 and 20 degrees C, 85-96% of the eggs did not hatch even by 30th day. Loss of infectivity and death of the preparasites occurred faster at higher temperatures. The 50% survival durations of preparasites at 20 and 35 degrees C were 105.8 and 10.6 hr respectively. They retained 50% infectivity up to 69.7 and 30.3 hr. The duration of the parasitic phase increased as temperature decreased. Low temperature favoured production of a higher proportion of females which were also larger in size. The maximum time taken for the juveniles to become adults was 14 days at 20 degrees C and the minimum was 9 days at 35 degrees C. Oviposition began earlier at higher temperature than at lower temperature. However, its fecundic period was shorter at 20 degrees C than at 35 degrees C indicating enhanced rate of oviposition at 20 degrees C. Fecundity was adversely affected at 20 degrees C and 35 degrees C. It is shown that the temperature range of 25 degrees-30 degrees C favours optimum development of R. iyengari.

The oviposition pattern of Romanomermis iyengari (Nematoda: Mermithidae) in relation to the moisture level in the soil was studied by seeding postparasites into two sets of 12 cm soil columns. While one set of the soil column was allowed to dry over a period of 60 days, the other set was maintained at a constant level of 15-20% moisture throughout the soil column. In the soil column maintained at constant moisture level, the postparasites oviposited primarily in the upper 3 cm layer (80-97%). In the soil column which was allowed to dry the percentages of eggs laid in the upper 3 cm layer were 96, 24 and 6 respectively on D20, D30 and D40 after seeding and the moisture levels on respective days were 12, 5 and 3%. On corresponding days the postparasites laid 1, 56 and 42% of the eggs in the lower most layer (9-12 cm) and the moisture levels were respectively 15, 10 and 9%. The results indicate that the adult nematodes migrated towards the bottom layer of the soil and laid eggs as the moisture of the upper layers decreased.

The parasitic nematodes Romanomermis culicivorax and R. iyengari, were evaluated as a means of bloodsucking mosquito larvae control in Apsheron Peninsula (the city of Baku) and Lenkoran Lowland (Masally District) in Azerbaijan. Being once brought, mermithids got acclimatized in 12 shallow freshwater nonpolluted pools with aquatic flora and inhibited the emergence of 13-97% of mosquitoes during the summer period. R. culicivorax successfully survived the temporary drying (up to 2 weeks) in 3 and spent a winter in 2 water reservoirs. The possibility of R. iyengari settling via transmission of infected mosquito larvae from a "nursery" pool to a mermithid free water pool was proved. The larvae Cx. modestus, Cx. theileri, Ae. caspius and Ur. unguiculata are first registered as hosts of R. culicivorax and An. sacharovi, Cx. theileri and that of R. iyengari. Both mermithid species may be considered as a means of prolonged inhibition of mosquito emergence in the shallow nonpolluted fresh-water reservoirs in the Azerbaijan SSR.

The imaginal discs of chironomid larvae infected with mermithid nematodes provided a useful model for the study of parasitic effects upon undifferentiated tissue. Specific aberrations concerning mitotic rate, cell death, organogenesis, cell pattern differentiation and cell differentiation are described and discussed.

Penetrating infective juveniles of Romanomermis culicivorax usually killed first-stage larvae of Chironomus maturus Johann., Chironomus sp., Simulium damnosum Theo., and S. venustum Say. Nematodes were melanized and died after they entered fourth stage larvae of 2 chironomid species, but no host reaction was evident after entry into fourth-stage blackfly larvae. In contrast, the nematodes initiated development in the latter hosts, which died before the nematodes completed their development.

The efficacy of the parasitic nematode Romanomermis culcivorax in controlling anopheline larvae, including malaria vectors, was studied in Fars and Baluchistan provinces. Twenty sites were treated once with the preparastic stage of R. culicivorax in 1984 with 3,000, 5,000 or 10,000 preparasites per m2 surface area, depending on larval density. The average parasitism of anopheline larvae ranged from 56 to 69% based on 24 hr posttreatment dissections. No correlation was found between the level of parasitism and the density of mosquito larvae present in a site. About 61% parasitism was obtained when different rates of preparasites were released in 14 larval breeding sites in 1985. No apparent difference was observed in the rate of parasitism in 10 sites receiving one treatment compared with four sites receiving two treatments, with a seven day interval between each treatment. Romanomermis culcivorax was established in the release sites but caused only minor reductions in anopheline larval populations. The nematode would be of limited use in antimalaria campaigns in southern Iran.

Empidomermis riouxi n. sp. is described from Aedes (Ochlerotatus) detritus, in Camargue. The species differs from E. cozi, which is the only other species of the genus, by having a sligtly larger size and a very little pronounced cephalic constriction. Males differ also from those of E. cozi by the number and disposition of the caudal papillae. The postparasitic larvae escape from adult mosquitoes, rarely from pupae. In laboratory conditions, the nematodes enter the moist sand in the bottom of Petri dishes where, after about 20 days, they molt, mate and oviposit. All the larval stages of A. detritus can be found infected. The presence of parasites induce the arrest of the ovarian development of the females and inhibits the mating activity of males. The rate of parasitism is very high and its fluctuations seem closely related to the host life cycle.