Population ecology of the commensal chironomid Epoicocladius flavens on its mayfly host Ephemera danica

SUMMARY. 1. Epoicocladius flavens (Diptera: Chironomidae) has a 1-year life cycle in the River Tud in eastern England. Pupation and adult emergence occurred from the end of May until the beginning of September. Larval growth was rapid during the summer but thereafter almost ceased until the following April.1 Unlike other chironomids, the population density of E. flavens was very stable and remained at around 1000m ⁻² all through the year.
2. Ephemera danica (Ephemeroptera: Ephemeridae), the host of E. flavens , has a 2-year life cycle. The proportion infested was about 90% for the older host year-class and less than 30% for the younger year-class from autumn until the following summer. A rapid increase in infestation occurred during the summer as a result of hatching of the E. flavens eggs. Of those infested, the older year-class hosts harboured two or three larvae, on average, whereas the younger ones had only one.
3. When host year-classes were treated separately, the frequency distribution of the number of E. flavens per host each month did not depart significantly from a Poisson distribution. Younger instars departed from a Poisson distribution more frequently than older instars.
4. The majority of E. flavens larvae attached to the gills of their hosts. As the instars progressed, increasing proportions were found on the thorax, legs and tails. The thorax was the preferred site for fourth instar larvae.
5. Enhanced chance of survival, rather than the supply of detrital food particles which collect on the host's body, is considered as an important benefit to accrue to E. flavens from this commensalism.     

The occurrence of ixodid ticks on wild mink Mustela visonin England and Wales

Abstract. Four species of ticks found to infest 1391 American mink (Mustela vison) in Britain in five years were, in declining order of frequency, Ixodes hexagonus, Ixodes canisuga, Ixodes ricinus and Ixodes acuminatus. Ixodes hexagonus and I.canisuga occurred on 40% and 2.5% of mink respectively. Infestation rates (the proportion of infested mink) of adult females, nymphs and larvae were similar and tended to be lower in summer. The distribution of infestation size (the number of ticks per host) for adult females describes a negative binomial. The mean infestation size of nymphs varied with the sex of the host and was 5.2 for males and 4.2 for females. Mink are competent hosts for /. hexagonus.     

Some factors affecting the location of Arthritica crassiformis (Bivalvia: Leptonacea) commensal upon Anchomasa similis (Bivalvia: Pholadidae).

Arthritica crassiformis Powell is found in New Zealand waters commensally associated with the pholad Anchomasa similis (Gray). Its location on the host is highly specific and is occasioned by the complex needs of the commensal and the facilities and limitations imposed by the tunnelling activity of the host. Other species of pholad because of different morphological adaptations do not serve as hosts. With increasing size of the host, the number of commensals/host increases which is due to (a) retention of young individuals within the parental burrow and (b) by colonization from the external environment.
A. crassiformis typically, however, occurs in pairs and is probably a protandric consecutive hermaphrodite. The young are incubated in the interlamellar spaces of the ctenidia of the parental female. This anatomical adaptation together with others including strong rejectory tracts suit the commensal for a life with an active tunnelling host.
The ciliary currents of the mantle cavity of A. similis are described.     

The infection of nymphal Baetisbicaudatus by the mermithid nematode Gasteromermis sp.

1. This study reports the infection in nymphs of a bivoltine mayfly host ( Baetis bicaudatus ) in a high-elevation watershed by the mermithid nematode Gasteromermis sp. Infection by Gasteromermis causes mortality in two ways. Fifty per cent of the infections do not successfully develop beyond the initial stage of penetration and result in the early death of both host and parasite.
2. Infected hosts that survive this initial stage are rendered completely sterile by the infection (reproductively dead). In addition to complete sterility, the emergence size of parasitized nymphs is reduced and development time lengthened compared with unparasitized nymphs.
3. Parasite infection levels are stable from year to year at one site, but with a higher incidence of infection in the mayfly summer generation. Size differences between the generations at the time of infection may account for their different susceptibilities.
4. Within a year infection levels vary seasonally and spatially from 1 to 71%. Seasonally, there is a condensation of parasitized hosts towards the end of development as unparasitized nymphs emerge earlier. Spatially, infection levels show a downstream decline that may result from upstream dispersal by infected hosts or differential parasite survivorship at different elevations.     

Pupation, emergence and fecundity of phoretic Epoicocladius ephemerae (Chironomidae)

The importance of the presence of nymphs of Ephemera danica for the emergence and survival of phoretic Epoicocladius ephemerae was tested in the laboratory. Only a few midges survived when hosts were absent. The synchronization of emergence of the host and its associate was examined in the field. Fecundity and total egg volume per unit wing length was compared between E. ephemerae and several free-living chironomids. The number of eggs was about the same in E. ephemerae as in the other species, but total egg volume was comparatively large. A comparison is also made between the host-related adaptations and feeding habits of E. ephemerae and of the parasitic Symbiocladius rhithrogenae. Some larvae of E. ephemerae use the fullgrown nymphs as pupation sites, probably because the emerging hosts carry the pupae to the water surface. The reason why not all midge larvae make use of nymphs for this purpose is probably because they are unable to adapt to the exact emergence time of the nymphs. Analysis of the emergence data of the midge and of the mayfly support this hypothesis.     

The probing behaviour of nymphs of Vanduzeea arquata and Enchenopa binotata (Homoptera: Membracidae) on host and non-host plants

Abstract. 1. Nymphs of Vanduzeea arquata Say have been found to be more host-specific in nature and to show a higher degree of selectivity in host discrimination experiments than nymphs of Enchenopa binotata (Say), It was hypothesized that this differential selectivity would be reflected in the probing behaviour of individuals placed on twigs of host and non-host plants. Probing behaviour was examined by direct observation of nymphs and by sectioning and staining the probed plant tissues.
2. All nymphs probed readily and for extended periods on both host and non-host twigs. E.binotuta nymphs showed no consistent differences in probing behaviour on hosts versus non-hosts, but V.atquuta nymphs were more likely to withdraw their stylets within 60 s when on non-host twigs and produced honeydew only when on their host species. V.urquatu nymphs reached the phloem sieve elements only when on host twigs and broke many cells in peripheral plant tissue layers while probing. E.binotata nymphs broke few cells and often reached the phloem of non-host as well as host plants.
3. Nymphs of V.arquata always reject non-host plants, apparently in the course of probing and prior to encountering the phloem sap. Chemical compounds released from ruptured parenchyma cells may act as probing stimulants or inhibitors. E.binotura nymphs often feed on non-host plants in a non-choice situation; their preferential settling on host twigs in discrimination experiments may reflect a tendency to abandon non-host twigs more readily than host twigs.     

Heterogeneity of resource utilization in a population of the Australian reptile tick, Aponomma hydrosauri (Denny)

Abstract. 1. The distribution of adult males, nymphs and larvae of the reptile tick Aponoma hydrosauri on their host, Trachydosaurus rugosus , in a study area near Tickera, South Australia, did not follow the Poisson distribution predicted if hosts were infested at random.
2. The non-random distribution of males can be accounted for by geographical heterogeneity in distribution. Adult females also showed geographical heterogeneity in their distribution pattern, but infested their hosts at random over all.
3. Larvae and nymphs show distribution patterns which change both with the time and the locality. However, when these biases are removed the distribution of larvae and nymphs on their hosts is still non-random.
4. Larger lizards tend to have larger numbers of ticks, but this cannot alone account for the clumped distribution.
5. A model is proposed relating the tick distribution pattern to the particular problems of a resource which is hard to find, but plentiful once found.     

A 9-year study of the life cycle of Ephemera danica Müll. (Ephemeridae: Ephemeroptera) in the River Lambourn, England

Abstract. 1. The life cycle of the mayfly, Ephemera danica , was studied on two contrasted sites on the River Lambourn between 1971 and 1979.
2. Quantitative samples of nymphs were taken on the five major biotopes of the river bed, gravel, silt, Ranunculus, Berula and Callitriche . Exposed areas of gravel and silt held significantly lower densities of nymphs than the three macro-phytes with their underlying substrata. The sandy substratum underlying beds of Berula frequently held significantly higher densities than the other macrophytes.
3. Monthly samples from March 1971 to April 1972 followed by samples in June and December from 1972 to 1979 indicated that the nymphal phase normally took 2 years in the River Lambourn.
4. All odd-numbered years from 1971 to 1979 produced weak year-classes. In contrast, all even-numbered years between 1970 and 1978, with the exception of 1972, produced relatively strong year-classes.
5. Meteorological data for the period of flight activity in E.danica indicated that conditions were colder and damper in 1972 than in the other even-numbered years of the study, when recruitment was more successful.     

Host size induced variation in progeny sex ratio of an aphid parasitoid Lysiphlebia mirzai

A study aimed at determining the effect of host size on the pattern of sex allocation by an aphid parasitoid Lysiphlebia mirzai Shuja-Uddin (Hymenoptera: Braconidae: Aphidiinae) was undertaken to test the validity of the following three hypotheses: (1) larger hosts will yield a greater proportion of daughters, (2) differential mortality of the sexes of the parasitoid during development is a function of host-size, and (3) daughters emerging from larger hosts are more fecund. The results suggested a propensity in the parasitoid to deposit fertilised (diploid) eggs in large hosts (third instar nymph) and unfertilised (haploid) eggs in small hosts (first and second instar nymphs). Unpreferred fourth instar nymphs and apterous aphid adults also received more haploid eggs despite being larger in size than the preferred third instar nymphs. However, the perception of host size by the mother was dependent on the extent of temporal variation in the host size distribution and on her previous experience of host size. Developmental period, longevity, mating potential, fecundity and progeny sex ratio of L. mirzai emerging from small versus large hosts indicated that the host size affected the fitness of the daughters more than that of sons. No differential mortality of the sexes during development of L. mirzai was observed in small versus large hosts. This shows that L. mirzai, while ovipositing in growing stages of the host, adjusts progeny sex ratio according to the host size and by doing so she tends to contribute maximally to progeny fitness without knowing about the future host quality.     

Species richness and regional distribution of myrmecophilous beetles

Four major hypotheses have been put forward to explain local species richness of commensal or parasitic species. The resource distribution hypothesis predicts that regionally widespread host species are able to support higher local species richness of commensals or parasites. On the other hand, the resource size hypothesis predicts that larger hosts can support more species than smaller hosts, and comparably, the resource abundance hypothesis predicts that hosts that offer more resources are able to support more species. Finally, the resource concentration hypothesis predicts that hosts that occur in high-density patches support higher species richness. In this study, we tested the first three of the above hypotheses with myrmecophilous beetles and their host ants. In addition to species richness of myrmecophilous beetles, we also applied the above hypotheses to explain the distribution of the beetles. Our data are exclusively based on an extensive literature survey. Myrmecophilous beetles live in naturally fragmented environments composed of host ant colonies and they are exclusively dependent on ants. We found that the distribution of the host ants and the colony size of the host ants had a positive effect on both the species richness and the distribution of myrmecophilous beetles. In the same way, we found that myrmecophilous beetle species that are generalists, i.e. have more than one host ant species, and thus have more abundant resources, were more widely distributed than specialist species. Thus, we found support for the hypothesis that resource distribution, resource size and resource abundance have an effect on species richness and on the distribution of species.     

Oviposition behaviour, influence of experience on host size selection, and niche overlap of the solitary Leptomastix epona and the gregarious Pseudaphycus flavidulus, two endoparasitoids of the mealybug Pseudococcus viburni

Oviposition behaviour and host size selection of the solitary parasitoid Leptomastix epona(Walker) and the gregarious Pseudaphycus flavidulus(Brèthes) [both Hymenoptera: Encyrtidae] were examined on five size classes of the mealybug Pseudococcus viburni(Signoret) [Hemiptera: Pseudococcidae]. The host size classes mostly consisted of one stage (first, second, third instar nymph, young adult and preovipositing adult) and were presented together to wasps of either parasitoid species. Both parasitoid species locate the host by drumming the surface of the patch with the antennae. Leptomastix eponaseems to use mainly the antennae to examine the host but P. flavidulusmay accept or reject a host for oviposition after antennation or insertion of the ovipositor. Leptomastix eponaattempts oviposition in all the host stages from second instar nymphs but P. flavidulusincludes first instar. Both parasitoid species select mainly larger hosts (>1 mm, third instar nymphs) to oviposit but P. flavidulusis able to parasitize more second instar nymphs compared to L. epona. Female wasps of L. eponamay host feed on small mealybugs (second and third instar nymphs) that they do not use for oviposition. Oviposition experience of either parasitoid species for 24 hours does not influence host size selection on patches with hosts of similar mixed sizes. Oviposition decisions are independent of the host sizes of the preceding ovipositions. Implications about stability of a single parasitoid – host system and the success of biological control of the mealybug were discussed in respect of the developmental refugia of the two parasitoid species. Niche overlap of the two parasitoid species was discussed with a view to giving an insight into a single or multiple introduction.     

Effects of larval movement behavior and density on emergence success and adult body size in a commensal midge

Changes in larval density and movement behavior of a commensal midge, Nanocladius (Plecopteracoluthus) sp. #4, were monitored for 26 weeks in recirculating laboratory streams. Adults were captured at emergence, sexed, and weighed to assess the effect of larval density and movement behavior on emergence success and adult size. The density of midges on hosts declined with time and coincided with a springtime increase in larval movement frequency. Midges residing on hosts with high spring densities emerged significantly less than midges on hosts with low densities. Resident midge density on hosts did not influence the likelihood of successful colonization by commensals and colonizers showed no preference for initial attachment site on hosts. However, colonizing midges emerged significantly less than resident midges. Similarly, successful emergers changed tube positions significantly less often as larvae relative to non-emergers. There was no difference in adult body weight of resident midges and colonists/movers, but adult males which emerged from commensal-laden (high density) hosts were significantly smaller than males from low density hosts. These data indicate larval density and movement behavior may have strong fitness consequences for commensal midges.     

Habitat utilization by sympatric arctic charr Salvelinus alpinus L. and brown trout Salmo trutta L. in Lake Atnsjø, south-east Norway

SUMMARY. 1. Habitat utilization, as well as inter- and intraspecific relations of different size groups of arctic charr (Salvelinus alpinus (L.)) and brown trout (Salmo trutta L.) in Lake Atnsjø, south-east Norway, were investigated by analysing food and spatial niches from monthly benthic and pelagic gillnet catches during June-October 1985.
2. Small individuals (150–230 mm) of both arctic charr and brown trout occurred in shallow benthic habitats. However, they were spatially segregated as arctic charr dominated at depths of 5–15 m and brown trout at depths of 0–5 m.
3. Larger (>230 mm) arctic charr and brown trout coexisted in the pelagic zone. Both species occurred mainly in the uppermost 2-3 m of the pelagic, except in August, when arctic charr occurred at high densities throughout the 0–12 m depth interval. On this occasion, arctic charr were segregated in depth according to size, with significantly larger fish in the top 6 m. This was probably due to increased intraspecific competition for food.
4. The two species differed in food choice in both habitats, Arctic charr fed almost exclusively on zooplankton, whereas brown trout had a more variable diet, consisting of surface insects, zooplankton. aquatic insects and fish.
5. The data suggest that the uppermost pelagic was the more favourable habitat for both species. Large individuals having high social position occupied this habitat, whereas small individuals lived in benthic habitat where they were less vulnerable to agonistic behaviour from larger individuals and less exposed to predators. The more aggressive and dominant brown trout occupied the more rewarding part of the benthic habitat.     

Effect of the primary host for production of both sexes on the mating interaction in an autoparasitoid species

Encarsia sophia (Girault and Dodd) is an autoparasitoid in the hymenopteran family Aphelinidae. The females develop as primary parasitoids on whitefly nymphs (primary hosts), whereas the males develop as hyperparasitoids on their own species or on other primary parasitoid species (secondary hosts). The autoparasitoids not only parasitise whiteflies but also kill them with strong host-feeding capacity. In this study, female and male E. sophia were reared on the primary hosts Trialeurodes vaporariorum and Bemisia tabaci ‘Q’, and the host-feeding and parasitism of wasps on both whitefly species were determined for the four possible different mating combinations: (i) E. sophia females reared on B. tabaci (ESF-BT) mated with E. sophia males from B. tabaci (ESM-BT), (ii) E. sophia females reared on T. vaporariorum (ESF-TV) mated with E. sophia males from T. vaporariorum (ESM-TV), (iii) ESF-BT mated with ESM-TV, and (iv) ESF-TV mated with ESM-BT. ESF-TV mated with ESM-TV killed the largest percentage of whitefly nymphs through host feeding. The ESF-TV with larger body size mating with larger ESM-TV killed more whitefly nymphs through host feeding than those mating with smaller ESM-BT. Whether B. tabaci or T. vaporariorum were used as hosts, ESF-TV mated with ESM-TV and ESM-BT and ESF-BT mated with ESM-BT significantly parasitised more whitefly nymphs than ESF-BT mated with ESM-TV. In general, ESF-BT mated with ESM-TV killed significantly fewer whitefly nymphs through parasitism and host feeding than the other three mating combinations on both whitefly species. These results indicated that the performance of autoparasitoids on insect pests was not only dependent on females but was also affected by mating with males from different primary host species.     

Observations on the occurrence and maturation of the nematode Rhabdochona denudata (Dujardin, 1845) in Chub, Leuciscus cephalus (L.), of the Rokytná River, Czechoslovakia

From October 1985 until July 1987, the seasonal dynamics of Rhabdochona denudata in its principal definitive host, the chub (Leuciscus cephalus), was studied in the Rokytná River (the Danube basin), Czechoslovakia. Prevalence (overall prevalence 77%) and intensity of infection (1-59 nematodes per fish) in the fish were high all year, with both abiotic and biotic factors influencing the fluctuations. Maximum values of prevalence (82-87%) were found in the fish with body length exceeding 20 cm; the mean intensity was highest in the size-group of fish 20-25 cm long, whereas its values gradually decreased in larger fish, this being due to the choice of food by fish of different size-groups. In contrast to the situation in other localities, an annual maturation cycle was not evident and egg-producing female nematodes were present in fish throughout the year. New infections in fish were most common in summer but occurred all year round. This life history pattern seems to be mainly due to the availability of infected ephemeropteran intermediate hosts during all seasons, in this case the nymphs of Ephemera danica, a species noted for its two-year development. In this locality, specifically not determined Rhabdochona larvae were recorded from the mayfly nymphs Ephemera danica (0.4%), Ecdyonurus aurantiacus (7%), and Caenis macrura (10%). The diet of the chub included mayfly nymphs at all seasons.     

Specificity and host predictability: a comparative analysis among monogenean parasites of fish

1. This article compares generalist (parasite species found on two or more host species) and specialist (found on only one host species) monogenean parasite species of fish. The reduction of the host range – that is an increase in host specificity – may correspond with a better adaptation of the parasite to a more predictable host environment. A more predictable environment may allow the parasite species to develop specific adaptations.
2. We assume that the more predictable host environment can be evaluated by host body size, since numerous life-traits, such as longevity, are positively correlated with size.
3. We found that specialist parasites parasitize larger hosts species than generalist parasites. We also found a good relationship between host body size and parasite body size for specialist parasite species.
4. An adaptation to the mechanical problems encountered in the host's gill chamber may lead to an increase in parasite body size. The infection of a larger part of the host population in order to decrease the chances of local extinction due to fluctuations of host abundance may be another adaptive mechanism.
5. We found a negative correlation between parasite body size and prevalence for generalist parasite species. This relationship disappeared when using the comparative method controlling for phylogeny, which proved that it was a phylogenetic effect.     

The influence of host distribution, sex, and size on the level of parasitism by Itoplectis conquisitor (Hymenoptera: Ichneumonidae)

Abstract. 1. Parasitism of the bagworm, Thyridopteryx ephemeraeformis Haw. (Lepidoptera: Psychidae), on red cedar was studied at Manassas, Va., U.S.A.
2. Fifteen parasitoids, five of which were hyperparasitoids, attacked the pupal stage of the bagworm. The Ichneumonid Itoplectis conquisitor alone accounted for 58% of the parasitized bagworms.
3. Parasitism by I conquisitor (Say) was inversely related to host size, but independent of host distribution within a tree. Male bagworms experienced disproportionately higher levels of parasitism than females. We conclude that it is small size that renders bagworms susceptible to parasitism rather than sex per se.
4. Ovipositor lengths of I.conquisitor were insufficient to penetrate the larger bagworm hosts (>57 mm); and in fact, as bagworm size increased, the proportion of the I.conquisitor population capable of penetrating the pupa declined abruptly. We suggest that the mechanical difficulties with oviposition, and perhaps the defensive capabilities of larger hosts, are responsible for the relationship between host size and per cent parasitism.
5. I.conquisitor could be an important selective agent for bagworm size at pupation, but it is not likely to act as a significant control agent of population density.     

Changes in prevalence and intensity of infection of <Emphasis Type="Italic">Profilicollis altmani</Emphasis> (Perry, 1942) cystacanth (Acanthocephala) parasitizing the mole crab <Emphasis Type="Italic">Emerita analoga</Emphasis> (Stimpson, 1857): an El Niño cascade effect?

Prevalence and intensity changes in cystacanths of the acanthocephalan Profilicollis altmani parasitizing the mole crab Emerita analoga under El Niño (EN) and non-El Niño (non-EN) conditions are analyzed. Both, mean intensity and prevalence of infection by P. altmani differ significantly for the whole size range and for each size class of 10 mm intervals (except prevalence for size classes exceeding 18 mm carapace length) between EN (1998) and non-EN (2002) years, without observed size distribution differences in the intermediate host E. analoga under either condition. Significant difference in infestation rates of the intermediate host E. analoga is discussed as being an EN cascade effect on predators such as sea birds (i.e., Larus spp. and Calidris sp.), acting as definitive hosts of P. altmani, and which are known to decrease significantly in abundance during EN.     

The evolutionary ecology of biotic association in a megadiverse bivalve superfamily: sponsorship required for permanent residency in sediment

Background

Marine lineage diversification is shaped by the interaction of biotic and abiotic factors but our understanding of their relative roles is underdeveloped. The megadiverse bivalve superfamily Galeommatoidea represents a promising study system to address this issue. It is composed of small-bodied clams that are either free-living or have commensal associations with invertebrate hosts. To test if the evolution of this lifestyle dichotomy is correlated with specific ecologies, we have performed a statistical analysis on the lifestyle and habitat preference of 121 species based on 90 source documents.

Methodology/Principal Findings

Galeommatoidea has significant diversity in the two primary benthic habitats: hard- and soft-bottoms. Hard-bottom dwellers are overwhelmingly free-living, typically hidden within crevices of rocks/coral heads/encrusting epifauna. In contrast, species in soft-bottom habitats are almost exclusively infaunal commensals. These infaunal biotic associations may involve direct attachment to a host, or clustering around its tube/burrow, but all commensals locate within the oxygenated sediment envelope produced by the host’s bioturbation.

Conclusions/Significance

The formation of commensal associations by galeommatoidean clams is robustly correlated with an abiotic environmental setting: living in sediments (). Sediment-dwelling bivalves are exposed to intense predation pressure that drops markedly with depth of burial. Commensal galeommatoideans routinely attain depth refuges many times their body lengths, independent of siphonal investment, by virtue of their host’s burrowing and bioturbation. In effect, they use their much larger hosts as giant auto-irrigating siphon substitutes. The evolution of biotic associations with infaunal bioturbating hosts may have been a prerequisite for the diversification of Galeommatoidea in sediments and has likely been a key factor in the success of this exceptionally diverse bivalve superfamily.     

Host selection and egg allocation behaviour by Aphytis melinus and A.lingnanensis: comparison of two facultatively gregarious parasitoids

Abstract. 1. The ovipositional and egg allocation behaviour of individual females of Aphytis melinus DeBach and A.lingnunensis Compere were compared.
2. Both Aphytis species exhibit the same behavioural sequence during oviposition.
3. Aphytis melinus laid most of its female eggs on the dorsum of a scale-insect beneath its cover, and most of its male eggs under the scale-insect's body. Aphytis lingnanensis also oviposited both dorsally and ventrally on scale-insect hosts, but female and male progeny arose with equal frequency from eggs laid in both locations.
4. Both A.melinus and A. lingnanensis are facultatively gregarious parasitoids. The degree of gregariousness depends on host size, i.e. the larger the host, the more the Iikelihood that several eggs will be deposited at each visit by the parasitoid.
5. When two eggs were laid during the same host visit, both A.melinus and A.lingnanensis laid one female and one male egg more often than would be expected under an assumption of random allocation of sexes.
6. Because A.melinus successfulIy utilize smaller hosts than A.lingnanensis to produce progeny, these parasitoids should not be considered ecological homo-logues, as suggested by DeBach & Sundby (1963).