Individual Specialization to Non-Optimal Hosts in a Polyphagous Marine Invertebrate Herbivore

Factors determining the degree of dietary generalism versus specialism are central in ecology. Species that are generalists at the population level may in fact be composed of specialized individuals. The optimal diet theory assumes that individuals choose diets that maximize fitness, and individual specialization may occur if individuals'' ability to locate, recognize, and handle different food types differ. We investigate if individuals of the marine herbivorous slug Elysia viridis, which co-occur at different densities on several green macroalgal species in the field, are specialized to different algal hosts. Individual slugs were collected from three original algal host species (Cladophora sericea, Cladophora rupestris and Codium fragile) in the field, and short-term habitat choice and consumption, as well as long-term growth (proxy for fitness), on four algal diet species (the original algal host species and Chaetomorpha melagonium) were studied in laboratory experiments. Nutritional (protein, nitrogen, and carbon content) and morphological (dry weight, and cell/utricle volume) algal traits were also measured to investigate if they correlated with the growth value of the different algal diets. E. viridis individuals tended to choose and consume algal species that were similar to their original algal host. Long-term growth of E. viridis, however, was mostly independent of original algal host, as all individuals reached a larger size on the non-host C. melagonium. E. viridis growth was positively correlated to algal cell/utricle volume but not to any of the other measured algal traits. Because E. viridis feeds by piercing individual algal cells, the results indicate that slugs may receive more cytoplasm, and thus more energy per unit time, on algal species with large cells/utricles. We conclude that E. viridis individuals are specialized on different hosts, but host choice in natural E. viridis populations is not determined by the energetic value of seaweed diets as predicted by the ODT.     

Characterization of the Parasitic Dinoflagellate Amoebophrya sp. Infecting Akashiwo sanguinea in Coastal Waters of China

The endoparasitic dinoflagellate Amoebophrya infects a number of free‐living marine dinoflagellates, including harmful algal bloom species. The parasitoid eventually kills its host and has been proposed to be a significant loss factor for dinoflagellate blooms in restricted coastal waters. For several decades, the difficulties of culturing host‐parasitoid systems have been a great obstacle for further research on the biology of Amoebophrya. Here, we established an Akashiwo sanguinea‐Amoebophrya sp. coculture from Chinese coastal waters and studied the parasitoid's generation time, dinospore survival and infectivity, as well as its host specificity. The lifespan of Amoebophrya sp. ex. A. sanguinea was approximately 58 h. The infective dinospores can survive up to 78 h in ambient waters but gradually lose their infectivity. The parasitoid was unable to infect other dinoflagellate species, its infection rate reached as high as 91% when the ratio of dinospores to host cells was 20:1. The high infectivity of dinospores suggests that the Amoebophrya strain was capable of removing a considerable fraction of host biomass within a short period, but that it is probably unable to maintain high infection levels under nonbloom conditions of its host, due to limited survival and time constraints in encountering host cells.     

Genetic covariation of the marine fungal symbiont Haloguignardia irritans (Ascomycota, Pezizomycotina) with its algal hosts Cystoseira and Halidrys (Phaeophyceae, Fucales) along the west coast of North America

The fungal endophyte Haloguignardia irritans induces gall formation on the brown algal genera Cystoseira and Halidrys occurring from Oregon to Baja California, Mexico. Here we examine genetic covariation and compare rDNA phylogenies to investigate the coevolutionary histories of H. irritans and its algal hosts. Despite recognition of H. irritans as a single morphological species, internal transcribed spacer rDNA sequences representative of its geographic range are characterized by sequence variation at the intraspecific to intrageneric levels. An assessment of parallel cladogenesis between endophyte and host phylogenies provides evidence for a combination of independent fungal divergence and host jumping, similar to that observed in terrestrial lichens. Our results suggest that reduced gene flow due to geographic isolation is a major contributing factor to more concerted covariation observed at one island site, rather than to differences among algal host species alone. Because geography and its effects on gene flow can create heterogeneous mosaics of coevolution for symbioses in terrestrial environments, our results support the notion that conservation efforts toward the maintenance of genetic diversity in marine environments should likewise consider geographic complexity and its effects on coevolving marine species.     

Brandtia ciliaticola gen. et sp. nov. (Chlorellaceae,Trebouxiophyceae) a common symbiotic green coccoid of various ciliate species

Many freshwater protists harbor unicellular green algae within their cells and these host‐symbiont relationships slowly are becoming better understood. Recently, we reported that several ciliate species shared a single species of symbiotic algae. Nonetheless, the algae from different host ciliates were each distinguishable by their different genotypes, and these host‐algal genotype combinations remained unchanged throughout a 15‐month period of sampling from natural populations. The same algal species had been reported as the shared symbiont of several ciliates from a remote lake. Consequently, this alga appears to play a key role in ciliate‐algae symbioses. In the present study, we successfully isolated the algae from ciliate cells and established unialgal cultures. This species is herein named Brandtia ciliaticola gen. et sp. nov. and has typical ‘Chlorella‐like’ morphology, being a spherical autosporic coccoid with a single chloroplast containing a pyrenoid. The alga belongs to the Chlorella‐clade in Chlorellaceae (Trebouxiophyceae), but it is not strongly connected to any of the other genera in this group. In addition to this phylogenetic distinctiveness, a unique compensatory base change in the SSU rRNA gene is decisive in distinguishing this genus. Sequences of SSU‐ITS (internal transcribed spacer) rDNA for each isolate were compared to those obtained previously from the same host ciliate. Consistent algal genotypes were recovered from each host, which strongly suggests that B. ciliaticola has established a persistent symbiosis in each ciliate species.     

Parasites and phytoplankton, with special emphasis on dinoflagellate infections

Planktonic members of most algal groups are known to harbor intracellular symbionts, including viruses, bacteria, fungi, and protozoa. Among the dinoflagellates, viral and bacterial associations were recognized a quarter century ago, yet their impact on host populations remains largely unresolved. By contrast, fungal and protozoan infections of dinoflagellates are well documented and generally viewed as playing major roles in host population dynamics. Our understanding of fungal parasites is largely based on studies for freshwater diatoms and dinoflagellates, although fungal infections are known for some marine phytoplankton. In freshwater systems, fungal chytrids have been linked to mass mortalities of host organisms, suppression or retardation of phytoplankton blooms, and selective effects on species composition leading to successional changes in plankton communities. Parasitic dinoflagellates of the genus Amoebophrya and the newly described Perkinsozoa, Parvilucifera infectans, are widely distributed in coastal waters of the world where they commonly infect photosynthetic and heterotrophic dinoflagellates. Recent work indicates that these parasites can have significant impacts on host physiology, behavior, and bloom dynamics. Thus, parasitism needs to be carefully considered in developing concepts about plankton dynamics and the flow of material in marine food webs.     

Evidence of photosynthate translocation in an algal-acoel symbiotic system: An in vivo, qualitative approach

Convolutriloba retrogemma, like other members of the genus and other symbiotic acoels, does not lend itself well to established methods for the detection and quantification of photosynthate translocation from symbiont to host. We describe a novel method for indirectly detecting evidence of photosynthate movement, in vivo, utilizing differential weight change in animals subjected to light and dark treatments without holozoic feeding. Unfed acoels kept in light lost 28% of their original weight while those kept in the dark lost 41%. Also described is a refined method for the separation of algal symbionts from host tissue in these species, as well as a method for determining accurate wet-weight of this and other soft-bodied, invertebrate species.     

Host-symbiont associations of polycystine Radiolaria: epifluorescence microscopic observation of living Radiolaria

In all 29 polycystine radiolarian species were obtained from surface seawater on May 28, 1999, using a plankton-net at one station (Site 990528; 26°37′18″N, 127°47′35″E) approximately 5 km northwest of Okinawa Island, Japan. In most polycystine radiolarians of the orders Nassellarida and Spumellarida symbiotic algae were observed under light microscopy. The light microscopic (LM) images of the symbionts, however, varied in clarity among individuals because of the variations in microanatomy of the host radiolarian cells. On the other hand, epifluorescence microscopic (EFM) observation easily detected and confirmed the existence of the algal symbionts within the host cytoplasm even in radiolarians such as Dictyocoryne truncatum (Ehrenberg) that include algal symbionts in the depth of the cytoplasm. The chloroplasts of the algal symbionts emitted autofluorescence in ultraviolet irradiation and they appeared red. That is, the autofluorescence images of the chloroplasts can be used to recognize the existence of the algal symbionts within the host radiolarians. Moreover, staining of the symbiont cells with 4′,6-diamido-2-phenylindle permitted visualization of the nucleus in the center of the symbiont cell, confirming the existence of living endosymbiotic algae within the polycystine radiolarians. Both the LM and EFM observations of eight polycystine radiolarian species revealed the specific patterns of various host-symbiont associations. (1) The investigated polycystine radiolarians all possess algal symbionts, except for one species, i.e. Dictyocoryne profunda Ehrenberg. (2) The size of the algal symbionts depends on the radiolarian species. The symbionts are largely classified into two types based on the size of their diameters, i.e. about 8–10 μm for the larger group and about 5 μm for the smaller one. (3) The algal symbionts show a variety of locations within the host radiolarian cytoplasm. The types of distribution of algal symbionts may be a useful characteristic for radiolarian taxonomy.     

Molecular approaches to diagnosing nutritional physiology in harmful algae: Implications for studying the effects of eutrophication

Every year harmful algal blooms (HABs) cause serious impacts to local economies, coastal ecosystems, and human health on a global scale. It is well known that nutrient availability can influence important aspects of harmful algae biology and ecology, such as growth, toxin production, and life cycle stage, as well as bloom initiation, persistence and decline. Increases in the rate of supply of organic matter to ecosystems (eutrophication) carries many possible ramifications to coastal systems, including the potential for nutrient enrichment and the potential for stimulation of harmful algal blooms. Traditional studies on algal nutrition typically use either cultured isolates or community level assays, to examine nutrient uptake, nutrient preference, elemental composition, and other metrics of a species’ response to nutrients. In the last decade, technological advances have led to a great increase in the number of sequences available for critical harmful species. This, in turn, has led to new insights with regards to algal nutrition, and these advances highlight the promise of molecular technologies, and genomic approaches, to improving our understanding of algal nutrient acquisition and nutritional physiological ecology, in both cultures and field populations. With these developments increased monitoring of nutritional physiology in field populations of harmful algae will allow us to better discriminate how eutrophication impacts these groups.     

Patterns of environmental variability influence coral‐associated bacterial and algal communities on the Mesoamerican Barrier Reef

A coral's capacity to alter its microbial symbionts may enhance its fitness in the face of climate change. Recent work predicts exposure to high environmental variability may increase coral resilience and adaptability to future climate conditions. However, how this heightened environmental variability impacts coral‐associated microbial communities remains largely unexplored. Here, we examined the bacterial and algal symbionts associated with two coral species of the genus Siderastrea with distinct life history strategies from three reef sites on the Belize Mesoamerican Barrier Reef System with low or high environmental variability. Our results reveal bacterial community structure, as well as alpha‐ and beta‐diversity patterns, vary by host species. Differences in bacterial communities between host species were partially explained by high abundance of Deltaproteobacteria and Rhodospirillales and high bacterial diversity in Siderastrea radians. Our findings also suggest Siderastrea spp. have dynamic core bacterial communities that likely drive differences observed in the entire bacterial community, which may play a critical role in rapid acclimatization to environmental change. Unlike the bacterial community, Symbiodiniaceae composition was only distinct between host species at high thermal variability sites, suggesting that different factors shape bacterial versus algal communities within the coral holobiont. Our findings shed light on how domain‐specific shifts in dynamic microbiomes may allow for unique methods of enhanced host fitness.     

Viral control of phytoplankton populations--a review

Phytoplankton population dynamics are the result of imbalances between reproduction and losses. Losses include grazing, sinking, and natural mortality. As the importance of microbes in aquatic ecology has been recognized, so has the potential significance of viruses as mortality agents for phytoplankton. The field of algal virus ecology is steadily changing and advancing as new viruses are isolated and new methods are developed for quantifying the impact of viruses on phytoplankton dynamics and diversity. With this development, evidence is accumulating that viruses can control phytoplankton dynamics through reduction of host populations, or by preventing algal host populations from reaching high levels. The identification of highly specific host ranges of viruses is changing our understanding of population dynamics. Viral-mediated mortality may not only affect algal species succession, but may also affect intraspecies succession. Through cellular lysis, viruses indirectly affect the fluxes of energy, nutrients, and organic matter, especially during algal bloom events when biomass is high. Although the importance of viruses is presently recognized, it is apparent that many aspects of viral-mediated mortality of phytoplankton are still poorly understood. It is imperative that future research addresses the mechanisms that regulate virus infectivity, host resistance, genotype richness, abundance, and the fate of viruses over time and space.     

PATTERNS IN HOST RANGE FOR TWO STRAINS OF AMOEBOPHRYA (DINOPHYTA) INFECTING THECATE DINOFLAGELLATES: AMOEBOPHYRA SPP. EX ALEXANDRIUM AFFINE AND EX GONYAULAX POLYGRAMMA1

The endoparasitic dinoflagellate Amoebophrya ceratii (Koeppen) Cachon uses a number of its free‐living relatives as hosts and may represent a species complex composed of several host‐specific parasites. Two thecate host–parasite systems [Amoebophrya spp. ex Alexandrium affine (Inoue and Fukuyo) Balech and ex Gonyaulax polygramma Stein], were used to test the hypothesis that two strains of Amoebophrya have a high degree of host specificity. To test this hypothesis, a series of cross‐infection experiments were conducted, with 10 thecate and three athecate dinoflagellate species as potential hosts. Surprisingly, the two strains of Amoebophrya lacked host specificity and had wider host ranges than previously recognized. Among the host species tested, Amoebophrya sp. ex Alexandrium affine was capable of infecting only species of genus Alexandrium (Alexandrium affine, Alexandrium catenella, and Alexandrium tamarense), while the parasite from Gonyaulax polygramma infected species covering five genera (Alexandrium, Gonyaulax, Prorocentrum, Heterocapsa, and Scripsiella). In the context of previous reports, these results suggest that host specificity of Amoebophrya strains varies from extremely species‐specific to rather unspecific, with specificity being stronger for strains isolated from athecate hosts. Information on host specificity of Amoebophrya strains provided here will be helpful in assessing the possibility of using these parasites as biological control agents for harmful algal blooms, as well as in defining species of Amoebophrya in the future.     

Green paramecia as an evolutionary winner of oxidative symbiosis: a hypothesis and supportive data

A single cell of the green paramecia (Paramecium bursaria) harbors several hundreds of endo-symbiotic Chlorella-like algae in its cytoplasm. Removal of algae from the host organism and re-association of ex-symbiotic host paramecia with ex-symbiotic algae can be experimentally demonstrated in the laboratory. However, the mechanism precisely governing the alga-protozoan association is not fully understood, and the origin of symbiosis in the evolutionary view has not been given. Here, we propose the possible biochemical models (models 1 and 2) explaining the co-evolution between Paramecium species and algal symbionts by pointing out that algal photosynthesis in the host paramecia plays a dual role providing the energy source and the risk of oxidative damage to the host. Model 1 lays stress on the correlation between the (re)greening ability of the paramecia and the tolerance to oxidative stress whereas model 2 emphasizes the cause of evolutionary selection leading to the emergence of Paramecium species tolerant against reactive oxygen species.     

Epibacterial community patterns on marine macroalgae are host-specific but temporally variable

Marine macroalgae are constantly exposed to epibacterial colonizers. The epiphytic bacterial patterns and their temporal and spatial variability on host algae are poorly understood. To investigate the interaction between marine macroalgae and epiphytic bacteria, this study tested if the composition of epibacterial communities on different macroalgae was specific and persisted under varying biotic and abiotic environmental conditions over a 2-year observation time frame. Epibacterial communities on the co-occurring macroalgae Fucus vesiculosus, Gracilaria vermiculophylla and Ulva intestinalis were repeatedly sampled in summer and winter of 2007 and 2008. The epibacterial community composition was analysed by denaturing gradient gel electrophoresis (DGGE) and 16S rRNA gene libraries. Epibacterial community profiles did not only differ significantly at each sampling interval among algal species, but also showed consistent seasonal differences on each algal species at a bacterial phylum level. These compositional patterns re-occurred at the same season of two consecutive years. Within replicates of the same algal species, the composition of bacterial phyla was subject to shifts at the bacterial species level, both within the same season but at different years and between different seasons. However, 7-16% of sequences were identified as species specific to the host alga. These findings demonstrate that marine macroalgae harbour species-specific and temporally adapted epiphytic bacterial biofilms on their surfaces. Since several algal host-specific bacteria were highly similar to other bacteria known to either avoid subsequent colonization by eukaryotic larvae or to exhibit potent antibacterial activities, algal host-specific bacterial associations are expected to play an important role for marine macroalgae.     

Fate of Parasite and Host Organelle DNA during Cellular Transformation of Red Algae by Their Parasites

The transfer of a nucleus into a cytoplasm of a genetically foreign cell and its subsequent multiplication in the cytoplasm of this cell characterize most parasitic red algal species and their interactions with specific red algal hosts. Nuclei enter the host's cytoplasm upon cell fusion of parasite and host cell; here, they replicate, are spread to contiguous host cells, and ultimately are packaged into spores that reinfect other host thalli. In this study, we examined whether the proplastids and mitochondria that occur in these red algal adelphoparasites are acquired from their host or whether they are unique to the parasite and are brought into the host along with the parasite nucleus. To establish their origins and fates, plastid and mitochondrial restriction fragment length polymorphisms (RFLPs) of parasite cells were compared with those of their host plastid and mitochondrial DNA in three host and parasite pairs. For plastids, no RFLP differences were found between hosts and parasites, supporting an earlier conclusion, based on microscopic studies, that the proplastids of parasites are acquired from their hosts. For mitochondria, characteristic RFLP differences were detected between host and parasite for two of the pairs of species but not for the third. Evidence of the evolutionary difference between hosts and their parasites was shown by RFLP differences between nuclear ribosomal repeat regions.     

The importance of shore height and host identity for amphipod assemblages

The spatial distribution of organisms associated with marine intertidal macroalgae may be a direct result of their tolerance to air exposure or an indirect consequence of the distribution of their host. We compared amphipod assemblages from five intertidal macroalgae to investigate their relationship with algal identity. To test the effect of height regardless of algal characteristics, we transplanted coralline algal turfs to three different levels within the intertidal zone and compared amphipod assemblages after 1 and 14 days. Interstitial volume was positively correlated to the abundance of amphipods, suggesting that this attribute may correspond better to the potential space for their occupation when compared to algal biomass, thallus volume or the ratio between thallus and interstitial volume. Algal level determined the structure of the amphipod assemblages. Upper-level (Acanthophora spicifera and Caulerpa racemosa) and intermediate-level (coralline) algae host similar amphipod assemblages dominated by Apohyale media, but different from lower-level algae (Padina gymnospora and Sargassum cymosum), which were dominated by Hyale niger. Ten of the 15 amphipod species reported from natural communities were found in the transplanted plots. Distinct pools of amphipod species colonized coralline transplants at upper and lower levels after 1 day. However, regardless of the position on the shore, transplanted coralline turfs supported similar assemblages after 14 days, indicating that algal identity is also important for species assemblages. Our results suggest that both height on the shore and host identity combine to determine the vertical structure of amphipod assemblages in the rocky intertidal.     

Cryptic diversity hides host and habitat specialization in a gorgonian‐algal symbiosis

Shallow water anthozoans, the major builders of modern coral reefs, enhance their metabolic and calcification rates with algal symbionts. Controversy exists over whether these anthozoan–algae associations are flexible over the lifetimes of individual hosts, promoting acclimative plasticity, or are closely linked, such that hosts and symbionts co‐evolve across generations. Given the diversity of algal symbionts and the morphological plasticity of many host species, cryptic variation within either partner could potentially confound studies of anthozoan‐algal associations. Here, we used ribosomal, organelle and nuclear sequences, along with microsatellite variation, to study the relationship between lineages of a common Caribbean gorgonian and its algal symbionts. The gorgonian Eunicea flexuosa is a broadcast spawner, composed of two recently diverged, genetically distinct lineages largely segregated by depth. We sampled colonies of the two lineages across depth gradients at three Caribbean locations. We find that each host lineage is associated with a unique Symbiodinium B1/184 phylotype. This relationship between host and symbiont is maintained when host colonies are reciprocally transplanted, although cases of within phylotype switching were also observed. Even when the phylotypes of both partners are present at intermediate depths, the specificity between host and symbiont lineages remained absolute. Unrecognized cryptic diversity may mask host‐symbiont specificity and change the inference of evolutionary processes in mutualistic associations. Symbiotic specificity thus likely contributes to the ecological divergence of the two partners, generating species diversity within coral reefs.     

Effect of Japanese Paramecium bursaria extract on photosynthetic carbon fixation of symbiotic algae

To investigate the relationship between the Japanese Paramecium bursaria host and its symbiont, we studied the effect of a host cell-free extract on carbon fixation and photosynthate release of the symbiont. The host extract enhanced symbiotic algal carbon fixation about 3-fold at an increased concentration; however, release of photosynthate hardly changed. Since the enhancing effect was not affected by elimination of carbon dioxide from the host extract, the existence of a host factor that stimulates algal carbon fixation was made clear. The host factor is a heat-stable, low molecular weight substance. In relation to the pH dependence, the extract improved carbon fixation at acidic and neutral pH and showed almost no effect at pH 9.0. Therefore, the stimulation of carbon fixation by the host factor is unlikely to be caused by intracellular pH change. The extract also improved carbon fixation of several Chlorella species, symbiotic and free-living, and apparently exhibited no species specificity. Therefore, the host seems to regulate the photosynthesis of the symbiont via a specific compound.     

THE EVOLUTION OF PARASITES FROM THEIR HOSTS: A CASE STUDY IN THE PARASITIC RED ALGAE

Morphological similarities of many parasites and their hosts have led to speculation that some groups of plant, animal, fungal, and algal parasites may have evolved directly from their hosts. These parasites, which have been termed adelphoparasites in the botanical literature, and more recently, agastoparasites in the insect literature, may evolve monophyletically from one host and radiate secondarily to other hosts or, these parasites may arise polyphyletically, each arising from its own host. In this study we compare the internal transcribed spacer regions of the nuclear ribosomal repeats of species and formae specialis (host races) included in the red algal parasite genus Asterocolax with its hosts, which all belong to the Phycodrys group of the Delesseriaceae and with closely related nonhost taxa of the Delesseriaceae. These analyses reveal that species of Asterocolax have evolved polyphyletically. Asterocolax erythroglossi from the North Atlantic host Erythroglossum laciniatum appears to have evolved from its host, whereas taxa included in the north Pacific species Asterocolax gardneri have had two independent origins. Asterocolax gardneri from the host Polyneura latissima probably arose directly from this host. In contrast, all other A. gardneri formae specialis appear to have originated from either Phycodrys setchellii or P. isabelliae and radiated secondarily onto other closely related taxa of the Phycodrys group, including Nienburgia andersoniana and Anisocladella pacifica. Gamete crossing experiments confirm that A. gardneri from each host is genetically isolated from both its host, and from other A. gardneri and their hosts. Cross-infection experiments reveal that A. gardneri develops normally only on its natural host, although some abberrant growth may occur on alternate hosts. The ability of red algal parasites to radiate secondarily to other red algal taxa, where they may become isolated genetically and speciate, suggests that this process of speciation is not a “genetic dead end” but one that may give rise to related clusters of parasite species.     

Development of the red algal parasite Vertebrata aterrimophila sp. nov. (Rhodomelaceae,Ceramiales) from New Zealand

Parasitic red algae grow only on other red algae and have over 120 described species. Developmental studies in red algal parasites are few, although they have shown that secondary pit connections formed between parasite and host and proposed that this was an important process in successful parasitism. Furthermore, it was recorded that the transfer of parasite nuclei by these secondary pit connections led to different host cell effects. We used developmental studies to reconstruct early stages and any host cell effects of a parasite on Vertebrata aterrima. A mitochondrial marker (cox1) and morphological observations (light and fluorescence microscopy) were used to describe this new red algal parasite as Vertebrata aterrimophila sp. nov. Early developmental stages show that a parasite spore connects via secondary pit connections with a pericentral host cell after cuticle penetration. Developmental observations revealed a unique connection cell that grows into a ‘trunk-like’ structure. Host cell transformation after infection by the parasite included apparent increases in both carbohydrate concentrations and nuclear size, as well as structural changes. Analyses of molecular phylogenies and reproductive structures indicated that the closest relative of V. aterrimophila is its host, V. aterrima. Our study shows a novel developmental parasite stage (‘trunk-like’ cell) and highlights the need for further developmental studies to investigate the range of developmental patterns and host effects in parasitic red algae.     

Host-parasite interaction of blood-feeding dipterans in health and productivity of mammals

For simplicity of study and quantification, parasitism is generally considered as an individual relationship between a species and its host. Systematic analysis of host-parasite interaction in models for the blood-feeding dipterous parasite complex demonstrates that a large number of species influence common behavioural and physiological functions in the host. Such systems are complicated, however, by inter-dependence of both graded and quantal types of host response and by mediation of the interdependence within a hierarchical pattern of animalbehaviour. These systems are also subject to host-parasite regulation at population as well as physiological levels. Therefore, impacts of parasitism are not directly related to numbers of parasites or to absolute measures of infestation even in the extremes of acute parasitic disease, morbidity, or death associated with some species.A significant implication of these systems in host-parasite relations is that the influence of a single species of parasite on host productivity is modified significantly by its interrelation and common mode of interaction with other parasitic species of the same complex in the same host environment. As a result of this relationship, the physiologically adapted species at the right extreme of host-parasite interaction have tended to be underestimated in their impact on health and productivity of the host, particularly in terms of suboptimum levels of productivity. A major problem in parasite management within the blood- feeding dipterous parasite complex is assessment of the economic importance of numerous combinations of species that are mutually reinforcing in their impact on the health and productivity of the host.