Species associations among larval helminths in an amphipod intermediate host

Larval helminths that share the same intermediate host may or may not also share the same definitive hosts. If one or more of these helminth species can manipulate the phenotype of the intermediate host, there can be great advantages or severe costs for other helminths resulting from co-occurring with a manipulator, depending on whether they have the same definitive host or not. Among 2372 specimens of the amphipod Echinogammarus stammeri collected from the river Brenta, northern Italy, there was a positive association between two acanthocephalan species with the same fish definitive hosts, the relatively common Pomphorhynchus laevis and the much less prevalent Acanthocephalus clavula.The number of cystacanths of P. laevis per infected amphipod, which ranged from one to five, did not influence the likelihood that the amphipod would also host A. clavula. A third acanthocephalan species, Polymorphus minutus,which matures in birds, showed no association with either of the two other species. These results show that associations among helminth species in intermediate hosts are not random, and are instead the product of selection favouring certain pathways of transmission.     

Intra- and interspecific competition among helminth parasites: effects on Coitocaecum parvum life history strategy, size and fecundity

Larval helminths often share intermediate hosts with other individuals of the same or different species. Competition for resources and/or conflicts over transmission routes are likely to influence both the association patterns between species and the life history strategies of each individual. Parasites sharing common intermediate hosts may have evolved ways to avoid or associate with other species depending on their definitive host. If not, individual parasites could develop alternative life history strategies in response to association with particular species. Three sympatric species of helminths exploit the amphipod Paracalliope fluviatilis as an intermediate host in New Zealand: the acanthocephalan Acanthocephalus galaxii, the trematode Microphallus sp. and the progenetic trematode Coitocaecum parvum. Adult A. galaxii and C. parvum are both fish parasites whereas Microphallus sp. infects birds. We found no association, either positive or negative, among the three parasite species. The effects of intra- and interspecific interactions were also measured in the trematode C. parvum. Both intra- and interspecific competition seemed to affect both the life history strategy and the size and fecundity of C. parvum. Firstly, the proportion of progenesis was higher in metacercariae sharing their host with Microphallus sp., the bird parasite, than in any other situation. Second, the intensity of intraspecific competition apparently constrained the ability of metacercariae to adopt progenesis and limited both the growth and egg production of progenetic individuals. These results show that the life history strategy adopted by a parasite may be influenced by other parasites sharing the same host.     

Fitness costs of intrinsic competition in two egg parasitoids of a true bug

Intrinsic competition in insect parasitoids occurs when supernumerary larvae develop in the same host as consequence of multiple ovipositions by females of the same species (intra-specific competition) or by females of different species (inter-specific competition). Studies on intrinsic competition have mainly focused on understanding the factors that play a role in the outcome of competition, while fitness-related effects for the parasitoid surviving the competition have been poorly investigated, especially in egg parasitoids. Interestingly, even the winning parasitoid can experience fitness costs due to larval development in a host in which multiple factors have been injected by the ovipositing females or released by their larvae. In this paper we studied fitness-related traits associated with intra- and inter-specific competition between Trissolcus basalis (Wollaston) and Ooencyrtus telenomicida (Vassiliev), the main egg parasitoids associated with the southern green stink bug Nezara viridula (L.) in Italy. We investigated the impact of intrinsic competition for the surviving parasitoid in terms of body size, developmental time, number and size of oocytes. Our results indicated that T. basalis adults did not experience fitness-related costs when surviving intra-specific competition; however, adults were smaller, took longer to develop and females produced fewer oocytes after surviving inter-specific competition. A different outcome was found for O. telenomicida where the emerging females were smaller, produced fewer and smaller oocytes when suffering intra-specific competition whereas no fitness costs were found when adults survived inter-specific competition. These results support the hypothesis that the impact of intrinsic competition in egg parasitoids depends on the severity of the competitive interaction, as fitness costs were more pronounced when the surviving parasitoid interacted with the most detrimental competitor.     

Do different parasite species interact in their effects on host fitness? A case study on parasites of the amphipod Paracalliope fluviatilis

There is a gap in our understanding of the relative and interactive effects of different parasite species on the same host population. Here we examine the effects of the acanthocephalan Acanthocephalus galaxii, an unidentified cyclophyllidean cestode, and the trematodes Coitocaecum parvum and Microphallus sp. on several fitness components of the amphipod Paracalliope fluviatilis, using a combination of infection surveys and both survival and behavioural trials. In addition to significant relationships between specific parasites and measures of amphipod survival, maturity, mating success and behaviour, interactions between parasite species with respect to amphipod photophilia were also significant. While infection by either A. galaxii or C. parvum was associated with increased photophilia, such increases were negated by co-infection with Microphallus sp. We hypothesize that this is due to the more subtle manipulative effect of A. galaxii and C. parvum being impaired by Microphallus sp. We conclude that the low frequency at which such double infections occur in our sampled population means that such interactions are unlikely to be important beyond the scale of the host individual. Whether or not this is generally true, implying that parasitological models and theory based on single parasite species studies do generally hold, requires cross-species meta-analytical studies.     

Patterns of intermediate host use and levels of association between two conflicting manipulative parasites.

For many parasites with complex life cycles, manipulation of intermediate host phenotypes is often regarded as an adaptation to increase the probability of successful transmission. This phenomenon creates opportunities for either synergistic or conflicting interests between different parasite species sharing the same intermediate host. When more than one manipulative parasite infect the same intermediate host, but differ in their definitive host, selection should favour the establishment of a negative association between these manipulators. Both Polymorphus minutus and Pomphorhynchus laevis exploit the amphipod Gammarus pulex as intermediate host but differ markedly in their final host, a fish for P. laevis and a bird for P. minutus. The pattern of host use by these two conflicting manipulative parasites was studied. Their incidence and intensity of infection and their distribution among G. pulex were first examined by analysing three large samples of gammarids collected from the river Tille, Eastern France. Both parasites had low prevalence in the host population. However, temporal fluctuation in the level of parasitic infection was observed. Overall, prevalence of both parasite species was higher in male than in female G. pulex. We then assessed the degree of association between the two parasites among their intermediate hosts, using two different methods: a host-centred measure and a parasite-centred measure. Both measures gave similar results; showing random association between the two acanthocephalan species in their intermediate hosts. We discuss our results in relation to the selective forces and ecological constraints that may determine the pattern of association between conflicting manipulative parasites.     

Differential influence of Pomphorhynchus laevis (Acanthocephala) on the behaviour of native and invader gammarid species.

Although various species of acanthocephalan parasites can increase the vulnerability of their amphipod intermediate hosts to predation, particularly by altering their photophobic behaviour, their influence on the structure of amphipod communities and the success of invader species has so far received little attention. We compared the prevalence and behavioural influence of a fish acanthocephalan parasite, Pomphorhynchus laevis, in two species of amphipods, Gammarus pulex and Gammarus roeseli in sympatry in the river Ouche (Burgundy, eastern France). There, G. pulex is a resident species, whereas G. roeseli is a recent coloniser. Both uninfected G. pulex and G. roeseli were strongly photophobic, although less so in the invading species. However, there was no significant difference in reaction to light between infected and uninfected G. roeseli, whereas infected G. pulex were strongly photophilic. We discuss our results in relation to the parasite's ability to manipulate invading host species, the possibility that resistant individuals have been selected during the invasion process, and the role that acanthocephalan parasites can play in shaping the structure of amphipod communities.     

高阶相互作用对宿主-寄生群落动态的影响

物种间相互作用是影响生物群落稳定性和多样性的重要因素。基于Lotka-Volterra竞争模型,通过构建多宿主种群的种内和种间高阶相互作用模型,研究宿主种群的间接竞争效应对寄生群落动态的影响机制。为有效地揭示高阶作用对种群动态的影响,通过对比宿主-寄生群落的现象模型以及机制模型,利用机制模型产生的合理数据集对现象模型中高阶项的参数进行拟合,进而探讨了高阶相互作用在群落动态中的作用。结果显示,完整的高阶相互作用模型在描述多宿主-寄生系统的群落动态中表现最优,而直接相互作用模型对群落动态的描述相对较差,即同时考虑种间和种内的高阶相互作用模型更加符合机制模型所描述的群落动态。此外,种内高阶作用和种间高阶作用产生不对称效应,宿主间的种间高阶作用对群落产生的影响较种内高阶作用更为显著。该研究结果在一定意义上丰富了宿主-寄生生物群落的稳定性研究,为理解物种间相互作用的多样性研究提供了依据。     

Differential influence of Pomphorhynchus laevis (Acanthocephala) on the behaviour of native and invader gammarid species

Although various species of acanthocephalan parasites can increase the vulnerability of their amphipod intermediate hosts to predation, particularly by altering their photophobic behaviour, their influence on the structure of amphipod communities and the success of invader species has so far received little attention. We compared the prevalence and behavioural influence of a fish acanthocephalan parasite, Pomphorhynchus laevis, in two species of amphipods, Gammarus pulex and Gammarus roeseli in sympatry in the river Ouche (Burgundy, eastern France). There, G. pulex is a resident species, whereas G. roeseli is a recent coloniser. Both uninfected G. pulex and G. roeseli were strongly photophobic, although less so in the invading species. However, there was no significant difference in reaction to light between infected and uninfected G. roeseli, whereas infected G. pulex were strongly photophilic. We discuss our results in relation to the parasite's ability to manipulate invading host species, the possibility that resistant individuals have been selected during the invasion process, and the role that acanthocephalan parasites can play in shaping the structure of amphipod communities.     

Spiracanthus bovichthys n. gen. n. sp. (Acanthocephala: Arhythmacanthidae), a parasite of littoral fishes of the central south coast of Chile

A new acanthocephalan species, Spiracanthus bovichthys n. gen. n. sp., is described. Samples were taken from 26 marine fish species between 23 and 53 degrees S of Chile. The parasite was found in the intestine of 6 species and only between 36 and 40 degrees S, especially in those fish that prey on small crustaceans in the upper and sublittoral zone. The parasite was found in Bovichthys chilensis (Reagan). Auchenionchus variolosus (Valenciennes), Calliclinus genigutattus (Valenciennes), Sindoscopus australis (Fowler and Bean), Myxodes cristatus Valenciennes, and Gobiesox marmoratus (Jenyns). However, only the first species is a suitable host for the parasite as evidenced by the presence of mature females. This acanthocephalan belongs to Arhythmacanthidae, but it does not correspond to any genus described. Spiracanthus bovichthys is different in the number and spiralled distribution of its hooks in the proboscis. In the short proboscis, 3 groups of hooks are distinguished according to size, summing up to 150-190 hooks. The group of largest hooks are found in the apical part of the proboscis, and there are 10 diagonal rows of small hooks from the prebasal to basal proboscis. Its trunk is covered partially by small spines. Amphipod and isopods were the prey items shared among the host fish species and are the best candidates to be the intermediate hosts of S. bovichthys.     

中亚热带濒危植物毛红椿和南方红豆杉种内与种间竞争差异

濒危植物是全球生物多样性的重要组成部分,为了弄清中亚热带地区不同生长型濒危乔木树种的竞争模式及其机制,研究了江西九岭山国家级自然保护区毛红椿和南方红豆杉的种内与种间竞争差异。结果表明: 阳性落叶树种毛红椿种群的竞争压力主要来自于同种个体,种内竞争指数占总竞争指数的66.4%,耐阴常绿树种南方红豆杉种群主要受种间竞争的影响,种间竞争指数占总竞争指数的68.7%。2种濒危植物受到的种内、种间以及总竞争强度均随着林木径级的增大而逐渐减小,表明竞争压力由幼林向成熟林逐渐降低。在生长过程中,毛红椿以种内竞争引起的自疏作用为主导,南方红豆杉则由种间竞争引起的他疏作用为主导。毛红椿和南方红豆杉种群在更新过程中,小径级个体都要经历强度较大的种内或种间竞争选择才能发育为成树。由于主要竞争压力的来源差异,对于生活在中亚热带同一区域内不同生长型的濒危植物应该采取不同的生物多样性保育措施。     

择伐对思茅松天然林乔木种间与种内关系的影响

通过分析不同择伐强度下思茅松(Pinus kesiya)天然林乔木种群的生态位、种间联结、种内与种间竞争和群落稳定性, 探讨干扰强度对思茅松天然林乔木种群种间与种内关系的影响。结果表明: (1)择伐干扰改变了大多数乔木树种在群落中的优势地位, 思茅松优势地位未受影响, 随着择伐干扰强度的增加, 思茅松在群落中的生态位宽度逐渐变大, 在重度干扰的群落中与红木荷(Schima wallichii)一起成为生态幅度最大的树种。(2)通过种间联结与生态位重叠分析发现, 随着干扰强度的加剧, 乔木种群的种间关系发生变化, 中度干扰群落中物种之间的生态位重叠明显高于未干扰和重度干扰群落, 而在三类群落中, 思茅松与其他种群之间的重叠值较小。随着干扰强度增加, 种群间正联结与负联结的种对所占的比例逐渐增加。(3) Hegyi竞争指数表明, 三类群落中思茅松种群的竞争压力主要来自种间, 而干扰强度进一步增加了思茅松种群的种间竞争强度, 其种内平均竞争指数随着对象木胸径的增加而减少。(4)林窗是思茅松种群种间和种内关系变化的主要原因, 三类群落都处于较不稳定的阶段, 但中度干扰群落的物种丰富度与稳定性都高于未干扰和重度干扰的群落。择伐原则依据森林管理的目的而确定, 如以木材培育为主或森林生态系统服务功能为主。     

Differential influence of Pomphorhynchus laevis (Acanthocephala) on brain serotonergic activity in two congeneric host species

The physiological mechanisms by which parasites with complex life cycles manipulate the behaviour of their intermediate hosts are still poorly understood. In Burgundy, eastern France, the acanthocephalan parasite Pomphorhynchus laevis inverses reaction to light in its amphipod host Gammarus pulex, but not in Gammarus roeseli, a recent invasive species. Here, we show that this difference in manipulation actually reflects a difference in the ability of the parasite to alter brain serotonergic (5-HT) activity of the two host species. Injection of 5-HT in uninfected individuals of both host species was sufficient to inverse reaction to light. However, a difference in brain 5-HT immunocytochemical staining levels between infected and uninfected individuals was observed only in G. pulex. Local adaptation of the parasite to the local host species might explain its inability to manipulate the behaviour and nervous system of the invasive species.     

Parasite-induced suppression of aggregation under predation risk in a freshwater amphipod: Sociality of infected amphipods

Recent findings suggest that grouping with conspecifics is part of the behavioural defences developed by amphipod crustaceans to face predation risk by fish. Amphipods commonly serve as intermediate hosts for trophically transmitted parasites. These parasites are known for their ability to alter intermediate host phenotype in a way that promotes predation by definitive hosts, where they reproduce. If aggregation in amphipods dilutes the risk to be preyed on by fish, then it may dilute the probability of transmission for the parasite using fish as definitive hosts. Using experimental infections, we tested whether infection with the fish acanthocephalan Pomphorhynchus laevis alters attraction to conspecifics in the amphipod intermediate host Gammarus pulex. We also measured G. pulex's activity and reaction to light to detect potential links between changes in aggregation and changes in other behaviours. The attraction to conspecifics in the presence of predator cue, a behaviour found in uninfected gammarids, was cancelled by the infection, while phototaxis was reversed and activity unchanged. We found no correlation between the three behaviours in infected amphipods, while activity and aggregation were negatively correlated in uninfected individuals after the detection of predation cue. The physiological causes and the adaptive value of aggregation suppression are discussed in the context of a multidimensional manipulation.     

Larval helminths in intermediate hosts: does competition early in life determine the fitness of adult parasites?

Density-dependent effects on parasite fitness have been documented from adult helminths in their definitive hosts. There have, however, been no studies on the cost of sharing an intermediate host with other parasites in terms of reduced adult parasite fecundity. Even if larval parasites suffer a reduction in size, caused by crowding, virtually nothing is known about longer-lasting effects after transmission to the definitive host. This study is the first to use in vitro cultivation with feeding of adult trematodes to investigate how numbers of parasites in the intermediate host affect the size and fecundity of adult parasites. For this purpose, we examined two different infracommunities of parasites in crustacean hosts. Firstly, we used experimental infections of Maritrema novaezealandensis in the amphipod, Paracalliope novizealandiae, to investigate potential density-dependent effects in single-species infections. Secondly, we used the crab, Macrophthalmus hirtipes (Ocypodidae), naturally infected by the trematodes, M. novaezealandensis and Levinseniella sp., the acanthocephalan, Profilicollis spp., and an acuariid nematode. These four helminths all develop and grow in their crustacean host before transmission to their bird definitive host by predation. In experimental infections, we found an intensity-dependent establishment success, with a decrease in the success rate of cercariae developing into infective metacercariae with an increasing dose of cercariae applied to each amphipod. In natural infections, we found that M. novaezealandensis-metacercariae achieved a smaller volume, on average, when infrapopulations of this parasite were large. Small metacercariae produced small in vitro-adult worms, which in turn produced fewer eggs. Crowding effects in the intermediate host thus were expressed at the adult stage in spite of the worms being cultured in a nutrient-rich medium. Furthermore, excystment success and egg-production in M. novaezealandensis in naturally infected crabs were influenced by the number of co-occurring Profilicollis cystacanths, indicating interspecific interactions between the two species. Our results thus indicate that the infracommunity of larval helminths in their intermediate host is interactive and that any density-dependent effect in the intermediate host may have lasting effects on individual parasite fitness.     

Parasitic infection and developmental stability: fluctuating asymmetry in Gammarus pulex infected with two acanthocephalan species

Several studies have reported a negative association between developmental stability and parasitic infection. However, the host-parasite associations examined so far consist only of a limited number of parasite taxa, and developmental stability was appraised on definitive hosts. The present study examines the association between infection by 2 acanthocephalan parasites. Pomphorhynchus laevis and Polymorphus minutus, and the developmental stability of their common intermediate host Gammarus pulex. Developmental stability was estimated from the fluctuating asymmetry (FA) levels of 6 morphological traits. A positive association was found between FA and infection. Infected gammarids tended to be more asymmetrical than the noninfected ones for an index generated by combining FA scores from 2 characters out of the 6 studied, even though no significant relationships were found between FA levels and parasitic loads. The simultaneous presence of both acanthocephalan species in the same host seems to be associated with increased FA levels of gammarids, but this trend was not statistically significant. For the same characters, males exhibited higher levels of FA than females.     

Metazoan parasites of the thumb grenadier Nezumia pulchella, from the south-eastern Pacific, off Chile, and their use for discrimination of host populations

The parasite communities of Nezumia pulchella across its geographical range in the south-eastern Pacific were described, from 217 fish captured from four localities in northern and central Chile (24° S to 33° S). Five ectoparasites and 13 endoparasites were recorded and the highest prevalence and intensity of infection was found for the copepods Jusheyhoea macrura and Clavella sp. 1, for the monogenean Diclidophora sp. and for the larval acanthocephalan Corynosoma australe . Parasite species richness decreased with latitude. Multivariate discriminant analysis and correspondence analysis supported the use of parasites of N. pulchella to discriminate populations of this host, suggesting the existence of three well-defined host populations in the geographical range examined. Jusheyhoea macrura , C. australe , Lophoura sp., Diclidophora sp., Capillaria sp. and Proleptus sp. were the most important species for discriminant analysis.     

Nearest-neighbour analysis and the prevelance of woody plant competition in South African savannas

Inter- and intra-specific competition has long been regarded as asignificant determinant of the structure and function of woody plantcommunitiesin African savannas. The most widely used approach to detect the presence orabsence of competition has been the use of nearest-neighbour methods. Althoughseveral studies have been published reporting the presence of competition inAcacia dominated communities, less than 20 plots have beensampled across all these studies. Results from broad-leaved communities arevariable, and also based on a small sample number. Consequently, this studysought to assess the prevalence of competition from a large number of savannasites (45), and to identify abiotic and biotic factors characterising siteswithcompetition relative to those without. Using the nearest-neighbour method onlyfour sites (9.3%) indicated the presence of inter-specific competition; twowereAcacia dominated communities (18.2% ofAcacia sites) and two were broad-leaved communities(7.7%).These four sites had a significantly higher mean annual rainfall and fewerwoodyspecies than sites without inter-specific competition. There were alsosignificant relationships between the inter-specific competition index and thedominance index and the number of species at a site. In terms of intra-specificcompetition only ten of the 31 sites tested revealed a significant correlationbetween nearest-neighbour distance and summed canopy volume of the twoneighbours. Seven were for Acacia species (55.6% of theAcacia species comparisons) and three were forbroad-leavedspecies (21.4% of the broad-leaved species comparisons). Sites lacking evidenceof intra-specific competition were at a significantly lower slope position andhad a smaller proportion of small stems. The lower prevalence of competition inSouth African savannas detected in this study using the same method as previousstudies is discussed, and the appropriateness of nearest-neighbour analysis isexamined.     

Metechinorhynchus salmonis infection and diet in the river-spawning whitefish of the Bothnian Bay

A total of 206 river-spawning whitefish Coregonus lavaretus (L.) from the Bothnian Bay was investigated for food objects and acanthocephalans in the years 1975–78. The only acanthocephalan species found was Metechinorhynchus salmonis (Müller 1780), the total number of which was 1295.
The whitefish is shown to be highly susceptible to infection by M. salmonis , with as many as 88·9% of specimens infected in a sample in June 1978, although the mean intensity was only 8·9 parasites per host. The river-spawning whitefish does not favour the intermediate host, Pontoporeia affinis , in its summer diet in the Bothnian Bay preferring to feed on eggs and molluscs, so that the incidence of M. salmonis infection decreases to 33·3% by the middle of summer. This is further reduced in the late summer by the fast associated with the migration to their upriver spawning grounds. The fact that the other acanthocephalan inhabiting the river-spawning whitefish in the southern Gulf of Bothnia, Echinorhynchus gadi Müller, 1776 is not to be found in the Bothnian Bay population suggests that the fish captured in spring have remained in the Bothnian Bay throughout the winter.     

灰飞虱与褐飞虱种内和种间密度效应比较

在人工气候室内比较研究了灰飞虱和褐飞虱在汕优63和秀水11上的种内和种间密度效应对其主要生物学参数的影响,结果表明两种稻飞虱均存在显著的种内竞争,主要表现为随着密度增加,若虫发育历期延长、若虫羽化率下降、雌成虫寿命缩短、每雌产卵量下降。种内密度效应与稻飞虱种类和寄主品种有显著互作关系,灰飞虱种内竞争较褐飞虱明显,二者在适宜寄主上种内竞争更为明显。同时,两种稻飞虱存在着显著的种间密度效应,主要表现为异种存在时的促进作用,即异种共存时若虫历期缩短、若虫羽化率提高、雌虫寿命延长、每雌产卵量增加。种间密度效应也与稻飞虱种类和寄主品种有显著互作关系,两种共存对灰飞虱的有利作用显著大于对褐飞虱的有利作用,在欠适宜寄主上种间互利效应更为明显。最后,本文对稻飞虱种内和种间密度效应的机制和进化意义进行了讨论。     

Seasonal and regional occurrence of <Emphasis Type="Italic">Acanthocephalus</Emphasis> sp. (Acanthocephala: Echinorhynchidae) in fishes and isopods (<Emphasis Type="Italic">Asellus hilgendorfi</Emphasis>) in a lake system in northern Japan

 The echinorhynchid acanthocephalan Acanthocephalus sp. was collected and described from four species of fishes (rainbow trout Oncorhynchus mykiss, Sakhalin huchen Hucho perryi, Japanese pond smelt Hypomesus nipponensis, and threespine stickleback Gasterosteus aculeatus) from a lake system, the Tsugaru-Jūniko Lakes, in Aomori Prefecture, northern Japan. In rainbow trout, the prevalence and intensity of infection markedly differed between lakes, and the fish were most frequently and most heavily infected in the lakes with a dense population of the isopod intermediate host Asellus hilgendorfi. In isopods, the prevalence of acanthocephalan larvae increased in the late winter and reached its highest level in March or April. In rainbow trout, male worms were abundant from winter to spring, and female worms were immature during these seasons. Gravid females were abundant in summer and autumn. These findings indicate that Acanthocephalus sp. is an annual species and its recruitment from the intermediate host to the fish occurs mainly in winter and spring. Received: January 9, 2002 / Accepted: April 18, 2002 Acknowledgments We thank Professor Shōichi Saito, Faculty of Education, Hirosaki University, for his encouragement during this study. Thanks are also due to many students of the Nature Study Laboratory, Faculty of Education, Hirosaki University, for their assistance in the field. We are greatly indebted to the Iwasaki Village Office and Fukaura Forestry Office for giving us permission for the survey. Correspondence to:A. Ohtaka