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1.
Parasitism is hypothesized to reduce reproductive success in heavily parasitized males because females may preferentially mate with less parasitized males (parasite-mediated sexual selection) or parasites may compromise male competitiveness. In marine systems, this hypothesis is largely unexplored. This paper provides the first confirmed record of a copepod ectoparasite (Caligus buechlerae Hewitt 1964) on the common triplefin (Forsterygion lapillum) and evaluates the hypothesis that males parasitized with C. buechlerae experience lower reproductive success than unparasitized males (as determined by the presence and area of eggs within male nests). We found that 38 % of males we surveyed were infected with at least one C. buechlerae, with a median of two individuals per infected male. About 32 % of males were defending eggs, with 62.5 % of those males infected with at least one parasite. Males of greater total length (TL) were both more likely to be infected and more likely to be defending eggs. However, when statistically accounting for the effects of TL, parasite infection had no effect on the probability of defending eggs, or the average surface area of eggs when present. Positive covariation in fish length, the presence of eggs and parasite infection observed here potentially suggest that the importance of parasitic infection on reproductive success may depend upon the strength of selection for larger male body size. Our study is one of the few studies to investigate the effects of ectoparasites on reproductive success in reef fish and also provides a quantitative measure of infection for a widespread species within New Zealand.  相似文献   

2.
Insect social parasites, like other parasites, may benefit from inhibiting their host from reproducing (complete or partial parasitic castration) because they can then exploit more of the host’s resources for their own reproduction. In particular, social parasites that kill or expel the host queen need to prevent host workers from reproducing; this is a common worker response to the absence of their queen. Indeed, host workers would benefit from detecting the presence of the parasite and investing in direct and indirect fitness. Studying whether and how social parasites control host worker reproduction can provide information about the degree of integration of the parasite in the host colony and help identify factors regulating workers’ reproductive decisions in social insects. We investigated whether the paper wasp social parasite, Polistes sulcifer, suppresses Polistes dominula (host) worker reproduction as efficiently as the dominant host female does in queen-right colonies by comparing worker reproductive efforts in parasitized and non-parasitized (control) colonies. Our results show that 6 weeks after usurpation of their colony by the social parasite, parasitized workers (1) had more developed ovaries than control workers and (2) laid more eggs as soon as the opportunity arose. This reproductive readiness of parasitized workers was not apparent 2 weeks after colony usurpation. This suggests that P. dominula workers have evolved means to react to social parasitism, as occurs in some ants, and that the parasite has only limited control over host reproduction.  相似文献   

3.
Parasite-induced alterations in host behaviour have been reported in a large number of taxa. However, some parasites are better than others to exploit the resources offered by their hosts. To date, our understanding of the extent to which some obligate parasites exploit social insect colonies is still limited. In this study, we examined parasite-mediated behavioural alterations of Polistes biglumis wasps parasitized by the obligate social parasite Polistes atrimandibularis by comparing host female-activity in parasitized and non-parasitized colonies. Host foundresses foraged more and rested less in parasitized than in non-parasitized colonies (controlling for the number of larvae in the nest, the time of day, and the day in the season). Next, we used short-term parasite removal experiments to investigate how social parasites manipulate their hosts. We found that parasitized hosts foraged more and rested less when social parasites were on the nest rather than after their removal, and we tested which kind of interactions occurred between parasites and hosts. P. atrimandibularis parasites may use mainly non-aggressive interactions (such as antennation and trophallaxis) to manipulate host activities, rather than visual, acoustic or chemical signals as other parasites do.  相似文献   

4.
Summary The diet selected by three-spined sticklebacks (Gasterosteus aculeatus) depends on the degree of parasitization by one or both of two parasite species (Schistocephalus solidus, Glugea anomala). Uninfested fish prefer the more profitable of two different size classes of prey (Daphinia magna). Fish parasitized by Glugea or Schistocephalus attack both prey types equally often, whereas sticklebacks infested by both parasite species prefer to attack the less profitable prey. The diet selected is optimal under the condition that parasites decrease their host's competitive ability.  相似文献   

5.
Observations in the Indian Ocean near Shimoni (Kenya) have added the labride Thalassoma amblycephalus (Bleeker) to the list of fishes which regularly visit sea-anemones. If disturbed, the fish seeks protection near the column or under the oral disk of anemones. It normally visits the anemones spontaneously, inspects the tentacles and picks up small food particles caught by the anemones; this behavior characterizes T. amblycephalus as a food parasite. New observations, and the fact that T. amblycephalus is a labride and not a pomacentride like the species of Amphiprion, Dascyllus and Premnas-other fishes known to establish associations with anemones-throw new light on the ecological importance of anemone-fish associations.  相似文献   

6.
This study is to investigate the behavioural responses of zebrafish to the toxic effects of waterborne zinc and chromium(VI). Swimming performance of fish was tracked by a real-time toxicity bio-monitoring system and transformed into data to calculate individual and fish school behavioural endpoints. Behavioural responses under metals were in accordance with the Stepwise Behavioural Response model. Significant and concentration-dependent effects on swimming speed and turning times were found at concentrations equal to or higher than 3.5 mg L-1 for zinc and 0.7 mg L-1 for chromium. Swimming height significantly declined in concentrations equal to or higher than 3.5 mg L-1 under zinc exposure, while for the chromium treatment, it significantly increased in lower concentration exposures and decreased in higher concentration exposures. Average distance and dispersion of fish school decreased at the beginning but there were no obvious changes during the whole exposure period. Behavioural responses under metal exposures were time-dependent: avoidance (increased speed and turning times and decreased average distance and dispersion) might be the first response and followed by height changes. Changes in behavioural endpoints ‘swimming speed and height’ can be used as early stress responses for acute aquatic metal contamination as part of ecological risk assessment.  相似文献   

7.
D. Parker  A. J. Booth 《Marine Biology》2013,160(11):2943-2950
Cymothoa borbonica prevalence in the buccal cavity of Trachinotus botla was high, with 45 % of all fish sampled being infected. Smaller fish were more susceptible to infection with no parasites found in fish over 400 mm FL. The detrimental effects of parasite infection on their hosts include basihyal (the bone commonly known as the “tongue”) damage, a loss in buccal cavity volume as a result of female parasite attachment, and a severe impact on host growth. By combining short-term dietary analysis and medium-term stable isotope analysis, there was little evidence to suggest a modification in either the diet or feeding habits of infected fish where infected and uninfected fish occupied the same trophic niche. Inhibited growth in infected fish is hypothesized to be from respiratory distress from long-term oxygen deficiency through buccal obstruction.  相似文献   

8.
Chemical communication appears to be the primary mode of information transfer among woodland salamanders of the genus Plethodon. These pheromones have been shown to convey a wide variety of information, and are used to establish territorial boundaries. Responses to pheromones can include changes in both behavior and energy consumption. The ability of Ozark zigzag salamanders, Plethodon angusticlavius, to detect the sex of a conspecific through chemical substrate markings has not been assessed. In addition, some populations of P. angusticlavius are naturally infected with the ectoparasitic mite Hannemania eltoni. Parasitism can impact both the behavior and metabolism of the host as well as the behavior of others towards the parasitized individual. We examined whether sex and parasitism of the pheromone donor and receiver influenced behavioral and metabolic responses in P. angusticlavius. In laboratory trials, behavior of P. angusticlavius was influenced by the sex and parasite load of the pheromone donors and receivers. Behavioral responses were also influenced by whether the receiver was in its own territory or in the territory of the pheromone donor. Metabolic responses were not affected by the sex or parasite load of the pheromone donor, but males had higher metabolic rates than females, even after adjusting for differences in body size. Overall, our study indicates that both sex and parasitism can be detected via pheromones in this species, and the nature of the response depends on residency status, sex, and parasite load of the receiver.  相似文献   

9.

Background

Since the 1960s the scientific community has become increasingly aware of the need for integrative and holistic approaches to solve environmental issues. It has not only been recognized that changing environmental conditions can alter toxicological properties of pollutants but also that susceptibility of organisms to man-made chemicals strongly depends on environmental factors. This is especially true with respect to the endocrine control of reproductive processes in many aquatic species including fish, which are mainly influenced by changes in the surrounding environment such as day length, temperature, water flow, etc. Thus, it is astounding that the use of integrative approaches to address the question of the ecotoxicological relevance of the disruption of endocrine and related reproductive functions is rate. As a consequence of this shortcoming, there is increasing demand both among scientists and politicians regarding more holistic approaches in the endocrine disruptor research.

Aim

This contribution aims to critically discuss current strategies and approaches as well as to identify alternative strategies in context with the environmental risk assessment of endocrine disruptors. This will be exemplified using an ecotoxicological study with bream (Abramis brama) that was conducted along a major European river, the Elbe.

Results and Discussion

Local bream (Abramis brama) populations collected along the river Elbe were characterized by different magnitudes and/or types of endocrine alterations. Although there is a relatively extensive data set available regarding its contamination with wide range of chemicals in the Elbe, it has not been possible to conclusively relate the observed endocrine effects to the exposure to individual compounds or mixtures of chemicals. However, during the gross morphological evaluations of the fish it was observed that at some locations a great number of the individuals caught were parasitized with the tapeworm Ligula intestinalis. Furthermore, statistical analysis revealed that a relatively great percentage of the variation of the biological endpoints observed both among individuals and sampling sites could be explained by the occurrence of this parasite. Overall, the findings of these studies indicate that there is a complex interaction not only among the exposure to chemicals and parasitism but also of both of these aspects with the occurrence of biological effects in fish.

Conclusions

The gaps in our current knowledge as well as the findings from the few studies available indicate, the there is a need to revise our current approaches to characterize and evaluate exposure scenarios regarding their potential to cause endocrine disruption in wild organisms. Given the complexity of most environmental exposure scenarios, involving multiple factors including anthropogenic and natural influences and their interactions, I therefore see the urgent need to initiate a discussion on more integrative approaches in this field in favor of true ‘ecotoxicological studies’ that integrate both ecology and toxicology.  相似文献   

10.
The shiny cowbird (Molothrus bonariensis) is a generalist brood parasite that lays either white-immaculate or spotted egg morphs in eastern Argentina and Uruguay. Some hosts accept both morphs, others accept spotted eggs and reject the white morph, but no host has been found to accept white eggs and reject spotted ones. It has been suggested that the yellow-winged blackbird (Agelaius thilius) may be that type of host. The finding of a white acceptor-spotted rejector species would help to explain the occurrence and maintenance of the parasite egg polymorphism. We studied the incidence of shiny cowbird parasitism on this host, its costs for their reproductive success and the presence of antiparasitic defenses in the yellow-winged blackbird - shiny cowbird system. The parasite affected the reproductive success of the host in two ways. Cowbirds punctured host eggs causing a reduction in clutch size, and yellow-winged blackbirds deserted their nests whenever they suffered high egg loss. In addition, parasitized nests suffered higher predation during the nestling stage, but not during egg stages, indicating that the difference found was related to the presence of the cowbird chick, and not to higher exposure of parasitized nests to both parasites␣and predators. Despite the costs imposed by the parasite, yellow-winged blackbirds have not evolved antiparasitic defenses. This host did not reject any egg morph of the shiny cowbird nor desert parasitized nests unless it had suffered high egg loss. Current explanations for the host lack of defenses, the “time lag” and the “equilibrium” hypothesis, are discussed. Received: 29 August 1997 / Accepted after revision: 10 January 1998  相似文献   

11.
Macroalgal fields are a feature of the shallow tropical benthos, yet their importance for coral reef fish population dynamics remains poorly understood. The abundance of fish recruits was recorded using underwater visual census at six macroalgal and 11 coral reef sites in the Montebello and Barrow Islands. Surveys identified 6,935 individual recruit fish from 105 species, 54 genera and 20 families. Of these, 1,401 recruits from 48 species, 31 genera and 14 families were observed in macroalgal sites. Sixteen of the 105 recruit species (15.2 %) were observed exclusively at macroalgal sites. Forty-two (87.5 %) of these species have been observed as adults on adjacent coral reefs. Species composition of fish recruits differed significantly between the two habitats. Corallivore, small omnivore and zooplanktivore recruits had significantly higher numbers in the coral sites, while the results clearly demonstrate that juveniles, within the genera Lethrinus and Choerodon, as well as large algal croppers, are predominantly found at macroalgal (74–100 %) rather than coral-dominated sites. High-canopy macroalgae cover was positively correlated with abundance of these taxa, particularly Lethrinids (r 2 = 0.40). This study is the first to highlight the important attributes of tropical macroalgal fields and suggests that they have a similar role to seagrass meadows as essential juvenile habitat, thus warranting greater attention in conservation planning and ecological studies.  相似文献   

12.
Hosts of the common cuckoo (Cuculus canorus), an avian brood parasite, develop antiparasite defense mechanisms to increase their reproductive success. Ejection of the parasite egg and desertion of the parasitized nest are the most typical adaptations in response to brood parasitism, but nest desertion may also occur in response to partial clutch reduction, independently from parasitism. Some great reed warblers (Acrocephalus arundinaceus) showed both mechanisms in the same incidence of cuckoo parasitism: in 18% of successful ejections of the parasite eggs, they deserted their nests. We studied if such cases of post-ejection nest-desertion are caused by brood parasitism or reduced clutch value. We experimentally parasitized clutches consisting of five or three host eggs with two painted conspecific eggs to mimic parasitic eggs, as multiple parasitism is frequent in the area. Although hosts ejected these parasitic eggs in both clutch categories (100% and 67% for the larger and smaller inital clutch sizes, respectively), we found that after manipulation, post-ejection nest-desertion frequently occurred at small (3-egg) clutches (40%), but rarely at large (5-egg) clutches (17%). The same phenomenon also occurred when unparasitized 3-egg clutches were reduced by two eggs, but not when 5-egg clutches were reduced in the same way. A logistic regression model revealed that only initial clutch size affected nest desertion of parasitized nests in our experiments. Therefore, we conclude that post-ejection nest-desertion is not a second antiparasite mechanism, which might serve as a redundant antiparasite defense, but a reaction to typically small and further decreased clutch size.  相似文献   

13.
Background, aim, and scope Reports about declines or unusual structures of fish populations in native aquatic systems in Central Europe and North America are in sharp contrast to an obvious improvement of general water quality. The Neckar River may serve as an example of a formerly severely contaminated freshwater system in Southern Germany, the ecological situation of which could be substantially improved over the last three decades. Nevertheless, there are still deficits in the composition of the fish fauna, which cannot be explained by conventional chemical-analytical, hydromorphological and limnological methodologies. Therefore, in search of explanations for ecological deficits, ecotoxicological investigations with an increasing focus on sediment contamination have been performed along the Lower Neckar River over a period of 10 years. In addition to sediment tests, fish populations were screened for genotoxic and embryotoxic effects as well as alterations in the structure of central metabolic organs such as the liver. Materials and methods Roach (Rutilus rutilus) and gudgeon (Gobio gobio) from the Lower Neckar River were studied with respect to histo- and cytological alterations of the liver as well as the induction of genotoxicity in liver, gut, gills and blood cells by means of the comet and micronucleus assays. At the same time, both native sediments and acetonic sediment extracts were tested for toxicity to zebrafish (Danio rerio) embryos and permanent fish cell cultures. Results Massive disturbances of the liver ultrastructure indicate severe stress in the fish from the Lower Neckar River despite good supply of nutrition. Both cyto- and embryotoxicity tests document a considerable toxic potential of sediments from the Lower Neckar River, and results of both the comet assay and the micronucleus test provide evidence of the presence of genotoxic agents in the sediments and their effects in fish. There has been no decrease of genotoxicity over the last 10 years. Discussion Cytopathology in conjunction with general cyto- and embryotoxicity in fish from the Lower Neckar River contribute to a syndrome which cannot be neglected in the interpretation of abnormal compositions of fish populations. In comparison to weight-of-evidence studies at other river systems in Southern Germany, the Neckar is characterized by an elevated embryotoxic but by a relatively moderate genotoxic potential. Conclusions The ecotoxicological burden of sediments at the Lower Neckar River is mirrored in the condition of native fish populations. A potential contribution of chemical contamination to the phenomenon of fish declines in aquatic ecosystems cannot be ignored. Recommendations and perspectives The improvement of the ecological status of freshwater systems may not be abused as an argument to discontinue biological monitoring programs. Rather, continuous surveillance of the environment and in-depth investigations into the consequences of chemical contamination are indispensible. For an adequate appreciation of the ecological relevance of sublethal and/or specific biological effects, their impact on population structure and function remains to be elucidated further. For the identification of the chemical basis of biological effects, methodologies such as effect-directed analysis appear to be promising approaches.  相似文献   

14.
Summary In laboratory experiments, we tested the hypothesis that by living in larger shoals, juvenile threespine (Gasterosteus aculeatus) and blackspotted (G. wheatlandi) sticklebacks lower their risk of being parasitized by the crustacean ectoparasite Argulus canadensis. An increase in shoal size resulted in a lower average number of attacks received by individual fish, but had no negative effect on the attack performance (attack rate and attack success) of the parasites. In addition, more fish formed shoals and shoal sizes were larger in the presence of parasites. We conclude that ectoparasitism may have been a strong selective factor in the evolution of social behaviour in juvenile sticklebacks.  相似文献   

15.
E. Radha 《Marine Biology》1971,8(3):213-219
The fish Caranx kalla of different size groups (2 to 18 cm body length) is present during various calender months of the year off the Madras coast (S. India). C. kalla displays a definite progressive shift in growth during successive months; the fish grows from 2 cm (September) to about 18 cm (August) over a period of 12 months. The highest (47%) or lowest (11%) incidence of a monogenean gill parasite Gastrocotyle indica has been observed in the C. kalla size groups 2.0 to 3.9 cm or 16.0 to 17.9 cm; however, the maximum specific density of 10.9 parasite/infested fish was observed in the size group 14.0 to 15.9 cm. Small fish (2.0 to 3.9 cm) carry only larvae (58/100 fish); larger hosts (4.0 to 5.9 cm) larvae (45/100 fish) and juveniles (8/100 fish); individuals of the 6.0 to 15.9 cm size group harbour larval, juvenile and adult stages of G. indica. Fishes of more than 16 cm carry only adult parasites. The level of parasitization is higher on the male hosts (30% incidence; 5 parasites/infested male) than on the females (25% incidence; 4 parasites/infested female). The highest incidences of larval, juvenile and adult G. indica occur in September, February, and May, respectively; the parasite seems to live for 1 year. The host requires a period of about 8 months to acquire immunity against infestation by G. indica or copepod parasites.  相似文献   

16.
《Ecological modelling》2005,181(4):535-556
Observational models for the catch of fish at age a (or size) at time t are fundamental equations in fisheries science, linking a population model with data. The well known Baranov catch equation (which assumes that fishing and natural mortalities are constant over both age and time) is a nominal basis of those most commonly used in fish stock assessment and fish population models (which assume that fishing and natural mortalities vary with both age and time). But, what should a catch equation look like, if the instantaneous rates of fishing and natural mortalities of fish of age a at time t vary with age a and time t? Without answering this question, use of those catch equations in fish stock assessment and population models renders their results uncertain. In this paper, I derive a general catch in number or in biomass equation as observational models of an age- and time-dependent model for a fish population by Taylor series expansion of, and by directly manipulating, a general catch integral, reduce it to commonly used catch equations, and compare the performance of 11 of them using data on the western king prawn Penaeus latisulcatus. I show that the nominal generalization of the Baranov catch equation misses several terms. In so doing, I derive the catch equations more accurately and restore these missing terms. Although almost all approximations overestimate the catch per recruit for older prawns, all commonly used catch equations and their extensions perform worse than theoretically sound representations of the general catch equation and their approximations. The age-specific bias of all models is <2.5, <18 and <90% for a time interval of sampling of 1, 7 and 30 days, respectively. Such large biases even for moderate values of the length of the time interval of sampling highlight a need for assessing the utility of commonly used catch equations for individual species.  相似文献   

17.
When brood parasites are about to lay an egg, they have to decide which nest to parasitize. The best nest in which to lay will depend on the parenting ability of the host. We have studied selection of magpie (Pica pica) hosts by great spotted cuckoos (Clamator glandarius). Great spotted cuckoos preferentially parasitize large host nests. Nest volume in magpies is a good indicator of territory quality, since there is a negative relationship between magpie nest size and breeding date, and timing of breeding in magpies is known to be positively related to territory quality. Moreover, magpies occupying high-quality territories have high breeding success. Therefore, nest size is positively related to the quality of magpies. Parasitized magpie nests were of greater volume than the nearest neighbouring nest not parasitized by the great spotted cuckoo. In order to test whether the great spotted cuckoos might select high-quality magpie hosts, we manipulated pairs of parasitized and non-parasitized nests with identical laying dates and habitats, introducing into each of the nests the same number of parasitic and non-parasitic eggs. The number of fledglings reared (magpie plus great spotted cuckoo chicks) in naturally parasitized nests was higher than in experimentally parasitized nests. Thus, the probability of survival of the parasite chicks increased if cuckoo eggs were laid in the nests of high-quality hosts originally chosen by the parasite.  相似文献   

18.
Fundamental hydrodynamic and ecological processes of a lake or reservoir could be adequately depicted by one-dimensional (1D) numerical simulation models. Whereas, lakes with significant horizontal water quality and hydrodynamic gradients due to their complex morphometry, inflow or water level fluctuations require a three-dimensional (3D) hydrodynamics and ecological analyses to accurately simulate their temporal and spatial dynamics. In this study, we applied a 3D hydrodynamic model (ELCOM) coupled with an ecological model (CAEDYM) to simulate water quality parameters in three bays of the morphologically complex Lake Minnetonka. A considerable effort was made in setting up the model and a systematic parameterization approach was adopted to estimate the value of parameters based on their published values. Model calibration covered the entire length of the simulation periods from March 29 to October 20, 2000. Sensitivity analysis identified the top parameters with the largest contributions to the sensitivity of model results. The model was next verified with the same setup and parameter values for the period of April 25 to October 10, 2005 against field data. Spatial and temporal dynamics were well simulated and model output results of water temperature (T), dissolved oxygen (DO), total phosphorus (TP) and one group of algae (Cyanobacteria) represented as chlorophyll a (Chla) compared well with an extensive field data in the bays. The results show that the use of the model along with an accurate bathymetry, a systematic calibration and corroboration (verification) process will help to analyze the hydrodynamics and geochemical processes of the morphologically complex Lake Minnetonka. An example of an ecological application of the model for Lake Minnetonka is presented by examining the effect of spatial heterogeneity on coolwater fish habitat analysis in 3D and under a scenario where horizontal spatial heterogeneity was eliminated (1D). Both analyses captured seasonal fish habitat changes and the total seasonal averages differed moderately. However, the 1D analysis did not capture local and short duration variabilities and missed suitable fish habitat variations of as much as 20%. The experiment highlighted the need for a 3D analysis in depicting ecological hot spots such as unsuitable fish habitats in Lake Minnetonka.  相似文献   

19.
Despite that the existence of animal personalities is widely recognized, no consensus has been reached on the relative importance of different ecological factors behind their expression. Recently, it has been suggested that parasites may have a crucial role in shaping animal personalities, but only a very few studies have experimentally tested the idea. We infected Eurasian minnows (Phoxinus phoxinus) with the brain-encysted trematode parasite, Diplostomum phoxini, and studied whether infection could modify the personality of their hosts. Our results show that D. phoxini infection did not affect the mean levels of boldness, activity or exploration, but infected minnows showed higher repeatability in boldness and activity, and reduced repeatability in exploration. We also found that D. phoxini may be able to break the associations (behavioral syndromes) between behavioral traits, but that this effect may be dependent on parasite intensity. Furthermore, the effect of D. phoxini infection on personality of the hosts was found to be nonlinearly dependent on infection intensity. Taken together, our results suggest that D. phoxini parasites may shape the personality of their hosts, but that behavioral consequences of ecologically relevant infection levels may be rather subtle and easily remain undetected if only the mean trait expressions are compared.  相似文献   

20.
G. Anderson 《Marine Biology》1977,42(3):239-251
I quantified the effects of parasitism by the isopod Probopyrus pandalicola on energy flow through the host Palaemonetes pugio by comparing secondary production, metabolism, ingestion, and egestion by unparasitized laboratory shrimp populations to the same parameters for parasitized groups during 10 months. The effects of parasitism on host growth and metabolism vary from month to month. Temperature, season, host age, sex, and reproductive condition affect energetics for host-parasite systems. Probopyrus pandalicola has little effect on host assimilation efficiency. However, tissue growth efficiences during most study months were higher for control shrimp than parasitized shrimp. These differences between groups were of lesser magnitude when parasite production was considered in the calculations. Trophic level energy intake efficiency for parasites was of the order of 6 to 10% throughout much of the study—the highest values were calculated during the parasites' reproductive months. Through parasitic castration, P. pandalicola significantly affects host energetics. Significantly, parasite reproduction was often of the same magnitude as reproduction by unparasitized hosts, although parasite biomass accounts for only about 4% of the total host-parasite system biomass.  相似文献   

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