Nematode parasitism in a Danish drosophilid community: further evaluation of the disproportionate parasitism hypothesis

The nematodes Parasitylenchus diplogenus and Howardula aoronymphium (Allantonematidae) are parasites of drosophilids (Diptera). Nematodes were found in 3/14 drosophilid species sampled in Danish woodlands: Drosophila phalerata Meigen (3.5% parasitism) was parasitized by H. aoronymphium and D. obscura Fallén (0.5%) and D. subobscura Collin (2.1%) by P. diplogenus. Parasitism was generally rare, and few drosophilid species parasitized, compared to elsewhere in Europe. Parasitism was positively correlated with relative host abundance both within Denmark and across communities, suggesting that species diversity is promoted by a disproportionately high parasitism of more common host species. The prevalence of H. aoronymphium, but not P. diplogenus, parasitism is positively related to temperature across communities.     

Three-dimensional dispersion of drosophilid flies in a cool temperate forest of northern Japan

Three-dimensional dispersion of drosophilid flies was studied within a secondary broad-leaved forest in relation to forest structure. The survey area included the forest margin and old canopy gaps and varied in the foliage height profile from place to place. Using multivariate analyses on the data of drosophilid dispersion, five microhabitats which were different from one another for drosophilids were recognized: (i) canopy layer; (ii) middle layer; (iii) floor layer of forest interior; (iv) upper layer of forest margin; and (v) herbaceous layer of forest margin and gap. The height of living space of canopy species was remarkably lowered at the forest margin. The forest edge was richer in both numbers of individuals and species than the forest interior from the overlap of the grassland and the forest canopy subcommunities and the addition of invaders from other habitats. However, no ‘edge’ species, which were mostly restricted to or spend most of their time in ecotones, were found. It is hypothesized that the above-ground forest structure consists fundamentally of three zones: (i) the canopy; (ii) the floor; and (iii) the edge. A significant positive correlation was found between the foliage height diversity and the degree of vertical habitat segregation among drosophilid species. The patchiness of vegetation structure influential to the three-dimensional dispersion in a forest drosophilid community was estimated to be on the scale of 110–450 m². This scale of subjective habitat patchiness or ‘ecological neighbourhood’ corresponds well with the most prevalent size of canopy gaps occurring in various forests.     

Grazing in a porous environment. 2. Nematode community structure

The influence of soil matric potential on nematode community composition and grazing associations were examined. Undisturbed cores (5 cm diameter, 10 cm depth) were collected in an old field dominated by perennial grasses on a Hinckley sandy loam at Peckham Farm near Kingston, Rhode Island. Ten pairs of cores were incubated at −3, −10, −20 and −50 kPa matric potential after saturation for 21–28 or 42–58 days. Nematodes were extracted using Cobb's decanting and sieving method followed by sucrose centrifugal-flotation and identified to family or genus. Collembola and enchytraeids present were also enumerated because they are grazers that reside in air-filled spaces. Direct counts of bacteria and fungi were made to estimate biovolume using fluorescein isothiocyanate and fluorescein diacetate stains, respectively. Trophic diversity and maturity indices were calculated for nematode communities. Three patterns of matric potential effect were observed for nematode taxa. One, there was a consistent effect of matric potential for all seasons for Alaimus, Monhysteridae, Prismatolaimus, Paraxonchium and Dorylaimoides. Two, some effects of matric potential were consistent among seasons and other effects were inconsistent for Aphelenchoides, Aphelenchus, Cephalobidae, Coomansus, Eudorylaimus, Huntaphelenchoides, Panagrolaimidae, Paraphelenchus, Sectonema, and Tripyla. Third, effects of matric potential were always inconsistent among seasons for Aphanolaimus, Aporcelaimellus, Bunonema, Rhabditidae, and Tylencholaimus. As predicted, fungal and bacterial biomass responded oppositely to matric potential. Total bacterial biomass was greater at −3 kPa than −10, −20 and −50 kPa (P=0.0095). Total fungal biomass was greater at −50, −20 and −10 kPa than −3 kPa (P=0.0095). Neither bacterial-feeding, fungal-feeding nor predacious nematodes correlated significantly with bacterial or fungal biomass. Omnivorous and predacious nematodes correlated positively with number of bacterial-feeding nematodes; predacious nematodes also correlated positively with fungal-feeding nematodes. Numbers of Collembola and enchytraeids were more often correlated positively with microbial-grazing nematode numbers in drier than moist soils. From this study, we propose two mechanisms that may explain nematode community structure changes with matric potential: differential anhydrobiosis and/or enclosure hypotheses. The later suggests that drying of soil generates pockets of moisture in aggregates that become isolated from one another enclosing nematodes and their food in relatively high concentrations creating patches of activity separated by larger areas of inactivity. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effects of parasitoids on a mycophagous drosophilid community in northern Japan and an evaluation of the disproportionate parasitism hypothesis

In a host–parasitoid system comprising mycophagous drosophilids and their parasitoids, the drosophilid and parasitoid species assemblages, host use, and the prevalence of parasitism were assessed, and the “disproportionate parasitism hypothesis” was examined with consideration given to yearly variations. The mycophagous drosophilids, their fungal food resources and parasitoids were studied by carrying out an intensive census throughout the activity seasons of 4 years (2000–2003) in Hokkaido, northern Japan. Five hymenopterous parasitoid species, four braconids and one eucoilid, were found. Parasitoids of mycophagous drosophilids are reported for the first time from Asia. Most parasitism (99.2%) was by braconids, in contrast to the dominance of eucoilids in Europe. Parasitism was restricted to the summer, and the rate was high from early July to early August every year. There was considerable yearly variation in the composition of abundant fungus, drosophilid and parasitoid species, especially between 2000 and 2001. The alternation of dominant host species was coupled with the alternation of dominant parasitoid species that differed in host use. Despite the yearly variation in the system, the most dominant host species suffered disproportionately heavy parasitism by the correspondingly dominant parasitoid species every year. The parasitism rate was positively correlated with the relative host abundance. This thus indicates that the disproportionate parasitism mechanism may operate, via which species coexistence is promoted by a higher rate of parasitism of the dominant species.     

Effects of nematode parasitism on mycophagous drosophilid species in northern Japan

Mycophagous drosophilids and their nematode parasites were studied in an intensive census conducted throughout the active season over four years (2000–2003) in Hokkaido, northern Japan. Using census data, I assessed the prevalence of nematode parasitism through seasons, determined its effects on drosophilid female fertility and tested the “disproportionate parasitism hypothesis” to evaluate its effects on drosophilid community structure. Over the four‐year census, a total of 43 fungal species were found on the census route and 18 177 adult drosophilid flies were collected. The yearly parasitism rate on the total drosophilid community varied from 6.2% in 2000 to 5.1% in 2001 and 3.0% in 2002 and 2003. There was strong seasonal variation in the prevalence of parasitism. In addition, parasitism rates varied among drosophilid species. All parasitized females carried only small numbers of eggs. Thus, nematode parasitism had a strong deleterious effect on the fitness of infected flies. The relative abundance of drosophilid species and parasitism rate were not significantly correlated. Thus, the disproportionate parasitism hypothesis likely does not apply to the mycophagous drosophilid community in Japan.     

Coexistence of drosophilid flies: Aggregation, patch size diversity and parasitism

We carried out field experiments to investigate the coexistence of Drosophila species in domestic and forest areas on the basis of the aggregation model. Three cosmopolitan species Drosophila simulans Sturtevant, Drosophila melanogaster Meigen and Drosophila immigrans Sturtevant, and a native species, Drosophila auraria Peng, emerged abundantly from banana placed at the domestic station, while Drosophila immigrans and five native species, Drosophila lutescens Okada, Drosophila rufa Kikkawa and Peng, Drosophila bizonata Kikkawa and Peng, Drosophila sternopleuralis Okada and Kurokawa and Scaptodrosophila coracina (Kikkawa and Peng), were abundant at the forest station. The present analysis suggests that their coexistence was facilitated by the aggregation mechanism. In the cosmopolitan species, the density of individuals that emerged from patches increased with the increase of patch size, but the relationship between fly density and patch size was not clear in the native species. This difference in distribution patterns between the cosmopolitan and native species is likely to be due to the difference in the female visiting behavior. In the present analysis, however, it was not clear whether patch size diversity facilitated their coexistence or not. The effect of patch size diversity may have been masked, because the effect of aggregation was more prominent. The rate of parasitism by wasps was high in October at the domestic station, and in May and June at the forest station. The present result suggests that the rate of parasitism was density-dependent, at least at the domestic station, and therefore parasitism facilitates the coexistence of drosophilid species in domestic areas.     

Trade-offs between life-history traits in a coldwater fish in the Mediterranean Sea: the case of blue whiting Micromesistius poutassou

A combination of traditional and emerging methodologies was used to assess the trade-offs between several life-history traits (linked to reproduction and condition) and parasitism in a commercially-exploited cold-water species, blue whiting Micromesistius poutassou, in the Mediterranean Sea. The use of histological and gravimetric methods revealed conflicting evidence as to the fecundity type (indeterminate or determinate) of this species. Moreover, there seem to be condition-mediated compensations between egg quality and egg quantity. The effects of parasitism on reproduction and condition are species-specific and occur mainly at high intensities of infection; they include a lower batch fecundity (affecting reproductive potential), a higher hepatosomatic index and a higher spleen-somatic index. Considering the fact that larger fish spawn more eggs and that the minimum landing size is lower than the size at maturity, these results may have implications for the future management of M. poutassou stocks in the Mediterranean Sea. Local environmental conditions may account for geographical differences regarding infection in M. poutassou. Altogether, the results support the idea that the complex trade-offs between parasitism, reproduction and condition need to be considered in order to understand the status of cold-water species such as M. poutassou.     

Nematode species abundance patterns and their use in the detection of environmental perturbations

If the concepts of biological indices and biomonitoring at the multi-species level are to prove viable, then meiofauna and marine nematodes in particular should be an ideal group with which to test the hypothesis. Many attempts to assess the structure of species assemblages, such as the use of diversity indices, nematode: copepod ratios and the graphical method of log normal plots, have been shown to be theoretically unsound and/or impractical, especially for routine use by extension workers. A method of assessing shifts in dominance patterns which involves all the proportional species abundances is suggested as a better means of comparing diversity. A modified method of rapidly assessing Simpson's dominance-weighted diversity index is also advocated as being of practical use. In combination, they should enable the diversity aspect of the multi-species approach to biomonitoring to be rigorously and exhaustively evaluated.     

Coastal ecosystems on a tipping point: Global warming and parasitism combine to alter community structure and function

Mounting evidence suggests that the transmission of certain parasites is facilitated by increasing temperatures, causing their host population to decline. However, no study has yet addressed how temperature and parasitism may combine to shape the functional structure of a whole host community in the face of global warming. Here, we apply an outdoor mesocosm approach supported by field surveys to elucidate this question in a diverse intertidal community of amphipods infected by the pathogenic microphallid trematode, Maritrema novaezealandensis. Under present temperature (17°C) and level of parasitism, the parasite had little impact on the host community. However, elevating the temperature to 21°C in the presence of parasites induced massive structural changes: amphipod abundances decreased species‐specifically, affecting epibenthic species but leaving infaunal species largely untouched. In effect, species diversity dropped significantly. In contrast, four degree higher temperatures in the absence of parasitism had limited influence on the amphipod community. Further elevating temperatures (19–25°C) and parasitism, simulating a prolonged heat‐wave scenario, resulted in an almost complete parasite‐induced extermination of the amphipod community at 25°C. In addition, at 19°C, just two degrees above the present average, a similar temperature–parasite synergistic impact on community structure emerged as seen at 21°C under lower parasite pressure. The heat‐wave temperature of 25°C per se affected the amphipod community in a comparable way: species diversity declined and the infaunal species were favoured at the expense of epibenthic species. Our experimental findings are corroborated by field data demonstrating a strong negative relationship between current amphipod species richness and the level of Maritrema parasitism across 12 sites. Hence, owing to the synergistic impact of temperature and parasitism, our study predicts that coastal amphipod communities will deteriorate in terms of abundance and diversity in face of anticipated global warming, functionally changing them to be dominated by infaunal species.     

From micro- to macroevolution: brood parasitism as a driver of phenotypic diversity in birds

A fundamental question in biology is how diversity evolves and why some clades are more diverse than others. Phenotypic diversity has often been shown to result from morphological adaptation to different habitats. The role of behavioral interactions as a driver of broadscale phenotypic diversity has received comparatively less attention. Behavioral interactions, however, are a key agent of natural selection. Antagonistic behavioral interactions with predators or with parasites can have significant fitness consequences, and hence act as strong evolutionary forces on the phenotype of species, ultimately generating diversity between species of both victims and exploiters. Avian obligate brood parasites lay their eggs in the nests of other species, their hosts, and this behavioral interaction between hosts and parasites is often considered one of the best examples of coevolution in the natural world. In this review, we use the coevolution between brood parasites and their hosts to illustrate the potential of behavioral interactions to drive evolution of phenotypic diversity at different taxonomic scales. We provide a bridge between behavioral ecology and macroevolution by describing how this interaction has increased avian phenotypic diversity not only in the brood parasitic clades but also in their hosts.     

Higher gregarine parasitism often in sibling species of host damselflies with smaller geographical distributions

1. This study investigated inter‐specific variation in parasitism by gregarines (Eugregarinorida: Actinocephalidae), among sibling species of damselflies (Odonata: Zygoptera), in relation to relative size of geographical ranges of host species. 2. Gregarines are considered generalist parasites, particularly for taxonomically related host species collected at the same sites or area. Prevalence and median intensity of gregarine parasitism was obtained for 1338 adult damselflies, representing 14 species (7 sibling species pairs) across 3 families within the suborder Zygoptera. Damselflies were collected at three local sites in Southeastern Ontario, during the same periods over the season. 3. Five out of seven species pairs had significant differences in parasitism between sibling species. The less widespread host species was the more parasitised for three species pairs with significant differences in gregarine prevalence, and for two species pairs with differences in median intensity. The more widespread host had a higher intensity of infection as expected, in two species pairs. 4. Future studies on ecological determinants of parasitism among related species should examine robust measures of abundance of species and representation of species regionally.     

管氏肿腿蜂密度对其寄生及繁衍能力的影响

本研究以管氏肿腿蜂Sclerodermus guani雌成蜂为研究对象,拟探明寄生蜂密度对其寄生及繁衍能力的影响,为该寄生蜂行为生物学研究提供科学依据。本研究以松墨天牛Monochamus alternatus幼虫(0.350～0.450 g)为寄主,设置8种不同雌蜂密度(1～8头/寄主),对比分析不同雌蜂密度下,管氏肿腿蜂的寄生行为和效能,以及雌蜂生殖力和子代适合度变化。结果表明,随雌蜂密度增加,管氏肿腿蜂蛰刺发生前历时、寄主麻痹历时及产卵前历时均缩短。基于Hassell-Varley数据模型分析(E=0.9023×P^-0.9378),雌蜂的寻找效应随自身密度增加而逐渐变小,雌蜂间的干扰降低了寄生效能。雌蜂生殖力随其密度增加而降低,单雌平均产卵量及产雌量明显下降,雌蜂密度为1头/寄主时,单雌产卵量分别是4头/寄主和8头/寄主处理下的1.4倍和2.4倍。子代适合度也随雌蜂密度增加而变化,子代发育历期明显缩短,低龄幼虫存活率和单雌体重均下降。雌蜂密度为1头/寄主时,其子代发育历期最长,平均为26.7 d, 8头/寄主时最短,平均为24.0 d。雌蜂密度为1头/寄主时...     

Nest destruction elicits indiscriminate con‐ versus heterospecific brood parasitism in a captive bird

Following nest destruction, the laying of physiologically committed eggs (eggs that are ovulated, yolked, and making their way through the oviduct) in the nests of other birds is considered a viable pathway for the evolution of obligate interspecific brood parasitism. While intraspecific brood parasitism in response to nest predation has been experimentally demonstrated, this pathway has yet to be evaluated in an interspecific context. We studied patterns of egg laying following experimental nest destruction in captive zebra finches, Taeniopygia guttata, a frequent intraspecific brood parasite. We found that zebra finches laid physiologically committed eggs indiscriminately between nests containing conspecific eggs and nests containing heterospecific eggs (of Bengalese finches, Lonchura striata vars. domestica), despite the con‐ and heterospecific eggs differing in both size and coloration. This is the first experimental evidence that nest destruction may provide a pathway for the evolution of interspecific brood parasitism in birds.     

On the origin of brood parasitism in altricial birds

The probability that obligate interspecific brood parasitism(OP), among altricial birds evolved directly from the normalbreeding (no parasitism, NP) mode or indirectly through intraspecificnest parasitism (INP) was examined by using maximum-likelihoodand parsimony approaches. We examined the probability of ancestralstates at 24 key nodes in order to test our hypotheses. Thestate of the most basal node in a tree of 565 genera of altricialbirds is equivocal; however, the state probability of NP atthis node is about 5.5-fold more likely than the state of obligateparasite. A similar trend was observed for basal nodes of mostfamilies examined. The INP state was supported only in the Hirundinidae.The high incidence of INP among martins and swallows explainsthis finding. Contrary to our predictions, even in other groupswhere there is a high incidence of INP and OP, such as in thetribe Icteri and the Old World finches, the probability of NPbeing ancestral was very high. We conclude that in all casesbut one (Hirundinidae) obligate, and probably facultative, broodparasitism evolved directly from normal breeding mode ratherthan indirectly through some other form of parasitism.     

Ecological and Morphological Differentiation Among COI Haplotype Groups in the Plant Parasitic Nematode Species Mesocriconema Xenoplax

DNA barcoding with the mitochondrial COI gene reveals distinct haplotype subgroups within the monophyletic and parthenogenetic nematode species, Mesocriconema xenoplax. Biological attributes of these haplotype groups (HG) have not been explored. An analysis of M. xenoplax from 40 North American sites representing both native plant communities and agroecosystems was conducted to identify possible subgroup associations with ecological, physiological, or geographic factors. A dataset of 132 M. xenoplax specimens was used to generate sequences of a 712 bp region of the cytochrome oxidase subunit I gene. Maximum-likelihood and Bayesian phylogenies recognized seven COI HG (≥99/0.99 posterior probability/bootstrap value). Species delimitation metrics largely supported the genetic integrity of the HG. Discriminant function analysis of HG morphological traits identified stylet length, total body length, and stylet knob width as the strongest distinguishing features among the seven groups, with stylet length as the strongest single distinguishing morphological feature. Multivariate analysis identified land cover, ecoregion, and maximum temperature as predictors of 53.6% of the total variation (P = 0.001). Within land cover, HG categorized under “herbaceous,” “woody wetlands,” and “deciduous forest” were distinct in DAPC and RDA analyses and were significantly different (analysis of molecular variance P = 0.001). These results provide empirical evidence for molecular, morphological, and ecological differentiation associated with HG within the monophyletic clade that represents the species Mesocriconema xenoplax.     

Male parasitism and intrasexual competition in a burrowing barnacle

Summary In sexually dimorphic animals, large male body size is often associated with direct interference competition among males for access to females or resources used in reproduction. In constrast, small male body size may be associated with indirect scramble competition among males for temporal or spatial access to females. Minute, “parasitic” males of the acrothoracican barnacleTrypetesa lampas (Hancock) appear to compete with one another for permanent attachment sites on the external body of the female. Several spatial patterns suggest indirect male-male competition: 1) males were consistently aggregated on the anterior surface of the female ovarian disc; 2) the average distance from attached males to the site of insemination correlated positively with local male density; 3) average male body size on a female decreased as a function of male density; 4) the distribution of males on the left and right hand sides of the female ovarian disc was more even than expected, suggesting that males avoided crowded settlement sites. The number of males attached to a female increased with female body size and matched a null model in which males colonized female “targets” of differing areas. These results suggest that competition between males primarily affected settlement sites and male body sizes within, rather than among, females. Male parasitism may have evolved through both sexual selection for efficient access to females (Ghiselin 1974) and natural selection for reduced burrow density in a space-limited habitat (Turner and Yakovlev 1983).     

鸟类巢寄生伤害理论的进化

鸟类巢寄生的寄主无论是成乌还是雏鸟,对宿主都是极具伤害性的,因为它们降低了宿主的生育力,然而,无论是在寄主种内还是种间,其伤害性的差异变化很大。综述了以往对这种伤害性差异的各种解释,以往的解释在很大程度上集中在伤害性所带来的利益。认为寄主的伤害性行为可以像病原体的伤害性一样进行分类,伤害性在为寄主带来利益的同时,也伴随着代价,所以,由病原体伤害性进化研究衍生而来的平衡假说,适用于解释鸟类巢寄生伤害理论的进化。     

The evolution of obligate interspecific brood parasitism in birds

We present a simple analytical model to investigate the conditionsfor the evolution of obligate interspecific brood parasitismin birds, based on clutch size optimization, when birds canlay more eggs than their optimal clutch size. The results showthat once intraspecific parasitism has appeared (i.e., femalesstart to spread their eggs over their own and other nests) the evolutionarily stable number of eggs laid in its own nest decreases.Two possible ESSs exist: (1) either the evolutionarily stablenumber of eggs laid in its own nest is larger than zero, anda fraction of the total number of eggs is laid parasitically(i.e., intraspecific parasitism); and (2) either the evolutionarilystable number of eggs laid in its own nest is zero and all eggs are laid parasitically. Since all females lay parasitically,this could favor the evolution of obligate interspecific broodparasitism. The key parameter allowing the shift from intraspecificto obligate interspecific parasitism is the intensity of density-dependentmortality within broods (i.e., nestling competition). Strongnestling competition, as in altricial species, can lead toan ESS where all eggs are laid parasitically. Altricial speciesare, therefore, predicted to evolve more easily toward obligate interspecific parasitism than precocial species. These predictionsfit the observed distribution of brood parasitism in birds,where only one species out of 95 obligate interspecific parasitesexhibits a precocial mode of development. Different nestlingsurvival functions provided similar findings (i.e., obligatebrood parasitism is more likely to evolve in altricial species),suggesting that these results are robust with respect to themain assumption of the model.