Aging alters interspecific competition between two sympatric insect–parasitic nematode species

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Interspecific competition in Drosophila II. Competitive outcome in some 2–resource environments

Populations of Drosophila metanogaster and D. simulans were made to compete in a number of 2– resource environments. The resources used were all based on a standard Drosophila medium and differed from each other only in their concentrations of ethanol. Each experiment comprised three cages started at a species–frequency of 0.8 D. melanogaster and three at 0.2 D. melanogaster. This enabled screening for a wide range of equilibria.
Reversal of competitive superiority was achieved (unintentionally) by raising the alcohol level of one of the resources. Thus while D. simulans won in an environment with media containing 0 and 896 ethanol, D. melanogaster won if the ethanol concentrations were 0 and 10%, or 0 and 12%. This parallels the reversal of dominance reported earlier for single–resource systems. However, in none of the 2–resource systems studied was there a stable equilibrium in species–frequency–in fact there was no significant frequency–dependence at all.
An artificial selection experiment produced a stock of D. melanogaster ovipositing a higher proportion of eggs on the alcoholic resource. Thus, this stock was more divergent from D. simulans , in at least one aspect of resource use, than was the original stock of D. melanogaster. However, competition between stock D. simulans and selected D. melanogaster showed no frequency–dependence. Again, reversal of competitive dominance was unintentionally achieved.
The implications of these results are discussed, and the need for a comprehensive study of resource–utilization in a pair of competing species is stressed.     

Two non-linked genes for specific virulence of Leptopilina boulardi against Drosophila melanogaster and D. yakuba

The developmental success of Leptopilina boulardi parasitoids within host species of the melanogaster subgroup is determined mainly by their ability to suppress the host immune reaction (virulence). Host resistance and parasitoid virulence are genetically variable in both partners. A gene for specific resistance against L. boulardi (Rlb) has been identified in Drosophila melanogaster, and a gene for the immune suppression (IS) of D. melanogaster has been identified in L. boulardi. To understand the evolution of the IS gene, we determined its specificity regarding potential hosts of the melanogaster subgroup. It did not affect the virulence against any other species of the melanogaster subgroup and was called ISm for immune suppression of D. melanogaster. Another gene (ISy), non-linked to the gene ISm, was characterized for the specific immune suppression of D. yakuba. These results suggesting that natural selection for virulence against one host species does not influence the evolution of virulence against another will allow us to develop pertinent hypotheses concerning the evolution of this character which is expected to drive the evolution of the parasitoid toward narrow host specialization. This revised version was published online in July 2006 with corrections to the Cover Date.     

Evolutionary relationships to parasitism by seven species of the Drosophila melanogaster subgroup

Parasitic wasps are an important component of the niche of Drosophila species. The susceptibility to the Cynipid Leptopilina boulardi was estimated in the seven sibling species of Drosophila belonging to the melanogaster subgroup. Three categories of Hies can be distinguished, according to the level of cellular immune reaction and success of parasitism. Drosophila melanogaster and D. mauritiana belong to the category 1, specified by no encapsulative reaction and a high rate of successful parasitism. Category 2, characterized by a moderate encapsulation rate and a high mortality include D. simulans.5, D. erecta and D. orena. Category 3, with D. yakuba and D. tcissien, is specified by a very low rate or absence of successful parasitism due to a highly efficient immune cellular reaction. This classification parallels the phylogenic relationship based upon polytene chromosome banding sequences. Such specific ditferences in susceptibility to parasites may plan an important role in the competition between these species in Africa.     

Evolutionary relationships to parasitism by seven species of the Drosophila melanogaster subgroup

Parasitic wasps are an important component of the niche of Drosophila species. The susceptibility to the Cynipid Leptopilina boulardi was estimated in the seven sibling species of Drosophila belonging to the melanogaster subgroup. Three categories of Hies can be distinguished, according to the level of cellular immune reaction and success of parasitism. Drosophila melanogaster and D. mauritiana belong to the category 1, specified by no encapsulative reaction and a high rate of successful parasitism. Category 2, characterized by a moderate encapsulation rate and a high mortality include D. simulans.5, D. erecta and D. orena . Category 3, with D. yakuba and D. tcissien , is specified by a very low rate or absence of successful parasitism due to a highly efficient immune cellular reaction. This classification parallels the phylogenic relationship based upon polytene chromosome banding sequences. Such specific ditferences in susceptibility to parasites may plan an important role in the competition between these species in Africa.     

Effectively vertical transmission of a Drosophila‐parasitic nematode: mechanism and consequences

1. Long‐term control of insects by parasites is possible only if the parasite populations persist. Because parasite transmission rate depends on host density, parasite populations may go extinct during periods of low host density. Vertical transmission of parasites, however, is independent of host density and may therefore provide a demographic bridge through times when their insect hosts are rare. 2. The nematode Howardula aoronymphium, which parasitises mycophagous species of Drosophila, can experience both horizontal and effectively vertical transmission, relative rates of which depend, in theory at least, on the density of hosts at breeding sites. 3. A nine‐generation experiment was carried out in which nematodes were transmitted either exclusively vertically or primarily horizontally. This experiment revealed that these parasites can persist and exhibit positive population growth even when there is only vertical transmission. 4. Assays at the end of the experiment revealed that the vertically transmitted nematodes had suffered no inbreeding depression and that they were similar to the horizontally transmitted nematodes in terms of virulence, infectivity, within‐host growth rate, and fecundity. Thus, vertical transmission of H. aoronymphium did not appear to compromise the ability of these parasites to control Drosophila populations.     

Getting there and around: Host range oscillations during colonization of the Canary Islands by the parasitic nematode Spauligodon

Episodes of expansion and isolation in geographic range over space and time, during which parasites have the opportunity to expand their host range, are linked to the development of host–parasite mosaic assemblages and parasite diversification. In this study, we investigated whether island colonization events lead to host range oscillations in a taxon of host‐specific parasitic nematodes of the genus Spauligodon in the Canary Islands. We further investigated whether range oscillations also resulted in shifts in host breadth (i.e., specialization), as expected for parasites on islands. Parasite phylogeny and divergence time estimates were inferred from molecular data with Bayesian methods. Host divergence times were set as calibration priors after a priori evaluation with a global‐fit method of which individual host–parasite associations likely represent cospeciation links. Parasite colonization history was reconstructed, followed by an estimation of oscillation events and specificity level. The results indicate the presence of four Spauligodon clades in the Canary Islands, which originated from at least three different colonization events. We found evidence of host range oscillations to truly novel hosts, which in one case led to higher diversification. Contemporary host–parasite associations show strong host specificity, suggesting that changes in host breadth were limited to the shift period. Lineages with more frequent and wider taxonomic host range oscillations prior to the initial colonization event showed wider range oscillations during colonization and diversification within the archipelago. Our results suggest that a lineage's evolutionary past may be the best indicator of a parasite's potential for future range expansions.     

Geographical distributions,relative abundance and coexistence of Drosophila aldrichi and Drosophila buzzatii in Australia

Abstract Climatic data and collection records for the cactophilic Drosophila aldrichi and Drosophila buzzatii for 97 localities were used to examine the effects of geographical location, season, host plant species and climatic factors on their range and relative abundance. Temporal variation in relative abundance was assessed from monthly collections over 4 years at one locality. Effects of weather variables over the 28 days before each collection were examined. A generalized linear model of the spatial data showed significant geographical variation in relative abundance, and significant climatic effects, with the proportion of D. aldrichi higher in the warm season, and increasing as temperature variation decreased and moisture indices increased. The temporal data gave generally concordant results, as D. aldrichi proportion was higher in summer and autumn, and increased as maximum and minimum temperatures increased, and as variation in maximum temperature decreased. In a laboratory competition experiment, D. aldrichi eliminated D. buzzatii at 31°C, but was itself eliminated at 18°C and 25°C. The range of D. buzzatii is constrained only by availability of its host plant, Opuntia species, although its relative abundance is reduced in the northern part of its distribution. The range of D. aldrichi, from central Queensland to northern NSW, Australia, is entirely within that of D. buzzatii, and its relative abundance decreases from north to south. Both climate and weather, particularly temperature variability, have direct effects on the relative abundances of the two species, and both likely act indirectly by influencing the outcome of interspecific competition.     

A comparative study of competitive ability between two cactophilic species in their natural hosts

Abstract Competition is a major aspect of the ecology of insect communities exploiting ephemeral and fragmented resources. We analysed the effect of intraspecific (single species culture) and interspecific (mixed species culture) competition on larval viability, developmental time and wing length in the cactophilic Drosophila buzzatii and Drosophila koepferae (Diptera: Drosophilidae) reared in cultured media prepared with fermenting tissues of three common natural cactus hosts in nature at different densities. Our results show that all traits measured were affected by both intra‐ and interspecifc competition, although the effect of competition depended on the Drosophila species and the rearing cactus. In fact, flies tended to have a lower viability, shorter wing size and longer developmental time as a function of increasing density in single species culture in both D. buzzatii and D. koepferae (intraespecific competition). Besides, the performance of both species was seriously affected (shorter body size, slower developmental times, lower viability) by the presence of heterospecific competitors except in the case of D. koepferae reared in its primary host plant, Trichocereus terschekii. We also show that D. koepferae successfully utilized Opuntia quimilo, which is absent in most parts of its distribution range. We discuss the roles of intra‐ and interspecific competition as determinants of the relative abundance of these two species in the arid zones of Southern South America.     

Potential interspecific competition and host specificity: hispine beetles on Heliconia

Abstract. 1. Adults of rolled-leaf hispine beetles (Chrysomelidae, Coleoptera) spend their lives in the scrolls formed by immature leaves of Heliconia (Heli-coniaceae, Monocotyledonae) in Tropical America. As many as eight hispine species can intermingle in the host scrolls at a single site. Scrolls of single host species are invariably occupied by adults of more than one hispine species, and as many as five species can simultaneously occupy one scroll.
2. I made virgin scrolls, which had never contained any insects, by growing leaves under the cover of plastic bags.
3. Thirty-seven combinations of single hispine and Heliconia species were experimentally created in the virgin scrolls, at two sites and in two seasons, in Costa Rica. Combinations included all abundant host species at the sites.
4. All beetles left all leaves of the twenty-eight unnatural beetle—host combinations, within 24 h. On the other hand, none of the nine natural combinations was completely abandoned within the experimental period.
5. Thus, host specificity is not broader in the absence of similar hispine species that might be competitors, and interspecies competition does not affect this obvious aspect of resource utilization for these phytophagous insects.     

Testing for effects of climate change on competitive relationships and coexistence between two bird species

Climate change is expected to have profound ecological effects, yet shifts in competitive abilities among species are rarely studied in this context. Blue tits (Cyanistes caeruleus) and great tits (Parus major) compete for food and roosting sites, yet coexist across much of their range. Climate change might thus change the competitive relationships and coexistence between these two species. Analysing four of the highest-quality, long-term datasets available on these species across Europe, we extend the textbook example of coexistence between competing species to include the dynamic effects of long-term climate variation. Using threshold time-series statistical modelling, we demonstrate that long-term climate variation affects species demography through different influences on density-dependent and density-independent processes. The competitive interaction between blue tits and great tits has shifted in one of the studied sites, creating conditions that alter the relative equilibrium densities between the two species, potentially disrupting long-term coexistence. Our analyses show that long-term climate change can, but does not always, generate local differences in the equilibrium conditions of spatially structured species assemblages. We demonstrate how long-term data can be used to better understand whether (and how), for instance, climate change might change the relationships between coexisting species. However, the studied populations are rather robust against competitive exclusion.     

Interspecific transfer of P elements by crosses between Drosophila simulans and Drosophila mauritiana

Interspecific crosses were carried out between P element-transformed strains of D. simulans and a strain of D. mauritiana, a species devoid of this transposable element family. Four lines were established from hybrid females backcrossed with D. mauritiana males for four generations, and then maintained by intra-line mass mating. In situ hybridization of polytene chromosomes and southern blots showed that full-length and deleted P elements were present in all of the lines after 15 generations. We conclude that at least some of the P elements observed in two lines result from their transposition into D. mauritiana genome. Gonadal sterility, induced at 29°C in D. melanogaster by P elements also occurred with these two latter lines.     

Competitive interactions between two allelopathic algal species: Heterosigma akashiwo and Phaeodactylum tricornutum

ABSTRACT

Both Heterosigma akashiwo and Phaeodactylum tricornutum have been reported to produce allelochemicals capable of inhibiting the growth of co-occurring microalgae. Here, potential allelopathy between H. akashiwo and P. tricornutum was evaluated using bi-algal culture, cell-free culture filtrate and a no-contact co-culturing system in nutrient-replete media. Experiments were also conducted in the no-contact co-culturing system under nutrient-limited conditions. In nutrient-replete bi-algal culture, the growth of P. tricornutum and H. akashiwo each tended to be strongly suppressed when the other species was inoculated at high cell densities. A mathematical model was used to simulate the growth interactions of the two species in bi-algal culture and showed that P. tricornutum outcompeted H. akashiwo over time with different initial cell densities under nutrient-sufficient conditions, indicating that P. tricornutum was more potent in allelopathy. Heterosigma akashiwo growth was inhibited both in the P. tricornutum culture filtrate and no-contact co-culturing system. This confirmed that the extracellular allelopathic compounds released by P. tricornutum were one of the sources of the H. akashiwo growth inhibition. Nutrient deficiency did not increase the extent of allelopathic activity of allelochemicals.     

Coexistence between two fruit fly species is supported by the different strength of intra‐ and interspecific competition

1. Unravelling the strength and modes of interspecific interactions between resident and introduced species is necessary in order to understand the basis of their coexistence or the displacement of the former by the latter. In Argentina, the indigenous Tephritidae fly Anastrepha fraterculus overlaps its distribution and host fruit with the introduced species Ceratitis capitata. 2. This study focused on the relative strength of intra‐ and interspecific competition during the larval stage as a potential factor supporting coexistence. Classical competition experiments (addition and substitution) were conducted between larvae of the two species reared in artificial larval diet. The study evaluated whether a temporal separation between oviposition events affects the outcome of the competition. 3. When both species started to consume the resource at the same time, A. fraterculus experienced a negative effect in larval survival, pupal weight and duration of larval stage, while for C. capitata, pupal weight decreased. When A. fraterculus started feeding 1 day earlier than C. capitata, the negative effects became milder, and when the temporal separation increased, these effects were reversed. Substitution experiments showed an increase in pupal weight when larvae had to share the resource with heterospecific larvae, and showed negative effects suffered for both species when they shared the resource with conspecific individuals. 4. These results suggest that intraspecific competition is stronger than interspecific competition, and a differential oviposition preference could generate an asynchrony of these species in nature. Such mechanisms could favour coexistence between A. fraterculus and C. capitata in an environment previously occupied only by the former.     

Environmental factors affecting patterns of distribution and co‐occurrence of two competing raptor species

Habitat selection is a complex process, that is affected by several factors, including habitat characteristics, environmental conditions, and both intra‐ and interspecific interactions. We analysed habitat preferences of two top avian predators, Peregrine Falcon Falco peregrinus, a medium‐sized diurnal raptor, and Eagle Owl Bubo bubo, a large nocturnal raptor. These two species are known to compete for preferred nest‐sites, and proximity to cliffs with Eagle Owls may reduce Peregrine breeding output through predation of young Falcons. We investigated the environmental factors affecting occurrence and coexistence of the two species and the potential role of habitat suitability in favouring co‐occurrence in 3519 km² of the central pre‐Alps of Italy, where the two species breed on cliffs and sometimes co‐occur on the same cliff. Peregrines settled on long, steep and favourably orientated cliffs in woodland landscapes close to urban areas. Eagle Owls settled on topographically similar cliffs, but in lower rainfall areas compared with cliffs occupied by Peregrines and cliffs unoccupied by either species. Sites where the two species co‐occurred were characterized by more horizontally extended cliffs compared with sites of exclusive occurrence of each species. An analysis of relative habitat suitability revealed that sites where the two species co‐occurred had the highest predicted probability of occupancy for both species, suggesting that those sites should be regarded as high‐quality sites. Breeding productivity of Eagle Owls was negatively affected by the co‐occurrence of Peregrines, whereas the effect of Eagle Owl proximity on Peregrine productivity varied according to cliff suitability for the Peregrines. Habitat selection had fitness consequences for Eagle Owls because breeding productivity increased with cliff length. Environmental conditions, particularly climatic factors, could allow the widespread coexistence of these competing raptors at the landscape scale, whereas at the local scale co‐occurrence could take place only on larger cliffs. These were preferred sites for both species, presumably because breeding at such sites offsets the costs of settling close to the competitor species.