Can the tortoise beat the hare? A possible mechanism for the coexistence of competing mosquitoes in bamboo groves

We studied interspecific competition between the larvae of the two mosquitoesAedes albopictus andTripteroides bambusa, which are predominantly found in water-filled bamboo stumps in northern Kyushu, south-western Japan, using microcosms with dead bamboo leaves in the laboratory. We compared short-term competition between single cohorts of the two species and long-term competition involving four cohorts of each species, which were introduced at 6-day intervals. In the single cohort experiment,A. albopictus grew faster thanT. bambusa. However, in the multiple cohort experiment, although the first cohort ofA. albopictus grew faster and began to pupate earlier than that ofT. bambusa, molting rates of later cohorts ofA. albopictus, that were introduced on the 12th and the 18th day, were lower than those ofT. bambusa. The survival rate ofA. albopictus became lower than that ofT. bambusa after the 18th day. The cumulative number of the pupatedT. bambusa individuals exceeded that ofA. albopictus on the 96th day. The final pupation success was higher inT. bambusa than inA. albopictus, especially when additional leaves were supplied on the 48th and the 96th days. The reversed outcomes between short- and long-term interspecific competition and the variation in the lifespans of small aquatic sites may contribute to the coexistence of the two mosquito species in bamboo groves.     

Distributions of larval mosquitoes among bamboo-stump pools which vary in persistence and resource input

We studied the seasonal occurrence and the distribution patterns of larval mosquitoes in a bamboo grove in northern Kyushu, Japan. The number of pools was large from June to August and was small in winter. Deep stumps and semi-closed-type stumps held water more persistently than shallow and open-type stumps, respectively. Open-type stumps trapped more leaves than semi-closed-type stumps and the number of leaves trapped in the open-type stump was positively correlated with the area of the pool. The incidences ofTripteroides bambusa andAedes albopictus were low early in their breeding season and gradually increased thereafter. In August, the density ofT. bambusa larvae per pool was higher in the old (>2 months) pools than in the new pools and was positively correlated with the depth of the stumps. In contrast, the density ofA. albopictus did not differ significantly between the new-and old pools and was not significantly correlated with the depth of the stump. The density ofA. albopictus was positively correlated with the number of leaves that had been trapped in the stump, whereas the density ofT. bambusa was not. In addition to the seasonal fluctuation in habitat-patch availability, the variations in habitat persistence and resource input among bamboo stumps may facilitate the coexistence of the two mosquitoes that were differentially limited by these factors.     

Variability of intra- and interspecific competitions of bamboo stump mosquito larvae over small and large spatial scales

We carried out field experiments to examine the variability of interspecific competition of mosquito larvae among microcosms in a bamboo grove (small spatial scale) and between bamboo groves at two sites, with single and multiple mosquito species (large spatial scale). Four types of microcosms that differed in capacity and litter input were set. In the hillside bamboo grove, where multiple species occurred, succession of the predominant species from Aedes albopictus to Tripteroides bambusa was observed in control microcosms from which no mosquito larvae were removed. Weekly removal of competitive species resulted in increased pupation of A. albopictus and adult body weight under both rich and poor resource conditions. In the late period of the experiments, the effect of competitor removal on pupation of A. albopictus was greater in deep containers that never dried than in shallow containers that were dried in the laboratory. The number of eggs showed a slight difference between competitor‐excluded and deep control microcosms. These results indicate that interspecific competition limits pupation of A. albopictus more strongly in deep containers than in shallow and drought‐prone containers.
Compared with the hillside site, the larval density of A. albopictus attained a higher density in the bamboo grove in the plain where no competitive species occurred, due to a higher oviposition rate. Lower rate of pupation and lower adult weight at the plain site than at the hillside site indicated that resource limitation was more severe at the plain site. Populations of A. albopictus at hillside and plain sites appeared to suffer from strong inter‐ and intraspecific competition, respectively. At the hillside site, the intensity of interspecific competition appeared to increase later in the breeding season, with a high larval density of T. bambusa. In contrast, at the plain site, intensity of intraspecific competition appeared to be reduced later in the breeding season with decreasing larval density of A. albopictus.     

Habitat stability and the larval mosquito community in treeholes and other containers on a temperate Island

The temporal fluctuation of water levels and the presence of mosquito larvae were investigated for four types of small container habitats (treeholes, bamboo stumps, cemetery stone vases, and cemetery stone vessels) on Kabeshima Island in Kyushu, southwestern Japan. The probability that containers held water was positively correlated with the quantity of the preceding rainfall and with the depth and volume of the containers. It was estimated that dehydration occurred more regularly in autumn and winter than in summer. The probability that mosquito larvae were present in each type of container was positively correlated with habitat stability in terms of the probability of the existence of standing water and the coefficient of variation of the water level. Twelve species of mosquito larvae, including two rare predators, were found. Species composition differed between the different types of container. Although 4 to 10 species used each type of container, the median number of species per container was two for treeholes and bamboo stumps, and one for the others. The dominant species wasTripteroides bambusa in treeholes and bamboo stumps,Aedes albopictus in stone vases, andA. japonicus in stone vessels. The larval mosquito community, which lacked major predators, possessed the following features that may facilitate the coexistence of many species: (1) niche segregation amongst species in terms of their selection of container types; (2) an aggregated distribution of the individual species among containers of the same type; (3) high intraspecific mean crowding (and hence probably intense intraspecific competition) in the dominant species in each type of container; (4) independent species associations within the same type of container; and (5) low interspecific mean crowding (and hence probablynot intense interspecific competition) between species in the same type of container.     

Bamboo stumps as mosquito larval habitats in Darjeeling Himalayas, India: A spatial scale analysis

Bamboo stumps can be a congenial breeding habitat of the mosquitoes. In view of this, a preliminary assessment of the dipteran immatures inhabiting the stumps of bamboo groves in the Darjeeling Himalayas was carried out at a spatial scale. Of the 104 stumps of Dendrocalamus hamiltoni surveyed, 70 were found to host immatures of three dipteran species, the mosquitoes Aedes aegypti and Culex quinquefasciatus and the midges Chironomus sp. in varying densities. Though the stumps varied in diameter, in each stump on average 12. 1 immatures were found. The abundance of the immatures was positively correlated with the diameter of the stumps (r=+0.382; P < 0.001) but negatively with the pH of the water present in the stumps (r=–0.336; P < 0.01). The coefficient of association was found to be +8.4 for the Ae. aegypti and Chironomus immatures, while in the rest of the species pair the association seemed to be independent. Thus it can be concluded that the stumps in the bamboo groves of Darjeeling Himalayas provides a favourable habitat for the mosquito and chironomid immatures.     

Searching clusters of community composition along multiple spatial scales: a case study on aquatic invertebrate communities in bamboo stumps in West Timor

We examined spatial heterogeneity at multiple scales in composition of the aquatic invertebrate communities in bamboo stumps in a mountainous area of West Timor. We partitioned the study area (ca. 15,000 m²) at five levels of patchiness: two sites, four sub-sites, eight super-clumps, 14 clumps, and 84 stumps. Similarity of community composition between stumps varied more than expected from independent occurrence of each taxon in comparisons within any levels of patches. Negative association was frequently detected among taxa. These results indicate heterogeneity in community composition at a stump level. At higher levels, similarity among stumps within each site was greater than expected from null models which assumed no spatial heterogeneity, and similarities among super-clumps, sub-sites and between sites in a whole area were lower than expected from the null models. The observed patterns in similarities among subsets of the community and distribution of each taxon were mostly consistent with the models which assumed site-level heterogeneity. Therefore, we conclude that the community in this area was spatially heterogeneous at stump and site levels. The relationship between mean intra- and inter-specific crowding suggested that the site level habitat heterogeneity might reduce the chance of encounters between two predators, the larvae of the Toxorhynchites mosquito and the Brachyceran fly.     

Distribution of <Emphasis Type="Italic">Rigidoporus lignosus</Emphasis> genotypes in a rubber plantation,as revealed by somatic compatibility

A study was conducted to determine the spatial distribution of somatic incompatibility groups (SIGs) in Rigidoporus lignosus populations responsible for white root disease in a south Sumatera rubber plantation. Most isolates colonizing the stumps were somatically compatible with isolates from either a nearby stump or a living rubber tree, which suggested evidence of stump-to-tree or stump-to-stump clonal growth of the fungus. SIGs of R. lignosus occupied an area territorially in which the minimum size estimate for the largest genotype was 25m in length.     

Chronological location analyses of giant bamboo (Phyllostachys pubescens) groves and their invasive expansion in a satoyama landscape area,western Japan

Since the 1980s, giant bamboo groves consisting mainly of naturalized Phyllostachys pubescens have been expanding in Japanese rural landscapes (satoyama). Using aerial photographs, I evaluated changes in bamboo grove distribution between 1947 and 2006 at Mount Hachiman, Ohmi‐hachiman City in western Japan, where there is an area of satoyama landscape adjacent to the suburbs. I also quantitatively determined the natural and artificial factors that accelerated bamboo grove expansion using a multiple logistic regression analysis with stepwise procedure. I attempted to describe the positions of bamboo groves in Japanese satoyama landscapes. The area of bamboo groves in the survey area expanded after 1967. The bamboo groves are located at the base of mountain slopes and on alluvial fans adjacent to residential areas, where they are most susceptible to human interferences (mainly disturbance). They mark the boundary between agricultural fields and secondary forests. The model analysis indicated that the abandonment of lands adjacent to bamboo groves caused bamboo‐grove expansion. As forests and agricultural land were abandoned, bamboo, mainly P. pubescens, invaded these areas by rhizomatous clonal growth. Bamboo grove expansion in Japanese satoyama landscapes is dependent on the type of land use, topography, and existing vegetation as well as the invasive life‐history traits of P. pubescens.     

Use of mires and food habits of sika deer in the Oze Area,central Japan

Sika deer Cervus nippon entered the Oze Area, a snowy area in central Japan, in the 1990s. The use of mires and the food habits of this species were studied in 1999 and 2000. Deer used the mires immediately after snow melted and did not use the mires in mid summer, although they did use them lightly in autumn. The summer food habits differed from those of the deer population living in Nikko in the lower area: non-Sasa graminoids and other monocotyledonous leaves occupied more (approximately 30%) of the feces in the former population, whereas Sasa nipponica, a dwarf bamboo, was the dominant food (60%) of the latter population. Seasonal changes in fecal composition in the Oze deer were that mire graminoids occupied a considerable portion of their food in spring (11.6%) and summer (17.0%), whereas the amount of dwarf bamboo increased in fall (26.0%). Despite the small size of the mires, Oze deer appear to prefer mires to forests. Nitrogen concentrations of the mire plants did not differ from those of the forest plants; however, foraging efficiency would be greater in the mires and this may explain the preference observed. Despite the small total biomass, leaf biomass of the mire was equivalent to that of the adjacent forest floor. In addition, biomass was concentrated near the ground, or the biomass concentration was greater than that of the forest floor.     

Chromosomal localization and genomic organization of cloned repetitive DNA fragments in mosquitoes (Diptera: Culicidae)

The chromosomal localization and genomic organization of three cloned repetitive DNA fragments (viz., H-76, H-61, and H-19) isolated from theAedes albopictus genome have been examined inAe. albopictus and six otherAedes species:Ae. aegypti, Ae. seatoi, Ae. flavopictus, Ae. polynesiensis, Ae. alcasidi andAe. katherinensis. The results fromin situ and Southern hybridization analyses show that the sequences homologous to cloned repetitive DNA fragments are dispersed throughout the genome in each species. The sequences homologous to these cloned repetitive DNA fragments are also found inHaemagogus equinus, Tripteroides bambusa andAnopheles quadrimaculatus and are dispersed in their genomes. Data indicate divergence in the amount and the structural organization of sequences homologous to these cloned fragments among mosquito species.     

A simple method to estimate the rate of the bamboo expansion based on one-time measurement of spatial distribution of culms

Moso bamboo (Phyllostachys edulis) groves have been expanding in Japanese forests, thus an assessment of the rate of bamboo grove expansion is a critical issue. Previous studies used time-series data which span typically more than a few years in order to determine the expansion of bamboo groves. This method cannot be used when time-series data were not available. In this study, we developed and tested a simple indicator based on one-time measurement of spatial distribution of culms to estimate the rate of bamboo grove expansion. Among 19 locations at four sites, the distribution patterns of culms from the pure bamboo stands to the expanding front were highly correlated with the actual edge expansion rate (E) based on the aerial photographs collected over ca. 15- and 30-year periods. In contrast, the culm density in the original stand itself was not correlated with E. These results suggest that one-time measurement of the distribution patterns of living culms (but not stand density) could be a useful indicator for estimating the expansion rates of bamboo groves.     

Seasonal migration patterns of female sika deer in eastern Hokkaido,Japan

Fifty-seven female sika deer (Cervus nippon yesoensis), captured at the wintering area in the Shiranuka Hills in eastern Hokkaido, Japan, were radio-tracked during 1997–2001 to examine the factors affecting seasonal migration at the individual-landscape level. Ten of the 57 deers migrated between low-altitude summer home ranges and intermediate-altitude winter home ranges (the upward migrants). Twenty-nine migrated between high-altitude summer home ranges and intermediate-altitude winter home ranges (the downward migrants). Twelve used the intermediate-altitude home ranges all year round (the non-migrants). The remaining six were unknown. The summer home ranges of deer were widely scattered over an area of 5734km². Migration distances ranged between 7.2 and 101.7km. Deer showed high site fidelities to their seasonal home ranges. The upward migrants wintered in areas of less snow, higher quality of bamboo grass, and more coniferous cover than their summer home ranges. The downward migrants wintered in areas of less snow, higher quality of bamboo grass, higher winter temperature, and more southern slopes, but less coniferous cover than their summer home ranges. The non-migrants used year-round ranges with little snow, high quality of bamboo grass, and sufficient coniferous cover. We suggest that snow cover and bamboo grass are the factors affecting seasonal migration of the population and that coniferous cover is another factor for the upward migration.     

Spatial patterns in pupal mortality in urban palm caterpillars

Mortality factors were determined in over 20000 pupal skins of Brassolis sophorae and Opsiphanes invirae, collected from 55 sites in the urban area of Campinas, São Paulo, southeast Brazil. Almost 50% of B. sophorae pupae and 30% of O. invirae pupae were killed by various identifiable factors, which may have an appreciable influence in determining adult population size. Mortality was related to type of attachment site, distance from the center of the city, area occupied by buildings and houses, vegetation cover, and other urban variables. Exposed pupation sites such as outside walls had a lower frequency of successful eclosions than darker or more sheltered sites, such as offset skirting boards, garages, or mailboxes, which act as partial refuges against parasitoids. Mortality levels were higher in regions of the city with high vegetation cover. Vegetation cover and distance from the city center were the best predictors of parasitism level of B. sophorae by the chalcidid wasp Spilochalcis morleyi. The viability of S. morleyi (data from parasitized B. sophorae) did not differ significantly among different regions of the city. These data do not support an exclusive model of escape from parasites (or other natural enemies) for explaining the abundance of urban butterflies.This paper was presented at the VI INTECOL, Manchester, UK, in September 1994     

Effects of capacity on resource input and the aquatic metazoan community structure in phytotelmata

Aquatic metazoan communities in treeholes and bamboo stumps were studied on Tsushima Island, Japan, in March and June in relation to the habitat size and the resouce amount. The amount of leaf detritus increased with an increase in the capacity (volume) of the phytotelma. The input rate of leaves depended upon the opening area of the phytotelma. Twenty-one metazoan inhabitant taxa consisted of 17 dipteran saprophages, a helodid beetle larva, a nematoda, a naidid, and a predator,Toxorhynchites towadensis mosquito larva. Treeholes harbored 17 taxa and bamboo stumps 20; 16 taxa were common in both phytotelmata. The number of taxa and the biomass of metazoa per phytotelma was positively correlated with the capacity. The densities of dominant saprophages such as helodid, chironomid, andCulicoides larvae were positively correlated with the capacity of phytotelmata, and the variation in their abundance contributed much to the variation in the species composition in individual phytotelmata. With an increase in capacity, treeholes harbored more helodids andCulicoides, whereas bamboo stumps harbored more chironomids andDasyhelea. Thus, the amount of resource, species richness and biomass per phytotelma largely depended upon the capacity, and the difference in community structure between treeholes and bamboo stumps was attributable to the differential colonization patterns of different taxa.     

Identification of novel LTR retrotransposons in the genome of Aedes aegypti

We have detected seventy-six novel LTR retrotransposons in the genome of the mosquito Aedes aegypti by a genome wide analysis using the LTR_STRUC program. We have performed a phylogenetic classification of these novel elements and a distribution analysis in the genome of A. aegypti. These mobile elements belong either to the Ty3/gypsy or to the Bel family of retrotransposons and were not annotated in the mosquito LTR retrotransposon database (TEfam). We have found that  1.8% of the genome is occupied by these newly detected retrotransposons that are distributed predominantly in intergenic genomic sequences and introns. The potential role of retrotransposon insertions linked to host genes is described and discussed. We show that a retrotransposon family belonging to the Osvaldo lineage has peculiar structural features, and its presence is likely to be restricted to the A. aegypti and to the Culex pipiens quinquefasciatus genomes. Furthermore we show that the ninja-like group of elements lacks the Primer Binding Site (PBS) sequence necessary for the replication of retrotransposons. These results integrate the knowledge on the complicate genomic structure of an important disease vector.     

Conservation of a highly repeated DNA family of Aedes albopictus among mosquito genomes (Diptera: Culicidae)

Summary The genomic organization and chromosomal localization of a cloned 0.79-kb highly repeated DNA fragment, H-115, isolated from Aedes albopictus has been examined. The cloned fragment is a part of a larger unit of 1.86 kb that is tandemly repeated in the Ae. albopictus genome. The H-115 family of sequences are located at the intercalary position on chromosome 1 in Ae. albopictus. Similar patterns of in situ and Southern blot hybridization results are obtained in Ae. aegypti, Ae. seatoi, Ae. flavopictus, Ae. polynesiensis, Ae. Alcasidi, and Ae. katherinensis. The H-115 sequences are widely conserved in Culicidae and are found in Haemagogus equinus, Tripteroides bambusa, and Anopheles quadrimaculatus by hybridization under high stringency conditions. The H-115 sequences are also tandemly repeated in Hg. equinus with a monomer unit of 1.86 kb and in Tp. bambusa with a slightly diverged monomer unit of 1.90kb. In Anopheles quadrimaculatus, the H-115 sequences are dispersed throughout the genome. Partial sequence analysis shows that the H-115 insert is 62% AT and contains two perfect inverted repeats and numerous perfect direct repeats. The occurrence of inverted repeats with potential to form intrastrand palindromic structure suggests that the H-115 family of sequences may be involved in chromatin condensation.     

Delineating fine-scale genetic units in amphibians:Probing the primacy of ponds

The population structure of pond-breeding amphibians is shaped by their distinct breeding foci, but it is unclear to what extent this is reflected in the fine-scale distribution of genetic diversity. We used microsatellite genotypes to investigate the genetic signatures of 24 populations of European newts, Triturus cristatus and T. marmoratus, inhabiting 21 ponds in a confined study area (7.5×3.5km) in western France. Employing a Bayesian clustering approach based on individual genotypes that minimises departures from Hardy–Weinberg equilibrium and linkage disequilibrium, no evidence was found for within-pond substructuring. Subjecting all sampled ponds simultaneously to this procedure revealed a clear signal of partitioning, with the most likely number of clusters however below the actual number of ponds (seven in T. cristatus, three in T. marmoratus). A more hierarchical Bayesian approach, with pond as analysis unit, was achieved to separate ponds from genetically more meaningful units, and reduced the T. cristatus populations to 11 clusters, and the T. marmoratus populations to five clusters. We were unable to specify a minimum nearest-neighbour distance where ponds are separate units, probably due to both historical and current demographic processes. The implications for strategies to manage and conserve endangered amphibians in human-altered landscapes are discussed.     

Effects of deer,mice and dwarf bamboo on the emergence,survival and growth of <Emphasis Type="Italic">Abies homolepis</Emphasis> (Piceaceae) seedlings

In the subalpine mixed forest of Mt. Ôdaigahara, mid-western Japan, the understory is dominated by dwarf bamboo (Sasa nipponica), which is the major forage of overly populous sika deer (Cervus nippon). In the present study, we monitored the survival and growth of Abies homolepis seedlings over 5years to determine how they responded to the experimental removal of dwarf bamboo and to the exclusion of sika deer and mice (Apodemus argenteus and A.speciosus). Deer and dwarf bamboo reduced the survival of seedlings but had different effects on growth. The stems of seedlings were shorter in the presence of deer, indicating that taller seedlings were apt to be browsed by deer, whereas the diameters of seedlings were smaller in the presence of dwarf bamboo, mainly owing to its shading effect. The presence of mice decreased the number of seedlings germinating in a particular site, but had no effect on seedling survival after germination. There was no significant indirect effect whereby the survival of seedlings was predicted to be facilitated by the decreased biomass of bamboo because of grazing by deer. We supposed that this might be because the direct negative effect of deer was so large as to conceal the positive indirect effect.     

How mosquitoes fly to man

1. A model is presented to estimate the mosquito density and radius (r) of the attraction range of man as the bait.
2. Man walked to a spot in the woods and sampled mosquitoes flying to him with a net continuously. The arrival pattern was constant in each mosquito species, and in all species, the number of mosquitoes sampled per five min became constant after ten minutes sampling.
3. The mosquitoes gathered to man were composed of D: those waiting alight, I: those intruding the attraction range and F: those following man to the spot. They were estimated by means of continuous sampling sets and densities of D and I were calculated.
4. r s of female and male of A. albopictus and A. japonicus were 4.0, 5.6 and 9.0 m respectively. In these species, most individuals were D, although, in T. bambusa all individuals were I, flying to search for hosts.
5. How mosquitoes and hosts encounter, is discussed.

     

Utilized and potentially utilizable seaweeds on the Namibian coast: biogeography and accessibility

Of the 1500 km coast of Namibia, only 80 km is predominantly rocky, the remaining 1420 km being sandy with only minor rocky outcrops. At present two species are utilized, Gracilaria verrucosa for agar and Laminaria schinzii for human consumption. Other potentially utilizable seaweeds occurring on the coast are Ecklonia maxima for alginates, Suhria vittata for agar, Gigartina radula, G. stiriata and Aeodes orbitosa for carrageenans and Porphyra capensis for human consumption. Laminaria schinzii also can be used for alginate production. Due to the diamond-mining areas around the 80 km rocky area at Luderitz (26° 39 S), only 20 km are accessible; hence, at present the seaweeds are being exploited almost to their full potential. The species currently not being utilized could be used, however, to support the industry that already exists at Luderitz. To increase production, access to the diamond areas would have to be gained or a cultivation program initiated.