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.     

Home range and mating system of two sympatric field mouse species,Apodemus speciosus andApodemus argenteus

The size and spatial distribution of home ranges in two sympatric field mouse species,Apodemus speciosus andApodemus argenteus, were revealed by the capture-recapture method in a temperate deciduous forest from June 1987 to June 1990. InA. speciosus, the home ranges of males were significantly larger tha those of females, and overlapped with those of other males and females during the breeding season. InA. argenteus, the home range of each male overlapped only that of a single female throughout the year. These results suggested thatA. speciosus was promiscuous or polygynous andA. argenteus was monogamous.     

Interspecific competition and coexistence of the two chrysomelids,Gastrophysa atrocyanea motschulsky andGalerucella vittaticollis baly (Coleoptera: Chrysomelidae), under limited food resource conditions

Field experiments were conducted in order to investigate the mode of exploitation of food resources and the mechanism of coexistence of mixed larval populations of the two chrysomelids,Gastrophysa atrocyanea andGalerucella vittaticollis, under limited food resource conditions. The larval survival rates seemed high enough to assure coexistence when hatchlings of the two species were released in 1∶1 and 1∶3 ratios on a host plant. However, the survival rate became almost nil for both species when a 3∶1 ratio was employed, suggesting asymmetrical interspecific competition. Wasted food consumption was much higher inG. atrocyanea larvae. The population ofG. atrocyanea seemed to be regulated more by intraspecific competition, while on the other hand, the population ofG. vittaticollis was considered to be more likely affected by the interspecific competition withG. atrocyanea, depending on the initial ratio and density of the two species.     

Competition and habitat selection in Namib desert tenebrionid beetles

Summary We tested whether intra- and interspecific competition could affect habitat selection in the two most abundant tenebrionid beetles,Physadesmia globosa andOnymacris rugatipennis, in a dry riverbed in the Namib desert. The spatial distributions of these beetles at the microhabitat scale were negatively correlated. We performed a removal experiment, progressively removing first 25% and then a further 25% of the population of the most abundant species,P. globosa, under the trees where most of the preferred food of both species is concentrated. There was no response ofO. rugatipennis to this removal in the tree habitat. In the open, barely-vegetated habitat where mostO. rugatipennis are found, the number of this species caught in pitfall traps increased following both removals and decreased followingP. globosa replacement under the trees. It appears that intraspecific competition forces someP. globosa to occupy the open habitat. Interspecific competition betweenP. globosa andO. rugatipennis in the open habitat reduces the number ofO. rugatipennis that can co-exist withP. globosa there. Removal ofP. globosa under the trees allows conspecifics in the open habitat to move under the trees, releasingO. rugatipennis in the open habitat from competition. This then results in an increase in the numbers ofO. rugatipennis in the open habitat as a result of immigration from neighbouring areas. We found that differences in foraging efficiency, measured as giving-up times in artificial food patches, create a likely mechanism of co-existence that explains the distinct preferences of these two species for tree and open habitats.     

Hoarding behaviors of two wood mouse species: Different preference for acorns of two Fagaceae species

Two species of wood mouse, Apodemus argenteus and A. speciosus, were observed consuming and hoarding acorns of Quercus serrata and Castanopsis cuspidata. When each species of acorn was supplied individually, both species of mice used each species of acorn for eating and hoarding. When both species of acorn were supplied, A. argenteus consumed or hoarded only C. cuspidata, whereas A. speciosus tended to eat C. cuspidata acorns at the feeding site, and disperse or hoard Q. serrata acorns. Apodemus speciosus is unlikely to disperse C. cuspidata acorns (their utilization was biased towards consumption) when Q. serrata acorns are also available. Apodemus argenteus will make almost no contribution to the dispersal of Q. serrata when the two acorn species coexist.     

Amensalism via webs causes unidirectional shifts of dominance in spider mite communities

Competitive displacement is considered the most severe consequence of interspecific competition; if a superior competitor invades the habitat of an inferior species, the inferior species will be displaced. Most displacements previously reported among arthropods were caused by exotic species. The lack of investigation of displacement among native species may be due to their apparently harmonious coexistence, even if it is equivalent to an outcome of interspecific association. A seasonal change in the species composition of spider mites, from Panonychus ulmi to Tetranychus urticae, is observed in apple trees worldwide. Previous laboratory experiments have revealed amensal effects of T. urticae on P. ulmi via their webs. Using manipulation experiments in an orchard, we tested whether this seasonal change in species composition occurred as the result of interspecific competition between these spider mites. Invasion by T. urticae prevented an increase in P. ulmi densities throughout the experimental periods. Degree of overlap relative to the independent distribution on a leaf-surface basis (ω _S) changed from positive to negative with increasing density of T. urticae. T. urticae invasion drove P. ulmi toward upper leaf surfaces (competitor-free space). The niche adjustment by P. ulmi occurred between leaf surfaces but not among leaves. Our findings show that asymmetrical competition between T. urticae and P. ulmi plays an important role in this unidirectional displacement and that the existence of refuges within a leaf produces the apparently harmonious coexistence of the mites and obscures their negative association.     

Expansion of the geographical distribution of an exotic ladybird beetle, Adalia bipunctata (Coleoptera: Coccinellidae), and its interspecific relationships with native ladybird beetles in Japan

The first record of the exotic ladybird beetle, Adalia bipunctata (Coleoptera; Coccinellidae), in Japan was in 1993 at Osaka Nanko Central Park. Since that time, studies on the life history and geographical distribution of A. bipunctata have been ongoing, and its establishment in the Osaka Nanko area has been confirmed. A. bipunctata is a predacious ladybird beetle and a member of a guild that overlaps in habitat and prey with that of native ladybird beetles such as Harmonia axyridis and Menochilas sexmaculatus. We investigated the distribution of A. bipunctata and its interspecific relationships with native predacious ladybird beetles. In some areas, A. bipunctata was dominant in interspecific relationships with native ladybird beetles. For the first 10 years after A. bipunctata was discovered, it occurred only in the Osaka Nanko area, but the present geographical distribution indicates that it has expanded its range. Though the population density of this species was highest at the area recorded first, and tended to decrease in inverse proportion to the distance from Osaka Nanko Central Park, a satellite occurrence was observed in a remote area. The numbers of aphid and tree species (leaf shelter for aestivation and over-wintering) utilized by A. bipunctata have recently increased. Such increases will cause the rate of distribution of A. bipunctata to accelerate. Interspecific competition between H. axyridis and A. bipunctata, which occurs earlier than H. axyridis, may be avoided by desynchronization of the occurrence seasons, and another common predacious ladybird beetle, Coccinella septempunctata, may escape interspecific competition by habitat segregation.     

Habitat segregation in a salt marsh among adult sticklebacks (Gasterosteidae)

Synopsis The spatial distribution, seasonal abundance and diel activity of four sticklebacks,Gasterosteus aculeatus, G. wheatlandi, Pungitius pungitius, andApeltes quadracus. coexisting in a St. Lawrence salt marsh were examined to see how these closely related species share their habitat. While all four species breed in the Riviè des Vases, a tidal creek, only three species are found in the adjacent salt marsh pools,A. quadracus being absent. Results are interpreted in terms of avoidance of interspecific competition for space during the relatively short breeding season at this latitude.     

The life cycle and growth ofIsoperla grammatica andI. difformis (Plecoptera) in southernmost Sweden: intra- and interspecific considerations

Studies on the life cycle and growth ofIsoperia grammatica andI. difformis from eight southern Swedish streams demonstrated that the former species shows greater between-stream differences than the latter species.I. grammatica larvae had typically two periods of rapid growth, autumn and spring, whileI. difformis larvae grew mainly in autumn-winter. In one of the streams, however,I grammatica grew rapidly throughout all winter. Maximum larval size differed between streams, and so did the timing of the first appearance of nymphs and nymphal maturation. Temperature was probably the most influential factor explaining all these differences. While sexual dimorphism inI. grammatica is slight but significant, it is very prominent inI. difformis. The sizes of larvae overlapped little between species as well as sexes suggesting that intra- and interspecific competition was, or had been, important. InI. grammatica this reduced overlap was expressed in several size-classes of males and females successively replacing each other, resulting in a meandering pattern of the size distribution charts. There were additional indications of possible interactions, such as a significant negative correlation between the densities of the two species.     

The dietary basis for temporal partitioning: food habits of coexisting Acomys species

Two rodent species of the genus Acomys coexist on rocky terrain in the southern deserts of Israel. The common spiny mouse (A. cahirinus) is nocturnally active whereas the golden spiny mouse (A. russatus) is diurnally active. An early removal study suggested that competition accounts for this pattern of temporal partitioning: the golden spiny mouse is forced into diurnal activity by its congener. Theoretically, temporal segregation should facilitate coexistence if the shared limiting resources differ at different times (primarily among predators whose prey populations have activity rhythms), or if they are renewed within the period of the temporal segregation. We studied food preferences of the two Acomys species in a controlled cafeteria experiment in order to assess resource overlap and the potential for competition for food between the two species. We found no significant difference in food preferences between species. The dietary items preferred by both were arthropods. We also carried out a seasonal study of the percentage and identity of arthropods taken in the field by individuals of the two species. Individuals of both species took on annual average a high percentage of arthropods in their diets. Seasonal diet shifts reflect seasonal abundance of arthropods at Ein Gedi during day and night. Diurnal activity may also reduce interspecific interference competition between A. russatus and A. cahirinus. However, the strong interspecific dietary overlap in food preference, the heavy reliance on arthropods in spiny mouse diets, and the seasonal and circadian differences in arthropod consumption suggest that prey partitioning may be a viable mechanism of coexistence in this system. Received: 6 July 1998 / Accepted: 10 May 1999     

Relative effects of climate change, isolation and competition on body-size evolution in the Japanese field mouse, Apodemus argenteus

Aim This paper examines body size variation in both recent and Quaternary populations of the Japanese field mouse Apodemus argenteus in order to assess the relative effects on body size of climate change, isolation and competitive interactions with its congeneric A. speciosus. Both temporal (since the Last Glacial Maximum, LGM) and spatial (over the Japanese archipelago) scales are considered. Location The small field mouse is widespread in Japan, and the specimens examined were collected from 10 localities on islands of widely differing area (from 4 km² to 230,510 km²) and at latitudes ranging from 30.3° N to 45.1° N. Methods The effects of geographical factors such as latitude and island area on the size variation of A. argenteus were investigated, using the lower incisor size. In addition, the size of some specimens from two Quaternary localities was compared with the size of the extant specimens. Evolutionary rates of size change since the LGM were calculated in darwins. Hutchinson size ratios were used to examine the pattern of variation of the size segregation between the two Japanese field mice, A. argenteus and A. speciosus, in relation to time and space. Results There was a negative relationship between size and latitude among living A. argenteus populations. In addition, there was no effect of island area on body size, especially at higher latitudes. At lower latitudes, A. argenteus were larger on smaller islands, although this trend was not statistically significant. Quaternary specimens of A. argenteus were smaller in size than their living representatives. The interspecific size ratio between the two Japanese Apodemus was larger on smaller islands and at higher latitudes, and there has been a decrease in the size ratio between the two Apodemus since the LGM. Lastly, in accordance with the theory of character displacement, the small A. argenteus was larger in allopatry than in sympatry, whereas the large A. speciosus was smaller in allopatry than in sympatry. Main conclusions These results indicate that A. argenteus does not conform to Bergmann's rule or to the island rule. The variation in size for the small Japanese field mouse at both spatial and temporal scales may be related to climate change, with an additional effect of competition with the large field mouse, especially on smaller islands. The size convergence between the two Japanese Apodemus observed over the last 21,000 years may be explained by the diminution of available food resources due to the reduction of land mass areas following the LGM. It may also be the result of an evolution towards an optimal body size; a hypothesis previously proposed to explain the evolution of body size in island mammals. Lastly, the evolutionary rates of body size calculated for A. argenteus since the LGM are typical of rates calculated for other Quaternary mainland mammals, thus suggesting that the evolution in this species was not particularly rapid, as is often thought for island mammals.     

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.     

Coexistence of three differentDrosophila species by rescheduling their life history traits in a natural population

We present evidence for coexistence of three differentDrosophila species by rescheduling their life history traits in a natural population using the same resource, at the same time and same place.D. ananassae has faster larval development time (DT) and faster DT(egg-fly) than other two species thus utilizing the resources at maximum at both larval and adult stages respectively. Therefore,D. ananassae skips the interspecific competition at preadult stage but suffers more from intraspecific competition. However,D. melanogaster andD. biarmipes have rescheduled their various life history traits to avoid interspecific competition. Differences of ranks tests for various life history traits suggest that except for DT(egg-pupa), the difference of ranks is highest for the combination ofD. melanogaster andD. ananassae for all other life history traits. This difference is maintained by tradeoffs between larval development time and pupal period and between pupal period and DT(egg-pupa) inD. ananassae.     

The evolution of competing species of terrestrial salamanders: niche partitioning or interference?

Summary One of the central assumptions of evolutionary ecology is that interspecific competition is a potent evolutionary force acting on coexisting species. There are few animal species that provide an opportunity for an experimental analysis of the evolutionary consequences of the phenomenon. We have taken advantage of the fact that two species of terrestrial salamander,Plethodon glutinosus andP. jordani, have different altitudinal distributions on two mountain ranges in North Carolina. Field removal experiments showed that interspecfic competition was much stronger in the Great Smoky Mountains than in the Balsam Mountains, and transplant experiments between the two mountain ranges showed that neither species from the Balsam Mountains had a measurable effect on its congener in the Smokies, although both species from the Smokies had strong negative effects on the Balsam congeners. Other experiments were conducted on the behavioral and ecological changes that have (or have not) evolved in the two areas. Our studies show that increased interspecific interference was the major evolutionary response of these largePlethodon species to interspecific competition, and that partitioning of food or microhabitat was not involved.     

Lack of habitat segregation and no interspecific territoriality in three syntopic cryptic species of the golden‐spectacled warblers Phylloscopus (Seicercus) burkii complex

It is widely accepted that exploitative competition prevents the coexistence of any two or more closely‐related species unless differences exists in their ecological niches and resource use. In sibling bird species, exploitative competition is reduced mainly by spatial segregation of competing species. Spatial segregation can be achieved in two basic ways: by using different habitats or microhabitats that each species is predominately exploiting, or by interspecific territoriality. To our knowledge, either habitat segregation or interspecific territoriality or both have been found in all dyads and groups of sympartic sibling bird species studied so far. In this study, we investigated the coexistence of three cryptic sibling species of the golden‐spectacled warblers Phylloscopus burkii complex in Hunan Province, China: Ph. tephrocephalus, Ph. omeiensis and Ph. valentini. We analyzed their habitat preferences, and spatial interrelations between the species. In order to analyze either presence or absence of interspecific territoriality, we additionally performed con‐ and heterospecific playback experiments. Contrary to expectations, we found no evidences for either habitat segregation or interspecific territoriality in these three species.     

Reproductive seasonality and nest-site differentiation in three closely related armoured catfishes (Siluriformes: Callichthyidae)

Synopsis Three sympatric, closely related armoured catfishes showed a similar, bimodal breeding season in coastal plain swamps in Suriname (South America). The bimodal pattern of reproduction inCallichthys callichthys,Hoplosternum littorale andHoplosternum thoracatum was correlated to the annual distribution of rainfall. Floating bubble nests were constructed throughout the period of swamp inundation (rainy season). Close inspection of the timing of reproduction ofH. littorale in northern South America revealed the relationship between reproduction and rainfall, but also unexpected differences in the length of the breeding season. Strong interspecific competition among the three species was probably avoided through differentiation of nest sites with respect to water depth, distance to the nearest tree, distance to the edge of the swamp, and cover above the nest. Nests ofH. littorale were built in herbaceous swamps, whileC. callichthys andH. thoracatum build their nests in swamp-forest. Nests ofC. callichthys were observed in extremely shallow water or in holes. Few nests of the three species were observed in canals. Over large parts of their geographical rangeC. callichthys andH. thoracatum are found in small rainforest streams. Several aspects of the reproductive ecology and behaviour of both species are probably related to the unpredictability of the stream habitat.     

Food patch structure and plant resource partitioning in interspecific associations of amazonian tamarins

ávila-Pires’ saddle-back tamarins (Saguinus fuscicollis avilapiresi) and red-cap moustached tamarins (S. mystax pileatus), coexisting in highly stable mixed-species groups, overlapped considerably in their use of plant food resources at an Amazonian terra firme forest site. Overlap between food types consumed by the two species was particularly high during periods of lowest fruit availability, when they resorted to a common food supply, primarily the pod exudates of two emergent species of legume trees (Parkia nitida andParkia pendula) and nectar ofSymphonia globulifera. Within-group interspecific competition did not covary with independent measures of resource availability, contrary to predictions based on resource partitioning models. A greater number of both saddle-back and moustached tamarins were able to feed for longer patch residence periods within larger and more productive food patches, whereas small and clumped patches could be monopolized by the socially and numerically dominant moustached tamarins to the physical exclusion of the smaller-bodied saddle-back tamarins. Overall rates of interspecific aggression were extremely low, however, partly because patches that could be monopolized contributed with a minor proportion of either species’ diet. Saddle-backs foraged at lower levels in the understory and encountered smaller food patches more often, whereas moustached tamarins foraged higher and encountered more larger patches in the middle canopy. Although the two species led one another to differently-sized patches, moustached tamarins initiated most feeding bouts and encountered significantly larger and more productive patches that tended to accommodate the entire mixed-species group. Disadvantages of exploitative and interference feeding competition over plant resources, and advantages of shared knowledge of food patches, are but one component of the overall cost-benefit relationship of interspecific associations in tamarins.     

A field test for competitive effects of<Emphasis Type="Italic"> Aedes albopictus</Emphasis> on<Emphasis Type="Italic"> A. aegypti</Emphasis> in South Florida: differences between sites of coexistence and exclusion?

We tested whether interspecific competition from Aedes albopictus had measurable effects on A. aegypti at the typical numbers of larval mosquitoes found in cemetery vases in south Florida. We also tested whether the effect of interspecific competition from A. albopictus on A. aegypti differed between sites where A. aegypti either persists or went extinct following invasion by A. albopictus. Similar experiments manipulating numbers of A. albopictus in cemetery vases were conducted at three sites of A. aegypti persistence and three sites where A. aegypti was apparently extinct. The experiments were done using numbers of larvae that were determined by observed numbers of larvae for each site, and with resources (leaf detritus) that accumulated in experimental vases placed into each field site. In both the early rainy season (when number of mosquito larvae was low) and the late rainy season (when number of mosquito larvae was high), there was a significant effect of treatment on developmental progress of experimental A. aegypti. In the late rainy season, when numbers of larvae were high, there was also a significant effect of treatment on survivorship of A. aegypti. However, the competition treatment × site type (A. aegypti persists vs extinct) interaction was never significant, indicating that the competitive effect of A. albopictus on A. aegypti did not differ systematically between persistence versus extinction sites. Thus, although competition from A. albopictus is strong under field conditions at all sites, we find no evidence that variation in the impact of interspecific competition is associated with coexistence or exclusion. Interspecific competition among larvae is thus a viable explanation for exclusion or reduction of A. aegypti in south Florida, but variation in the persistence of A. aegypti following invasion does not seem to be primarily a product of variation in the conditions in the aquatic environments of cemetery vases.     

Structural differentiation of microhabitats of sympatricmacaca fascicularis andM. nemestrina in East Kalimantan,Indonesia

Macaca nemestrina andM. fascicularis coexist through much of their geographic ranges in Southeast Asia, but locally they segregate into drier hilly terrain and wetter alluvial riverine terrain respectively. SinceM. fascicularis travel arboraally andM. nemestrina, travel terrestrially, structural characteristics of habitats on the two different substrates may explain local segregation of these two species. This paper reports measures comparing habitat structure ofM. fascicularis fascicularis andM. nemestrina nemestrina at the Mentoko site in East Kalimantan, Indonesia. Results show that (1) ground cover is thicker; (2) there are fewer gaps in the canopy; and (3) the foliage of the canopy is more dense in habitat occupied by arborealM. f. fascicularis than in habitat occupied byM. n. nemestrina. These differences in habitat structure appear to explain segregation of the two species at the Mentoko. Similar differences in habitat may permit coexistence of the two species throughout their geographical ranges.