The consequences of multiple indirect pathways of interaction for species coexistence

Theoretical Ecology - Species in diverse communities typically have direct interactions with a small subset of other species, yet indirect effects can be traced between all of the species in a...     

Long run coexistence in the chemostat with multiple species

In this work we analyze the transient behavior of the dynamics of multiple species competing in a chemostat for a single resource, presenting slow/fast characteristics. We prove that coexistence among a subset of species, with growth functions close to each other, can last for a substantially long time. For these cases, we also show that the proportion of non-dominant species can be increasing before decreasing, under certain conditions on the initial distribution.     

Dynamical mechanism for coexistence of dispersing species.

Dispersal of organisms may play an essential role in the coexistence of species. Recent studies of the evolution of dispersal in temporally varying environments suggest that clones differing in dispersal rates can coexist indefinitely. In this work, we explore the mechanism permitting such coexistence for a model of dispersal in a patchy environment, where temporal heterogeneity arises from endogenous chaotic dynamics. We show that coexistence arises from an extreme type of intermittent behavior, namely the phenomenon known as on-off intermittency. In effect, coexistence arises because of an alternation between synchronized and de-synchronized dynamical behaviors. Our analysis of the dynamical mechanism for on-off intermittency lends strong credence to the proposition that chaotic synchronism may be a general feature of species coexistence, where competing species differ only in dispersal rate.     

Stability of a diamond-shaped module with multiple interaction types

Indirect interactions among species emerge from the complexity of ecological networks and can strongly affect the response of communities to disturbances. To determine these indirect interactions and understand better community dynamics, ecologists focused on the interactions within small sets of species or modules. Thanks to their analytical tractability, modules bring insights on the mechanisms occurring in complex interaction networks. So far, most studies have considered modules with a single type of interaction although numerous species are involved in mutualistic and antagonistic interactions simultaneously. In this study, we analyse the dynamics of a diamond-shaped module with multiple interaction types: two resource species sharing a mutualist and a consumer. We describe the different types of indirect interaction occurring between the resource species and the conditions for a stable coexistence of all species. We show that the nature of indirect interactions between resource species (i.e. apparent facilitation, competition or antagonism), as well as stable coexistence, depend on the species generalism and asymmetry of interactions, or in other words, on the distribution of interaction strengths among species. We further unveil that a balance between mutualistic and antagonistic interactions at the level of resource species favours stable coexistence, and that species are more likely to coexist stably if there is apparent facilitation between the two resource species rather than apparent competition. Our results echo existing knowledge on the trophic diamond-shaped module, and confirm that our understanding of communities combining different interaction types can gain from module analyses.     

Maladaptation and mass effects in a metacommunity: consequences for species coexistence

Metacommunity theories predict multispecies coexistence based on the interplay between local species interactions and regional migration. To date, most metacommunity models implicitly assume that evolution can be ignored. Yet empirical studies indicate a substantial potential for contemporary evolution. I evaluate how evolution alters species diversity in a simulated mass-effects (sink-source) metacommunity. Populations inhabiting source habitats became locally adapted, while subordinate competitors became maladapted because of assumed ecological and phenotypic trade-offs between habitats. This maladaptation decreased and leveled relative abundances among subordinate populations. These two effects produced two regions of departure from nonevolutionary predictions. Assuming low proportional migration, maladaptation reduced local species richness via an overall reduction in reproductive rates in sink populations. With intermediate proportional migration, a greater absolute reduction of reproductive rates in intermediate competitors leveled reproductive rates and thereby enhanced local species richness. Although maladaptation is usually viewed as a constraint on species coexistence, simulations suggest that its effects on diversity are manifold and dependent on interpatch migration and community context. Hence, metacommunity predictions often may profit from an evolutionary perspective. Results indicate that modifications of community connectivity, such as might occur during habitat fragmentation, could elicit rapid shifts in communities from regions of high to low biodiversity.     

Occurrence of a winged perispore in a new world species of polypodium

Spores ofPolypodium chnoodes Spreng. were found to possess a large winged perispore surrounding the exospore. This character had hitherto been found only in the closely related families of the aspidioid-dennstaedtioid complex. Its presence in a species ofPolypodium is another example of the remarkable parallelisms which occur in ferns.     

Adaptation of sympatric Achromatium spp. to different redox conditions as a mechanism for coexistence of functionally similar sulphur bacteria

Changes in the abundance of sympatric Achromatium spp. in response to the artificial manipulation of redox conditions in sediment microcosms was determined by fluorescence in situ hybridization (FISH). Adaptation to different redox conditions was shown to be one mechanism that supported the coexistence of functionally similar Achromatium spp. In sediment microcosms, in which the overlying water was oxygenated, Achromatium community size and composition remained unchanged over time. However, imposition of anoxic conditions induced changes in community structure. Anoxia caused a reduction in the relative abundance of Achromatium sp. RY8 (72 +/- 4% to 49 +/- 2%) and an increase in Achromatium sp. RY5 (19 +/- 5% to 32 +/- 3%) and a newly identified Achromatium sp., RYKS (14 +/- 4% to 27 +/- 2%). In anoxic microcosms supplemented with a single addition of nitrate at different initial concentrations the relative decline in Achromatium sp. RY8 was dependent on the initial nitrate concentration. In these experiments nitrate was rapidly removed. In contrast, when high levels of nitrate were maintained by periodic replacement of the overlying water with nitrate supplemented anoxic water, the composition of the Achromatium community remained stable over time. This suggested that all of the coexisting Achromatium spp. are obligate or facultative anaerobes, but, Achromatium sp. RY8 was more sensitive to sediment redox conditions than the other Achromatium species. Given the heterogeneous nature of sedimentary environments, redox-related niche differentiation may promote coexistence of sympatric Achromatium spp.     

Ants on swollen-thorn acacias: species coexistence in a simple system

 On the black cotton soils of the Laikipia ecosystem in Kenya, two swollen-thorn acacia species support nine ant species, four of which are apparently obligate plant-ants. Among the ants, there are five species of Crematogaster, two species of Camponotus, and one each of Tetraponera and Lepisota. Acacia drepanolobium is host to four ant species that are both common and mutually exclusive. These four ant species, and an additional non-exclusive ant species, tend to occur on trees of different sizes, implying a succession of ant occupants. Nonetheless, all four exclusive species occur in substantial proportions on trees of intermediate size. There is direct evidence that an early successional ant species (Tetraponera penzigi) is actively evicted by two late successional ant species in the genus Crematogaster. There was also some evidence of height differentiation among ant species resident on A. seyal. Different acacia-ant species had different direct effects on A. drepanolobium. Extrafloral nectaries were eaten and destroyed only on trees inhabited by Tetraponera. Axillary shoots were eaten only on trees inhabited by C. nigriceps (potentially another early successional ant). This was associated with more new terminal shoots and healthier leaves than other trees, but also the virtual elimination of flowering and fruiting. Different resident acacia-ant species also had characteristic relationships with other insects. Among the four mutually exclusive ant species, only Crematogaster sjostedti was associated with two species of Camponotus, at least one of which (C. rufoglaucus) appears to be a foraging non-resident. A. drepanolobium trees occupied by C. sjostedti were also far more heavily infested with leaf galls than were trees occupied by other ant species. A. drepanolobium trees occupied by C. mimosae and C. sjostedti uniquely had tended adult scale insects. This diversity of ant inhabitants, and their strikingly different relationships with their hosts and other insect species, are examples of coexisting diversity on an apparently uniform resource. Received: 13 November 1995 / Accepted: 16 May 1996     

Vertical habitat segregation as a mechanism for coexistence in sympatric rodents

Coexistence has been widely studied in small mammals and frequently is assumed to be facilitated by habitat segregation. Using live trapping and spool-and-line experiments, we analyzed habitat selection and segregation across multiples scales for Peromyscus leucopus and Ochrotomys nuttalli. At the habitat scale of a forest stand (∼1–100s ha) P. leucopus co-occurred at all sites where O. nuttalli was found, and we did not detect evidence of positive or negative associations or habitat segregation. However, O. nuttalli was restricted to early successional forests, and P. leucopus had significantly lower abundances in early successional forests than in other habitats. We found similar patterns at the mesohabitat scale of the study site (400 m²). O. nuttalli abundance increased with increasing shrub and tree densities, while increases in P. leucopus abundance were associated with open understories. At the microhabitat scale of the individual movement trail, we found vertical segregation. Movement trails for O. nuttalli were at significantly higher elevation (mean height = 142.93 ± 37.10 cm) than P. leucopus trails (mean height = 15.4 ± 4.98 cm; F-value = 35.29, p < 0.001). We concluded that microhabitat segregation was driven by differential use of vertical space for movement and foraging. We suggest O. nuttalli is superior to P. leucopus in its ability to acquire food especially in shrubby subcanopies where few acorns are available. However, P. leucopus is superior in its ability to forage and avoid predators.     

Seeds redistribution in sand dunes: a basis for coexistence of two rodent species

Spatial and temporal heterogeneity is a major factor structuring communities and contributing to coexistence of the species they contain. In this study we examine a critical aspect of environmental heterogeneity that is assumed to promote coexistence in two gerbil species of the Western Negev Desert. Previous studies assumed that temporal partitioning, in activity time, is the result of daily redistribution of seeds that the dominant species is the first to utilize while the sub-ordinate and efficient species is being pushed to use the later and poorer part of the night. We tested the assumption that daily afternoon winds generating spatial and temporal heterogeneity in seed availability by the redistribution of sand and seeds. This was done by comparing plots experiencing normal wind condition with manipulated plots where wind action was diminished by a shade-cloth fence. Our results show that considerable amount of sand and seeds are redistributing regularly on a time scale of a single day. Our results also show that gerbil foraging behavior is strongly related to the pattern of the redistribution dynamics of the seeds. When we prevented redistribution of seeds, gerbil foraging activity was reduced considerably. However, both seed redistribution and gerbil activity did not change much on control plots. Furthermore, the two gerbil species responded differently to the reduction in seed redistribution. The larger Gerbillus pyramidum was shown to be more sensitive to the reduction than the smaller G. a. allenbyi . Daily variability in the availability of seed resources is probably the niche axis which, together with the trade-off in foraging efficiency of the species, forms the mechanism for the coexistence of the two gerbil species in the semi-stabilized sands.     

Eimeria from bats of the world: a new species in Tomopeas ravus from Peru

Two of 17 (12%) crevice bats, Tomopeas ravus (Chiroptera: Vespertilionidae), from Peru had coccidian oocysts in their feces when examined. Sporulated oocysts of Eimeria tomopea n. sp. are ellipsoid to subspheroid, 30.6 X 24.6 (26-34 X 20-28) micron with ovoid sporocysts 13.9 X 9.0 (12-15 X 8-10) micron. A micropyle and substieda body are absent, but polar bodies, oocyst and sporocyst residua, and small, indistinct Stieda bodies are present. The oocyst wall is thick, approximately equal to 1.5 micron and consists of 2 layers, the outermost being rough and mammillated. This is only the 13th eimerian to be described from bats worldwide. The currently recognized species of Eimeria are listed, and a possible New World-Old World phyletic split within the genus is described.     

Cryptosporidium species found in cattle: a proposal for a new species

Humans and animals are infected worldwide by apicomplexan parasites of the genus Cryptosporidium. Yet, parasitologists are continuously surprised by the expanding complexity of this genus. Over the past 20 years, cattle were identified as being a reservoir host for taxa transmitted from animals to humans. However, a remarkable assemblage of species affects cattle, including both cattle-specific, in addition to a zoonotic, species. To clarify species classification, Cryptosporidium pestis n. sp. is proposed for the species formerly recognized as the bovine genotype of C. parvum. The observed increasing complexity of Cryptosporidium species, along with recent advances in knowledge, should be reconsidered in the context of past records, and not vice versa. In this way, the gaps in our understanding of Cryptosporidium species can be identified and addressed in a scientific manner.     

Intraspecific niche flexibility facilitates species coexistence in a competitive community with a fluctuating environment

We analyzed the role of niche usage flexibility (i.e. niche width) in promoting species coexistence in competitive communities in a one-dimensional niche space. We included two types of stochasticity, namely, a random sampling effect of community founding and environmental fluctuation. Fluctuation was further divided into two categories: niche-independent fluctuation (synchronized over the niche space) and niche-dependent fluctuation (variable among individual niche positions). In the analysis, two types of genetic and inheritance systems of individual niche position were considered, i.e. sexual reproduction with multiple loci and asexual reproduction with phenotypic plasticity. We found that niche usage flexibility promoted species diversity only under restricted situations when the environment was constant, but it generally promoted diversity when the environment fluctuated. In particular, under niche-independent fluctuation, niche usage flexibility significantly enhanced species diversity. In contrast, the analysis also predicted that when niche flexibility was constant, species diversity decreased with increasing environmental correlation between neighboring niches.     

Biodiversity in model ecosystems, I: coexistence conditions for competing species

This is the first of two papers where we discuss the limits imposed by competition to the biodiversity of species communities. In this first paper, we study the coexistence of competing species at the fixed point of population dynamic equations. For many simple models, this imposes a limit on the width of the productivity distribution, which is more severe the more diverse the ecosystem is (1994, Theor. Popul. Biol. 45, 227-276). Here we review and generalize this analysis, beyond the "mean-field"-like approximation of the competition matrix used in previous works, and extend it to structured food webs. In all cases analysed, we obtain qualitatively similar relations between biodiversity and competition: the narrower the productivity distribution is, the more species can stably coexist. We discuss how this result, considered together with environmental fluctuations, limits the maximal biodiversity that a trophic level can host.     

Spatial mechanisms for coexistence of species sharing a common natural enemy

We examine the conditions under which spatial structure can mediate coexistence of apparent competitors. We use a spatially explicit, host-parasitoid metapopulation model incorporating local dynamics of Nicholson-Bailey type and global dispersal. Depending on the model parameters, the resulting system displays a plethora of asynchronous dynamical behaviors for which permanent or transient coexistence is observed. We identify a number of spatially mediated tradeoffs which apparent competitors can utilize and demonstrate that the dynamics of spatial coexistence can typically be understood from consideration of two and three patch systems. The phase relationships of species abundances are different for our model than for some other mechanisms of spatial coexistence. We discuss the implications of our findings relative to issues of community organization and biological conservation.     

Dynamics of species coexistence: maintenance of a plant-ant competitive metacommunity

The metacommunity approach is an adequate framework to study coexistence between interacting species at different spatial scales. However, empirical evidence from natural metacommunities necessary to evaluate the predictive power of theoretical models of species coexistence remains sparse. We use two African ant species, Cataulacus mckeyi and Petalomyrmex phylax , symbiotically associated with the myrmecophyte Leonardoxa africana africana , to examine spatio-temporal dynamics of species coexistence and to investigate which environmental and life-history parameters may contribute to the maintenance of species diversity in this guild of symbiotic ants. Using environmental niche partitioning as a conceptual framework, we combined data on habitat variation, social structure of colonies, and population genetics with data from a colonisation experiment and from observation of temporal dynamics. We propose that the dynamics of ant species colonisation and replacement at local and regional scales can be explained by a set of life history traits for which the two ants exhibit hierarchies, coupled with strong environmental differences between the different patches in the level of environmental disturbances. The role of the competition–colonisation tradeoff is discussed and we propose that interspecific tradeoffs for traits related to dispersal and to reproduction are also determinant for species coexistence. We therefore suggest that species-sorting mechanisms are predominant in the dynamics of this metacommunity, but we also emphasise that there may be many ways for two symbionts in competition for the same host to coexist. The results speak in favour of a more complete integration of the various metacommunity models in a single theoretical framework.     

Cachers, scavengers, and thieves: a novel mechanism for desert rodent coexistence

A biologically explicit simulation model of resource competition between two species of seed-eating heteromyid rodent indicates that stable coexistence is possible on a homogeneous resource if harvested food is stored and consumers steal each other's caches. Here we explore the coexistence mechanisms involved by analyzing how consumer phenotypes and presence of a noncaching consumer affect the competitive outcome. Without cache exchange, the winning consumer is better at harvesting seeds and produces more offspring per gram of stored food. With cache exchange, coexistence is promoted by interspecific trade-offs between harvest ability, metabolic efficiency, and ability to pilfer defended caches of heterospecifics or scavenge undefended caches of dead conspecifics or heterospecifics. Cache exchange via pilferage can equalize competitor fitnesses but has little stabilizing effect and leads to stable coexistence only in the presence of a noncaching consumer. In contrast, scavenging is both equalizing and stabilizing and promotes coexistence without a third consumer. Because body size affects a heteromyid rodent's metabolic rate, seed harvest rate, caching strategy, and ability to steal caches, interspecific differences in body size should produce the trade-offs necessary for coexistence. The observation that coexisting heteromyids differ in body size therefore indicates that cache exchange may promote diversity in heteromyid communities.