Population models of sperm-dependent parthenogenesis

Organisms that reproduce by sperm-dependent parthenogenesis are asexual clones that require sperm of a sexual host to initiate egg production, without the genome of the sperm contributing genetic information to the zygote. Although sperm-dependent parthenogenesis has some of the disadvantages of sex (requiring a mate) without the counterbalancing advantages (mixing of parental genotypes), it appears amongst a wide variety of species. We develop initial models for the density-dependent dynamics of animal populations with sperm-dependent parthenogenesis (pseudogamy or gynogenesis), based on the known biology of the common Enchytraeid worm Lumbricillus lineatus. Its sperm-dependent parthenogenetic populations are reproductive parasites of the hermaphrodite sexual form. Our logistic models reveal two alternative requirements for coexistence at density-dependent equilibria: (i) If the two forms differ in competitive ability, the form with the lower intrinsic birth rate must be compensated by a more than proportionately lower competitive impact from the other, relative to intraspecific competition, (ii) If the two forms differ in their intrinsic capacity to exploit resources, the sperm-dependent parthenogen must be superior in this respect and must have a lower intrinsic birth rate. In general for crowded environments we expect a sperm-dependent parthenogen to compete strongly for limiting resources with the sexual sibling species. Its competitive impact is likely to be weakened by its genetic uniformity, however, and this may suffice to cancel any advantage of higher intrinsic growth rate obtained from reproductive investment only in egg production. We discuss likely thresholds of coexistence for other sperm-dependent parthenogens. The fish Poeciliopsis monacha-lucida likewise obtains an intrinsic growth advantage from reduced investment in male gametes, and so its persistence is likely to depend on it being a poor competitor. The planarian Schmidtea polychroa obtains no such intrinsic benefit because it produces fertile sperm, and its persistence may depend on superior resource exploitation.     

Evolution of cross-diffusion and self-diffusion

This article is concerned with the evolution of certain types of density-dependent dispersal strategy in the context of two competing species with identical population dynamics and same random dispersal rates. Such density-dependent movement, often referred to as cross-diffusion and self-diffusion, assumes that the movement rate of each species depends on the density of both species and that the transition probability from one place to its neighbourhood depends solely on the arrival spot (independent of the departure spot). Our results suggest that for a one-dimensional homogeneous habitat, if the gradients of two cross- and self-diffusion coefficients have the same direction, the species with the smaller gradient will win, i.e. the dispersal strategy with the smaller gradient of cross- and self-diffusion coefficient will evolve. In particular, it suggests that the species with constant cross- and self-diffusion coefficients may have competitive advantage over species with non-constant cross- and self-diffusion coefficients. However, if the two gradients have opposite directions, neither of the two dispersal strategies wins as these two species can coexist.     

Competition between the nymphs of two regionally co-occurring species of Notonecta (Hemiptera: Notonectidae)

1. Two species of freshwater invertebrate predator, Notonecta maculata and N. obliqua , showed a negative association in a series of small, man-made ponds in the Peak National Park, Derbyshire, U.K. The present study examines the potential role of interspecific interactions among nymphs on this regional distribution pattern.
2. The survival, development and feeding efficiency of nymphs were examined in laboratory and field mesocosm experiments with intra- and interspecific competition and contrasting environmental complexity.
3. Survival to adulthood and mean lifespan varied significantly in interspecific competition treatments in both laboratory and field experiments, with N . maculata showing higher survival in the simple environment and N. obliqua higher survival in the complex environment.
4. Variations in feeding efficiency were consistent with the survival trends: N. maculata had a higher efficiency in the simple environment, whereas N. obliqua had greater efficiency in the complex environment. There was evidence of a developmental response in feeding efficiency, with differences between species increasing with age.
5. These results suggest that the relative competitive abilities of the two species are affected by habitat complexity, and that competition between species may modify the species distribution where they co-occur.     

Fat-tailed gene flow in the dioecious canopy tree species Fraxinus mandshurica var. japonica revealed by microsatellites

Pollen flow, seed dispersal and individual reproductive success can be simultaneously estimated from the genotypes of adults and offspring using stochastic models. Using four polymorphic microsatellite loci, gene flow of the wind-pollinated and wind-seed-dispersed dioecious tree species, Fraxinus mandshurica var. japonica, was quantified in a riparian forest, in northern Japan. In a 10.5-ha plot, 74 female adults, 76 male adults and 292 current-year seedlings were mapped and genotyped, together with 200 seeds. To estimate dispersal kernels of pollen and seeds, we applied normal, exponential power, Weibull, bivariate t-distribution kernels, and two-component models consisting of two normal distribution functions, one with a small and one with a large variance. A two-component pollen flow model with a small contribution (26.1%) from short-distance dispersal (sigma = 7.2 m), and the rest from long-distance flow (sigma = 209.9 m), was chosen for the best-fitting model. The average distance that integrated pollen flows inside and outside the study plot was estimated to be 196.8 m. Tree size and flowering intensity affected reproduction, and there appeared to be critical values that distinguished reproductively successful and unsuccessful adults. In contrast, the gene flow model that estimated both pollen and seed dispersal from established seedlings resulted in extensive seed dispersal, and the expected spatial genetic structures did not satisfactorily fit with the observations, even for the selected model. Our results advanced small-scale individual-based parentage analysis for quantifying fat-tailed gene flow in wind-mediated species, but also clarified its limitations and suggested future possibilities for gene flow studies.     

Coexistence with similar life cycles in two species of freshwater copepods (Crustacea)

Two species of freshwater cyclopoid copepods, Cyclops scutifer Sars and Mesocyclops leuckarti (Claus), with different ecological requirements, coexist in three humic ponds near the timberline in boreal forests in south-central Norway. The life cycle in both species is basically regulated by complete depletion of oxygen during the long winter. There is a short period of reproduction in spring followed by a rapid development during summer and autumn to advanced instars, capable of diapause when oxygen depletion starts in early winter. Both species have an exclusive 1-yr cycle.The long winter under anoxic conditions is passed in diapause as large copepodids (late developmental instars) with usually no individuals left in the plankton.Even if co-occurring in the same habitats with similar life cycles, there is a temporal and spatial segregation.     

Sex ratio and gender stability in the dioecious plants of Israel

The hypothesis that in dioecious plants genders are usually stable and that the sex ratio is 1:l was tested in the dioecious species of Israel. The flora of Israel comprises some 2500 wild species, among which 43 are dioecious, one gynodioecious and one androdioecious. Forty-one out of these 45 species were examined during 1995-97. These species were examined in 120 populations comprising 7019 individuals, at 81 different localities. The majority of the species (33/41) manifested the expected 1:1 sex ratio. Of the eight species deviating from this ratio, a significant bias was repeatedly scored in five of them. Three other species of the willow family manifested sometimes strongly biased sex ratios or even monomorphic (unisexual) stands, apparently as a result of extensive vegetative cloning. The stability of the male and female morphs was also examined in the 41 species. Only eight individuals (from seven species) out of the 7019 individuals surveyed (0.1%) were found to be bisexual. In addition, in 27 populations belonging to 17 species, 909 males were tagged and revisited in fruiting time, and only three of them (0.3%) were found to set fruits. In addition not a single case of gender reversal between seasons was found. The field survey suggests that in dioecious plants sex determination and expression are usually effectively maintained. The very slight deviations from the 1:1 sex ratio and from the distinct male and female morphs suggest that sexual dimorphism in dioecious plants is usually stabile.     

Permanent coexistence in general models of three interacting species

We address the question of the long term coexistence of three interacting species whose dynamics are governed by the ordinary differential equations x _i = X _i f _i (i = 1, 2, 3). In order for any theory in this area to be useful in practice, it must utilize as little information as possible concerning the forms of the f _i , in view of the great difficulty of determining these experimentally. Here we obtain, under rather general conditions on the equations, a criterion for judging whether the species will coexist in a biologically realistic manner. This criterion depends only on the behaviour near the one or two species equilibria of the two dimensional subsystems, the behaviour there being relatively easy to examine experimentally. We show that with the exception of one class of cases, which is a generalization of a classical example of May and Leonard [21], invasibility at each such equilibrium suitably interpreted is both necessary and sufficient for a strong form of coexistence to hold. In the exceptional case, a single additional condition at the equilibria is enough to ensure coexistence.     

Fire controls population structure in four dominant tree species in a tropical savanna

The persistence of mesic savannas has been theorised as being dependent on disturbances that restrict the number of juveniles growing through the sapling size class to become fire-tolerant trees. We analysed the population structures of four dominant tropical savanna tree species from 30 locations in Kakadu National Park (KNP), northern Australia. We found that across KNP as a whole, the population size structures of these species do not exhibit recruitment bottlenecks. However, individual stands had multimodal size-class distributions and mixtures of tree species consistent with episodic and individualistic recruitment of co-occurring tree species. Using information theory and multimodel inference, we examined the relative importance of fire frequency, stand basal area and elevation difference between a site and permanent water in explaining variations in the proportion of sapling to adult stems in four dominant tree species. This showed that the proportion of the tree population made up of saplings was negatively related to both fire frequencies and stand basal area. Overall, fire frequency has density-dependent effects in the regulation of the transition of saplings to trees in this Australian savanna, due to interactions with stem size, regeneration strategies, growth rates and tree–tree competition. Although stable at the regional scale, the spatiotemporal variability of fire can result in structural and floristic diversity of savanna tree populations.     

雌雄异株树种黄连木种群性比及空间分布

研究了100m×140m固定样地内黄连木种群的性比格局和空间分布。结果表明,调查样地中dbh≥4cm的黄连木植株共有2116株,其中包括526株雌树,1200株雄树,性别未确定植株390株。黄连木性比(雄/雌=2.28)显著偏雄(P0.001)。雌树和雄树平均胸径分别为7.34和7.81cm,雄树胸径显著大于雌树胸径(P0.05)。黄连木幼树、雌树和雄树均呈显著聚集性分布。黄连木幼树与雌树、幼树与雄树均在较小尺度上表现为相互吸引;雌树与雄树则在空间上相互排斥,即雌树与雄树存在空间分离现象;黄连木不同大小以及不同性别植株之间主要表现为相互排斥。     

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.     

Testing mechanisms of coexistence among two species of frugivorous primates

1. We examined mechanisms of coexistence between two congeneric species of frugivorous primates, the blue monkey (Cercopithecus mitis) and the red-tailed monkey (C. ascanius). 2. We used giving-up densities (the amount of food which animals leave in a patch) in fruit trees to measure foraging efficiency and to evaluate possible mechanisms of coexistence. Animals with higher giving-up densities are less likely to persist in the company of those with lower giving-up densities because the former are not able to exploit food patches used previously by the latter. We climbed trees to estimate giving-up densities by counting the fruit which primates left behind. 3. We tested five possible mechanisms of coexistence. Three mechanisms proposed that each frugivorous species has a lower giving-up density than the other in at least one of the following: (1) different tree species, (2) within-tree foraging zones or (3) seasons. The fourth mechanism predicted that the socially dominant species exploits resources first and that the subordinate species has lower giving-up densities. The final mechanism predicted that one species would find resources more quickly than the other, which would in turn have a lower giving-up density. 4. Four of the five mechanisms received no support from our data. Only a trade-off between interspecific dominance and giving-up densities was supported. 5. We discuss the generality of our results and possible interactions with other factors.