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Kinds of Species Pairs Selected AbstractsPatterns of interspecific associations of stem gallers on willowsDIVERSITY AND DISTRIBUTIONS, Issue 6 2003Jens-Peter Kopelke Abstract., The pattern of interspecific associations of three stem-galling sawfly species (Euura atra, E. elaeagnos, E. purpureae) and three stem-galling gallmidge species (Rabdophaga sp. 3,5) was investigated on five willow taxa (Salix alba, S. fragilis, S. × rubens, S. elaeagnos, S. purpurea) at five natural sites in Central Europe. The willow species harboured specific species associations of two stem gallers, each pair consisting of one Euura and one Rabdophaga species. The stem gallers were patchily distributed and their densities varied significantly among willow host plant species, host plant individuals, and host plant sexes. Four of the six species showed a significant increase in galling rate with shoot length. The other two species were the sawfly and cecidomyiid pair that induce galls on S. purpurea. The preference of stem gallers to longer shoots was generally not related to higher larval performance in terms of survival. Only one species, Rabdophaga sp. 5, was found to be more abundant on male plants. The correlation of densities of the species pairs of stem gallers was independent of willow sexes. Species pairs of stem gallers co-occurring on the same willow species tended to attack different shoots within the same host plant individual. When species pairs co-occurred on shoots they were usually found in similar densities as when occurring alone on shoots. The stem-galling sawflies usually formed galls at the basal part of a shoot, whereas the gallmidge R. sp. 5 (R. sp. 3 and R. sp. 4 showed no clear tendency) preferred the middle or distal part of a shoot. This is interpreted with differences of their phenology and oviposition period. [source] Satellite species in lampreys: a worldwide trend for ecological speciation in sympatry?JOURNAL OF FISH BIOLOGY, Issue 2 2003V. Salewski Amongst several theories of speciation, sympatric speciation has been the most controversial but it is now widely accepted that populations can become reproductively isolated without being separated geographically. One problem with the acceptance of the theory of sympatric speciation, however, has been the lack of supporting empirical data and it is still believed that geographical isolation is responsible for the majority of speciation events. Here the example of species pairs in lampreys suggests that sympatric speciation in a whole taxonomic group could occur throughout its worldwide range. Lampreys occur in two ecologically distinct forms: parasitic mostly anadromous species that forage on tissue and body fluids of host fishes, and non-parasitic forms that, apart from a short adult life when they cease feeding, spend their entire life as filter feeders in the substratum of stream beds. Both forms occur in sympatric species pairs throughout the range of lampreys that occur in Eurasia, North America and Australia and it is widely acknowledged that non-parasitic forms derive from parasitic forms. The larvae of both forms can be distinguished by their potential fecundity and therefore, it is argued that the mode of life is not a consequence of different ecological conditions. Furthermore, as lampreys prefer to choose mates of similar sizes and fertilization success decreases with increasing difference in body size, there is a strong disruptive selection between the two forms and they are therefore reproductively isolated. Besides theoretical aspects, the similarity of the species pairs, together with their occurrence in sympatry, the occurrence of forms with intermediate characteristics, and examples where speciation might be in progress, hints at the possibility that speciation also occurred in sympatry. The difference between lampreys and other examples of sympatric speciation is that there seems to be a trend towards sympatric speciation events throughout the worldwide range of lampreys which is neither restricted to relatively small localities nor caused by human disturbance. Species pairs in lampreys therefore offer a unique possibility of studying the process of sympatric speciation on a large scale. [source] Null model analysis of communities on gradientsJOURNAL OF BIOGEOGRAPHY, Issue 6 2004James G. Sanderson Abstract Aim, I employed a novel null model and metric to uncover unusual species co-occurrence patterns in a herpetofaunal assemblage of 49 species collected at discrete elevations along a gradient. Location, Mount Kupe, Cameroon. Methods, Using a construction algorithm that started from a matrix of 0s, a sample null space of 25,000 unique null matrices was generated by simultaneously conserving (1) the number of occurrences of each species, (2) site richness and (3) species range spans derived from the observed incidence matrix. I then compared the number of times each pair of confamilial species co-occurred in the null space with the same number derived from the observed incidence matrix. Two cases dealing with embedded absences in species ranges were tested: (1) embedded absences were maintained, and (2) embedded absences were assumed to be sampling omissions and were replaced by presences. Results, In the observed absence/presence assemblage there were 147 possible confamilial species pairs. Therefore, 5% or eight were expected by chance alone to have co-occurrence patterns that differed from chance expectations by chance alone. Of these confamilial species pairs, 38 were congeneric and so 5% or two were expected to differ from chance expectations. For case (1) 16, and for case (2) 17 confamilial species pairs' co-occurrence patterns differed significantly from chance expectations. For case (1) nine congeneric species pairs, and for case (2) 10 congeneric pairs differed significantly from chance expectations. For case (1) four, and for case (2) five congeneric species pairs formed checkerboards (patterns of mutual exclusion). Results from case (1) were a proper subset of case (2) indicating that sampling omissions did not alter greatly the results. Main conclusions, I have demonstrated that null models are valuable tools to analyse ecological communities provided that proper models are employed. The choice of the appropriate null space to analyse distributions is critical. The null model employed to analyse birds on islands of an archipelago can be adapted to analyse species along gradients provided an additional range constraint is added to the null model. Moreover, added precision to results can be obtained by analysing each species pair separately, particularly those in the same family or genus, as opposed to applying a community-wide metric to the faunal assemblage. My results support some of the speculations of previous authors who were unable to demonstrate their suspicions analytically. [source] Genetic and ecological divergence of a monophyletic cichlid species pair under fully sympatric conditions in Lake Ejagham, CameroonMOLECULAR ECOLOGY, Issue 6 2001Ulrich Schliewen Abstract Although there is mounting evidence that speciation can occur under sympatric conditions, unambiguous examples from nature are rare and it is almost always possible to propose alternative allopatric or parapatric scenarios. To identify an unequivocal case of sympatric speciation it is, therefore, necessary to analyse natural settings where recent monophyletic species flocks have evolved within a small and confined spatial range. We have studied such a case with a cichlid species flock that comprises five Tilapia forms endemic to a tiny lake (Lake Ejagham with a surface area of approximately 0.49 km2) in Western Cameroon. Analysis of mitochondrial D-Loop sequences shows that the flock is very young (approximately 104 years) and has originated from an adjacent riverine founder population. We have focused our study on a particular pair of forms within the lake that currently appears to be in the process of speciation. This pair is characterized by an unique breeding colouration and specific morphological aspects, which can serve as synapomorphic characters to prove monophyly. It has differentiated into a large inshore and a small pelagic form, apparently as a response to differential utilization of food resources. Still, breeding and brood care occurs in overlapping areas, both in time and space. Analysis of nuclear gene flow on the basis of microsatellite polymorphisms shows a highly restricted gene flow between the forms, suggesting reproductive isolation between them. This reproductive isolation is apparently achieved by size assortative mating, although occasional mixed pairs can be observed. Our findings are congruent with recent theoretical models for sympatric speciation, which show that differential ecological adaptations in combination with assortative mating could easily lead to speciation in sympatry. [source] Compounds with a `stuffed' anti-bixbyite-type structure, analysed in terms of the Zintl,Klemm and coordination-defect conceptsACTA CRYSTALLOGRAPHICA SECTION B, Issue 1 2009Angel Vegas The bixbyite structure (Mn2O3) () is often described as a distorted face-centered cubic (f.c.c.) array of Mn atoms, with O atoms occupying 3/4 of the tetrahedral holes. The empty M4 tetrahedra are centred at 16c. In anti-bixbyite structures (Mg3N2), cation vacancies are centred in empty N4 tetrahedra. If 16 hypothetical atoms were located at this site they would form the structure of ,-Si. This means that anti-bixbyite structures are ideally prepared to accommodate Si(Ge) atoms at these holes. Several compounds (Li3AlN2 and Li3ScN2) fully satisfy this expectation. They are really anti-bixbyites `stuffed' with Al(Sc). The presence of these atoms in 16c is illuminated in the light of the extended Zintl,Klemm concept (EZKC) [Vegas & García-Baonza (2007). Acta Cryst. B63, 339,345], from which a compound would be the result of `multiple resonance' pseudo-structures, emerging from electron transfers between any species pair (like or unlike atoms, cations or anions). The coordination-defect (CD) concept [Bevan & Martin (2008). J. Solid State Chem.181, 2250,2259] is also consistent with the EZKC description of the pseudo-structures. A more profound insight into crystal structures is gained if one is not restricted to the contemplation of classical anions and cations in their conventional oxidation states. [source] Patterns of interspecific associations of stem gallers on willowsDIVERSITY AND DISTRIBUTIONS, Issue 6 2003Jens-Peter Kopelke Abstract., The pattern of interspecific associations of three stem-galling sawfly species (Euura atra, E. elaeagnos, E. purpureae) and three stem-galling gallmidge species (Rabdophaga sp. 3,5) was investigated on five willow taxa (Salix alba, S. fragilis, S. × rubens, S. elaeagnos, S. purpurea) at five natural sites in Central Europe. The willow species harboured specific species associations of two stem gallers, each pair consisting of one Euura and one Rabdophaga species. The stem gallers were patchily distributed and their densities varied significantly among willow host plant species, host plant individuals, and host plant sexes. Four of the six species showed a significant increase in galling rate with shoot length. The other two species were the sawfly and cecidomyiid pair that induce galls on S. purpurea. The preference of stem gallers to longer shoots was generally not related to higher larval performance in terms of survival. Only one species, Rabdophaga sp. 5, was found to be more abundant on male plants. The correlation of densities of the species pairs of stem gallers was independent of willow sexes. Species pairs of stem gallers co-occurring on the same willow species tended to attack different shoots within the same host plant individual. When species pairs co-occurred on shoots they were usually found in similar densities as when occurring alone on shoots. The stem-galling sawflies usually formed galls at the basal part of a shoot, whereas the gallmidge R. sp. 5 (R. sp. 3 and R. sp. 4 showed no clear tendency) preferred the middle or distal part of a shoot. This is interpreted with differences of their phenology and oviposition period. [source] Predicting competition coefficients for plant mixtures: reciprocity, transitivity and correlations with life-history traitsECOLOGY LETTERS, Issue 4 2001R.P. Freckleton There are few empirical or theoretical predictions of how per capita or per individual competition coefficients for pairs of plant species should relate to each other. In contrast, there are a considerable number of general hypotheses that predict competitive ability as a function of a range of ecological traits, together with a suite of increasingly sophisticated models for competitive interactions between plant species. We re-analyse a data set on competition between all pairwise combinations of seven species and show that competition coefficients relate strongly to differences between the maximum sizes, root allocation, emergence time and seed size of species. Regressions suggest that the best predictor of competition coefficients is the difference in the maximum size of species and that correlations of the other traits with the competition coefficients occur through effects on the maximum size. We also explore the patterns of association between coefficients across the competition matrix. We find significant evidence for coefficient reciprocity (inverse relationships between the interspecific coefficients for species pairs) and transitivity (numerically predictable hierarchies of competition between species) across competition matrices. These results therefore suggest simple null models for plant community structure when there is competition for resources. [source] THE BIOLOGY OF SPECIATIONEVOLUTION, Issue 2 2010James M. Sobel Since Darwin published the "Origin," great progress has been made in our understanding of speciation mechanisms. The early investigations by Mayr and Dobzhansky linked Darwin's view of speciation by adaptive divergence to the evolution of reproductive isolation, and thus provided a framework for studying the origin of species. However, major controversies and questions remain, including: When is speciation nonecological? Under what conditions does geographic isolation constitute a reproductive isolating barrier? and How do we estimate the "importance" of different isolating barriers? Here, we address these questions, providing historical background and offering some new perspectives. A topic of great recent interest is the role of ecology in speciation. "Ecological speciation" is defined as the case in which divergent selection leads to reproductive isolation, with speciation under uniform selection, polyploid speciation, and speciation by genetic drift defined as "nonecological." We review these proposed cases of nonecological speciation and conclude that speciation by uniform selection and polyploidy normally involve ecological processes. Furthermore, because selection can impart reproductive isolation both directly through traits under selection and indirectly through pleiotropy and linkage, it is much more effective in producing isolation than genetic drift. We thus argue that natural selection is a ubiquitous part of speciation, and given the many ways in which stochastic and deterministic factors may interact during divergence, we question whether the ecological speciation concept is useful. We also suggest that geographic isolation caused by adaptation to different habitats plays a major, and largely neglected, role in speciation. We thus provide a framework for incorporating geographic isolation into the biological species concept (BSC) by separating ecological from historical processes that govern species distributions, allowing for an estimate of geographic isolation based upon genetic differences between taxa. Finally, we suggest that the individual and relative contributions of all potential barriers be estimated for species pairs that have recently achieved species status under the criteria of the BSC. Only in this way will it be possible to distinguish those barriers that have actually contributed to speciation from those that have accumulated after speciation is complete. We conclude that ecological adaptation is the major driver of reproductive isolation, and that the term "biology of speciation," as proposed by Mayr, remains an accurate and useful characterization of the diversity of speciation mechanisms. [source] EVOLUTION OF REPRODUCTIVE ISOLATION WITHIN A PARASITIC FUNGAL SPECIES COMPLEXEVOLUTION, Issue 7 2007Mickael Le Gac Despite important advances in the last few years, the evolution of reproductive isolation (RI) remains an unresolved and critical gap in our understanding of speciation processes. In this study, we investigated the evolution of RI among species of the parasitic fungal species complex Microbotryum violaceum, which is responsible for anther smut disease of the Caryophyllaceae. We found no evidence for significant positive assortative mating by M. violaceum even over substantial degrees of genetic divergence, suggesting a lack of prezygotic isolation. In contrast, postzygotic isolation increased with the genetic distance between mating partners when measured as hyphal growth. Total RI, measured as the ability of the pathogen to infect and produce a diploid progeny in the host plant, was significantly and positively correlated with genetic distance, remaining below complete isolation for most of the species pairs. The results of this study, the first one on the time course of speciation in a fungus, are therefore consistent with previous works showing that RI generally evolves gradually with genetic distance, and thus presumably with time. Interestingly, prezygotic RI due to gamete recognition did not increase with genetic distance, in contrast to the pattern found in plants and animals. [source] CALIBRATING A MOLECULAR CLOCK FROM PHYLOGEOGRAPHIC DATA: MOMENTS AND LIKELIHOOD ESTIMATORSEVOLUTION, Issue 10 2003Michael J. Hickerson Abstract We present moments and likelihood methods that estimate a DNA substitution rate from a group of closely related sister species pairs separated at an assumed time, and we test these methods with simulations. The methods also estimate ancestral population size and can test whether there is a significant difference among the ancestral population sizes of the sister species pairs. Estimates presented in the literature often ignore the ancestral coalescent prior to speciation and therefore should be biased upward. The simulations show that both methods yield accurate estimates given sample sizes of five or more species pairs and that better likelihood estimates are obtained if there is no significant difference among ancestral population sizes. The model presented here indicates that the larger than expected variation found in multitaxa datasets can be explained by variation in the ancestral coalescence and the Poisson mutation process. In this context, observed variation can often be accounted for by variation in ancestral population sizes rather than invoking variation in other parameters, such as divergence time or mutation rate. The methods are applied to data from two groups of species pairs (sea urchins and Alpheus snapping shrimp) that are thought to have separated by the rise of Panama three million years ago. [source] PERSPECTIVE: GENE DIVERGENCE, POPULATION DIVERGENCE, AND THE VARIANCE IN COALESCENCE TIME IN PHYLOGEOGRAPHIC STUDIESEVOLUTION, Issue 6 2000ScottV. Abstract Molecular methods as applied to the biogeography of single species (phylogeography) or multiple codistributed species (comparative phylogeography) have been productively and extensively used to elucidate common historical features in the diversification of the Earth's biota. However, only recently have methods for estimating population divergence times or their confidence limits while taking into account the critical effects of genetic polymorphism in ancestral species become available, and earlier methods for doing so are underutilized. We review models that address the crucial distinction between the gene divergence, the parameter that is typically recovered in molecular phylogeographic studies, and the population divergence, which is in most cases the parameter of interest and will almost always postdate the gene divergence. Assuming that population sizes of ancestral species are distributed similarly to those of extant species, we show that phylogeographic studies in vertebrates suggest that divergence of alleles in ancestral species can comprise from less than 10% to over 50% of the total divergence between sister species, suggesting that the problem of ancestral polymorphism in dating population divergence can be substantial. The variance in the number of substitutions (among loci for a given species or among species for a given gene) resulting from the stochastic nature of DNA change is generally smaller than the variance due to substitutions along allelic lines whose coalescence times vary due to genetic drift in the ancestral population. Whereas the former variance can be reduced by further DNA sequencing at a single locus, the latter cannot. Contrary to phylogeographic intuition, dating population divergence times when allelic lines have achieved reciprocal monophyly is in some ways more challenging than when allelic lines have not achieved monophyly, because in the former case critical data on ancestral population size provided by residual ancestral polymorphism is lost. In the former case differences in coalescence time between species pairs can in principle be explained entirely by differences in ancestral population size without resorting to explanations involving differences in divergence time. Furthermore, the confidence limits on population divergence times are severely underestimated when those for number of substitutions per site in the DNA sequences examined are used as a proxy. This uncertainty highlights the importance of multilocus data in estimating population divergence times; multilocus data can in principle distinguish differences in coalescence time (T) resulting from differences in population divergence time and differences in T due to differences in ancestral population sizes and will reduce the confidence limits on the estimates. We analyze the contribution of ancestral population size (,) to T and the effect of uncertainty in , on estimates of population divergence (,) for single loci under reciprocal monophyly using a simple Bayesian extension of Takahata and Satta's and Yang's recent coalescent methods. The confidence limits on , decrease when the range over which ancestral population size , is assumed to be distributed decreases and when increases; they generally exclude zero when /(4Ne) > 1. We also apply a maximum-likelihood method to several single and multilocus data sets. With multilocus data, the criterion for excluding = 0 is roughly that l/(4Ne)> 1, where l is the number of loci. Our analyses corroborate recent suggestions that increasing the number of loci is critical to decreasing the uncertainty in estimates of population divergence time. [source] A molecular approach to detect hybridisation between crucian carp (Carassius carassius) and non-indigenous carp species (Carassius spp. and Cyprinus carpio)FRESHWATER BIOLOGY, Issue 3 2005B. HÄNFLING Summary 1. Releases of non-native fish into the wild is an increasing problem posing considerable ecological and genetic threats through direct competition and hybridisation. 2. We employed six microsatellite markers to identify first generation hybrids and backcrosses between native crucian carp (Carassius carassius) and introduced goldfish (C. auratus) and common carp (Cyprinus carpio) in the U.K. We also investigated the genetic characteristics of the taxonomically controversial gibel carp (Carassius spp.) from sites across Europe. 3. Natural hybridisation between goldfish and crucian carp occurs frequently, although hybrids between all other species pairs were observed. Only 62% of British crucian carp populations (n = 21) consisted exclusively of pure crucian carp. In some populations hybrids were so frequent, that no pure crucian carp were caught, indicating a high competitive ability of hybrids. 4. Most hybrids belonged to the F1 generation but backcrossing was evident at a low frequency in goldfish × crucian carp hybrids and goldfish × common carp hybrids. Furthermore, some local populations had high frequencies of backcrosses, raising the opportunity for introgression. 5. Gibel carp from Germany and Italy belonged to two triploid clonal lineages that were genetically closely related to goldfish, whereas all individuals identified from British populations proved to be crucian carp × goldfish hybrids. 6. Our study suggests that the release of closely related exotic cyprinids not only poses a threat to the genetic integrity and associated local adaptations of native species, but may also contribute to shifts in community structure through competitive interactions. [source] Competition in variable environments: experiments with planktonic rotifersFRESHWATER BIOLOGY, Issue 6 2002KEVIN L. KIRK 1.,In a constant environment, competition often tends to reduce species diversity. However, several theories predict that temporal variation in the environment can slow competitive exclusion and allow competing species to coexist. This study reports on laboratory competition experiments in which two pairs of planktonic rotifer species competed for a phytoplankton resource under different conditions of temporal variability in resource supply. 2.,For both species pairs, Keratella cochlearis dominated under all conditions of temporal variability, and the other species (Brachionus calyciflorus or Synchaeta sp.) almost always went extinct. Increasing temporal variation in resource supply slowed competitive exclusion but did not change competitive outcome or allow coexistence. 3.,Rotifers show a gleaner,opportunist trade-off, because gleaner species have low threshold resource levels (R*) and low maximum population growth rates, while opportunist species have the opposite characteristics. In the competition experiments, the gleaner always won and the opportunists always lost. Thus, a gleaner,opportunist trade-off was not sufficient to facilitate coexistence under conditions of resource variability. Instead, the winning species had both the lowest R* and the greatest ability to store resources and ration their use during times of extreme resource scarcity. [source] Geographical range size heritability: what do neutral models with different modes of speciation predict?GLOBAL ECOLOGY, Issue 3 2007David Mouillot ABSTRACT Aim, Phylogenetic conservatism or heritability of the geographical range sizes of species (i.e. the tendency for closely related species to share similar range sizes) has been predicted to occur because of the strong phylogenetic conservatism of niche traits. However, the extent of such heritability in range size is disputed and the role of biology in shaping this attribute remains unclear. Here, we investigate the level of heritability of geographical range sizes that is generated from neutral models assuming no biological differences between species. Methods, We used three different neutral models, which differ in their speciation mode, to simulate the life-history of 250,000 individuals in a square lattice of 50 × 50 cells. These individuals can speciate, reproduce, migrate and die in the metacommunity according to stochastic events. We ran each model for 3000 steps and recorded the range size of each species at each step. The heritability of geographical range size was assessed using an asymmetry coefficient between range sizes of sister species and using the coefficient of correlation between the range sizes of ancestors and their descendants. Results, Our results demonstrated the ability of neutral models to mimic some important observed patterns in the heritability of geographical range size. Consistently, sister species exhibited higher asymmetry in range sizes than expected by chance, and correlations between the range sizes of ancestor,descendant species pairs, although often weak, were almost invariably positive. Main conclusions, Our findings suggest that, even without any biological trait differences, statistically significant heritability in the geographical range sizes of species can be found. This heritability is weaker than that observed in some empirical studies, but suggests that even here a substantial component of heritability may not necessarily be associated with niche conservatism. We also conclude that both present-day and fossil data sets may provide similar information on the heritability of the geographical range sizes of species, while the omission of rare species will tend to overestimate this heritability. [source] Mixed-species bird flocks in dipterocarp forest of north-central Burma (Myanmar)IBIS, Issue 4 2001DAVID I. KING We studied the bird community in deciduous, dipterocarp forest of north-central Burma (Myanmar) during December 1994, March 1996, and January 1997 and 1999. Most members of this community participated in mixed-species flocks. Seventy-three flocks were encountered during our study, containing 52 species. Of these, 25 species occurred in more than 10% of flocks, and were included in our analyses. There were 26 significant correlations among species pairs, 25 of which were positive. Cluster analysis indicated that there were three principal types of flocks: one consisting mostly of small passerines and picids, commonly including Common Wood-Shrike, Small Minivet and White-browed Fantail, among others; a second type consisting mainly of sylviids, e.g. Arctic, Dusky and Radde's Warblers; and a third type which generally centred around Greater and Lesser Necklaced Laughingthrushes. Bird-eating hawks were numerous at these sites, and we witnessed several attacks on flocks during the study. Thus we infer that enhanced protection from predation is an important benefit conferred by flock membership. In contrast, there was little overlap in foraging behaviour among species, suggesting that foraging facilitation is a relatively minor benefit enjoyed by flock members, although we did observe White-browed Fantails and Greater Racket-tailed Drongos kleptoparasitizing other species on occasion. [source] Are ectoparasite communities structured?JOURNAL OF ANIMAL ECOLOGY, Issue 6 2006Species co-occurrence, null models, temporal variation Summary 1We studied temporal variation in the structure of flea communities on small mammalian hosts from eastern Slovakia using null models. We asked (a) whether flea co-occurrences in infracommunities (in the individual hosts) in different hosts as well as in the component communities (in the host species) demonstrate a non-random pattern; (b) whether this pattern is indicative of either positive or negative flea species interactions; (c) whether this pattern varies temporally; and (d) whether the expression of this pattern is related to population size of either fleas or hosts or both. 2We constructed a presence/absence matrix of flea species for each temporal sample of a host species and calculated four metrics of co-occurrence, namely the C -score, the number of checkerboard species pairs, the number of species combinations and the variance ratio (V -ratio). Then we compared these metrics with the respective indices calculated for 5000 null matrices that were assembled randomly using two algorithms, namely fixed-fixed (FF) and fixed-equiprobable (FE). 3Most co-occurrence metrics calculated for real data did not differ significantly from the metrics calculated for simulated matrices using the FF algorithm. However, the indices observed for 42 of 75 presence/absence matrices differed significantly from the null expectations for the FE models. Non-randomness was detected mainly by the C -score and V -ratio metrics. In all cases, the direction of non-randomness was the same, namely the aggregation, not competition, of flea species in host individuals and host species. 4The inclusion or exclusion of the uninfested hosts in the FE models did not affect the results for individual host species. However, exclusion of the uninfested host species led to the acceptance of the null hypothesis for only six of 13 temporal samples of the component flea communities for which non-randomness was detected when the uninfested hosts were included in the analysis. 5In most host species, the absolute values of the standardized size effect of both the C -score and V -ratio increased with an increase in host density and a concomitant decrease in flea abundance and prevalence. 6Results of this study demonstrated that (a) flea assemblages on small mammalian hosts were structured at some times, whereas they appeared to be randomly assembled at other times; (b) whenever non-randomness of flea co-occurrences was detected, it suggested aggregation but never segregation of flea species in host individuals or populations; and (c) the expression of structure in flea assemblages depended on the level of density of both fleas and hosts. [source] Migration strategies of sylviid warblers: chance patterns or community dynamics?JOURNAL OF AVIAN BIOLOGY, Issue 1 2000Peter Howlett The effects of community dynamics in birds on the optimisation of their migratory strategies is a neglected area. For three years, we captured migrating warblers on autumn passage at a coastal site in western Britain. We used canonical correspondence analysis (CCA) to assess spatio-temporal patterns of occurrence, and principal components analysis (PCA) to assess morphological variation. We calculated Euclidean distance in ordination and morphological space to assess separation between species pairs, and used Monte-Carlo simulations to assess the probability of pattern occurring by chance. Ordination revealed five species-groups separated by habitat type and time of passage. Reed Warbler Acrocephalus scirpaceus and Sedge Warbler A. schoenobaenus (Group 1) occurred in wet habitats and peaked simultaneously. In drier habitats with scrub, a first wave of Blackcap Sylvia atricapilla (Group 2) significantly preceded Grasshopper Warbler Locustella naevia, Willow Warbler Phylloscopus trochilus, Whitethroat Sylvia communis and Lesser Whitethroat Sylvia curruca (Group 3), which in all but one case (Lesser Whitethroat) significantly preceded Garden Warbler Sylvia borin (Group 4); peak numbers of Chiffchaffs Phylloscopus collybita and a second wave of Blackcaps (Group 5) occurred later still. Age effects were found only in Acrocephalus, with adults peaking before juveniles. For seven out of eight pairings within genera, separation in time of passage increased significantly in species that were morphologically similar. The only exception was Blackcap and Lesser Whitethroat which differed substantially in both passage time and morphology. Monte-Carlo simulations showed that chance was unlikely to be responsible for ordination patterns, nor for inter-specific variation in passage time and its relationship with species morphology. These data provide annually consistent evidence that migrating sylviid warblers are separated ecologically by habitat use, time of passage and morphology: we cannot refute the hypothesis that community dynamics have influenced niche use and autumn migratory strategy. We call for further tests of the ,migrant interaction' hypothesis in other geographical locations and taxa, particularly where migrants are allopatric and interact ecologically only on migration. [source] A probability-based analysis of temporal and spatial co-occurrence in grassland birdsJOURNAL OF BIOGEOGRAPHY, Issue 12 2006Joseph A. Veech Abstract Aim, To test for non-random co-occurrence in 36 species of grassland birds using a new metric and the C -score. The analysis used presence,absence data of birds distributed among 305 sites (or landscapes) over a period of 35 years. This analysis departs from traditional analyses of species co-occurrence in its use of temporal data and of individual species' probabilities of occurrence to derive analytically the expected co-occurrence between paired species. Location, Great Plains region, USA. Methods, Presence,absence data for the bird species were obtained from the North American Breeding Bird Survey. The analysis was restricted to species pairs whose geographic ranges overlapped. Each of 541 species pairs was classified as a positive, negative, or non-significant association depending on the mean difference between the observed and expected frequencies of co-occurrence over the 35-year time-span. Results, Of the 541 species pairs that were examined, 202 to 293 (37,54%) were positively associated, depending on which of two null models was used. However, only a few species pairs (<5%) were negatively associated. An additional 89 species pairs did not have overlapping ranges and hence represented de facto negative associations. The results from analyses based on C -scores generally agreed with the analyses based on the difference between observed and expected co-occurrence, although the latter analyses were slightly more powerful. Main conclusions, Grassland birds within the Great Plains region are primarily distributed among landscapes either independently or in conjunction with one another. Only a few species pairs exhibited repulsed or segregated distributions. This indicates that the shared preference for grassland habitat may be more important in producing coexistence than are negative species interactions in preventing it. The large number of non-significant associations may represent random associations and thereby indicate that the presence/absence of other grassland bird species may have little effect on whether a given focal species is also found within the landscape. In a broader context, the probability-based approach used in this study may be useful in future studies of species co-occurrence. [source] Null model analysis of communities on gradientsJOURNAL OF BIOGEOGRAPHY, Issue 6 2004James G. Sanderson Abstract Aim, I employed a novel null model and metric to uncover unusual species co-occurrence patterns in a herpetofaunal assemblage of 49 species collected at discrete elevations along a gradient. Location, Mount Kupe, Cameroon. Methods, Using a construction algorithm that started from a matrix of 0s, a sample null space of 25,000 unique null matrices was generated by simultaneously conserving (1) the number of occurrences of each species, (2) site richness and (3) species range spans derived from the observed incidence matrix. I then compared the number of times each pair of confamilial species co-occurred in the null space with the same number derived from the observed incidence matrix. Two cases dealing with embedded absences in species ranges were tested: (1) embedded absences were maintained, and (2) embedded absences were assumed to be sampling omissions and were replaced by presences. Results, In the observed absence/presence assemblage there were 147 possible confamilial species pairs. Therefore, 5% or eight were expected by chance alone to have co-occurrence patterns that differed from chance expectations by chance alone. Of these confamilial species pairs, 38 were congeneric and so 5% or two were expected to differ from chance expectations. For case (1) 16, and for case (2) 17 confamilial species pairs' co-occurrence patterns differed significantly from chance expectations. For case (1) nine congeneric species pairs, and for case (2) 10 congeneric pairs differed significantly from chance expectations. For case (1) four, and for case (2) five congeneric species pairs formed checkerboards (patterns of mutual exclusion). Results from case (1) were a proper subset of case (2) indicating that sampling omissions did not alter greatly the results. Main conclusions, I have demonstrated that null models are valuable tools to analyse ecological communities provided that proper models are employed. The choice of the appropriate null space to analyse distributions is critical. The null model employed to analyse birds on islands of an archipelago can be adapted to analyse species along gradients provided an additional range constraint is added to the null model. Moreover, added precision to results can be obtained by analysing each species pair separately, particularly those in the same family or genus, as opposed to applying a community-wide metric to the faunal assemblage. My results support some of the speculations of previous authors who were unable to demonstrate their suspicions analytically. [source] Inter-ocean dispersal is an important mechanism in the zoogeography of hakes (Pisces: Merluccius spp.)JOURNAL OF BIOGEOGRAPHY, Issue 6 2001W. Stewart Grant Aim To present new genetic data and to review available published genetic data that bear on the phylogeny of hakes in the genus Merluccius. To construct a zoogeographical model from a summary phylogenetic tree with dated nodes. To search for an explanation of antitropical distributions in hakes. To assess peripheral isolate, centrifugal and vicariance models of speciation in view of the molecular phylogeny and zoogeography of hakes. Locations Northern and southern Atlantic Ocean, eastern Pacific Ocean, South Pacific Ocean. Methods Electrophoretic analysis of 20 allozyme loci in 10 species of hakes. Phylogenetic tree construction with parsimony and bootstrap methods. Reanalysis of previous genetic data. Analysis of zoogeographical patterns with geographical distributions of molecular genetic markers. Results Phylogenetic analyses of new and previous allozyme data and previous mitochondrial DNA data indicate a deep genetic partition between Old- and New-World hakes with genetic distances corresponding to 10,15 Myr of separation. This time marks a widening rift between Europe and North America and a rapid drop in ocean temperatures that subdivided an ancestral population of North Atlantic hake. Two Old-World clades spanning the equator include pairs of sister taxa separated by tropical waters. Divergence times between these pairs of sister-taxa variously date to the early Pliocene and late Pleistocene. Amongst New-World hakes, pairs of sister taxa are separated by equatorial waters, by the Southern Ocean, and by the Panama Isthmus. These genetic separations reflect isolation by the rise of the Isthmus 3,4 Ma and by Pliocene and Pleistocene dispersals. Pairs of species occurring in sympatry or parapatry in six regions do not reflect sister-species relationships, but appear to reflect allopatric divergence and back dispersals of descendent species. Some geographically isolated regional populations originating within the last few hundreds of thousands of years merit subspecies designations. Conclusions Vicariance from tectonic movement of continental plates or ridge formation cannot account for the disjunct distributions of most hake sister taxa. Molecular genetic divergences place the origin of most hake species diversity in the last 2,3 Myr, a period of negligible tectonic activity. Distributions of many hake species appear to have resulted from dispersals and back dispersals across both warm equatorial waters and cool waters in the Southern Ocean, driven by oscillations in climate and ocean temperatures. Genetic and ecological divergence prevents hybridization and competitive exclusion between sympatric species pairs in six regions. Sister-taxa relationships and estimates of divergence are consistent with the modified peripheral isolate model of speciation in which vicariances, range expansions and contractions, dispersals and founder events lead to isolated populations that subsequently diverge to form new species. [source] Ecological selection against hybrids in natural populations of sympatric threespine sticklebacksJOURNAL OF EVOLUTIONARY BIOLOGY, Issue 6 2007J. L. GOW Abstract Experimental work has provided evidence for extrinsic post-zygotic isolation, a phenomenon unique to ecological speciation. The role that ecological components to reduced hybrid fitness play in promoting speciation and maintaining species integrity in the wild, however, is not as well understood. We addressed this problem by testing for selection against naturally occurring hybrids in two sympatric species pairs of benthic and limnetic threespine sticklebacks (Gasterosteus aculeatus). If post-zygotic isolation is a significant reproductive barrier, the relative frequency of hybrids within a population should decline significantly across the life-cycle. Such a trend in a natural population would give independent support to experimental evidence for extrinsic, rather than intrinsic, post-zygotic isolation in this system. Indeed, tracing mean individual hybridity (genetic intermediateness) across three life-history stages spanning four generations revealed just such a decline. This provides compelling evidence that extrinsic selection plays an important role in maintaining species divergence and supports a role for ecological speciation in sticklebacks. [source] Genetic divergence between sympatric Arctic charr Salvelinus alpinus morphs in Gander Lake, Newfoundland: roles of migration, mutation and unequal effective population sizesJOURNAL OF FISH BIOLOGY, Issue 8 2008D. Gomez-Uchida A suite of 10 microsatellite loci was used to examine genetic divergence between two sympatric morphs of Arctic charr Salvelinus alpinus (,dark' and ,pale') inhabiting Gander Lake, Newfoundland. Results can be summarized as follows: (1) the morphs are strongly reproductively isolated , gene flow,migration estimates were consistently low in long and short-term evolutionary timescales of analysis; (2) intermorph divergence based on allele size (RST) was significantly larger than those based on allele state (,) implying a cumulative effect of stepwise-like mutations; (3) historical (coalescent) and current (linkage disequilibrium) point estimates of effective population size (Ne) were consistently higher for dark than for pale S. alpinus. The first and second findings lend support to the hypothesis that divergence between forms may have preceded the last glacial period (ending c. 12 000 years bp). The third finding argues for significant differences in habitat quantity and quality between morphs, which were emphasized in a previous study. Overall, these analyses underscore the importance of genetic assessment and monitoring in the conservation of fish diversity, with emphasis on ,rare' or under-represented forms among temperate species pairs. [source] Satellite species in lampreys: a worldwide trend for ecological speciation in sympatry?JOURNAL OF FISH BIOLOGY, Issue 2 2003V. Salewski Amongst several theories of speciation, sympatric speciation has been the most controversial but it is now widely accepted that populations can become reproductively isolated without being separated geographically. One problem with the acceptance of the theory of sympatric speciation, however, has been the lack of supporting empirical data and it is still believed that geographical isolation is responsible for the majority of speciation events. Here the example of species pairs in lampreys suggests that sympatric speciation in a whole taxonomic group could occur throughout its worldwide range. Lampreys occur in two ecologically distinct forms: parasitic mostly anadromous species that forage on tissue and body fluids of host fishes, and non-parasitic forms that, apart from a short adult life when they cease feeding, spend their entire life as filter feeders in the substratum of stream beds. Both forms occur in sympatric species pairs throughout the range of lampreys that occur in Eurasia, North America and Australia and it is widely acknowledged that non-parasitic forms derive from parasitic forms. The larvae of both forms can be distinguished by their potential fecundity and therefore, it is argued that the mode of life is not a consequence of different ecological conditions. Furthermore, as lampreys prefer to choose mates of similar sizes and fertilization success decreases with increasing difference in body size, there is a strong disruptive selection between the two forms and they are therefore reproductively isolated. Besides theoretical aspects, the similarity of the species pairs, together with their occurrence in sympatry, the occurrence of forms with intermediate characteristics, and examples where speciation might be in progress, hints at the possibility that speciation also occurred in sympatry. The difference between lampreys and other examples of sympatric speciation is that there seems to be a trend towards sympatric speciation events throughout the worldwide range of lampreys which is neither restricted to relatively small localities nor caused by human disturbance. Species pairs in lampreys therefore offer a unique possibility of studying the process of sympatric speciation on a large scale. [source] Environmental and neighbourhood effects on tree fern distributions in a neotropical lowland rain forestJOURNAL OF VEGETATION SCIENCE, Issue 1 2007Mirkka M. Jones Abstract Questions: To what extent are the distributions of tropical rain forest tree ferns (Cyatheaceae) related to environmental variation, and is habitat specialization likely to play a role in their local coexistence? Location: Lowland rain forest at La Selva Biological Station, Costa Rica. Methods: Generalized linear (GLM) and generalized additive (GAM) logistic regression were used to model the incidence of four tree fern species in relation to environmental and neighbourhood variables in 1154 inventory plots regularly distributed across 6 km2 of old-growth forest. Small and large size classes of the two most abundant species were modelled separately to see whether habitat associations change with ontogeny. Results: GLM and GAM model results were similar. All species had significant distributional biases with respect to micro-habitat. Environmental variables describing soil variation were included in the models most often, followed by topographic and forest structural variables. The distributions of small individuals were more strongly related to environmental variation than those of larger individuals. Significant neighbourhood effects (spatial autocorrelation in intraspecific distributions and non-random overlaps in the distributions of certain species pairs) were also identified. Overlaps between congeners did not differ from random, but there was a highly significant overlap in the distributions of the two most common species. Conclusions: Our results support the view that habitat specialization is an important determinant of where on the rain forest landscape tree ferns grow, especially for juvenile plants. However, other factors, such as dispersal limitation, may also contribute to their local coexistence. [source] Release vocalizations in neotropical toads (Bufo): ecological constraints and phylogenetic implicationsJOURNAL OF ZOOLOGICAL SYSTEMATICS AND EVOLUTIONARY RESEARCH, Issue 1-2 2001Di Tada The release vocalizations of four nominal neotropical toad taxa (Bufo achalensis, Bufo limensis, Bufo spinulosus, Bufo arenarum) which compose three sympatric species pairs, were quantitatively analysed and homologous call types statistically compared. The first three taxa are closely related members of the mainly Andean Bufo spinulosus species group in which advertisement calls are absent. The specific vocal repertoire consisted of a uniform, unpulsed release call and one (in B. arenarum, two) pulsed release trill(s) which were given either singly or in series of up to three single calls. The uniform call was similar in structure and dominant (= basic) frequency in all taxa and probably represents an unspecific acoustic signal which is used to avoid heterospecific amplexus between sympatric toads. The release trills significantly differed in most call features among the taxa and, using discriminant analyses, even single calls were correctly assigned to sympatric pairs of species. In contrast, calls of allopatric pairs of species were confounded at rates of up to 36% indicating that selection towards species-specific signals increases when different species live in sympatry. The release trills of two allopatric populations (Perú, Argentina) assigned to B. spinulosus were similar in structure, but nevertheless features of pulse train permitted an unequivocal distinction, suggesting a long-lasting independent evolution. The taxonomic significance of this finding remains to be evaluated in further investigations. The complex vocal repertoire also offered the opportunity to assess phylogenetic relationships among the taxa. The extra-Andean B. achalensis seems to be closer related to the Andean B. spinulosus than this species is to the widely sympatric B. limensis– a species with several primitive character states indicating an early separation from the ancestral stock. Befreiungsrufe neotropischer Kröten (Bufo): Ökologische Einflüsse und phylogenetische Bedeutung Die Befreiungsrufe vier nomineller neotropischer Krötentaxa (Bufo achalensis, Bufo limensis, Bufo spinulosus, Bufo arenarum), die drei sympatrisch lebende Paare bilden, wurden quantitativ untersucht und homologe Ruftypen statistisch verglichen. Die drei erstgenannten Taxa sind nahverwandte Mitglieder der hauptsächlich andinen Bufo spinulosus Artengruppe, die keinen Paarungsruf besitzt. Das artspezifische Rufrepertoire umfaßte einen gleichförmigen, nicht-gepulsten Befreiungsruf und einen (bei B. arenarum zwei) aus Impulsen bestehenden Befreiungstriller. Diese Rufe werden entweder einzeln oder als Rufserien mit bis zu drei Einzelrufen abgegeben. Der gleichförmige Ruf war bei allen Taxa strukturell ähnlich und ist wahrscheinlich ein nichtartspezifischer Teil des Kommunikationssytems, der zur Vermeidung heterospezifischer Amplexus zwischen sympatrischen Kröten dient. Die Triller unterschieden sich in den meisten Merkmalen signifikant zwischen den Arten und auch Einzelrufe wurden bei sympatrischen Arten mittels Diskriminanzanlyse korrekt klassifiziert. Hingegen betrugen bei allopatrischen Arten die Fehlklassifikationen bis zu 36%. Dies weist auf einen starken Selektionsdruck zu artspezifischen Signalen bei Sympatrie hin. Die Triller zweier allopatrischer Populationen (Perú, Argentina) von B. spinulosusähnelten sich zwar strukturell, liessen sich aber ohne Fehlklassifikation voneinander unterscheiden, wahrscheinlich ein Hinweis auf eine längere unabhängige Evolution. Zur Klärung der taxonomischen Bedeutung dieses Befundes sind jedoch weitere Untersuchungen nötig. Der komplexe Aufbau der Rufe eröffnete auch die Möglichkeit, mittels bioakustischer Merkmale die phylogenetischen Beziehungen zwischen den Taxa abzuleiten. Die extraandine B. achalensis scheint mit der andinen B. spinulosus näher verwandt zu sein als diese Art mit der weitgehend sympatrischen B. limensis. Letztere Art zeigt eine Reihe primitiver Merkmale, die auf eine frühe Trennung vom der ancestralen Evolutionlinie hinweisen. [source] The transcriptomics of life-history trade-offs in whitefish species pairs (Coregonus sp.)MOLECULAR ECOLOGY, Issue 7 2008J. ST-CYR Abstract Despite the progress achieved in elucidating the ecological mechanisms of adaptive radiation, there has been little focus on documenting the extent of adaptive differentiation in physiological functions during this process. Moreover, a thorough understanding of the genomic basis underlying phenotypic adaptive divergence is still in its infancy. One important evolutionary process for which causal genetic mechanisms are largely unknown pertains to life-history trade-offs. We analysed patterns of gene transcription in liver tissue of sympatric dwarf and normal whitefish from two natural lakes, as well as from populations reared in controlled environments, using a 16 006-gene cDNA microarray in order to: (i) document the extent of physiological adaptive divergence between sympatric dwarf and normal species pairs, and (ii) explore the molecular mechanisms of differential life history trade-offs between growth and survival potentially involved in their adaptive divergence. In the two natural lakes, 6.45% of significantly transcribed genes showed regulation either in parallel fashion (2.39%) or in different directions (4.06%). Among genes showing parallelism in regulation patterns, we observed a higher proportion of over-expressed genes in dwarf relative to normal whitefish (70.6%). Patterns observed in controlled conditions were also generally congruent with those observed in natural populations. Dwarf whitefish consistently showed significant over-expression of genes potentially associated with survival through enhanced activity (energy metabolism, iron homeostasis, lipid metabolism, detoxification), whereas more genes associated with growth (protein synthesis, cell cycle, cell growth) were generally down-regulated in dwarf relative to normal whitefish. Overall, parallelism in patterns of gene transcription, as well as patterns of interindividual variation across controlled and natural environments, provide strong indirect evidence for the role of selection in the evolution of differential regulation of genes involving a vast array of potentially adaptive physiological processes between dwarf and normal whitefish. Our results also provide a first mechanistic, genomic basis for the observed trade-off in life-history traits distinguishing dwarf and normal whitefish species pairs, wherein enhanced survival via more active swimming, necessary for increased foraging and predator avoidance, engages energetic costs that translate into slower growth rate and reduced fecundity in dwarf relative to normal whitefish. [source] Immigration, species radiation and extinction in a highly diverse songbird lineage: white-eyes on Indian Ocean islandsMOLECULAR ECOLOGY, Issue 12 2006BEN H. WARREN Abstract Molecular phylogenetic hypotheses of species-rich lineages in regions where geological history can be reliably inferred may provide insights into the scale of processes driving diversification. Here we sample all extant or recently extinct white-eye (Zosterops) taxa of the southwest Indian Ocean, combined with samples from all principal continental lineages. Results support a high dispersal capability, with at least two independent continental sources for white-eyes of the region. An early (within 1.8 million years ago) expansion into the Indian Ocean may have originated either from Asia or Africa; the three resulting lineages show a disparate distribution consistent with considerable extinction following their arrival. Africa is supported as the origin of a later expansion into the region (within 1.2 million years ago). On two islands, a pair of Zosterops species derived from independent immigrations into the Indian Ocean co-occur or may have formerly co-occurred, providing strong support for their origin by double-island colonization rather than within-island (sympatric or microallopatric) speciation. On Mauritius and La Réunion, phylogenetic placement of sympatric white-eyes allow us to rule out a scenario in which independent within-island speciation occurred on both islands; one of the species pairs must have arisen by double colonization, while the other pair is likely to have arisen by the same mechanism. Long-distance immigration therefore appears to be responsible for much of the region's white-eye diversity. Independent immigrations into the region have resulted in lineages with mutually exclusive distributions and it seems likely that competition with congeneric species, rather than arrival frequency, may limit present-day diversity. [source] Spawning times, reproductive compatibilities and genetic structuring in the Acropora aspera group: evidence for natural hybridization and semi-permeable species boundaries in coralsMOLECULAR ECOLOGY, Issue 8 2002Madeleine J. H. Van Oppen Abstract Species boundaries among five sympatric coral species of the Indo-Pacific Acropora aspera group were examined by a combination of in vitro breeding trials, comparisons of spawning times and DNA sequence analysis of ribosomal DNA internal transcribed spacer (rDNA ITS) and 5.8S regions. The breeding trials showed that reproductive compatibility exists between at least some colonies of all the species pairs tested, suggesting a large potential for natural hybridization and introgression. The Acropora ITS regions exhibited extremely high levels of variability (up to ,62% for ITS1, ,11% for 5.8S and ,43% for ITS2), but most of the variation was shared among four of the five species, A. millepora, A. papillare, A. pulchra and A. spathulata, consistent with extensive introgression. Phylogenetic analyses did not resolve these four species as distinct clusters across a wide biogeographic region stretching from the southern Great Barrier Reef to Papua New Guinea. However, most colonies of the fifth species, A. aspera, constituted a distinct clade in phylogenetic analyses. This is consistent with our observations of a semi-permeable temporal barrier involving differences in spawning times between this and the other four species. Although the majority of colonies of all five species generally spawned within 90 min of each other, in two out of four years, gametes were absent prior to mass spawning episodes from at least some A. aspera colonies. Hence, our data suggest that transient reproductive barriers may be the result of year-to-year variation in the date of spawning and that this difference in spawning time contributes to the genetic structure detected among Acropora species in this group. Occasional leakage through the reproductive barrier was confirmed by the observation of A. aspera ×A. pulchra F1 hybrids, identified based on additivity of ITS sequences. [source] Competition as a structuring force in leaf miner communitiesOIKOS, Issue 6 2009Ayco J. M. Tack The role of competition in structuring communities of herbivorous insects is still debated. Despite this, few studies have simultaneously investigated the strength of various forms of competition and their effect on community composition. In this study, we examine the extent to which different types of competition will affect the presence and abundance of individual leaf miner species in local communities on oak trees Quercus robur. We first use a laboratory experiment to quantify the strength of intra- and interspecific competition. We then conduct a large-scale field experiment to determine whether competition occurring in one year extends to the next. Finally, we use observational field data to examine the extent to which mechanisms of competition uncovered in the two experiments actually reflect into patterns of co-occurrence in nature. In our experiment, we found direct competition at the leaf-level to be stronger among conspecific than among heterospecific individuals. Indirect competition among conspecifics lowered the survival and weight of larvae of T. ekebladella, both at the branch and the tree-level. In contrast, indirect competition among heterospecifics was only detected in one out of three species pairs examined. In the field experiment, the presence of a given moth species in one year affected the relative abundance of leaf miner species in the next year. Nevertheless, patterns of competition observed in these experiments did not translate into repulsion among free-ranging leaf miners: conspecific larvae of four leaf-mining species were aggregated on the same trees, shoots and leaves. In contrast, heterospecific larvae were only aggregated at the tree-level. We propose that despite the fact that leaf miners do compete and that such effects extend through time, the incidence and strength of competition is relatively small at realistic densities. Hence, competition will likely be of minor importance in shaping the distribution of leaf miners in a natural setting. [source] Nearest-neighbor tree species combinations in tropical forest: the role of chance, and some consequences of high diversityOIKOS, Issue 3 2007Milton Lieberman In three permanent inventory plots comprising 12.4 ha of undisturbed forest at La Selva, Costa Rica, all stems ,10 cm dbh were mapped and identified to species. There were 1628, 1478 and 1954 trees in the plots, representing 168, 166 and 171 species respectively. We determined the species of each nearest-neighbor pair of trees, and asked whether the occurrence of species pairs conforms to a simple random mixing model. If trees are randomly mixed in terms of species, the expected frequency of any nearest neighbor species combination is a function of the relative abundance of the two species. Departures from random mixing could arise from species interactions, differential responses to habitat, or both. The number of possible ij species combinations increases approximately as the square of the number of species. For the 168 species in plot 1, for example, there are 14 196 possible combinations. We compared the expected frequency of each species combination in the three plots (42,736 combinations in all) with observed frequencies. Over 98% of the combinations had observed frequencies of zero and expected frequencies close to zero. A consequence of high diversity is low density of most individual species, and exceedingly low frequencies of the vast majority of species combinations. For each of the 805 combinations with observed frequencies >0, we used simulation to generate a distribution of expected frequencies. We used a t-test to compare the observed frequency with the mean of the simulated distribution for each combination. Only 40 combinations (0.09% of the possible species combinations in the plots) departed from expected frequencies; 39 combinations were more common, and one less common than expected. The overwhelming majority of nearest neighbor species combinations occur at frequencies predictable from their individual abundances. [source] |