Examination of the “null” model of connor and simberloff for species co-occurrences on Islands

Summary Among birds of the Bismarck Archipelago, only certain combinations of the species in a guild coexist on islands, and some species that are very similar ecologically have mutually exclusive distributions. Diamond (1975) interpreted these patterns as biologically significant, involving effects such as competition. Connor and Simberloff (1979) claimed such patterns to be not recognizably different from random, because they were scarcely distinguishable from those generated by a null distribution supposedly not incorporating competition.On examining the analysis by Connor and Simberloff, we find that it actually yields the opposite conclusion: three faunas tested have grossly non-random structures, while their test is unworkable for the fourth fauna. The method of Connor and Simberloff for generating a null distribution has many fatal weaknesses: dilution of relevant data from guilds with irrelevant data from the whole species pool; hidden incorporation of effects of competition into the constraints; inability to recognize a checkerboard, the extreme example of a distribution produced by competition; reliance on inefficient Monte Carlo simulations; severe constraints that sometimes make generation of a null distribution impossible; failure to weight species combinations; and failure to identify the direction of non-randomness or the species combinations most responsible.Finally, we use other recent studies by Simberloff and coleagues to examine the value of constructing null hypotheses in community ecology.     

Factors contributing to non-randomness in species Co-occurrences on Islands

Summary There has been dispute whether patterns of species co-occurrence on islands are largely random. We present a new method for testing this question; this method lets one not only examine whether a whole fauna is non-randomly structured, but also identify in which direction and by how much each particular species combination deviates from expectations based on randomness. Application of this method to the whole Bismarck and New Hebridean avifaunas, and to two particular guilds, shows that some pairs of species have more exclusive distributions than expected for random placement of species, because of competition, differing distributional strategies, or different geographical orgins. Other pairs of species have more coincident distributions than expected, because of shared habitat, single-island endemism, shared distributional strategies, or shared geographical origins. Much of the information about non-random co-occurrence is contained in the incidence graphs for occurrence of individual species. Finally, our present understanding of assembly rules is summarized.     

Testing multiple assembly rule models in avian communities on islands of an inundated lake,Zhejiang Province,China

Aim A fundamental question in community ecology is whether general assembly rules determine the structure of natural communities. Although many types of assembly rules have been described, including Diamond’s assembly rules, constant body‐size ratios, favoured states, and nestedness, few studies have tested multiple assembly rule models simultaneously. Therefore, little is known about the relative importance of potential underlying factors such as interspecific competition, inter‐guild competition, selective extinction and habitat nestedness in structuring community composition. Here, we test the above four assembly rule models and examine the causal basis for the observed patterns using bird data collected on islands of an inundated lake. Location Thousand Island Lake, China. Methods  We collected data on presence–absence matrices, body size and functional groups for bird assemblages on 42 islands from 2007 to 2009. To test the above four assembly rule models, we used null model analyses to compare observed species co‐occurrence patterns, body‐size distributions and functional group distributions with randomly generated assemblages. To ensure that the results were not biased by the inclusion of species with extremely different ecologies, we conducted separate analyses for the entire assemblage and for various subset matrices classified according to foraging guilds. Results The bird assemblages did not support predictions by several competitively structured assembly rule models, including Diamond’s assembly rules, constant body‐size ratios, and favoured states. In contrast, bird assemblages were highly significantly nested and were apparently shaped by extinction processes mediated through area effects and habitat nestedness. The nestedness of bird assemblages was not a result of passive sampling or selective colonization. These results were very consistent, regardless of whether the entire assemblage or the subset matrices were analysed. Main conclusions Our results suggest that bird assemblages were shaped by extinction processes mediated through area effects and habitat nestedness, rather than by interspecific or inter‐guild competition. From a conservation point of view, our results indicate that we should protect both the largest islands with the most species‐rich communities and habitat‐rich islands in order to maximize the number of species preserved.     

Detecting the influence of climatic variables on species distributions: a test using GIS niche-based models along a steep longitudinal environmental gradient

Aim  To investigate the influence of climate variables in shaping species distributions across a steep longitudinal environmental gradient.
Location  The state of Oklahoma, south-central United States.
Methods  We used Geographical Information Systems (GIS) niche-based models to predict the geographic distributions of six pairs of closely related amphibian and reptile species across a steep longitudinal environmental gradient. We compared results from modelling with actual distributions to determine whether species distributions were primarily limited by environmental factors, and to assess the potential roles of competition and historical factors in influencing distributions.
Results  For all species pairs, GIS models predicted an overlap zone in which both species should occur, although in reality in some cases this area was occupied by only one of the species. We found that environmental factors clearly influence the distributions of most species pairs. We also found evidence suggesting that competition and evolutionary history play a role in determining the distributions of some species pairs.
Main conclusions  Niche-based GIS modelling is a useful tool for investigating species distribution patterns and the factors affecting them. Our results showed that environmental factors strongly influenced species distributions, and that competition and historical factors may also be involved in some cases. Furthermore, results suggested additional lines of research, such as ecological comparisons among populations occurring inside and outside predicted overlap zones, which may provide more direct insight into the roles of competitive interactions and historical factors in shaping species distributions.     

Forest patch size and isolation as drivers of bird species richness in Maputaland,Mozambique

Aim To examine the response of forest‐dependent and generalist bird assemblages and feeding guilds to patch characteristics of forest fragments in the context of island biogeography and metapopulation theories in an area that supports high levels of species diversity and endemism. Location Dune, riverine, sand and swamp forest fragments in southern Mozambique’s Maputaland. Methods We recorded 20 forest‐dependent and 69 generalist bird species at 220 survey points in 30 forest fragments that ranged in size from 5 to 7432 ha and that were 100 to 5100 m apart. We used linear regressions to relate the number of species to forest fragment size, isolation, perimeter, fractal dimension, shape and core area for frugivores, insectivores, granivores, nectarivores, carnivores and omnivores separately for forest‐dependent and generalist bird assemblages. We tested for modality in occupancy frequency distributions of species in forest fragments and used the binary matrix temperature calculator to determine whether the assemblages had a nested structure separately for forest‐dependent and generalist bird species. Results The number of forest‐dependent and generalist bird species categorized into the feeding guilds varied independently from forest patch parameter values. Occupancy models showed a random distribution for forest‐dependent birds, and generalists had a unimodal distribution. Both the forest‐dependent and generalist bird assemblages had a non‐nested structure. The presence of eight rare forest‐dependent and four endemic bird species in 22 of the 30 fragments contributed to the non‐nested structure of these bird assemblages. Main conclusions Fragmented forests did not induce the expected responses of the forest bird assemblages in that the number of species was not explained by patch characteristics, nor could we find evidence for metapopulation dynamics. Non‐nestedness may be caused by the few rare and endemic species occupying forest fragments with a wide range of patch characteristics. Knowledge of the response of species to forest fragmentation can benefit conservation planning catering for rare and endemic species. Our study suggests that all forest fragments in the region may have to be included in a conservation network to complement existing landscape‐based plans to protect the region.     

Temporal dynamics and nestedness of an oceanic island bird fauna

Aim To examine temporal variation in nestedness and whether nestedness patterns predict colonization, extinction and turnover across islands and species. Location Dahlak Archipelago, Red Sea. Method The distributions of land birds on 17 islands were recorded in two periods 30 years apart. Species and islands were reordered in the Nestedness Temperature Calculator, software for assessing degrees of nestedness in communities. The occupancy probability of each cell, i.e. species–island combinations, was calculated in the nested matrix and an extinction curve (boundary line) was specified. We tested whether historical and current nested ranks of species and islands were correlated, whether there was a relationship between occupancy probability (based on the historical data) and number of extinctions or colonizations (regression analyses) and whether the boundary line could predict extinctions and colonizations (chi‐square analyses). Results Historical and current nested ranks of islands and species were correlated but changes in occupancy patterns were common, particularly among bird species with intermediate incidence. Extinction and turnover of species were higher for small than large islands, and colonization was negatively related to isolation. As expected, colonizations were more frequent above than below the boundary line. Probability of extinction was highest at intermediate occupancy probability, giving a quadratic relationship between extinction and occupancy probability. Species turnover was related to the historical nested ranks of islands. Colonization was related negatively while extinction and occupancy turnover were related quadratically to historical nested ranks of species. Main conclusions Some patterns of the temporal dynamics agreed with expectations from nested patterns. However, the accuracy of the predictions may be confounded by regional dynamics and distributions of idiosyncratic, resource‐limited species. It is therefore necessary to combine nestedness analysis with adequate knowledge of the causal factors and ecology of targeted species to gain insight into the temporal dynamics of assemblages and for nestedness analyses to be helpful in conservation planning.     

Two decades of interaction between the MacArthur-Wilson model and the complexities of mammalian distributions

More than two decades after its publication, MacArthur and Wilson's equilibrium model of insular biogeography continues to provide the conceptual foundation for investigating the distribution of species on islands and the composition of insular biotas. During this period, studies of the distributions of mammals among insular habitats have tested, modified, and extended MacArthur and Wilson's simple formalism to enhance greatly our understanding of the complexities of biogeographic patterns and processes. The papers in this symposium summarize many of the past contributions of mammalian biogeographers and introduce important new data and ideas. The diversity of biological characteristics and associated distributional patterns exhibited by mammals has facilitated this endeavour. Some insular mammalian faunas appear to represent approximate equilibria between opposing rates of contemporary colonization and extinction. Other faunas are currently decreasing in diversity because of extinctions, owing either to natural habitat fragmentation that has occurred since the Pleistocene or to human activities within the last few centuries. Still other faunas have been increasing in diversity (at least until recent human impacts) because limiting rates of origination, both colonization and speciation, have been extremely low. The questions and analyses of island biogeography can also be applied to continents with comparable overall results: the distributions of continental faunas reflect the consequences of similar processes of colonization, speciation and extinction. Analyses of insular distributions show unequivocally that probabilities of extinction, colonization and speciation are highly deterministic and vary in predictable ways among different taxa and archipelagos. These findings have important implications for applying the theory and data of insular biogeography to the pressing practical problems of designing natural reserves to preserve native species.     

Influence of body size on coexistence of bird species

Theory suggests that body size is an important factor in determining interspecific competition and, ultimately, in structuring ecological communities. However, there is a lack of pragmatic studies linking body size and interspecific competition to patterns in ecological communities. The objective of the present study was to determine the effect of body size (mass) on competitive interactions between bird pairs and to investigate the influence of food guilds. Point-counts were carried out in nine sites every month from November 2002 to November 2003 in the Cuetzalan Region, Mexico, and we used presence/absence and abundance data for the analyses. To calculate the strength of competition we used the Angle Frequency Method to extract form factors from 20 pairwise interactions. A prototype competition interaction and random pairs were also constructed. We used clustering techniques (PCA) to calculate the dissimilarity scores (distances, D) of each of the pairwise interactions to the prototype competition and random pairs and one-way ANOVA to test for differences between the means of the random and competitive pairs. The ratio in body mass (lnBM) for each of the interacting pairs was calculated, and the association between the lnBM ratio and the strength of competition (D) was tested using a Pearson product-moment correlation coefficient. To test for the influence of foraging guilds we used a univariate general linear model. Our results demonstrate a significant negative relationship between bird body mass ratio and competition strength – i.e. competition strength increased when the body masses of the birds became more similar. We did not find a significant influence of foraging guild on the relationship between body mass ratio and competition strength. On the basis of these results, we suggest that high variation in body sizes amongst sympatric species promotes coexistence in communities.     

Species co-occurrence: the case of congeneric species and a causal approach to patterns of species association

Aim To test whether congeneric species are significantly associated with one another in space, either positively or negatively. Also, to provide a framework for a causal investigation of co‐occurrence patterns by a parallel comparison of interactions in geographical and ecological data matrices. Location For the analysis of congeneric species’ co‐occurrences we used 30 matrices covering a wide range of taxa and geographical areas, while for the causal investigation we used the distribution of 50 terrestrial isopod species on 20 islands and 264 sampling stations in the central Aegean archipelago, as well as a number of ecological variables for each sampling station. Methods We developed a software program (cooc ) that incorporates the species‐by‐species approach to co‐occurrence analysis using EcoSim's output of prior null model analysis of co‐occurrence. We describe this program in detail, and use it to investigate one of the most common assembly rules, namely, the decreased levels of co‐occurrence among congeneric species pairs. For the causal analysis, we proceed likewise, cross‐checking the results from the geographical and the ecological matrices. There is only one possible combination of results that can support claims for direct competition among species. Results We do not get any strong evidence for widespread competition among congeneric species, while most communities investigated do not show significant patterns of species associations. The causal analysis suggests that the principal factors behind terrestrial isopod species associations are of historical nature. Some exceptional cases are also discussed. Main conclusions Presence/absence data for a variety of taxa do not support the assembly rule that congeneric species are under more intense competition compared to less related species. Also, these same data do not suggest strong interactions among species pairs, regardless of taxonomic status. When significant species associations can be seen in such matrices, they mainly reflect the effects of history or of habitat requirements.     

EVOLUTIONARY AND BIOGEOGRAPHIC ORIGINS OF HIGH TROPICAL DIVERSITY IN OLD WORLD FROGS (RANIDAE)

Differences in species richness between regions are ultimately explained by patterns of speciation, extinction, and biogeographic dispersal. Yet, few studies have considered the role of all three processes in generating the high biodiversity of tropical regions. A recent study of a speciose group of predominately New World frogs (Hylidae) showed that their low diversity in temperate regions was associated with relatively recent colonization of these regions, rather than latitudinal differences in diversification rates (rates of speciation–extinction). Here, we perform parallel analyses on the most species-rich group of Old World frogs (Ranidae; ∼1300 species) to determine if similar processes drive the latitudinal diversity gradient. We estimate a time-calibrated phylogeny for 390 ranid species and use this phylogeny to analyze patterns of biogeography and diversification rates. As in hylids, we find a strong relationship between the timing of colonization of each region and its current diversity, with recent colonization of temperate regions from tropical regions. Diversification rates are similar in tropical and temperate clades, suggesting that neither accelerated tropical speciation rates nor greater temperate extinction rates explain high tropical diversity in this group. Instead, these results show the importance of historical biogeography in explaining high species richness in both the New World and Old World tropics.     

Null model analysis of communities on gradients

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.     

Non-interactive fish communities in the coastal streams of North-western France

1. Spatial patterns of freshwater fish species at regional and local scales were investigated to explore the possible role of interspecific interactions in influencing distribution and abundance within communities occupying coastal streams of North-Western France.
2. Nine sites from nine streams situated in the same biogeographical region were sampled annually over the 6-year period from 1990 to 1995.
3. Similar habitats (sites) with richer regional colonization pools exhibited proportionally richer local communities in terms of number of species, total density and total biomass of individuals. Furthermore, no negative relationships were found between density and biomass of each of the most common species and local species richness.
4. Results of dynamic regression models (applied to the above-mentioned species) suggest an absence of strong competition between all pairs of species.
5. The evidence on lack of density compensation for species-poor communities and absence of perceptible interspecific competition between species suggest that the communities studied are non-interactive.
6. Two main explanations can be advanced. First, the local abundance of species in the communities studied could be determined through differential responses to unpredictable environmental changes, rather than through biological interactions. Second, as a result of historical events, the communities studied are reduced in congeneric species which can limit, in turn, the influence of interspecific competition in structuring these communities.
7. These results underline the strong influence of regional processes in shaping local riverine fish communities and minimize the possible influence of species interactions in governing these communities.     

Miniaturizing landscapes to understand species distributions

Species’ geographic distributions shape global patterns of biodiversity and therefore have long been of interest to ecology and conservation. Theory has generated valuable hypotheses about how landscape structure, dispersal, biotic interactions and evolution shape range dynamics, but most predictions have not been tested on real organisms because key variables are difficult to isolate, replicate or manipulate in natural ecosystems. An exciting and rapidly emerging approach is to extend classical microcosm and mesocosm systems to create experimental ‘micro-landscapes’. By enabling researchers to manipulate geographic features of interest, replicate landscapes, control colonization and follow dynamics across evolutionary timescales, micro-landscapes allow explicit tests of the ecological and evolutionary underpinnings of species distributions. Here we review the micro-landscape systems being used to advance biogeography, the major insights they have generated thus far, and the features that limit their application to some scenarios. We end by highlighting important questions about species’ biogeography that are ripe for testing with experimental micro-landscapes, particularly those of immediate concern given rapid global change, such as range contractions and constraints to range expansion.     

Are land bird assemblages functionally saturated? An empirical test in Mexico

According to the model of ideal free distribution, few or no assemblages will contain unused resources as long as species are free to colonize areas where resources are available. Consequently, because several food resources for birds are available throughout Mexico (nectar, seeds and fruits, vertebrates, carrion and different types of invertebrates), we predicted that all assemblages from Mexico contain birds from the corresponding foraging guilds. However, severe climates may prevent colonization by certain birds. Thus, a second prediction was that, if assemblages with unused resources were found, these would be from localities with climates that pose extreme stress (either extremely high temperature and low rainfall, or extremely low temperature). We tested these predictions on bird assemblages from 77 localities that represent the entire range of climatic conditions found in Mexico. Only two assemblages containing unused resources were found. These were from the localities with lowest temperature and lowest rainfall, respectively. We observed that land bird assemblages in Mexico do not contain unused resources, except when climate selectively restricts colonization by species from certain foraging guilds. Therefore, we conclude that even assemblages with almost no species in common have a common guild structure based on food resources. Null models of competition would be more realistic if they were to incorporate such patterns in their regional species pools.     

Bird dietary guild richness across latitudes,environments and biogeographic regions

Aim To integrate dietary knowledge and species distributions in order to examine the latitudinal, environmental, and biogeographical variation in the species richness of avian dietary guilds (herbivores, granivores, frugivores, nectarivores, aerial insectivores, terrestrial/arboreal insectivores, carnivores, scavengers, and omnivores). Location Global. Methods We used global breeding range maps and a comprehensive dietary database of all terrestrial bird species to calculate guild species richness for grid cells at 110 × 110 km resolution. We assessed congruence of guild species richness, quantified the steepness of latitudinal gradients and examined the covariation between species richness and climate, topography, habitat diversity and biogeographic history. We evaluated the potential of current environment and biogeographic history to explain global guild distribution and compare observed richness–environment relationships with those derived from random subsets of the global species pool. Results While most guilds (except herbivores and scavengers) showed strong congruence with overall bird richness, covariation in richness between guilds varied markedly. Guilds exhibited different peaks in species richness in geographical and multivariate environmental space, and observed richness–environment relationships mostly differed from random expectations. Latitudinal gradients in species richness were steepest for terrestrial/arboreal insectivores, intermediate for frugivores, granivores and carnivores, and shallower for all other guilds. Actual evapotranspiration emerged as the strongest climatic predictor for frugivores and insectivores, seasonality for nectarivores, and temperature for herbivores and scavengers (with opposite direction of temperature effect). Differences in species richness between biogeographic regions were strongest for frugivores and nectarivores and were evident for nectarivores, omnivores and scavengers when present‐day environment was statistically controlled for. Guild richness–environment relationships also varied between regions. Main conclusions Global associations of bird species richness with environmental and biogeographic variables show pronounced differences between guilds. Geographic patterns of bird diversity might thus result from several processes including evolutionary innovations in dietary preferences and environmental constraints on the distribution and diversification of food resources.     

Revision of the genus Chalcasthenes Arrow (Coleoptera: Scarabaeidae: Dynastinae: Oryctoderini) from the Solomon Islands

The genus Chalcasthenes Arrow (Dynastinae: Oryctoderini), a scarab beetle genus endemic to the Solomon Islands, is reviewed. Based on examination of type specimens, the genus Strehlia Frey (Rutelinae: Rutelini: Parastasiina) is a new junior synonym of Chalcasthenes . The historical classification of these genera (either in the subfamily Dynastinae or Rutelinae) and character-based criteria for assigning the taxa to the Dynastinae are provided. We discuss character states that support the monophyly of members of the genus Chalcasthenes, comment on the distribution and biogeography of species in the genus and provide a key to species. The genus includes four species: Chalcasthenes divinus Endrödi, Chalcasthenes pulcher Arrow, Chalcasthenes squamigerus Frey new combination and Chalcasthenes styracoceros Jameson and Ratcliffe n. sp. Species hypotheses are corroborated based on evidence from Pleistocene geological reconstructions of the Solomon Islands, geographic variation of bird species in the region and development in scarab beetles.     

Oppositions in panbiogeography: Can the conflicts between selection,constraint, ecology,and history be resolved?

Abstract

Croizat’s radical attack on traditional modes of biological inquiry has produced strongly polarised responses. Instead of arguing for or against the panbiogeographic approach I attempt to analyse some basic assumptions shared by both its defenders and critics. Although their emphasis is quite different both sides rely on two central oppositions. In evolutionary arguments selective explanations are opposed to those emphasising phylogenetic constraint (orthogenesis). In biogeographic arguments ecological explanations are opposed to historical explanations. I discuss how the legendary opposition between nature and nurture was resolved by reformulating the way in which causation was viewed. I suggest that the selection/constraint opposition shares many features in common with the nature/nurture opposition and argue that it can be resolved in a similar manner. Conflicts between ecological and historical explanations in biogeography are explored by contrasting Diamond’s analysis of New Guinea bird distributions with that of Croizat’s. By again drawing on the resolution of the nature/nurture dispute a way of synthesising ecological and historical explanations is outlined.     

Origin and evolution of the Micronesian biota: Insights from molecular phylogenies and biogeography reveal long-distance dispersal scenarios and founder-event speciation

Micronesian islands taxa show high endemism rates, but very little is known about their biogeographical histories. The lack of systematic biogeography is mainly due to insufficient phylogenetic research in Micronesia. With the recent increase in published molecular biogeographic data, we were able to, for the first time, answer fundamental biogeography questions by reviewing and analyzing numerous geological, ecological, and evolutionary studies. This review, in addition to providing an overview of Micronesian geological history, confirmed the importance of long-distance dispersal mechanisms and founder-event speciation, and morphological and physiological adaptations of plant propagules to cross vast stretches of ocean by wind, ocean currents, bird, or bat dispersal. These adaptations to habitat and geological features, including reef types, determined colonization success as well as inland dispersal and speciation mechanisms. We further identified the source areas of the Micronesian biota and reconstructed historical dispersal scenarios: a dominant Austro-Melanesian dispersal scenario, an Indo-Malaysian connecting to the Austro-Melanesian dispersal scenario, and a Neotropical American and an African dispersal scenario toward Micronesia. Most generic origins were estimated between the Eocene and the Miocene and dispersed to Micronesia between the Miocene and the Pleistocene.     

Spatial processes that maintain biodiversity in plant communities

The composition of communities of sessile organisms, and the change in species diversity with time, is a spatially explicit phenomenon. Three spatial factors clearly affect diversity: (1) the structure and heterogeneity of the landscape that limits species immigration and ultimate community size; (2) neighborhood interactions that determine colonization and extinction rates and influence residence times of local populations; and (3) disturbances that open spatially contiguous areas for recolonization by less abundant species. The importance of these three factors was first reviewed and then examined with a spatially explicit, multi-species model of plant dispersal, competition and establishment, with an assumption of neutrality (all species had equivalent life histories) that reduced the initial dimensionality of the problem. The simulations assumed that the probability of immigration was a linear function of mainland abundance and distance to islands, similar to the equilibrium theory of island biogeography and the unified neutral theory of biodiversity. The rate of increase in species richness was not constant across island sizes, declining as island area became very large. This pattern was explained by the spatial dynamics of colonization and establishment, a non-random process that cannot be explained by passive sampling alone. Simulations showed that population establishment depended critically on rare long-distance dispersal events while population persistence was achieved by the formation of aggregated species distributions that developed through restricted dispersal and local competitive interactions. Nevertheless, species richness always declined to a single species in the absence of disturbances, while up to 40 species could persist to 10,000 years when spatially dependent mortality was added. Further explorations with spatially explicit models will be required to fully appreciate the consequence of land use change and altered disturbance regimes on patterns of species distribution and the maintenance of diversity.     

Survival, development and life tables of two congeneric ladybirds in aphidophagous guilds

Two congeneric aphidophagous ladybirds, Coccinella septempunctata and Coccinella transversalis, were reared on three aphid species, Lipaphis erysimi, Myzus persicae and Aphis nerii, to estimate the effect of prey quality and intra- and interspecific interactions on their survival and development of life stages. Mortality of first instar ladybirds of both species was highest feeding on A. nerii. Preimaginal mortafity was lowest when feeding on L. erysimi （C. septempunctata, 1.6% and C. transversalis, 3.2%）, and highest when feeding on A. nerii （ C. septempunctata, 6.2% and C. transversalis, 8.2%）. Comparatively higher weight and larger size of C. septempunctata along with the lower levels of mortality recorded suggested that it is more likely to have acted as an intraguild predator than C. transversalis. High recorded mortality of C. transversalis is attributed to probable intraguild predation on account of its smaller size. The major sources of mortality were probably cannibalism, intraguild predation and other unknown factors. Lower prey quality increased the incidence of cannibalism and intraguild predation, especially in C. transversalis. The investigation suggests an intrinsic competitive advantage for C. septempunctata over C. transversalis in guilds of three aphid species.