Bat assemblages on Neotropical land-bridge islands: nested subsets and null model analyses of species co-occurrence patterns

A fundamental goal of ecology is to understand whether ecological communities are structured according to general assembly rules or are essentially dictated by random processes. In the context of fragmentation, understanding assembly patterns and their mechanistic basis also has important implications for conservation. Using distribution data of 20 bat species collected on 11 islands in Gatún Lake, Panama, we tested for non‐randomness in presence–absence matrices with respect to nestedness and negative species co‐occurrence. We examined the causal basis for the observed patterns and conducted separate analyses for the entire assemblage and for various species submatrices reflecting differences in species’ trophic position and mobility. Furthermore, we explored the influence of weighting factors (area, isolation, abundance) on co‐occurrence analyses. Unweighted analyses revealed a significant negative co‐occurrence pattern for the entire assemblage and for phytophagous bats alone. Weighting analyses by isolation retained a pattern of species segregation for the whole assemblage but nullified the non‐random structure for phytophagous bats and suggested negative associations for animalivores and species with low mobility. Area‐ and abundance‐weighted analyses always indicated random structuring. Bat distributions followed a nested subset structure across islands, regardless of whether all species or different submatrices were analysed. Nestedness was in all cases unrelated to island area but weakly correlated with island isolation for incidence matrices of all species, phytophagous bats, and mobile species. Overall, evidence for negative interspecific interactions indicative of competitive effects was weak, corroborating previous studies based on ecomorphological analyses. Our findings indicate that bat assemblages on our study islands are most strongly shaped by isolation effects and species’ differential movement and colonization ability. From a conservation viewpoint this suggests that even in systems with high fragment–matrix contrast, a purely area‐based approach may be inadequate, and structural and functional connectivity among patches are important to consider in reserve planning.     

Direct and indirect effects in microcosm communities of protists

Increased complexity in biological communities can increase the variety of interactions among species, but the relative strengths and long-term consequences of various direct and indirect interactions require further investigation. I studied interactions among four species of protists by monitoring their population dynamics when they were cultured either together or in seven different subset communities. Two protists were bacterivores (Chilomonas and Tetrahymena) and two were predators (Actinosphaerium and Euplotes). Actinosphaerium was omnivorous, and could eat both predatory Euplotes and bacterivores. Three indirect effects occurred among the four species of protists, including indirect facilitation of one predator by the other, apparent competition between bacterivores, and indirect facilitation of one bacterivore by the omnivorous predator. Community structure and invasibility depended on both direct and indirect effects; thus both can be mechanisms for assembly rules.     

The myth of constant predator: prey ratios

Apparent constancy in the ratio of predator species to prey species has been offered as evidence that ecological communities are structured by interspecific interactions. If significantly different from random expectation, this effect would be one of the few sound pieces of evidence for community structure. The evidence was re-evaluated by using the data from previous studies to form species pools, and forming simulated communities by drawing species at random from these pools (with replacement). Using a correlation coefficient (number of predator species versus number of prey species), and also the statistic used by the original workers (where different), the observed predator:prey correlation was compared to that for the random communities. In five studies, the observed predator:prey ratio was not significantly different from random expectation. In the only two studies where there was significant departure from the null model, it was with more variation in the ratio than expected on a random basis. It is concluded that there is as yet no evidence for near-constant predator:prey ratios.     

立地条件差异对天然东北红豆杉(Taxus cuspidata)种间竞争的影响

立地因子空间梯度特征是影响种间竞争的重要因素。通过研究主要竞争树种及竞争强度随海拔、坡向、坡位和坡度等立地因子变化的规律,揭示不同立地条件对天然东北红豆杉种间竞争的影响,为濒危东北红豆杉种群的保护与恢复提供理论指导。研究结果表明：东北红豆杉的种间竞争压力主要来自紫椴、臭冷杉、色木槭、红松、青楷槭、枫桦、花楷槭等7个树种,主要竞争树种以小径级个体居多。立地因子差异对种间竞争强度和主要竞争树种的排序有明显影响。东北红豆杉在阴坡受到的种间竞争压力是阳坡的3.27倍,主要树种的竞争强度显著高于阳坡,竞争树种的排序发生更迭;随着坡位的提升,东北红豆杉受到的种间竞争压力明显增加,主要竞争种类和强度也发生变化;坡度增加,种间竞争强度增加,主要竞争树种排序发生变化;海拔对种间竞争的影响总体不大,但影响主要竞争树种的竞争强度和排序。     

Temporal patterns of diversity: assessing the biotic and abiotic controls on ant assemblages

In this study, we use 12 months of data from 11 ant assemblages to test whether seasonal variation in ant diversity is governed by either the structuring influences of interspecific competition or environmental conditions. Because the importance of competition might vary along environmental gradients, we also test whether the signature of competition depends on elevation. We find little evidence that competition structures the seasonal patterns of activity in the ant assemblages considered, but find support for the effects of temperature on seasonal patterns of diversity, especially at low-elevation sites. Although, in general, both competition and the environment interact to structure ant assemblages, our results suggest that environmental conditions are the primary force structuring the seasonal activity of the ant assemblages studied here.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 191–201.     

Large-scale patterns in morphological diversity and species assemblages in Neotropical Triatominae (Heteroptera: Reduviidae)

We analysed the spatial variation in morphological diversity (MDiv) and species richness (SR) for 91 species of Neotropical Triatominae to determine the ecological relationships between SR and MDiv and to explore the roles that climate, productivity, environmental heterogeneity and the presence of biomes and rivers may play in the structuring of species assemblages. For each 110 km x 110 km-cell on a grid map of America, we determined the number of species (SR) and estimated the mean Gower index (MDiv) based on 12 morphological attributes. We performed bootstrapping analyses of species assemblages to identify whether those assemblages were more similar or dissimilar in their morphology than expected by chance. We applied a multi-model selection procedure and spatial explicit analyses to account for the association of diversity-environment relationships. MDiv and SR both showed a latitudinal gradient, although each peaked at different locations and were thus not strictly spatially congruent. SR decreased with temperature variability and MDiv increased with mean temperature, suggesting a predominant role for ambient energy in determining Triatominae diversity. Species that were more similar than expected by chance co-occurred near the limits of the Triatominae distribution in association with changes in environmental variables. Environmental filtering may underlie the structuring of species assemblages near their distributional limits.     

Neotropical diversification: the effects of a complex history on diversity within the poison frog genus Dendrobates

Aim We study the Neotropical poison frogs of the genus Dendrobates Wagler, 1830 in order to clarify their phylogenetic relationships and biogeographical history. The genus Dendrobates is an excellent taxon for examining patterns of Neotropical diversification as the four major species groups appear to correspond roughly to distinct geographical regions: (1) trans‐Andean, (2) Andean foreland, (3) Brazilian Shield and (4) Guianan Shield/Central America. In order to test the agreement of five of the most prominent hypotheses of Amazonian diversification, phylogenetic patterns were examined for agreement with patterns predicted by these hypotheses. Location Central and South America Methods The phylogenetic relationships of the genus Dendrobates were examined from novel and existing (GenBank) sequences of four mitochondrial loci totalling c. 1400 bp from 40 specimens of 22 different species using maximum parsimony and Bayesian methods. Results were compared with traditional taxonomic arrangements by means of SH tests. Phylogenetic relationships and genetic distances were used to test the adequacy of various diversification hypotheses. Results Phylogenetic analyses support the restructuring of two species groups of Dendrobates and the creation of a new species group. Statistical tests of the traditional taxonomic arrangement indicate a significantly bad fit to the molecular data. This restructuring has important implications for the understanding of the historical biogeography of Dendrobates. Biogeographical patterns within this genus suggest that a complex interaction of biotic and abiotic factors since the Eocene have produced the diversity observed today. Main conclusions The current classification of Dendrobates into discrete species groups does not accurately reflect evolutionary history. Data presented here strongly support a monophyletic Brazilian Shield lineage whose members have previously been split among the quinquevittatus and tinctorius groups. Furthermore, previous attempts at elucidating the historical biogeography of this genus were compromised by incomplete sampling and conclusions drawn from a paraphyletic ingroup. Our findings demonstrate a role for numerous hypotheses of diversification (e.g. river, refuge, disturbance–vicariance) in the history of Dendrobates, supporting previous warnings about the dangers of over‐simplification in the study of Neotropical diversification.     

Occasional intraguild predation structuring small mammal assemblages: the marsupial Didelphis aurita in the Atlantic Forest of Brazil

The didelphid marsupial, Didelphis aurita, is suggested as an intraguild predator and as key‐species in small mammal assemblages of the Atlantic Forest of Brazil. The field experiments required to test this hypothesis are complex to implement, but the recent revival of regression methods offers a viable alternative. Here we use the dynamic and static regression methods to determine the importance of D. aurita as a competitor and intraguild predator. Capture–recapture data from two localities in the Rio de Janeiro State were used, Garrafão (municipality of Guapimirim), a coastal forest of the Serra do Mar, and Barra de Maricá, a costal sand dune vegetation. Population and microhabitat variables were monitored from April 1997 to April 2003 in Garrafão, and from January 1986 to July 1990 in Barra de Maricá. Microhabitat variables were related to Canopy, Plant, Litter and Rock covers, Obstruction from 0 to 1.5 m, and Number of logs. Exploitation competition was tested by the dynamic method, which models the effects of D. aurita on the per capita growth rate of a species. Interference by predation or competition was tested by the static method, where the abundance of D. aurita at trap stations was regressed against the abundance of other small mammals, after removal of any variation associated with microhabitat factors. Exploitation competition was not detected, but the interference of D. aurita was pervasive, affecting all small mammals studied in the two localities. The clear avoidance of D. aurita by all small mammals tested in two localities of different physiognomies indicates that it functions as an intraguild predator, even if actual predation by D. aurita is an occasional event.     

长苞铁杉天然林群落种内及种间竞争关系研究

通过各种竞争指数的比较筛选,得到较能客观反映长苞铁杉种内、种间竞争关系的竞争指数,定量地分析了长苞铁杉种内和种间竞争强度。结果表明:长苞铁杉种内竞争随胸径的增大而逐渐减少;种间与种内竞争关系顺序为:长苞铁杉-长苞铁杉>猴头杜鹃-长苞铁杉>青冈栎-长苞铁杉>其它树种-长苞铁杉。竞争强度与对象木的胸高直径服从双曲线回归关系,利用模型预测了长苞铁杉种内、种间的竞争强度。     

Influence of species and abiotic factors on extreme tree ring modulation:

 The influence of biotic (tree species) and abiotic factors (regional climate, altitude and stand aspect) on extreme tree-ring widths is dendroecologically investigated. Negative pointer years are observed when abrupt radial growth reductions (exceeding 40%) occurred synchronously in a given tree population. Pointer year intensity is expected to follow several theoretical models when trees are sampled along ecological gradients in various site conditions. Hypotheses are made about expected patterns of pointer year spatial distribution. They are compared with observed results obtained with Norway spruce [Picea abies (L.) Karst.] and silver fir (Abies alba Mill.) sampled in two French Alpine valleys (Tarentaise and Maurienne) with different regional climates, on two aspects and three altitude levels. Results reveal that extreme years follow expected patterns well. Thus, clear site modulations are observed in 1956 and 1986 (only trees located in Maurienne reacted). Moreover, species differences are observed, with typical cases of single species sensitivity (for spruce, occurring in 1962 and 1992). Abiotic factors such as altitude and aspect also involve clear pointer year patterns, such as narrow rings in 1976 due to summer drought that occurred especially at lower altitudes. However, some observed extreme years sometimes appear to be a combination of two (or three) of these one-factor theoretical models (e.g. 1921 with both lower altitude level and only observed in Tarentaise; 1948 involving both higher altitude and northern slopes). These results demonstrate the consistency of various tree-ring and climate relationships along ecological gradients.     

Non-native fishes and native species diversity in freshwater fish assemblages across the United States

The proliferation of non-native species in North American freshwater ecosystems is considered a primary threat to the integrity of native community structure. However, a general understanding of consistent and predictable impacts of non-native species on native freshwater diversity is limited, in part, because of a lack of broad-scale studies including data from numerous localities across multiple drainages. This study uses data from 751 localities collected during the United States Geological Survey (USGS) National Water Quality Assessment (NAWQA) program to examine the influence of non-native fish species on native freshwater fish assemblages across the United States. In general, no significant differences in native fish richness and diversity measures were detected between sites with only native species and sites containing non-native species. However, at sites with non-native species, the number of non-native species present was negatively correlated with native species richness and Shannon diversity and positively correlated with native evenness. Non-native piscivores were negatively correlated with native species richness and Shannon diversity and positively correlated with native evenness. Native piscivores were positively correlated with native richness and diversity and negatively correlated with native evenness at sites with only native species. Our results suggest that from a superficial perspective, native species richness and diversity are not different among sites with and without non-native species. However, when patterns of native species richness and diversity are examined at sites containing non-native species, correlations between non-native and native species richness and diversity imply the expected negative effect of invasive taxa. Additionally, non-native piscivores appear to have a significant negative effect on native taxa and possibly represent a novel selective force on naive native prey.     

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.     

When partitioning is not an option: Resource availability predicts intraguild interactions in two isolated Amazonian primate assemblages

Assemblage structure and acquisition of high-value resources will usually be affected by changes in resource availability and differential competitive abilities of assemblage members. In fragmented habitats where carrying capacity limits are exceeded due to high population densities and biomass, interspecific interactions can be expected to occur at a high frequency, potentially turning into an important cost for coexistence. We studied assemblage- and guild-level patterns of interspecific interactions in two highly diverse isolated primate assemblages in southern Amazonia. Specifically, we assessed the effects of temporal variation in fruit availability on the rates of interspecific interactions between gray woolly monkeys (Lagothrix lagotricha cana), one of the largest tree-dwelling mammals of the Amazon forests, and nine syntopic primate species. We found that fruit availability positively predicted rate of intraguild interactions in contrast to overall assemblage interaction rate. We did not find statistical evidence for the effect of fruit availability on the assemblage rates of type-dependent (i.e., agonistic or non-agonistic) and context-dependent (i.e., feeding or non-feeding) interactions. However, there was a clear trend toward increased feeding-related and agonistic-related interactions as fruit availability increased, with agonistic interactions mostly occurring at guild-level. These results provide support for a significant role of fruit availability in structuring spatial–temporal intraguild interactions at our study sites. Costly interspecific interactions and spatial habitat overlap can be expected to occur at a high frequency in highly diverse assemblages living in human-modified landscapes, which in turn, can have potentially negative impacts to the species involved.     

Asymmetric coexistence: bidirectional abiotic and biotic effects between goose barnacles and mussels

1. Species coexistence depends on the net effect of interacting species, representing the sum of multiple interaction components that may act simultaneously and vary independently depending on ambient environmental conditions. Consequently, for a comprehensive understanding of the compound nature of species interactions and coexistence, a mechanistic approach that allows a separate evaluation of each interaction component is required. 2. Two sessile filter-feeders, the goose barnacle Capitulum mitella and the mussel Septifer virgatus, coexist on moderately wave-exposed rocky shores in south-western Japan. In the upper intertidal, Capitulum positively influenced Septifer survivorship and growth through amelioration of thermal stress and of physical disturbance. On the other hand, these species are potential competitors as they have similar body sizes and modes of resource utilization. These opposite processes, facilitation and competition, are based on abiotic characteristics and biotic functions of the two species, respectively. 3. In order to quantify the bidirectional abiotic, biotic and net effects, a series of experimental manipulations was conducted involving the use of living neighbours with both abiotic and biotic effects, and artificial mimics to simulate abiotic effects without biotic effects. 4. Capitulum had strong positive abiotic effects on the mussel survivorship in most experimental periods, while the biotic effect was negligible or weakly negative, suggesting that the net effect of Capitulum on mussel survival was largely attributable to the abiotic effect. In contrast, a significantly negative biotic effect on the mussel growth rate was always present, though this was cancelled out by the larger, positive abiotic effect. In the case of Septifer, its abiotic and biotic effects on the survivorship of goose barnacles were negligible, while those on the growth rate showed temporal variation. 5. With respect to the relationship between species interaction and environmental conditions, the strength of abiotic facilitative effect of Capitulum on mussel survival increased with increasing abiotic stress, while the strength of biotic effect was negligible or weakly negative. As regards the effects of mussels on goose barnacles, our study indicated no obvious relationship. These results point to the importance of decomposing interaction for an accurate, mechanistic understanding of species relations and coexistence.     

Spatial and temporal variation in fish assemblages of drying stream pools: The role of abiotic and biotic factors

Droughts and summer drying create unusual temporary aquatic habitats in the form of isolated pools in many small streams around the world. To examine spatial and temporal variation in fish community structure of drying stream pools, their relation to abiotic environmental variables, and associations among species, fish were sampled during summer 1995 and 1996 from pools of four streams in the Ozark mountains, Arkansas, USA. Redundancy analysis of physical-chemical variables showed significant differences among stream sites, but no significant difference between years or stream site by year interaction. Stream sites separated consistently along axes one (habitat heterogeneity) and two (temperature/canopy cover) in both years. Redundancy analysis of fish species-size class densities showed a significant stream site by year interaction. Groupings of stream sites based on fish assemblages were not well explained by physical-chemical variables measured at the pool scale, but were related to location within the drainage basin, and these groupings differed between years. There were 27 (15.8%) and 10 (5.8%) significant associations found among fish species-size classes in 1995 and 1996, respectively, and all but two significant associations in 1995 were positive. Pool depth, habitat heterogeneity, pool size and dissolved oxygen/canopy cover were important local abiotic factors depending on response variables examined. In both years, large fish total density, large central stoneroller density (80 mm TL), and small sunfish (<80 mm TL) density were positively related to pool depth. Otherwise, there was no consistent relationship between physical-chemical variables and dependent variables (fish density and species richness) within a year or between years for a given dependent variable. These results support the hypothesis that local abiotic factors are important in structuring fish assemblages in harsh environments, but the importance of those factors varies temporally, and regional influences appear to override local abiotic conditions as factors structuring fish assemblages in drying stream pools. Predation by terrestrial vertebrates may also be an important factor structuring these fish assemblages that has been largely overlooked.