Modifying native and exotic species richness correlations: the influence of fire and seed addition.

An important goal in restoration is to increase the richness of native species while reducing exotic species. However, native species richness is often positively correlated with exotic species richness. In a grassland-savanna system in Michigan (USA), we show that management that focuses on changing the nature of the exotic-native richness relationship can be used to restore native communities. Native and exotic species richnesses were positively correlated, likely due to a shared coupling with aboveground live biomass (a surrogate for productivity). The addition of native seed shifted the exotic-native richness relationship from a linear positive to a monotonic relationship: in areas of intermediate levels of exotic species richness, seed addition increased native diversity without an associated effect on exotic diversity, but in areas of high or low exotic richness, it did not affect native species richness. Prescribed burning broke the correlation between native and exotic richness with no consistent effect on the richness of either group. However, when burning was combined with native-seed addition, the relationship between native and exotic richness was maintained and was shifted upwards, enhancing native recruitment. Although aboveground productivity was strongly related to species richness across the landscape, changes in productivity did not drive these shifts.     

Disrupting mycorrhizal mutualisms: a potential mechanism by which exotic tamarisk outcompetes native cottonwoods

The disruption of mutualisms between plants and mycorrhizal fungi is a potentially powerful mechanism by which invasives can negatively impact native species, yet our understanding of this mechanism's role in exotic species invasion is still in its infancy. Here, we provide several lines of evidence indicating that invasive tamarisk (Tamarix sp.) negatively affects native cottonwoods (Populus fremontii) by disrupting their associations with arbuscular mycorrhizal (AM) and ectomycorrhizal (EM) fungi. At a field site in the early stages of tamarisk invasion, cottonwoods with tamarisk neighbors had reduced EM colonization and altered EM fungal community composition relative to cottonwoods with native neighbors, leading to reductions in EM propagule abundance in the soil beneath tamarisk. Similarly, AM colonization of cottonwoods was reduced with a tamarisk neighbor, but there were no significant changes in AM fungal spore communities or propagule abundance. Root colonization by nonmycorrhizal fungi, including potential pathogens, was higher in cottonwoods with tamarisk neighbors. A greenhouse experiment in which AM and EM inoculation and plant neighbor were manipulated in a fully factorial design showed that cottonwoods benefited from mycorrhizas, especially EM, in terms of shoot biomass when grown with a conspecific, but shoot biomass was similar to that of nonmycorrhizal controls when cottonwoods were grown with a tamarisk neighbor. These results are partially explained by a reduction in EM but not AM colonization of cottonwoods by a tamarisk neighbor. Tamarisk neighbors negatively affected cottonwood specific leaf area, but not chlorophyll content, in the field. To pinpoint a mechanism for these changes, we measured soil chemistry in the field and the growth response of an EM fungus (Hebeloma crustuliniforme) to salt-amended media in the laboratory. Tamarisk increased both NO3- concentrations and electrical conductivity 2.5-fold beneath neighboring cottonwoods in the field. Salt-amended media did not affect the growth of H. crustuliniforme. Our findings demonstrate that a nonnative species, even in the early stages of invasion, can negatively affect a native species by disrupting its mycorrhizal symbioses. Some of these changes in mycorrhizal fungal communities may remain as legacy effects of invasives, even after their removal, and should be considered in management and restoration efforts.     

Quadratic diversity: Its maximization can reduce the richness of species

In recent decades numerous diversity indices have been introduced. Among them the quadratic entropy index Q expresses the mean difference between two individuals chosen from the community at random. Differing from diversity indices habitually employed, Q does not satisfy a property postulated earlier for those measures. Namely, the uniform distribution of species does not necessarily yield the maximal index value. Q is based on the difference matrix of species. For a given matrix one can seek for the vector yielding the maximum quadratic entropy. This task leads to a quadratic programming problem. Using the appropriate program of a program package, we determined the maximum vector for a genetic difference matrix of crane species, as published in the literature. We discovered that some components (frequencies) in the maximum vector are equal to zero. That is, by maximizing the quadratic diversity some species can be eliminated. We discuss briefly the possible implications of this observation. Moreover, even if all elements in the maximum vector are positive, the elements can differ.     

Local richness along gradients in the Siskiyou herb flora: R. H. Whittaker revisited

In his classic study in the Siskiyou Mountains (Oregon, USA), one of the most botanically rich forested regions in North America, R. H. Whittaker (1960) foreshadowed many modern ideas on the multivariate control of local species richness along environmental gradients related to productivity. Using a structural equation model to analyze his data, which were never previously statistically analyzed, we demonstrate that Whittaker was remarkably accurate in concluding that local herb richness in these late-seral forests is explained to a large extent by three major abiotic gradients (soils, topography, and elevation), and in turn, by the effects of these gradients on tree densities and the numbers of individual herbs. However, while Whittaker also clearly appreciated the significance of large-scale evolutionary and biogeographic influences on community composition, he did not fully articulate the more recent concept that variation in the species richness of local communities could be explained in part by variation in the sizes of regional species pools. Our model of his data is among the first to use estimates of regional species pool size to explain variation in local community richness along productivity-related gradients. We find that regional pool size, combined with a modest number of other interacting abiotic and biotic factors, explains most of the variation in local herb richness in the Siskiyou biodiversity hotspot.     

Experimental tree removal in tallgrass prairie: variable responses of flora and fauna along a woody cover gradient

Woody plant encroachment is a worldwide phenomenon in grassland and savanna systems whose consequence is often the development of an alternate woodland state. Theoretically, an alternate state may be associated with changes in system state variables (e.g., species composition) or abiotic parameter shifts (e.g., nutrient availability). When state-variable changes are cumulative, such as in woody plant encroachment, the probability of parameter shifts increases as system feedbacks intensify over time. Using a Before-After Control-Impact (BACI) design, we studied eight pairs of grassland sites undergoing various levels of eastern redcedar (Juniperus virginiana) encroachment to determine whether responses of flora and fauna to experimental redcedar removal differed according to the level of pretreatment redcedar cover. In the first year after removal, herbaceous plant species diversity and evenness, woody plant evenness, and invertebrate family richness increased linearly with pretreatment redcedar cover, whereas increases in small-mammal diversity and evenness were described by logarithmic trends. In contrast, increases in woody plant diversity and total biomass of terrestrial invertebrates were accentuated at levels of higher pretreatment cover. Tree removal also shifted small-mammal species composition toward a more grassland-associated assemblage. During the second year postremoval, increases in herbaceous plant diversity followed a polynomial trend, but increases in most other metrics did not vary along the pretreatment cover gradient. These changes were accompanied by extremely high growing-season precipitation, which may have homogenized floral and faunal responses to removal. Our results demonstrate that tree removal increases important community metrics among grassland flora and fauna within two years, with some responses to removal being strongly influenced by the stage of initial encroachment and modulated by climatic variability. Our results underscore the importance of decisive management for reversing the effects of woody plant encroachment in imperiled grassland ecosystems.     

Context matters: matrix vegetation influences native and exotic species composition on habitat islands

The extensive research on plant communities of natural-habitat islands has primarily focused on the "islands." The island analogy, however, potentially limits understanding of processes influencing composition on habitat islands because the nature of their matrix is overlooked. We determine how plant community structure of the surrounding matrix influences vegetation on volcanic outcrops in the modified landscape of Banks Peninsula, New Zealand. Our primary purpose is to address whether the matrix is more important for recently established exotic species than it is for well-established native species and whether such invasion by exotics has led to homogenization of the outcrop flora. To test this, we examined our data at three spatial scales: that of the entire outcrop flora, between individual outcrops and their immediate surrounding matrix, and between individual outcrop faces and the individual relevés of the immediate surrounding matrix. We found that 81% of the native flora and 90% of the exotic flora also occur in the matrix. This high level of species shared between the outcrop and matrix persists at the scale of individual outcrop faces (68% of the total flora of individual faces is shared with the matrix). We predicted that floras from different outcrops would vary in their distinctiveness from their immediate matrix. We found Bray-Curtis distance coefficient values to range from 0.26 to 0.64; these were even higher at the outcrop-face scale. Variability in outcrop distinctiveness relates primarily to the outcrop face properties of area, vegetation height, and soil depth, and matrix properties of vegetation structure and vegetation heterogeneity. The effect of the vegetation structure of the matrix is more pronounced on the exotic than on the native outcrop flora. The component of composition and structure of the matrix that was independent of outcrop properties and local environment accounts for similar levels of explainable variation in total and native composition (29-31%), but considerably more (40%) in composition of exotic species. Our results support our prediction that, as the surrounding matrix becomes more modified, invasion by exotics makes outcrop vegetation less distinct from its matrix.     

Aquatic plant community invasibility and scale-dependent patterns in native and invasive species richness

Invasive species richness often is negatively correlated with native species richness at the small spatial scale of sampling plots, but positively correlated in larger areas. The pattern at small scales has been interpreted as evidence that native plants can competitively exclude invasive species. Large-scale patterns have been understood to result from environmental heterogeneity, among other causes. We investigated species richness patterns among submerged and floating-leaved aquatic plants (87 native species and eight invasives) in 103 temperate lakes in Connecticut (northeastern USA) and found neither a consistently negative relationship at small (3-m2) scales, nor a positive relationship at large scales. Native species richness at sampling locations was uncorrelated with invasive species richness in 37 of the 60 lakes where invasive plants occurred; richness was negatively correlated in 16 lakes and positively correlated in seven. No correlation between native and invasive species richness was found at larger spatial scales (whole lakes and counties). Increases in richness with area were uncorrelated with abiotic heterogeneity. Logistic regression showed that the probability of occurrence of five invasive species increased in sampling locations (3 m2, n = 2980 samples) where native plants occurred, indicating that native plant species richness provided no resistance against invasion. However, the probability of three invasive species' occurrence declined as native plant density increased, indicating that density, if not species richness, provided some resistance with these species. Density had no effect on occurrence of three other invasive species. Based on these results, native species may resist invasion at small spatial scales only in communities where density is high (i.e., in communities where competition among individuals contributes to community structure). Most hydrophyte communities, however, appear to be maintained in a nonequilibrial condition by stress and/or disturbance. Therefore, most aquatic plant communities in temperate lakes are likely to be vulnerable to invasion.     

Invasion through quantitative effects: intense shade drives native decline and invasive success.

The effects of invasive nonnative species on community composition are well documented. However, few studies have determined the mechanisms by which invaders drive these changes. The literature indicates that many nonnative plant species alter light availability differently than natives in a given community, suggesting that shading may be such a mechanism. We compared light quantity (photosynthetically active radiation, PAR) and quality (red: far-red ratio, R:Fr) in riparian reaches heavily invaded by a nonnative tree (Acer platanoides) to that in an uninvaded forest and experimentally tested the effects of our measured differences in PAR and R:Fr on the survival, growth, and biomass allocation of seedlings of the dominant native species and Acer platanoides. Light conditions representative of the understory of Acer platanoides-invaded forest decreased survival of the native maple Acer glabrum by 28%; Amelanchier alnifolia by 32%; Betula occidentalis by 55%; Elymus glaucus by 46%; and Sorbus aucuparia by 52%, relative to seedlings growing in PAR similar to that of native understories. In contrast, Acer platanoides and the native shrub Symphoricarpos albus were not affected by reductions in PAR. Acer platanoides seedlings and saplings are uniquely adapted to shade relative to native species. Acer platanoides was the only species tested that decreased allocation to roots relative to shoots in the invaded forest vs. the native forest light conditions. Therefore it was the only species to demonstrate an adaptive response to the particular light environment associated with Acer platanoides invasion as predicted by optimal partitioning theory. The profound change in light quantity associated with Acer platanoides canopies appears to act as an important driver of native suppression and conspecific success in invaded riparian communities. Further research is necessary to determine whether the effect of nonnative plant-driven changes on light quantity and quality is a widespread mechanism negatively affecting resident species and facilitating invasion by nonnatives.     

Additive effects of aboveground polyphagous herbivores and soil feedback in native and range-expanding exotic plants

Plant biomass and plant abundance can be controlled by aboveground and belowground natural enemies. However, little is known about how the aboveground and belowground enemy effects may add up. We exposed 15 plant species to aboveground polyphagous insect herbivores and feedback effects from the soil community alone, as well as in combination. We envisaged three possibilities: additive, synergistic, or antagonistic effects of the aboveground and belowground enemies on plant biomass. In our analysis, we included native and phylogenetically related range-expanding exotic plant species, because exotic plants on average are less sensitive to aboveground herbivores and soil feedback than related natives. Thus, we examined if lower sensitivity of exotic plant species to enemies also alters aboveground-belowground interactions. In a greenhouse experiment, we exposed six exotic and nine native plant species to feedback from their own soil communities, aboveground herbivory by polyphagous insects, or a combination of soil feedback and aboveground insects and compared shoot and root biomass to control plants without aboveground and belowground enemies. We observed that for both native and range-expanding exotic plant species effects of insect herbivory aboveground and soil feedback added up linearly, instead of enforcing or counteracting each other. However, there was no correlation between the strength of aboveground herbivory and soil feedback. We conclude that effects of polyphagous aboveground herbivorous insects and soil feedback add up both in the case of native and related range-expanding exotic plant species, but that aboveground herbivory effects may not necessarily predict the strengths of soil feedback effects.     

Terrestrial vertebrates alter seedling composition and richness but not diversity in an Australian tropical rain forest

Although birds and mammals play important roles in several mechanisms hypothesized to maintain plant diversity in species-rich habitats, there have been few long-term, community-level tests of their importance. We excluded terrestrial birds and mammals from fourteen 6 x 7.5 m plots in Australian primary tropical rain forest and compared recruitment and survival of tree seedlings annually over the subsequent seven years to that on nearby open plots. We re-censused a subset of the plots after 13 years of vertebrate exclusion to test for longer-term effects. After two years of exclusion, seedling abundance was significantly higher (74%) on exclosure plots and remained so at each subsequent census. Richness was significantly higher on exclosure plots from 1998 to 2003, but in 2009 richness no longer differed, and rarefied species richness was higher in the presence of vertebrates. Shannon's diversity and Pielou's evenness did not differ in any year. Vertebrates marginally increased density-dependent mortality and recruitment limitation, but neither effect was great enough to increase richness or diversity on open plots relative to exclosure plots. Terrestrial vertebrates significantly altered seedling community composition, having particularly strong impacts on members of the Lauraceae. Overall, our results highlight that interactions between terrestrial vertebrates and tropical tree recruitment may not translate into strong community-level effects on diversity, especially over the short-term, despite significant impacts on individual species that result in altered species composition.     

滇西北地区裸子植物多样性的地理分布格局及其与区系分化之间的关系

研究利用大尺度的物种分布信息,探讨了长江上游滇西北地区裸子植物多样性的地理分布格局及其与区系分化之间的关系。结果表明,在属和种的水平上,裸子植物的多样性从南到北呈递增趋势,但在科的水平上无明显变化趋势。在科和属的水平上,裸子植物的区系分化强度随着纬度的升高呈递增趋势。较高的裸子植物多样性和区系分化强度主要出现在北部地区。区系分化强度与表征地形复杂度的单位面积海拔高差之间存在显著的相关性。区系分化强度明显影响研究区域内裸子植物多样性的地理分布格局,这可能与地质历史上滇西北地区地层的差异性抬升和地形格局有关。     

Plant-soil feedback: testing the generality with the same grasses in serpentine and prairie soils

Plants can alter soil properties in ways that feed back to affect plant performance. The extent that plant-soil feedback affects co-occurring plant species differentially will determine its impact on plant community structure. Whether feedback operates consistently across similar plant communities is little studied. Here, the same grasses from two eastern U.S. serpentine grasslands and two midwestern tallgrass prairie remnants were examined for plant-soil feedback in parallel greenhouse experiments. Native soils were homogenized and cultured (trained) for a year with each of the four grasses. Feedback was evaluated by examining biomass variation in a second generation of (tester) plants grown in the trained soils. Biomass was lower in soils trained by conspecifics compared to soils trained by heterospecifics in seven of 15 possible cases; biomass was greater in conspecific soils in one other. Sorghastrum nutans exhibited lower biomass in conspecific soils for all four grasslands, so feedback may be characteristic of this species. Three cases from the Hayden prairie site were explained by trainer species having similar effects across all tester species so the relative performance of the different species was little affected; plants were generally larger in soils trained by Andropogon gerardii and smaller in soils trained by S. nutans. Differences among sites in the incidence of feedback were independent of serpentine or prairie soils. To explore the causes of the feedback, several soil factors were measured as a function of trainer species: nutrients and pH, arbuscular mycorrhizal (AM) spore communities, root colonization by AM fungi and putative pathogens, and functional diversity in bacterial communities as indicated by carbon substrate utilization. Only variation in nutrients was consistent with any patterns of feedback, and this could explain the greater biomass in soils trained by A. gerardii at Hayden. Feedback at Nottingham (one of the serpentine sites) differed, most notably for A. gerardii, from that of similar past studies that used different experimental protocols. To understand the consequences of feedback for plant community structure, it is important to consider how multiple species respond to the same plant-induced soil variation as well as differences in the feedback detected between greenhouse and field settings.     

Effects of vegetation cover, presence of a native ant species, and human disturbance on colonization by Argentine ants

The spread of non-native invasive species is affected by human activity, vegetation cover, weather, and interaction with native species. We analyzed data from a 17-year study of the distribution of the non-native Argentine ant (Linepithema humile) and the native winter ant (Prenolepis imparis) in a preserve in northern California (U.S.A.). We conducted logistic regressions and used model selection to determine whether the following variables were associated with changes in the distribution of each species: presence of conspecifics at neighboring sites, distance to development (e.g., roads, buildings, and landscaped areas), proportion of vegetation cover taller than 0.75 m, elevation, distance to water, presence of both species at a site, temperature, and rainfall. Argentine ants colonized unoccupied sites from neighboring sites, but the probability of appearance and persistence decreased as distance to development, vegetation cover, and elevation increased. Winter ants appeared and persisted in sites with relatively high vegetation cover (i.e., highly shaded sites). Presence of the 2 species was negatively associated in sites with high vegetation cover (more winter ants) and sites near development (more Argentine ants). Probability of colonization of Argentine ants decreased where winter ants were most persistent. At sites near development within the preserve, abundant Argentine ant populations may be excluding winter ants. The high abundance of Argentine ants at these sites may be due to immigration from suburban areas outside the preserve, which are high-quality habitat for Argentine ants. In the interior of the preserve, distance from development, low-quality habitat, and interaction with winter ants may in combination exclude Argentine ants. Interactions among the variables we examined were associated with low probabilities of Argentine ant colonization in the preserve.     

On the relationship between regional and local species richness: a test of saturation theory

What are the local community consequences of changes in regional species richness and composition? To answer this question we followed the assembly of microarthropod communities in defaunated areas of moss, embedded in a larger moss "region." Regions were created by combining moss from spatially distinct sites, resulting in regional species pools that differed in both microarthropod richness and composition, but not area. Regional effects were less important than seasonality for local richness. Initial differences in regional richness had no direct effect on local species richness at any time along a successional gradient of 0.5-16 months. The structure of the regional pool affected both local richness and local composition, but these effects were seasonally dependent. Local species richness differed substantially between dates along the successional gradient and continued to increase 16 months after assembly began. To the best of our knowledge, this is the first critical test of saturation theory that experimentally manipulates regional richness. Further, our results failed to support the most important mechanisms proposed to explain the local richness-regional richness relationship. The results demonstrate that complicated interactions between assembly time, seasonality, and regional species pools contribute to structuring local species richness and composition in this community.     

Diving deep in a foraging hotspot: acoustic insights into bottlenose dolphin dive depths and feeding behaviour

To exploit resources in their environment, odontocete cetaceans have evolved sophisticated diving abilities to allow effective foraging. However, data on the diving behaviour and underwater foraging behaviour remains limited. This study made use of echolocation clicks and other calls to study the diving behaviour of bottlenose dolphins. Dolphins used the full water column and consistently dived to depths of around 50 m, close to the seabed. However, the majority of their time appeared to be spent within the surface layers of the water column. In addition, by localising calls that have been associated with prey capture events (Janik, Proc R Soc Lond Ser B 267:923–927, 2000a), it appeared that certain forms of feeding behaviour occurred primarily at depths of between 20 and 30 m. Furthermore, data on the depth of clicks made before and after these feeding calls suggested that during the minute before the calls, dolphins were consistently diving from the surface to depths close to the seabed, and were subsequently returning to the surface after the calls. This passive acoustic technique proved an accurate method for studying the depth distribution of dolphin vocalisations. By exploiting the natural sounds made by these wild odontocetes, this investigation provided a previously unavailable perspective on the the 3D nature of bottlenose dolphins foraging behaviour. It confirmed that while the dolphins spent the majority of time close to the surface, the full water column was exploited during foraging events.     

Breeding systems and seed size in a neotropical flora: testing evolutionary hypotheses

A well-known, but largely untested, prediction in plant reproductive ecology is that dioecious taxa should produce larger, more, higher-quality, or better-defended seeds than cosexual taxa. Using a data set composed of 972 species in 104 families, representing the flora of the Tambopata Wildlife Reserve (Madre de Dios, Peru), we evaluated the first component of this prediction, examining ecological and evolutionary relationships between breeding system and mean seed size with two kinds of tests. First, we conducted cross-species analyses to determine whether species with different breeding systems differed significantly with respect to mean individual seed size. Second, we used a hypothesized phylogeny to identify pairs of the most closely related taxa or clades within the Tambopata community that differed with respect to breeding system. Comparing pair members allowed us to determine whether evolutionary divergence in breeding system (between taxa with unisexual vs. cosexual individuals) was consistently associated with evolutionary change in seed size. In both analyses, we controlled for potentially confounding effects of growth form by examining these relationships within woody and nonwoody taxa. Cross-species analyses revealed that dioecious species produced larger seeds than cosexual species among woody species, shrubs, lianas (each growth form analyzed separately), and all species pooled, but not among trees. Phylogenetically independent contrasts upheld the significant association between breeding system and seed size among woody taxa, lianas, and all taxa pooled, but not among shrubs. We discuss the implications of our findings for evolutionary hypotheses regarding associations between dioecy and seed size.     

The influence of productivity on the species richness of plants: a critical assessment

Despite much scrutiny the relationship between productivity and species richness remains controversial, and there is little agreement about causal processes. We present the results of a survey of 159 productivity-plant species richness (P-PSR) relationships from 131 published studies. We critically assessed each study with respect to experimental design and for the appropriateness of the surrogates used for productivity. We were able to accept only 60 of the reported relationships as robust tests of the P-PSR relationship and a further 18 as robust tests of the biomass species richness relationship. Previous analyses have found that unimodal P-PSR relationships predominate. In contrast, we found that, in studies that used data of continental to global extent, all P-PSR relationships were positive regardless of grain, that almost all were also positive in data sets of regional extent, and that unimodal relationships were not dominant even in studies of fine grain or small spatial extent. Our results differ substantially from previous meta-analyses because previous studies have included a large number of studies that do not meet basic experimental design criteria for objectively testing P-PSR relationships. These results have important implications for theory that attempts to explain species richness patterns. We critically review four dominant theories in light of our results and develop new falsifiable predictions of relationship from these theories at both small and large spatial scales.     

Metacommunity models predict the local-regional species richness relationship in a natural system

Many natural communities exhibit positive relationships between local and regional species richness (LSR-RSR relationships), which can be either linear or curvilinear. Previous models have shown that the form of this relationship depends on the relative rates of colonization and extinction and the sensitivity of these rates to competition. We use simple models to show that the LSR-RSR relationship also depends on the type of metacommunity structure (Levins-like or mainland-island), and our models generate a wider range of realistic forms than do most previous models. We parameterize and test our models with two independent data sets for Daphnia in rock pools on islands in Finland and Sweden. We find that the Levins-like model with competition correctly predicts the observed LSR-RSR relationship and provides the best fit to the average local species richness per island. Simulations show that our models are robust to relaxing our assumption of identical species properties. Our study is one of the first to make and successfully test quantitative predictions for how a widely studied community pattern, the LSR-RSR relationship, arises from metacommunity dynamics.