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1.
Fronds of clonal seaweeds with extensive holdfasts relative to frond size are known not to self‐thin during growth, even in crowded stands. We tested whether frond self‐thinning would occur for such a seaweed since these traits are more similar to those of unitary seaweeds, which do self‐thin in crowded conditions. We used Sargassum lapazeanum Setch. et N. L. Gardner (Fucales, Phaeophyceae) from the Pacific coast of Mexico, for which we first confirmed its clonal nature by performing a regeneration experiment in culture tanks. During the growth season (winter to late spring), S. lapazeanum stand biomass increased, while frond density and size inequality (Gini coefficient for frond biomass) decreased. These results indicate that self‐thinning occurred at the frond level. We propose a conceptual model for frond dynamics for clonal seaweeds in general. In stands of clonal species with small fronds and relatively extensive holdfasts (particularly when holdfasts are perennial), frond dynamics would be determined mostly by intraclonal regulation, which seems to prevent excessive crowding from occurring. Such species display a positive biomass–density relationship during the growth season. On the contrary, in stands of clonal species with large fronds relative to holdfast size, frond dynamics would be determined mostly by interactions among genets. For such species, self‐thinning may be detected at the frond level in crowded stands, resulting in a negative biomass–density relationship during growth.  相似文献   

2.
Scrosati 《Ecology letters》2000,3(3):191-197
For crowded stands of terrestrial plants, ranging from mosses to trees, plant (or ramet, for clonal plants) density is negatively related to stand biomass. Stand biomass and ramet density were determined for Mazzaella cornucopiae and for Pterocladiella capillacea , two morphologically distinct intertidal clonal red seaweeds, to compare them with terrestrial plants. For these seaweeds, ramet densities were similar to the highest values reported for terrestrial plants (mosses, specifically). Stand biomass was higher than average values expected from the terrestrial interspecific biomass–density relationship, but lower than the limits expected from the terrestrial ultimate biomass–density line. These seaweeds show unexpectedly low ramet slenderness and high biomass packing per unit of volume, compared with the trend observed for terrestrial plants. Possible explanations for these differences are related to the particular physiology and habitat of intertidal clonal seaweeds.  相似文献   

3.
Seagrasses are important habitat-formers and ecosystem engineers that are under threat from bloom-forming seaweeds. These seaweeds have been suggested to outcompete the seagrasses, particularly when facilitated by eutrophication, causing regime shifts where green meadows and clear waters are replaced with unstable sediments, turbid waters, hypoxia, and poor habitat conditions for fishes and invertebrates. Understanding the situations under which seaweeds impact seagrasses on local patch scales can help proactive management and prevent losses at greater scales. Here, we provide a quantitative review of available published manipulative experiments (all conducted at the patch-scale), to test which attributes of seaweeds and seagrasses (e.g., their abundances, sizes, morphology, taxonomy, attachment type, or origin) influence impacts. Weighted and unweighted meta-analyses (Hedges d metric) of 59 experiments showed generally high variability in attribute-impact relationships. Our main significant findings were that (a) abundant seaweeds had stronger negative impacts on seagrasses than sparse seaweeds, (b) unattached and epiphytic seaweeds had stronger impacts than 'rooted' seaweeds, and (c) small seagrass species were more susceptible than larger species. Findings (a) and (c) were rather intuitive. It was more surprising that 'rooted' seaweeds had comparatively small impacts, particularly given that this category included the infamous invasive Caulerpa species. This result may reflect that seaweed biomass and/or shading and metabolic by-products like anoxia and sulphides could be lower for rooted seaweeds. In conclusion, our results represent simple and robust first-order generalities about seaweed impacts on seagrasses. This review also documented a limited number of primary studies. We therefore identified major knowledge gaps that need to be addressed before general predictive models on seaweed-seagrass interactions can be build, in order to effectively protect seagrass habitats from detrimental competition from seaweeds.  相似文献   

4.
Biological invasions are acknowledged among the main drivers of global changes in biodiversity. Despite compelling evidence of species interactions being strongly regulated by environmental conditions, there is a dearth of studies investigating how the effects of non‐native species vary among areas exposed to different anthropogenic pressures. Focusing on marine macroalgae, we performed a meta‐analysis to test whether and how the direction and magnitude of their effects on resident communities and species varies in relation to cumulative anthropogenic impact levels. The relationship between human impact levels and non‐native species impact intensity emerged only for a reduced subset of the response variables examined. Yet, there was a trend for the effects of non‐native species on community biomass and abundance and on species abundance to become less negative at heavily impacted sites. By contrast, the magnitude of negative effects of seaweed on community evenness tended to increase with human impact levels. The hypothesis of decreasing severity of invader’ impacts along a gradient of habitat degradation was also tested experimentally at a regional scale by comparing the effects of the removal of non‐native alga, Caulerpa cylindracea, on resident assemblages among rocky reefs exposed to different anthropogenic pressures. Assemblages at urban and pristine site did not differ when invaded, but did so when C. cylindracea was removed. Our results suggest that, despite the generally weak relationship between human impacts levels and non‐native species impacts, more negative impacts can be expected in less stressful environments (i.e. less degraded or pristine sites), where competitive interactions are presumably the driving force structuring resident communities. Implementing strategies for controlling the establishment of non‐native seaweeds should be, thus, considered a priority for preserving biodiversity in relatively pristine areas. On the other hand, control of invaders at degraded sites could be warranted to lessen their role as propagule sources. Synthesis Local anthropogenic stressors that severely alter biotic and abiotic conditions may underpin context‐dependency in the impacts of biological invasions. We used a meta‐analysis and an experimental test to examine the relationship between cumulative human impacts and ecological impact of non‐native seaweeds on resident assemblages. Our results suggest that more negative impacts of non‐native seaweeds on the abundance and biomass of resident assemblages can be expected in less degraded or pristine sites. Possibly, stronger impacts prevail at pristine sites, where assemblages are mainly structured by biotic interactions. Hence, management efforts should be mostly directed to prevent the establishment and spread of non‐native seaweeds in pristine areas. On the other hand, weak, but positive effects of seaweeds at the most degraded sites add to the ongoing debate on the role of non‐native species in rehabilitation plans.  相似文献   

5.
Scrosati  Ricardo 《Hydrobiologia》1996,326(1):259-265
The negative relationship between stand biomass and plant density observed in terrestrial plants was tested among fronds of a clonal red alga, Mazzaella cornucopiae. Stand biomass and frond density were estimated bimonthly for 1 year on 7 permanent quadrats. A positive linear correlation was found between biomass and density for the whole data set, suggesting the lack of self-thinning among fronds of this intertidal alga at natural densities. Higher frond densities could favor increased water retention among fronds, thus minimizing desiccation during low tides. In this way, stands could maintain higher production of biomass. Fronds may also be cushioned better against wave action at higher frond densities, thus decreasing the detachment of larger fronds. The temporal variation of the relationship between biomass and density was plotted separately for these 7 quadrats. Four quadrats showed positive linear correlation between both variables (the other three quadrats showed non-significant positive linear correlation). Their four slopes are statistically similar to that found for the entire data set. It is possible, then, that there is only one positive slope for the biomass-density relationship in this population. If this is true, standing biomass could be estimated from the density of fronds.  相似文献   

6.
Fronds from crowded stands of clonal seaweeds, particularly those in which holdfasts are mostly perennial and are the major source of new fronds every year, are thought not to undergo self-thinning during the growth season, unlike those from crowded stands of unitary seaweeds. For clonal seaweeds, it is not known, however, what happens at the very end of the growth season, when crowding is highest for the year. By sampling twice more frequently than previously done for similar species, the possible occurrence of frond self-thinning was tested for Mastocarpus papillatus (Rhodophyta, Gigartinales, Petrocelidaceae) from western Canada during the growth season (spring) of 2003. Initially, stand biomass increased together with frond density, as found previously for similar clonal seaweeds. Shortly before stand biomass peaked for the year (June), frond density remained statistically unchanged. Thus, the increased sampling precision of this study confirms that fronds of these clonal seaweeds do not undergo self-thinning, not even shortly before crowding is highest. Frond size inequality for M. papillatus remained statistically similar during the growth season, which is also consistent with a model of no self-thinning. There are similarities in biomass–density dynamics and in size inequality dynamics between clonal seaweeds and clonal vascular plants.  相似文献   

7.
Little is known about the dynamics and the ecological interactions among ramets (fronds) from populations of clonal red seaweeds. Small ramets are very difficult to tag, so their growth cannot be monitored directly. The temporal variation of the relationship between stand biomass and ramet density offers information on ramet performance. We calculated this relationship for an intertidal population of Pterocladiella capillacea (Gmelin) Santelices et Hommersand (Gelidiales) from Baja California, Mexico. Biomass and density were positively correlated on an annual basis, indicating that biomass accumulated without involving self-thinning among ramets. This contrasts with nonclonal seaweeds, for which self-thinning among individuals occurs during growth, but agrees with other clonal red seaweeds, such as Chondrus crispus Stackhouse and Mazzaella cornucopiae (Postels et Ruprecht) Hommersand (both Gigartinales). The growth pattern for these members of the Gelidiales and of the Gigartinales holds despite differences in holdfast morphology and ramet branching degree and despite differences in the capacity of coalescence during early stages, known only for the Gigartinales. The positive slope for the dynamic biomass–density relationship, on a bilogarithmic scale, was statistically steeper for M. cornucopiae than for P. capillacea and for C. crispus. This suggests that the addition of new ramets during the growth season may be relatively more beneficial for biomass accumulation rates for M. cornucopiae. This would be expected for high-intertidal species subjected to strong abiotic stress, for which ramet crowding constitutes a key protection. Pterocladiella capillacea occurs at the mid-intertidal zone and C. crispus at the subtidal zone, so ramets would be relatively less important in that respect.  相似文献   

8.
Abstract

The aim of the present study was to investigate carbon and nutrient cycling and the role of root dynamics in terrestrial ecosystems such as large abandoned pastures and natural grasslands present in the Prealps, for which below‐ground processes are currently enigmatic. In particular, we quantified root/leaf biomass and C:N ratio throughout two growing seasons. Additionally, root traits such as root length density (RLD), root mass density (RMD), and root diameter classes (RDC) were also investigated with the aim of understanding the spatial distribution of roots in the soil. In our samples, we found that the roots could be divided into three main diameter classes and hence quantified the presence of each class along the soil profile. With regard to total root biomass, we found the occurrence of two peaks of biomass accumulation during the growth season, and when biomass accumulation was compared with climatic data, it was impossible to obtain a clear indication of the root turnover rate. In fact, the strong influence of grazing on the above‐ground biomass could have affected, in turn, root biomass. In future, this possible complication will be avoided by repeating the measurements within enclosures to avoid grazing interference. We found that C:N ratio remained constant, with a single peak, suggesting a lower root decomposition during the warmest period (August 2006). The concentration of nitrogen in roots decreased with depth as a result of a decrease in roots with smaller diameters. The reverse was found for carbon content, which increased with depth, probably due to an increase in roots with larger diameters. This study represents the first attempt to estimate root turnover rates in this prealpine ecosystem, which have been analysed to date only for the above‐ground biomass.  相似文献   

9.
Latitudinal gradients in species abundance and diversity have been postulated for nearshore taxa but few analyses have been done over sufficiently broad geographic scales incorporating various nearshore depth strata to empirically test these gradients. Typically, gradients are based on literature reviews and species lists and have focused on alpha diversity across the entire nearshore zone. No studies have used a standardized protocol in the field to examine species density among sites across a large spatial scale while also focusing on particular depth strata. The present research used field collected samples in the northern hemisphere to explore the relationships between macroalgal species density and biomass along intertidal heights and subtidal depths and latitude. Results indicated no overall correlations between either estimates of species density or biomass with latitude, although the highest numbers of both were found at mid-latitudes. However, when strata were examined separately, significant positive correlations were found for both species numbers and biomass at particular strata, namely the intertidal ones. While the data presented in this paper have some limitations, we show that latitudinal macroalgal trends in species density and biomass do exist for some strata in the northern hemisphere with more taxa and biomass at higher latitudes.  相似文献   

10.
Explaining variability in the strength and sign of trophic interactions between primary consumers and plants is a long‐standing research challenge. Consumer density and body size vary widely in space and time and are predicted to have interactive effects on consumer–plant interactions. In a southern US salt marsh, we used replicate field enclosures to orthogonally manipulate the body size (mass) and density of a dominant consumer (a snail). We investigated impacts (leaf damage and biomass) on monocultures of cordgrass, the foundation species, over three months. Increasing consumer density and body size increased leaf damage additively and, as predicted, multiplicatively reduced plant biomass. Notably, size and density determined the sign of consumer impact on plants: low to medium densities of small consumers enhanced, while high densities of large consumers strongly suppressed, plant biomass. Finally, total consumer metabolic biomass (mass0.75) within an enclosure parsimoniously explained plant biomass response, supporting theoretical predictions and suggesting that multiplicative effects of density and body size resulted from their effects on total metabolic biomass. The consequences of changes in consumer density and body size resulting from anthropogenic perturbations may therefore be predicted based on metabolic biomass. Synthesis Consumer size, density and biomass can all strongly affect consumer–plant interactions. Though density and body size have been extensively studied as drivers of variation in interaction strength, the role of biomass as the ultimate driver has been less appreciated. We manipulated body size and density of a single consumer species and, based on metabolic theory, integrated these into a single variable: total metabolic biomass. Our results suggest that changes in interaction strength attributed to size or density may in fact be due to changes in metabolic biomass. This metric could thus serve as a useful tool in further understanding species interactions.  相似文献   

11.
Numerous species of seaweed have been successfully cultivated in the sea for commercial purposes. Although considerable experimental work has been done on on-shore cultivation systems, none of these has yet proved to be economically viable on a sustained basis; nevertheless, such cultivation systems offer the potential for productivities greater than can be achieved in other systems. In on-shore systems, factors other than light can be controlled and provided at saturation levels. As density of biomass is controllable, all the light entering the cultivation system is absorbed. This results in efficient conversion of light energy to biomass when only light is limiting; moreover, density, rather than growth rate, is the major factor determining productivity. As growth of seaweeds in on-shore systems is only vegetative, there is no interruption for reproduction or maturation of the plants, and all of the net production can be recovered. Seaweeds have, though, relatively low percentage carbon composition, compared with terrestrial plants, and this may result in apparent high productivities based on dry matter.  相似文献   

12.
Abstract House mice (Mus domesticus L.) have been present on sub‐Antarctic Marion Island since the early 1800s. Several authors have suggested that an increase in mice density as a result of a general warming trend in the sub‐Antarctic climate from the 1960s has led to a decline in invertebrate biomass and abundance. These suggestions have been supported by the observation that the invertebrates of nearby mouse‐free Prince Edward Island are apparently larger and more numerous than on Marion. Our experiment was designed to determine whether mice have a direct effect on invertebrate abundance, biomass and community structure, or an effect on the vegetation community and thus potentially an indirect effect on invertebrates. We constructed five wire‐mesh mouse‐free exclosures in one habitat type on Marion Island and recorded both the soil macro‐invertebrate community and the vegetation inside and outside each of the exclosures before the start of the experiment in 1996 and twice thereafter (1998 and 2000). Mice had no significant effect on any of the eight prey groups' abundance or biomass, or on community structure (diversity and composition). Four of the prey groups changed significantly over time in either biomass or abundance, independent of the presence of mice. Our results, which may have been affected by generally low statistical power, suggest that factors other than mice had a larger impact on invertebrates than mice alone.  相似文献   

13.
Our understanding of natural patterns of fertilization in seaweeds has increased substantially over the last 10 years due to new approaches and methods to characterize the nature and frequency of fertilization processes in situ, to recognize the conditions and mechanisms enhancing fertilization success, and to anticipate population and community consequences of the patterns of natural fertilization. Successful reproduction in many species depends on a delicate juxtaposition of abiotic and biotic conditions. Important abiotic factors are those triggering gamete release (e.g. single or interacting effects of light quality and water movement) and those affecting gamete viability or concentrations (e.g. salinity effects on polyspermy blocks; gamete dilution due to water movement). Examples of important biotic components are synchronous gamete release, efficiency of polyspermy‐blocking mechanisms, population density of sexually fertile thalli, interparent distances, and male‐to‐female ratios. Field data indicate fertilization frequencies of 70%–100% in broadcasting‐type seaweeds (e.g. fucoids) and 30%–80% in brooding‐type (red) algae. Red algal values are higher than previously thought and challenge presently accepted explanations for their complex life histories. Important population and community questions raised by the recent findings relate to the magnitude of gene flow and exchange occurring in many micropopulations that seemingly breed during periods of isolation, the physiological basis and population effects of male‐to‐male competition and sexual selection during fertilization of brooding seaweeds, and the effects of massive gamete release, especially in holocarpic seaweeds, on benthic and planktonic communities. Comparative studies in other algal groups are now needed to test the generality of the above patterns, to provide critical pieces of information still missing in our understanding of natural fertilization processes, and to elucidate the evolutionary consequences of the different modes of reproduction (e.g. brooders vs. broadcasters).  相似文献   

14.
Yue Lin  Franka Huth  Uta Berger  Volker Grimm 《Oikos》2014,123(2):248-256
Metabolic scaling theory (MST) predicts a ‘universal scaling law’ for plant mass–density relationships, but empirical observations are more variable. Possible explanations of this variability include plasticity in biomass allocation between the above‐ and belowground compartment and different modes of competition, which can be asymmetric or symmetric. Although complex interactions of these factors are likely to occur, so far the majority of modelling and empirical studies has focussed on mono‐factorial explanations. We here present a generic individual‐based model, which allows exploring the plant mass–density relationship in realistic settings by representing plasticity of biomass allocation and different modes of competition in the above‐ and belowground compartment. Plants grew according to an ontogenetic growth model derived from MST. To evaluate the behavior of the simulated plants related to the allocation patterns and to validate model predictions, we conducted greenhouse experiments with tree seedlings. The model reproduced empirical patterns both at the individual and population level. Without belowground resource limitation, aboveground processes dominated and the slopes of mass–density relationships followed the predictions of MST. In contrast, resource limitation led to an increased allocation of biomass to belowground parts of the plants. The subsequent dominance of symmetric belowground competition caused significantly shallower slopes of the mass–density relationship, even though the growth of individual plants followed MST. We conclude that changes in biomass allocation induced by belowground resource limitation explain the deviations from the mass–density relationship predicted by MST. Taking into account the plasticity of biomass allocation and its linkage to the above‐ and belowground competition is critical for fully representing plant communities, in particular for correctly predicting their response of carbon storage and sequestration to changing environmental conditions.  相似文献   

15.
Local disturbance history affects patchiness of benthic river algae   总被引:3,自引:0,他引:3  
1. Recent research has shown that high‐flow events in streams leave a small‐scale mosaic of bed patches that have experienced scouring, sediment deposition (fill), or remained stable. Few studies have investigated if this ‘local disturbance history’ contributes to the patchy distribution of benthic organisms in streams and rivers. 2. In the present research, we demonstrate that local disturbance history in a mid‐sized river can have both short‐ and long‐term effects on epilithic algae. Chains buried vertically in the substratum of the river bed (236 in a 800‐m reach) indicated that two floods (return periods ≤1 year) caused a mosaic of bed patches with different disturbance histories. Once after the first and twice after the second flood, we sampled epilithic algae (mainly diatoms) in replicate patches that had been scoured, filled, or remained stable during the respective event. Algal biomass and cell density per substratum area were determined. 3. Three months after the first flood, algal biomass, total diatom density, diatom taxon richness, and densities of six of nine most common taxa were highest in fill patches. Six days after the second flood, biomass was highest in stable patches, indicating a refugium function of these patches. The refugium patches consisted of average‐sized stones, in contrast to previous studies of flood refugia for benthic algae in which these refugia were always large and/or immobile substrata. Four weeks after the second flood, diatoms tended to be most abundant in scour patches. With one exception, these differences between patch types could not be attributed to differences in local near‐bed current velocity or water depth. 4. The effects of disturbance history were more complex than a simple refugium function of stable patches because algal patterns changed with time since the last disturbance, possibly depending on the successional state of the algal mats.  相似文献   

16.
Plant succession theory underpins the development of strategies for the conservation and regeneration of native communities. Current theory has been based largely on space‐for‐time rather than long‐term monitoring data, which have known limitations. There is general consensus that more site‐specific studies are needed to corroborate existing hypotheses. The target vegetation is a brigalow (Acacia harpophylla, Mimosaceae) forest in one of Australia's most endangered ecosystems, which was cleared and burnt in 1963. Forty quadrats were placed systematically within each of six 20 m × 20 m permanent plots. Presence, density and per cent canopy cover data were recorded for each species at 18 times over 46 years. Brigalow dominated the original vegetation, assumed dominance soon after clearing through massive root suckering and remained dominant throughout the study. It achieved maximum density within two years when severe intraspecific competition led to self‐thinning. After approximately 30 years, vacant niches appeared. Woody understorey species were slow to recolonise. Species richness and other diversity indices increased rapidly to a maximum after 2–4 years, declined until the 30th year when they again increased. This was the pattern of the species‐rich herbaceous layer; woody species showed a steady monotonic increase. The ‘hump‐shaped’ relationship between cover (biomass) and species richness was confirmed. This example fits the inhibition model for which few examples have been described. While the long‐term successional pattern is slightly confounded by climatic variability preceding sample surveys, this space‐for‐time study not only supports a bimodal pattern of diversity over time but also indicates that the relative species richness of the herbaceous and woody layers may explain the extreme variability reported in the literature.  相似文献   

17.
The seasonal variability in the extraction yield, physicochemical characteristics, and rheological properties of ulvan from two Ulva species contributing to Brittany “green tides” has been studied. These seaweeds were collected in the water column for Ulva armoricana Dion, de Reviers et Coat and on hard substrata for Ulva rotundata Bliding. The maximum ulvan extraction efficiency was not related to the maximum ulvan content in the seaweeds, but with the active growth period of the seaweeds. Ulvan chemical structure, macromolecular characteristics, and rheological properties were affected by both species and seasons. The proportion of high‐molecular‐weight ulvan was the major factor positively correlated with the gelling properties. Characteristics of ulvan from U. rotundata subjected to tides were more affected by seasons than ulvan from U. armoricana living in a more constant environment. These results point to several useful recommendations concerning Ulva sp. biomass collected with regard to ulvan characteristics and uses.  相似文献   

18.
海藻有性繁殖生态学研究进展   总被引:3,自引:0,他引:3  
邹定辉  夏建荣 《生态学报》2004,24(12):2870-2877
由于研究技术与方法的不断改进 ,人们对海藻 (seaweeds)早期生活史方面的研究兴趣日益增大。评述了近年来海藻有性繁殖过程中配子释放和受精生态学问题。到达一定生理状态的海藻 ,通过对环境暗示应答而诱导配子形成。海藻生殖器官感受另一些特定的环境条件 ,通过有关信号传递机制 ,触发配子释放。海藻配子释放的时间及其所需的环境条件 ,依不同的种类而变化。配子同步释放以及各种适当的环境条件提高了受精频率和受精成功率 ,有利于受精的各种生物的和非生物的因素组合 ,形成了“机会窗 (the window of opportunity)”的概念。近年来的研究表明 ,海藻 (特别是红藻 )的自然受精成功率比以前所认为的要高得多。受精后合子 (胚 )的散布、集落与附着显著地影响其生存以及种群动态。对今后值得进一步研究的方面作了展望  相似文献   

19.
Seaweeds are the primary producers of all aquatic ecosystems. Chemical constituents isolated from diverse classes of seaweeds exert a wide range of nutritional, functional and biological activities. Unique metabolites of seaweeds possess specific biological properties that make them potential ingredients of many industrial applications such as functional foods, pharmaceuticals and cosmeceuticals. Cosmeceuticals of natural origin are becoming more popular than synthetic cosmetics. Hence, the investigation of new seaweeds derived functional components, a different source of natural products, has proven to be a promising area of cosmeceutical studies. Brown seaweeds also produce a range of active components including unique secondary metabolites such as phlorotannins and many of which have specific biological activities that give possibilities for their economic utilization. Brown seaweeds derived active compounds have been shown various functional properties including, antioxidant, antiwrinkling, whitening, antiinflammatory and antiallergy. It is well-known that these kind of biological effects are closely associated with cosmeceutical preparations. This communication reviews the current knowledge on brown seaweeds derived metabolites with various biological activities and the potential use as cosmeceutical ingredients. It is hoped that the reviewed literature on multifunctional properties of brown seaweeds will improve access to the seaweed based natural products specially the ability to incorporate these functional properties in cosmeceutical applications.  相似文献   

20.
This paper evaluates both the ideas and empirical evidence pertaining to the control of species density in herbaceous plant communities. While most theoretical discussions of species density have emphasized the importance of habitat productivity and disturbance regimes, many other factors (e.g. species pools, plant litter accumulation, plant morphology) have been proposed to be important. A review of literature presenting observations on the density of species in small plots (in the vicinity of a few square meters or less), as well as experimental studies, suggests several generalizations: (1) Available data are consistent with an underlying unimodal relationship between species density and total community biomass. While variance in species density is often poorly explained by predictor variables, there is strong evidence that high levels of community biomass are antagonistic to high species density. (2) Community biomass is just one of several factors affecting variations in species density. Multivariate analyses typically explain more than twice as much variance in species density as can be explained by community biomass alone. (3) Disturbance has important and sometimes complex effects on species density. In general, the evidence is consistent with the intermediate disturbance hypothesis but exceptions exist and effects can be complex. (4) Gradients in the species pool can have important influences on patterns of species density. Evidence is mounting that a considerable amount of the observed variability in species density within a landscape or region may result from environmental effects on the species pool. (5) Several additional factors deserve greater consideration, including time lags, species composition, plant morphology, plant density and soil microbial effects.Based on the available evidence, a conceptual model of the primary factors controlling species density is presented here. This model suggests that species density is controlled by the effects of disturbance, total community biomass, colonization, the species pool and spatial heterogeneity. The structure of the model leads to two main expectations: (1) while community biomass is important, multivariate approaches will be required to understand patterns of variation in species density, and (2) species density will be more highly correlated with light penetration to the soil surface, than with above-ground biomass, and even less well correlated with plant growth rates (productivity) or habitat fertility. At present, data are insufficient to evaluate the relative importance of the processes controlling species density. Much more work is needed if we are to adequately predict the effects of environmental changes on plant communities and species diversity.  相似文献   

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