Plant coexistence in coastal heaths: Habitat segregation in the post-fire environment

Abstract Previous studies of heaths on Pleistocene coastal sands showed consistent variation in patterns of floristic composition between areas on ridges and slopes but not within them. A large wild-fire that swept this system provided an opportunity to observe temporal processes in habitat segregation of species. Ridges and slopes were found to differ in species richness of seedlings in 0.0625 m2 quadrats and there was no evidence that the species richness in the two habitats was converging over 3 years. This suggests that initially these processes differ in space between the two habitats with more species in the wet heath being packed into a smaller area than in the dry heath. Seed-banks of species did not saturate available space for recruitment in either habitat, but seedling densities differed asymmetrically between habitats across two pairs of species studied. Experimental manipulation of seeds among habitats also showed distinct differences in establishment and survival among representative species from different habitats. The presence of seed-banks of wet-heath species in dry heath suggested that recruitment of their seedlings may occur there, but experiments showed that seedlings that arise from them do not survive there. Conversely, although no seed-bank of either dry-heath species studied was found in wet heath, our manipulations showed that their seedlings could grow there. Thus, superficially, physiological tolerance appeared to limit regeneration of wet-heath species in dry heath. However, there appeared to be no physiological limit for dry-heath plants to survive in wet heath. This paper shows that segregation of species may operate at either of two stages of the life cycle: at dispersal when safe sites for establishment are required, and at recruitment after seedlings have established. These findings highlight the importance of the regeneration niche in structuring community composition in coastal heaths, and contrast with traditional explanations of species segregation invoking physiological tolerance and competition at later life-history stages.     

Plant coexistence in coastal heaths: Floristic patterns and species attributes

Abstract An investigation was made of plant coexistence in coastal heaths. A 4 km stretch of Pleistocene beach sand was selected, and patterns of variation in heath vegetation on ridges and slopes in the beach system were studied. The sampling design covered variation with geographic distance among transects, plots within transects and quadrats within plots which were orthogonal to the alignment of ridges and slopes. Cover of vascular plant species was scored within each quadrat and both multivariate and univariate analyses tested the hypothesis that variation in the data was associated with environmental difference and geographic distance. There was a strong flohstic variation between the ridge and slope habitats. However, within these habitats there was no significant variation with distance at scales of kilometres, nor at hundreds of metres within transects nor at tens of metres within plots in floristic composition, in numbers of species, and in cover. Though differing floristically, the vegetation of the slopes and ridges had similar curves of importance values and fairly similar spectra of life forms and forms of post-fire regeneration in their constituent species, although the ridges had slightly more obligate-seeding species and fewer species of sedges and herbs than the slopes. The two habitats shared several genera, however, within such genera, species with similar attributes mostly occurred in different habitats. Hypotheses were suggested to explain segregation of species between the two habitats and the coexistence of species within habitats. Since patterns of variation among habitats were consistent over both smaller and larger scales of distance examined, experiments to test hypotheses may be designed so that their findings might be applied generally to the whole system studied.     

Interacting coexistence mechanisms in annual plant communities: Frequency-dependent predation and the storage effect

We study frequency-dependent seed predation (FDP) in a model of competing annual plant species in a variable environment. The combination of a variable environment and competition leads to the storage-effect coexistence mechanism (SE), which is a leading hypothesis for coexistence of desert annual plants. However, seed predation in such systems demands attention to coexistence mechanisms associated with predation. FDP is one such mechanism, which promotes coexistence by shifting predation to more abundant plant species, facilitating the recovery of species perturbed to low density. When present together, FDP and SE interact, undermining each other’s effects. Predation weakens competition, and therefore weakens mechanisms associated with competition: here SE. However, the direct effect of FDP in promoting coexistence can compensate or more than compensate for this weakening of SE. On the other hand, the environmental variation necessary for SE weakens FDP. With high survival of dormant seeds, SE can be strong enough to compensate, or overcompensate, for the decline in FDP, provided predation is not too strong. Although FDP and SE may simultaneously contribute to coexistence, their combined effect is less than the sum of their separate effects, and is often less than the effect of the stronger mechanism when present alone.     

Quantifying the relative importance of variation in predation and the environment for species coexistence

Coexistence and food web theory are two cornerstones of the long‐standing effort to understand how species coexist. Although competition and predation are known to act simultaneously in communities, theory and empirical study of these processes continue to be developed largely independently. Here, we integrate modern coexistence theory and food web theory to simultaneously quantify the relative importance of predation and environmental fluctuations for species coexistence. We first examine coexistence in a theoretical, multitrophic model, adding complexity to the food web using machine learning approaches. We then apply our framework to a stochastic model of the rocky intertidal food web, partitioning empirical coexistence dynamics. We find the main effects of both environmental fluctuations and variation in predator abundances contribute substantially to species coexistence. Unexpectedly, their interaction tends to destabilise coexistence, leading to new insights about the role of bottom‐up vs. top‐down forces in both theory and the rocky intertidal ecosystem.     

Scaling the effects of predation and disturbance in a patchy environment

The effects of hydraulic disturbances on the impact of two predatory benthic invertebrates on their prey were examined in a stream at two distinct spatial scales. At the scale of small habitat patches (0.0625 m²), hydraulic patch type was an important determinant of the microdistribution of prey and predators. Prey abundances were similar across all patch types at baseflow, but local densities were higher in patches identified as low-flow refugia after periods of high and fluctuating flow. The microdistribution pattern of predatory larvae of a caddisfly, Plectrocnemia conspersa, was similar to that of its prey, whereas predatory larvae of an alderfly, Sialis fuliginosa, did not shift their microdistribution significantly with discharge and were always most abundant in lowflow refugia. There was little evidence of an aggregative response of predators with prey, even though both predators and prey are mobile. Both predator species showed similar patch-specific patterns of per capita consumption rates: uniform consumption rates across hydraulic patch types at low and moderate flows, but highest in flow refugia during high flows. Species-specific patterns, however, were apparent in the magnitude and direction of differences between consumption rates during disturbance events, and in comparable patches at base flow: At high flow, consumption rates for P. conspersa were exaggerated (3.9 times higher) in flow refugia but at par in other patches; for S. fuliginosa they were at par in flow refugia but reduced in other patches (up to 3.3. times lower). These differences may be related to species-specific foraging behaviours (search vs ambush predators) and the influence of prey movements on feeding success. Using the patch-scale results only, it is difficult to predict the effects of physical disturbance on predation intensity at the larger scales of whole habitats, populations or communities. At the large scale (>200 m²), net predator impacts were estimated over the stream reach, using a spatially explicit model that accounts, in an additive way, for habitat heterogeneity and patch-specific responses of predators and prey. The relationship between predator impact over the whole reach and hydraulic disturbance differed for the two predators. The predator impact of S. fuliginosa decreased with increasing hydraulic disturbance, as predicted by the harsh-benign hypothesis. There was no directional trend for P. conspersa, however, and maximum predator impact may occur at intermediate disturbance levels. For the prey community in this stream, predation pressure from S. fuliginosa appears to fluctuate directly with the discharge hydrograph, whereas predation from P. conspersa may be more persistent. Flow refugia may play a dual role in the sructure of stream communities by preventing catastrophic mortality of animals (predators and prey) from physical forces during disturbances, and by maintaining (or perhaps increasing) predation pressure. Summing the effects of species interactions in small habitat patches to the larger scale of a whole stream reach indicates that the scale of approach influences the observed patterns and their implied underlying process.     

Plant species richness and community productivity: why the mechanism that promotes coexistence matters

This paper stresses that the mechanism of coexistence is the key to understanding the relationship between species richness and community productivity. Using model plant communities, we explored two general kinds of mechanisms based on resource heterogeneity and recruitment limitation, with and without any trade-off between reproductive and competitive abilities. We generated different levels of species richness by changing model parameters, in particular the number of species in the regional pool, the degree of recruitment limitation, and the level of heterogeneity. Different diversity–productivity patterns are obtained with different coexistence mechanisms, indicating there is no reason to expect any general relationship between species richness and productivity. We discuss these results in the context of the within-site and across-site aspects of the relationship between species richness and productivity. Furthermore, we extend these results to hypothesize the relationship between species richness and productivity for other coexistence mechanisms not explicitly considered here.     

The effects of fire, local environment and time on ant assemblages in fens and forests

We investigated the effects of the abiotic environment, plant community composition and disturbance by fire on ant assemblages in two distinct habitat types in the Siskiyou Mountains in northern California and southern Oregon, USA. Sampling over 2 years in burned and unburned Darlingtonia fens and their adjacent upland forests, we found that the effects of disturbance by fire depended on habitat type. In forests, fire intensity predicted richness in ant assemblages in both years after the fire, and plant community composition predicted richness 2 years after the fire. No factors were associated with richness in the species‐poor fen ant assemblages. Species‐specific responses to both habitat type and disturbance by fire were idiosyncratic. Assemblage composition depended on habitat type, but not disturbance by fire, and the composition of each assemblage between years was more dissimilar in burned than unburned sites.     

Plant communities,species richness and their environmental correlates in the sandy heaths of Little Desert National Park,Victoria

Plant community composition and its likely environmental controls were investigated for 200 sample plots (each 100 m²) from Mediterranean-type vegetation in the Little Desert National Park, Victoria. TWINSPAN classification revealed four readily identifiable vegetation types; mallee-broombush, heathland, stringybark open woodland, and an assemblage intermediate between mallee-broombush and heathland referred to here as broom-heath. Mallee-broombush was found on Parilla Sands characterized by high Ca levels relative to heathland and stringybark open woodlands on unconsolidated Lowan Sands. The first axis of a 2 dimensional non-metric MDS ordination also divided heathlands (high axis scores) from mallee-broombush (low scores), while the second separated these vegetation types from stringybark woodlands and broom-heath. Vector-fitting revealed significant correlations between the locations of samples in ordination space and exchangeable soil Ca, soil colour, aspect and Shannon–Weiner diversity. Highest species richness/diversity was associated with the ecotonal area between Parilla and Lowan Sands (i.e. broom-heath) where a number of species characteristic of different assemblages had overlapping ranges. The fire-sensitive conifer, Callitris rhomboidea, was preferentially located in stringybark woodland and broom-heath vegetation types. Its presence was positively associated with high species richness and aspects having a southerly component. Four Callitris stands sampled for population structure were all > 40 years old and showed evidence of interfire recruitment from seeds released by old, serotinous cones. Overall, results suggest that variations in plant community composition and structure in the eastern block of the Little Desert are primarily due to variations in soil properties associated with the distribution of the two dominant substrate types, Parilla Sand and Lowan Sand. However, the interplay of topography and fire behaviour has probably been more important than substrate type in determining the distribution and population structure of longer-lived, fire-sensitive species such as Callitris rhomboidea.     

Do rabbits eat voles? Apparent competition,habitat heterogeneity and large‐scale coexistence under mink predation

Habitat heterogeneity is predicted to profoundly influence the dynamics of indirect interspecific interactions; however, despite potentially significant consequences for multi-species persistence, this remains almost completely unexplored in large-scale natural landscapes. Moreover, how spatial habitat heterogeneity affects the persistence of interacting invasive and native species is also poorly understood. Here we show how the persistence of a native prey (water vole, Arvicola terrestris ) is determined by the spatial distribution of an invasive prey (European rabbit, Oryctolagus cuniculus ) and directly infer how this is defined by the mobility of a shared invasive predator (American mink, Neovison vison ). This study uniquely demonstrates that variation in habitat connectivity in large-scale natural landscapes creates spatial asynchrony, enabling coexistence between apparent competitive native and invasive species. These findings highlight that unexpected interactions may be involved in species declines, and also that in such cases habitat heterogeneity should be considered in wildlife management decisions.     

Carrion fly community dynamics: patchiness, seasonality and coexistence

ABSTRACT. 1. An outdoor cage experiment was conducted to test the theory that localized interactions facilitate coexistence of species breeding in ephemeral habitats. The same amount of larval resource (50 g of liver) but divided into one, two, four, eight or sixteen pieces was placed into each of fifteen cages once a week from June to September for 4 years.
2. The cage populations were initially mixtures of thirteen species but after 4 years only Lucilia illustris (Meig.), Sarcophaga scoparia Pand. and S.aratrix Pand. remained. Lucilia illustris , by far the most abundant species in the field, predominated in every cage.
3. Sarcophaga went extinct in most cages, but they survived in five cages in which the larval resource was much divided, in agreement with the theoretical prediction.
4. Lucilia silvarum (Meig.) emerged about a week later than L.illustris in early summer; this phenological difference gave rise to a competitive asymmetry between the two species. This difference appears to explain why L.silvarum went extinct in the cages in 3 years, and it may explain why L.silvarum has consistently been less abundant than L.illustris in the natural community.
5. Other differences in phenology, voltinism and rate of development in the flies are described, and such differences are suggested to facilitate regional coexistence of many species in seasonal environments.     

Subthreshold and superthreshold coexistence of pathogen variants: the impact of host age-structure

It is well known that in the most general epidemic models with multiple pathogen variants a competitive exclusion principle is valid, such that the variant with the highest reproduction number eliminates the rest. Mechanisms such as super-infection, coinfection, and cross-immunity can lead to pathogen polymorphism where multiple strains coexist. It is also known that variability of infectivity with host age can destabilize the endemic equilibrium and cause oscillations. In this article we show that the hosts' chronological age can itself lead to coexistence of microparasites in the most basic model where competitive exclusion will occur without the age structure. Moreover, the host age-structure leads to multiple subthreshold dominance equilibria, and both weakly and strongly subthreshold coexistence. We find that the two pathogens cannot cooperate to persist subthreshold if neither one of them can persist subthreshold by itself. If, however, one of them can persist subthreshold by itself, it can cause the two pathogens to coexist in a strongly subthreshold equilibrium. The second strain that persists subthreshold through the mediation of the first always has a lower virulence. Our results show that age structure in infectivity can permit the coexistence of competing pathogens when the incidence is of proportionate mixing type (frequency-dependent transmission) and at least one of the strains is virulent.     

Community structure of branchiopods (Anostraca,Notostraca and Conchostraca) in the Banat province in Yugoslavia

A survey of large branchiopods in the Banat provincein Yugoslavia revealed a particularly diverse fauna in the vicinity of the village Melenci, 100 km north of Belgrade. The temporary ponds here include 10 species of branchiopods, 5 anostracans, 2 notostracans and 3conchostracans (all belonging to the Spinicaudata group). This diversity represents 53% of the species known in Yugoslavia. Community structure, spatial distribution and seasonal dynamics of the species were analyzed over the period 1991–1995. More than one species was found in about 67% of ponds. The maximum number of coexisting species was 7.Anostracans were the most common as a group, while the notostracan Triops cancriformis and the conchostracan Leptestheria saetosa were the most frequent species. A high frequency of coexistence was established. Regarding community structure, two significantly different periods during the season were distinguished: the first, from the end of February to mid-April, and the second, from mid-April to the end of July. This extensive pool longevity may underlie the high diversity in this region. This revised version was published online in August 2006 with corrections to the Cover Date.     

深圳杨梅坑山地植物群落演替动态

以深圳市杨梅坑山地两个植物群落为调查对象,分析其物种组成、物种多样性特征、重要值的变化,探讨不同阶段植物群落的演替动态规律。结果表明,两个植物群落经过7年演替后,植物群落的丰富度有所增加,且种类组成出现明显差异,两个植物群落在不同时期所呈现的自然景观不一样。群落科、属、种的数量差异明显。原鸭脚木-九节-铁线蕨群落2013年共有26科36属39种,2018年共有19科29属30种,2019年共有32科51属61种;原绒毛润楠-瓜馥木-团羽铁线蕨群落2013年共有25科30属33种,2018年共有24科33属33种,2019年共有42科67属83种。两个植物群落乔木层、灌木层、草本层三个层次的物种多样性指数和Pielou均匀度指数几乎都呈先下降再上升的趋势。演替系列中,乔木层、灌木层和草本层的物种组成均表现出耐荫种替代非耐荫种的趋势,且植物群落的层次由简单趋向复杂。     

Dynamics in a discrete two-species competition model: coexistence and over-compensation

The dynamic features of an over-compensating discrete two-species competition system with stable coexistence are recaptured, and it is shown how the probabilities of the different possible ecological scenarios, e.g. coexistence, may be calculated when the assumption of no over-compensation is loosened. A Bayesian methodology for calculating the probability that stable oscillations or chaos may occur in plant populations or communities is outlined. The methodology is exemplified using an experimental population of Arabidopsis thaliana. It is concluded that, when making ecological predictions it is preferable and possibly important to test for the possibility of chaotic population dynamics due to over-compensation rather than assuming a priori that over-compensation does not occur.     

植物竞争和昆虫取食调节入侵植物对土壤细菌群落和功能的影响

以入侵植物空心莲子草(Alternanthera philoxeroides(Mart.)Griseb)、本土近缘种莲子草(Alternan-thera sessilis(L.)DC)、生防昆虫莲草直胸跳甲(Agasicles hygrophila(Selman&Vogt))和本地昆虫虾钳菜披龟甲(Cassida pi...     

An idealized model for tree–grass coexistence in savannas: the role of life stage structure and fire disturbances

1.  We discuss a simple implicit-space model for the competition of trees and grasses in an idealized savanna environment. The model represents patch occupancy dynamics within the habitat and introduces life stage structure in the tree population, namely adults and seedlings. A tree can be out-competed by grasses only as long as it is a seedling.
2.  The model is able to predict grassland, forest, savanna and bistability between forest and grassland, depending on the different characteristics of the ecosystem, represented by the model's parameters.
3.  The inclusion of stochastic fire disturbances significantly widens the parameter range where coexistence of trees and grasses is found. At the same time, grass-fire feedback can induce bistability between forest and grassland.
4.   Synthesis . These results suggest that tree–grass coexistence in savannas can be either deterministically stable or stabilized by random disturbances, depending on prevailing environmental conditions and on the types of plant species present in the ecosystem.     

The enemy of my enemy is my friend: intraguild predation between invaders and natives facilitates coexistence with shared invasive prey

Understanding and predicting the outcomes of biological invasions is challenging where multiple invader and native species interact. We hypothesize that antagonistic interactions between invaders and natives could divert their impact on subsequent invasive species, thus facilitating coexistence. From field data, we found that, when existing together in freshwater sites, the native amphipod Gammarus duebeni celticus and a previous invader G. pulex appear to facilitate the establishment of a second invader, their shared prey Crangonyx pseudogracilis. Indeed, the latter species was rarely found at sites where each Gammarus species was present on its own. Experiments indicated that this may be the result of G. d. celticus and G. pulex engaging in more intraguild predation (IGP) than cannibalism; when the ‘enemy’ of either Gammarus species was present, that is, the other Gammarus species, C. pseudogracilis significantly more often escaped predation. Thus, the presence of mutual enemies and the stronger inter- than intraspecific interactions they engage in can facilitate other invaders. With some invasive species such as C. pseudogracilis having no known detrimental effects on native species, and indeed having some positive ecological effects, we also conclude that some invasions could promote biodiversity and ecosystem functioning.     

拉鲁湿地主要植物群落结构及物种多样性

2002年6~9月对拉鲁湿地植物种类、群落结构、生物量以及物种多样性进行了调查研究.结果表明:(1)拉鲁湿地共有植物30科52属85种,主要有芦苇[Phragmites australis(Cav.)Trin.ex Steud.](A)、尼伯尔酸模(RumexnepalensisSpreng)(B)、水葱(Scirpus validusVahl)(C)、穗状狐尾藻(MyriophyllumspicatumL.)(D)、灯芯草(Juncus effususL.)(E)、针叶苔草(CarexonoeiFranch.et Sav.)和黑褐苔草(CarexatrofuscaSchkuhr)(F)、藏北嵩草(KobresialittledaleiC.B.Clarke)(G)、藏蓟[Cirsiumlanatum(Roxb.ex Willd.)Spreng.](H)等8个主要植物群落.(2)在8个主要植物群落中,嵩草群落所含物种数最多,为44种,穗状狐尾藻群落最少,仅8种;芦苇群落生物量最高,为124.5~1 027.0 g/m2,藏蓟群落生物量最低,为124.5~220.5 g/m2;优势种芦苇的生物量最高,为183.5~660.0 g/m2,藏蓟生物量最低,为43.1~76.3 g/m2.(3)各群落的垂直结构特点是:芦苇群落、水葱群落、穗状狐尾藻群落具有挺水层、浮叶层和沉水层,其他群落只具有可分为高、低两层的草本层.(4)在各群落的相似性方面,嵩草群落与藏蓟群落、水葱群落与芦苇群落和尼伯尔酸模群落的相似性要明显高于其他群落,其Jaccard系数分别为0.549 0、0.538 5、0.500 0;芦苇群落、尼伯尔酸模群落、水葱群落、穗状狐尾藻群落与藏蓟群落以及芦苇群落、穗状狐尾藻群落与嵩草群落在建群种上没有相似之处,Jaccard系数为0.     

Early post-fire succession in California chaparral: Changes in diversity, density, cover and biomass

For four consecutive years, following the fires in November 1993, temporal variations in species richness, cover and biomass of component plant groups in early post-fire chaparral succession were monitored on different aspects at the Stunt Ranch Santa Monica Mountains Reserve, southern California. Plant groups were categorized based on growth form, life form, ability to fix nitrogen, geographic origin and regeneration strategies. North-facing slopes exhibited higher species richness, higher species turnover rate over time and faster vegetation recovery in terms of biomass accumulation and return to pre-fire species composition. This was probably due to higher species richness and biomass of nitrogen-fixing species found on north-facing slopes in comparison to south-facing slopes. On both north- and south-facing slopes, annuals had the highest species turnover rate, followed by herbaceous perennials and shrubs. In the first four post-fire years, annual species were the largest floristic group, but herbaceous perennials and shrubs were the major contributors to community biomass. Nitrogen-fixing species and exotics contributed significantly to early post-fire community structure. Although the general trends in post-fire succession are clear in terms of temporal changes in the relative proportions of different plant groups, environmental variation and the nature of plant life histories of component species, especially dominant species, could alter such trends significantly.