Trait‐divergence assembly rules have been demonstrated: Limiting similarity lives! A reply to Grime

The recent Forum contribution by Grime (2006) contrasts the MacArthur/Diamond assembly‐rule approach to studying plant communities with the study of environmental trait gradients. Both are valid and useful. In doing so, Grime declares that the assembly rules model, in which negative interactions between plants act with limiting similarity to cause local trait divergence, is “not supported by empirical study of plant communities”. This is, he says, the agony of community ecology. I show that there is now abundant evidence for assembly rules, and no agony.     

甘南高寒草甸土壤线虫营养功能群 的地统计学分析

土壤线虫是土壤生态系统的重要指示生物。研究高寒草甸土壤线虫营养功能群空间分布格局特征,有助于揭示土壤线虫分布对高寒草甸生态因子变化的响应。2019年7月中旬,采用网格法（1.0 m × 0.5 m）研究5.0 m × 4.0 m尺度土壤层（0 ~ 20 cm）土壤线虫营养功能群空间特征。采用改良湿漏斗法,共获得土壤线虫1 830条,隶属于49属,土壤线虫个体平均密度以50 g干土计量为63.2条,其中,绕线属（Plectus）和拟丽突属（Acrobeloides）为优势类群。土壤线虫以食细菌线虫、植物寄生线虫为主。土壤线虫营养功能群变异系数介于64.26% ~ 107.69%（n = 38）之间,存在较强的变异性。食真菌线虫与食细菌线虫（P < 0.001）、食真菌线虫与植物寄生线虫（P < 0.001）间空间分布均存在显著相关性。地统计分析显示,土壤线虫营养功能群的空间相关范围存在明显差异,有效变程介于0.50 ~ 27.07 m之间,其67.26% ~ 99.79%的变异与空间自相关过程有关。食细菌土壤线虫分布格局呈较均匀的斑块镶嵌结构,斑块较小;非食细菌土壤线虫呈斑块连接或斑块镶嵌结构,斑块较大且斑块间过渡明显,表明土壤线虫小尺度明显的斑块结构可能是高寒草甸受干扰后的一种常见分布模式,并受多种因素影响。     

Disentangling patch and landscape constraints of nested assemblages

The role of the matrix for the functionality of residual patches and therefore for species persistence has been recently recognised. However its influence on assemblage structure and composition is still unclear. We studied the effect of both patch and matrix attributes on community assemblage structure and composition in a fragmented landscape of central Italy. We used bird assemblages as a study model. Birds were sampled in 24 wood patches and distribution patterns were analysed considering both assemblage and guild-level responses. Patch composition and structure together with landscape patterns quantified with GIS were used as explanatory variables. The assemblages appeared to be strongly influenced by the landscape context, both in terms of composition and configuration. The functional connectivity of the patches, in terms of amount of neighbouring borders between woods and hedgerows, the shape of the woods in the matrix, together with the age of the patches, expressed as tree diameter, acted as the essential determinants, and even if the assemblages presented a significant nested structure, seven out of 40 species showed an idiosyncratic distribution. The study of the association between functional guilds and environmental parameters highlighted the influence of ecological constraints on the occurrence of unexpected presences and absences, suggesting the ability of some species to perceive and exploit matrix habitats. The matrix holds the capability to at least mitigate the effects of isolation and habitat loss, and the semi-natural elements of the mosaic may represent a biodiversity reservoir in an otherwise deprived landscape.     

A null model of guild proportionality,applied to stratification of a New Zealand temperate rain forest

Summary Much ecological theory assumes that the number of species that can coexist (by species packing) is limited, because competitive exclusion occurs when any pair of species within a guild is too similar — species saturation or niche limitation. If such niche limitation occurs, the proportion of species in each guild should be relatively constant — guild proportionality. This concept is applied to the guilds represented by strata in a forest. A method is produced, and used to examine a New Zealand temperate rain forest. Most strata showed no deviation from a null model of no niche limitation, i.e. no tendency to guild proportionality. The proportion of lianes was more variable than in the null model, tending to be inversely related to the proportion of epiphytes, Canopy tree proportion was significantly more constant than in the null model, but this could be interpreted as a limit caused by the size of a canopy tree individual.     

Evidence for constraint on species coexistence in vegetation of the Park Grass experiment

Repeated patterns, of a type that would be expected to result from limitations to species coexistence (i.e. assembly rules) were sought in the Park Grass experiment. This classical grassland experiment was sampled in two years, using replicated biomass samples. Variance in a number of measures was examined, and compared to the variance expected under appropriate null models, the latter based on assumptions of no interactions between species. In each case, an assembly rule would result in low variance. Examining variance in species richness between quadrats within a treatment, there was no indication of constraint on species co-occurrences; variance in richness was actually greater than expected under the null model, attributable to environmental variation or perhaps positive interactions between species. However, there was control on biomass, evidenced by variance in total biomass (i.e. over all species) within a treatment being significantly lower than expected under the null model. There was no indication of community structure based on guilds (i.e. functional types). Although there was in 1991 some, non-significant, indication of a constant proportion of species from the legume guild, there was no sign of such an effect in 1992. Searches for intrinsic guilds failed to converge. There was no indication at all of constancy in the proportional representation of guilds by biomass. Thus, there is good evidence for competitive control on plant growth, but none for control of species occurrences. There is no convincing evidence for guild structure in this community at the scale sampled. Possible conflict is discussed between the existence of evidence for temporal stability but the absence of evidence for spatial uniformity. It is concluded that most of the mechanisms proposed for temporal stability will not necessarily lead to control on spatial variation. For many mechanisms, this would depend on the spatial scale examined.     

The influence of habitat structure on arthropod diversity in Argentine semi-arid Chaco forest

Abstract. The Argentine Chaco is a mosaic of grassland and open forest habitats maintained by natural disturbance activities such as fire. Since the introduction of domestic livestock and other human activities, the balance of this mosaic has been significantly altered, both in plant species and structural composition. This study focuses on the impact of such changes on the diversity of ground-dwelling arthropods within semi-arid Chaco forest. Quantitative measures of habitat structure and arthropod diversity were taken in forest areas previously subjected to grazing, logging and ploughing. Results indicated that arthropod diversity was smaller on sites with reduced structural complexity, with marked changes in arthropod family composition. The habitat components relating to plant architectural and vertical diversity were particularly influential on arthropod diversity. The guild size ratio of predatory to non-predatory arthropods also differed significantly between habitats suggesting a change in the resource base available to some groups. The latter suggests a shift in the functional organisation of the forest ecosystem which could have important repercussions for the diversity of other trophic levels.     

Prairie plant guilds: a multivariate analysis of prairie species based on ecological and morphological traits

An ecomorphological analysis of the tallgrass prairie of central North America divided representative species of the native grassland flora into eight guilds or groups of species with similar life-form, phenology, and ecology. The guilds, segregated by multivariate analysis, are: (1) warm-season graminoids with Kranz anatomy and the Hatch-Slack photosynthetic pathway (C4 grasses); (2) cool-season graminoids without Kranz anatomy, but with the common Calvin or C3 photosynthetic pathway (C3 grasses and sedges); (3) annuals and biennial forbs; (4) ephemeral spring forbs; (5) spring forbs; (6) summer/fall forbs; (7) legumes; and (8) woody shrubs. The study was based on 158 plant species indigenous to three upland prairie sites in northeastern Kansas. Each species was scored for 32 traits which fall into five broad categories: plant habit, leaf characteristics, stem structures, root structures, and reproductive traits, including phenology. A multivariate, detrended correspondence analysis sorted the 158 species into the eight principal groups or guilds. These groups were further supported by a cluster analysis and discriminant function analysis of the same data set. The discriminant function analysis determined that 94.3% of the species were correctly classified in their respective guilds, and that the guilds were statistically different. Results indicate that guild analysis offers a basis for detailed classification of grassland vegetation that is more ecologically focused than species composition, as the myriad of species (about 1,000 prairie species on the central plains of North America) vary in presence, cover, and importance with their individualistic distribution.Abbreviations C3= C3 photosynthesis - C4= C4 photosynthesis - LSD= least significant difference     

Coexistence in a guild of filter feeding caddis larvae: do different instars act as different species?

Summary Guild structure and coexistence of filter feeding caddis larvae was studied in three lake-outlet streams in northern Finland. Natural stones were used as sampling units, and for each stone measurements of six microhabitat variables were taken. Species and species instar relationships in the multivariate microhabitat space were analyzed by a method based on Principal Components Analysis. Seven species of filtering caddis larvae were found at the study sites, but detailed analysis was restricted to the four most abundant ones (Polycentropus flavomaculatus, Hydropsyche angustipennis, H. pellucidula, H. saxonica). P. flavomaculatus was the microhabitat generalist of the guild, whereas all the hydropsychids were narrowly specialized in at least one larval instar. In all species, different instars utilized the microhabitat somewhat differentially. Moss content and Froude number were significantly different between instar microhabitats in all comparisons. Ontogenetic niche changes not only separated species instars from each other, but also profoundly changed the biotic as well as abiotic environment of the species at different phases of larval development. This may have important effects on the dynamics of interacting species and, consequently, on the guild structure. While some of the more classical models of coexistence may also be relevant for understanding coexistence in this guild of sessile stream insects, ontogenetic niche changes clearly need to be taken into consideration in future studies of guild structure and species interactions of filter feeding caddisfly larvae.     

Structural analysis of the phytophagous insect guilds associated with the roots of Centaurea maculosa Lam. C. diffusa Lam., and C. vallesiaca Jordan in Europe:

Summary During extensive field surveys in central and eastern Europe, 21 herbivorous root insect species were found on Centaurea maculosa ssp. rhenana Boreau, 12 species on C. diffusa Lam. and 11 species on C. vallesiaca Jordan, representing 12 families in 4 orders. The large geographic distribution (species-area function), the high number of Centaurea spp. present (host speciation rate), and the high apparency of the rosettes and the rich food resources offered by the roots during winter, together with their poor accessibility, correlate with the high number of specialist feeders associated with the roots of C. maculosa and C. diffusa. The members of the taxonomically diverse root entomofauna exploit specific structures of the tap root (food niches). Interspecific competition among members of food niches, as well as species-specific responses to different phenological stages (for oviposition) and tissues (for larval development) are thought to be responsible for the high predictability in guild structure. The relatively low levels of host plant attack (two thirds of the roots were unattacked) and the fact that food niches remained unoccupied in most of the regions suggest, however, that the majority of the studied guilds do not represent equilibrium assemblages. Ecological (different habitats), climatic (transitional zone) and historical (ancient pre-Pleistocene communities) factors could account for the highest values of species diversity, infestation levels, species packing and food niche utilization, which are found on C. maculosa in E. Austria/NW. Hungary, compared to other regions. A positive correlation between species packing (number of root-feeding species per population) and infestation rates (percent of roots attacked) was only found for the more stable, semi-natural habitats. A comparative analysis of the regional root insect guilds of C. maculosa with corresponding data for the phytophagous insects associated with the flower heads revealed distinct taxonomical differences, but a high degree of numerical and structural similarity. The different geographical regions are similarly ranked for host plant attack, herbivore pressure, average species packing and level of food niche utilization.     

Plant functional types – a strategic perspective

Abstract. This paper discusses the use of Plant Functional Types to describe vegetation types – from plant communities to formations – in order to understand the ecological constraints of plants and plant communities on a higher level of abstraction. The terminology around plant functional types, traits and strategies is in need of further clarification and unification. The use of the term plant functional type is recommended. We are in need of a total environmental framework in which, hierarchically or reticulately, plant functional types can be compared and interpreted.