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.     

Limitations to species coexistence: evidence for competition from field observations,using a patch model

Abstract. Plant community structure in four dune slacks is examined for evidence of competition, expressed as local constraint on species biomass. Such constraint would result in a low variance in total quadrat biomass, compared to a null model. A method of analysis is introduced, using a patchbased null model, which is intended to discount most of the effects of environmental heterogeneity. The method is applied to data in which previous methods had failed to find such structure. There was significant biomass constraint (i.e. low variance in biomass) examining total plant-community biomass in Site 1, and there was a trend in this direction also in Site 2. When guilds defined on morphology were examined, significant biomass constraint was found for the Rhizomatous guild in Site 2. There was a trend to low variance for this guild also in Site 4, but it was significant only in a one-tailed test. After a consideration of other mechanisms, it is concluded that the most likely explanation for this biomass constraint is interspecies competition. Site 2, which showed the strongest evidence of biomass constraint in the present analyses, was also the site that demonstrated a significant deficit of variance in richness in earlier work, and was the most species-rich of the four sites and the least disturbed. The advantages of the new method, for obtaining evidence on competition, are discussed in comparison to pot experiments and field-perturbation experiments. We conclude that community structure can be seen in the pattern of species biomasses provided that a suitable method of analysis is used, i.e. examining small-scale effects and excluding the effects of environmental heterogeneity. In some cases such effects can be related to guild membership. We hypothesise that constraint on biomass is likely to be stronger in more species-rich and less disturbed communities.     

Time and space in the community structure of a species-rich limestone grassland

Abstract. The community structure of a species-rich grassland was investigated at a small spatial scale (0.001 m²), to determine whether evidence suggesting assembly rules could be found in temporal or spatial variation in either species richness or guild proportions. The community was alvar limestone grassland on the island of Öland, Sweden. Three sites were sampled: two lightly grazed, the other recently ungrazed. Plots with and without fertilizer were compared. Evidence was sought for restriction on the ability of species to co-occur within a limited spatial area. Restriction due to a limited number of niches available,‘niche limitation’, could be manifest as lower variance in quadrat richness than expected under a null model (i.e. RV_r, the ratio of observed: expected variance in richness, would be < 1.0). In several cases, RV_r values were significantly < 1.0, even using a patch model to allow for possible spatial variation in the environment. Low RV_r values were found only at the smallest square quadrat size, 10 cm². On Fertilized plots in the years immediately after application of fertilizer, low RV_r could not be demonstrated. Explanations of low RV_r other than niche limitation are considered, such as environmental heterogeneity (present and/or historical) and limitations to the co-occurrence of individual plant modules. Assembly rules based on guild membership were sought by looking at the variance across quadrats in the proportions of species from morphological guilds. An assembly rule would be seen as relatively constant proportion, estimated via RV_gp, the ratio of observed: expected variances in guild proportions. Significant guild proportionality was found in some cases. There was no evidence of guild proportionality in the years after the application of fertilizer. The significant effects in RV_r were more numerous than expected on a random basis, though not observed in every site in every year. Similar trends were seen in RV_gp. At the space/time scales examined, the species in a plant community may be constrained by assembly rules only intermittently, e.g. when resources are more limiting (Wiens 1977). Under this concept, when competition is relaxed, such as following fertilizer application, there is a temporary microhabitat ‘waterhole’ in which more species can coexist, and the assembly rules break down, at least temporarily whilst the species composition adjusts. There was some indication of a return to more deterministic community structure four years after fertilization commenced. Variants of van der Maarel's Carousel model were tested. A Niche-limited Carousel Model (i.e. a model in which there is some limitation in the number of species that can occupy a microsite) would imply restricted variation in richness through time for a single quadrat (temporal RV_r). Overall differences between years in species richness were demonstrated, and their effect removed; after this adjustment there was support for the Niche-limited Carousel Model. The extent of this limitation varied between sites. There were also consistent differences between quadrats in species richness. There was little evidence for constancy of guild proportions through time. The site that showed the strongest community structure in time and space, least year-to-year variation in mean species richness, and least response to fertilizer perturbation, is that on the shallowest soil. Possibly the thin soil results in greater resource limitation, supporting suggestions that assembly rules are stronger when resources are more limited.     

Community Structure (Niche Limitation and Guild Proportionality) in Relation to the Effect of Spatial Scale, in a

A Nothofagus-dominated rainforest in eastern Fiordland, New Zealand, was sampled by shoot frequency in contiguous 1 x 1 m quadrats, along a topologically-circular transect. The data were analysed at five scales up to 5 x 1 m, to search for assembly rules, i.e., generalised restrictions on species co- occurrences. There was no evidence of niche limitation in terms of the whole community, at any scale examined. Rather, variance in species richness was greater than expected from a null model, suggesting environmental heterogeneity. This conclusion was confirmed by using a patch-model. Guild structure was examined in terms of synusiae, based on vertical stratification. The proportion of species in a quadrat that were from the Herb guild was significantly more constant than expected under the null model. However, using a patch-model this effect was reduced, and no longer significant. This suggests pools of species adapted to different micro- environments, but with parallel guild composition. Liane guild proportions were more variable than predicted by the null model; although this effect was not significant, it parallels reports from other South Island forests. The work confirms conclusions from previous work, that plant assembly rules are to be found only at small spatial scales. Processes leading to high and low variance in species richness and in guild proportions are discussed. Selection of a uniform site in the present work, and use of a patch-model, make it less likely that effects are due to environmental or historical effects. The use of guild proportionality as the criterion eliminates the effect of limitation on the number of individuals that can co-occur locally.     

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.     

Generating and testing predictions about community structure: Which theory is relevant and can it be tested with observational data?

Three issues are discussed relevant to the controversy over using null models and observational data on guild structure to test community-level predictions based on limiting similarity theory. First, I argue that most limiting similarity theory is not based on reasonable assumptions for plants and that the theory that is relevant does not generate any predictions about expected guild proportionality on a small spatial scale. Therefore, regardless of adequacy of the statistical methods, the predictions being tested by the body of literature using null models to test for niche limitation are unlikely to be relevant in most plant comunities. Second, assuming that the predictions are after all worth being tested, I argue that most tests using the guild approach do not provide adequate explanations of how the defined guilds could lead to greater competition within vs. between guilds. If this is not true for the particular guilds used, the predictions of guild proportionality or size constancy will not be valid. Third, I address the controversy over whether field experiments can provide more solid evidence than observational data about the role of competition in determining community structure by (1) suggesting methods of dealing with potential drawbacks of field experiments, and (2) suggesting alternative experimental approaches for directly addressing issues about community structure.     

Obituaries

Abstract

It has been suggested that bryophyte communities differ from those of higher plants, in that species coexistence is not limited by inter-specific competition. To test this hypothesis, bryophyte ‘lawns’ were-sampled at six locations in southern New Zealand. At each site, 625 quadrats, each ca 1 X 1cm, were arranged on a contiguous grid.

Variance in quadrat richness was significantly less than expected on a random basis at four sites, with the same trend in a fifth, i.e. quadrat richness was relatively constant. Since a patch model was used to reduce the effects of micro-environmental variation and spatial autocorrelation, this suggests that competition was restricting coexistence. The degree of restriction was similar to that found previously in grass lawn communities.

Species association was calculated on a patch basis, examining only small spatial-scale deviations from expectations. At four- sites this gave only negative associations. At the other two sites there were both positive and negative associations, but the pattern of associations gave no indication of Root/Pianka-type guilds.

Examination of the variance in guild proportions, using taxonomic (moss vs liverwort) and morphological (prostrate vs erect) guilds, gave- no indication of significant guild structure for any site (i.e. in the overall analyses for each site, index RV_gp was very close to the null-model expectation of 1.0).

Searches using the Wilson- Roxburgh method failed to reveal any significant intrinsic guild classification. That is, there was no indication of groups of species within a community that tended to be mutually exclusive because of similarity in resource use. Thus, lack of guild proportionality using a priori guilds was not due to the use of an inappropriate guild classification, but to lack of a guild structure in the communities.

It is concluded that there is community structure among bryophytes, in that species exclude each other to the same degree as higher-plants do in their communities. However, there is no evidence of structuring of these bryophyte communities into guilds; it seems that bryophyte species all form one guild.     

Oribatid mites reveal that competition for resources and trophic structure combine to regulate the assembly of diverse soil animal communities

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Ecological comparisons of robber fly species (Diptera: Asilidae) coexisting in a neotropical forest

Summary Data regarding seasonal abundance, microhabitat preference, and diet were collected over 3 field seasons for adults of 15 robber fly species on Barro Colorado Island, Panama. The species comprised 2 distinct thermal guilds; light-seeking (hereafter LS) species foraged in sunlit areas, while shade-seeking (SS) species foraged only in deep shade. All species were rare during the dry season. During the rainy months, most SS species had flight periods of 2–3 months, and no temporal segregation was apparent. In contrast, most LS species had flight periods of only 4–6 weeks, and a distinct sequence of occurrence was evident during 2 years of censusing. Most SS and LS species displayed a pronounced specificity for perches of a particular substrate type. However, the ranges of perching heights utilized varied considerably among species. Dietary comparisons revealed that mean and maximum prey sizes increased with increasing robber fly size, while minimum prey sizes were constant. Robber fly species <20 mg fed primarily upon nematocerous Diptera, whereas larger species generally fed upon a wide variety of prey types. For each thermal guild, the actual mean overlap for a particular niche dimension was compared to mean overlaps generated by randomly assigning species to thermal guilds. No significant differences from the random null hypothesis were found for the SS guild. However, niche complementarity between dietary and spatial overlaps and dietary overlap was apparent among the 5 large LS species.     

Guild structure of arthropods from Bornean rain forest trees

ABSTRACT. 1. 23275 arthropods collected by insecticide fogging from ten Bornean lowland rain forest trees were sorted to approximately 3000 species and assigned to guilds using two sets of criteria.
2. The rank order for proportions of guilds of species but not individuals is similar in tropical and temperate canopy samples.
3. Misplacement in the guild assignments of a few species can cause important differences in the proportions of certain guilds. This can seriously affect the results of comparisons of different samples, and views on proportional constancy of guilds of species on different trees.
4. Guilds of arthropod species such as chewers, suckers, all phytophages, predators, ants and tourists, occur in constant proportions in samples from different tropical trees and this constancy of proportion is similar in samples from groups of'closely related'and'distantly related'trees.
5. The guild concept of community structure and problems in assigning species to guilds are considered with respect to arthropod samples. The conclusions on constancy of proportion for guilds of species are tempered by remarks on the problems of guild composition.     

Variance deficit is not reliable evidence for niche limitation

Variance deficit in the number of species per sampling unit, as compared to a null model with species occurring independently of each other has been considered by various authors to be evidence for niche limitation. I show that simpler mechanisms could explain this phenomenon. Of those, the competition of individuals for space limits the number of individuals per sampling unit, resulting in decrease of variance in species richness. Testing the proportions of species belonging to various guilds seems to be more ecologically interesting, although here also the variance deficit can hardly be taken as reliable evidence for niche limitation.     

Global patterns of guild composition and functional diversity of spiders

The objectives of this work are: (1) to define spider guilds for all extant families worldwide; (2) test if guilds defined at family level are good surrogates of species guilds; (3) compare the taxonomic and guild composition of spider assemblages from different parts of the world; (4) compare the taxonomic and functional diversity of spider assemblages and; (5) relate functional diversity with habitat structure. Data on foraging strategy, prey range, vertical stratification and circadian activity was collected for 108 families. Spider guilds were defined by hierarchical clustering. We searched for inconsistencies between family guild placement and the known guild of each species. Richness and abundance per guild before and after correcting guild placement were compared, as were the proportions of each guild and family between all possible pairs of sites. Functional diversity per site was calculated based on hierarchical clustering. Eight guilds were discriminated: (1) sensing, (2) sheet, (3) space, and (4) orb web weavers; (5) specialists; (6) ambush, (7) ground, and (8) other hunters. Sixteen percent of the species richness corresponding to 11% of all captured individuals was incorrectly attributed to a guild by family surrogacy; however, the correlation of uncorrected vs. corrected guilds was invariably high. The correlation of guild richness or abundances was generally higher than the correlation of family richness or abundances. Functional diversity was not always higher in the tropics than in temperate regions. Families may potentially serve as ecological surrogates for species. Different families may present similar roles in the ecosystems, with replacement of some taxa by other within the same guild. Spiders in tropical regions seem to have higher redundancy of functional roles and/or finer resource partitioning than in temperate regions. Although species and family diversity were higher in the tropics, functional diversity seems to be also influenced by altitude and habitat structure.     

A comparative analysis of food-niche relationships and trophic guild structure in two assemblages of vertebrate predators differing in species richness: causes,correlations, and consequences

Summary We conducted a comparative analysis of foodniche relationships and trophic guild structure in two assemblages of vertebrate predators (including hawks, owls, snakes, and carnivores) living in similar habitats of Chile and Spain, which differed in species richness (11 and 25 predator species, respectively), to explore how the structure of predator assemblages reflects an increase (or decrease) in the number of coexisting species. Our results indicate that the Spanish assemblage appears enriched by the symmetrical addition of species with disparately large and small values of body weight, diet breadth, and mean prey size, around the same median value found in Chile. Or, alternatively, the Chilean assemblage appears impoverished by the symmetrical loss of peripheral species — those at both tails of the frequency distribution of the above trophic estimators. Spanish predators overlap less extensively among themselves, and with smaller variance, than Chilean predators. Consequently, the higher predator richness in Spain is not ascribable to their narrower diet breadths or more extensive diet overlaps as predicted by some theoretical models. Instead, it is associated with a more spread-out use of the available resources. Trophic guilds can be recognized in the two countries, but in Spain guilds are less tightly packed. Some taxomically unrelated species are close diet analogues between Spain and Chile, whereas some closely related species cannot be matched well between countries. Circumstantial evidence suggests that the presence of some trophic guilds and of some specialist predators in either country is based on the high abundance attained by some particular prey types.     

Guilds and assembly rules in lawn communities

Abstract. The community structure of 11 lawn sites in New Zealand and Fiji was examined in terms of guilds, seeking assembly rules based on guild proportionality. First, associations were analysed, using a new patch model which examined the mean of associations within patches of about 4 cm x 4 cm. As expected from the previously-demonstrated existence of niche limitation at this scale in lawns, the majority of associations between individual species were negative. Even in a lawn only four months old there were significant associations (mostly negative, and one positive). At some sites those species with the most negative associations were those whose morphology might be expected to restrict co-existence with other species. Much ecological theory is based on the assumption that there are limitations to coexistence, related to the resource usage of the species, such that species that are too similar in resource use are less likely to coexist. This theory was tested by looking for evidence of guild proportionality. We defined two guilds, graminoids and forbs. Using these guilds, three of the sites showed significant evenness in proportional representation from the two guilds in quadrats containing four species, i.e. there was less variation in the graminoid:forb ratio than expected on a random basis. Inclusion of species-poor quadrats in a joint analysis over all richness categories overwhelmed this effect, though there was still a strong trend for two of the New Zealand sites: the two species-rich grass lawns. Preliminary analyses of alternative guild delimitations gave no indication that other guild delimitations would have revealed stronger community structure. There was a strong and significant tendency in most sites for the graminoid guild to be more strongly represented in species-poor quadrats, an effect caused by species frequencies.     

Spatial structure of community of darkling beetles (Coleoptera: Tenebrionidae) in the Negev Highlands, Israel

The distribution of niches in resource space of darkling beetles of Negev desert, Israel were studied using discriminant analysis Seventeen environmental variables (substrate structure and vegetation) were measured at 375 plots within 15 grids The first three discriminant axes account for 77% of the variance and reflect a soil structure gradient from sand to clay (the first axis), a gradient of increasing productivity (the second axis), and a type of vegetation architecture (the third axis) Abundance and biomass of tenebrionids increased linearly along the first, the second and the third axes Species richness and diversity had a slanting parabola shape along the substrate axis and increased along the productivity axis There were two distinct spatial guilds of tenebrionid species One guild was composed by species that preferred sandy-gravel soils, and another guild consisted of species that preferred clay soils Niche overlapping of the "sandy-gravel" species was lower than that of the "clay" species Niches of the "sandy-gravel" species were randomly distributed in the resource space Those of the "clay" species formed a tight cluster with most niches placed in the center of the space, representing the centrifugal type of organization The conformity of the guild structure with the different types of organization was supported by compansons between empirical data and stochastic models The border between these two guilds matches with proposed earlier biogeographical boundary (Ramon-Zohar line)     

Niche relations among dung-inhabiting beetles

Summary The whole dung-inhabiting (adult) beetle community living in southern Finland was studied with reference to the seasonal (6 summer months), macrohabitat (open field, half-open pine forest and closed spruce forest) and successional (30 days) gradients. The material comprised 50 coprophagous and 129 carnivorous species, represented by 26,650 and 35,850 individuals, respectively. The most important characteristics of each species are given in an Appendix.In the coprophages the species-abundance relations fitted the lognormal distribution well, but in the carnivores the distribution was strikingly less even. A great number of other differences (see below) apparent between the two trophic groups were explained as the result of a difference in the degree of specialization, apart from the difference in their trophic position: coprophages are microhabitat (dung) and food specialists, while carnivores are microhabitat generalists (and therefore their critical niche space was not completely covered in this study).In the coprophages, two important species guilds were observed: the first occurred at the very geginning of the succession (mean position 2.5 days), and consisted mainly of true dung specialists; the second guild contained species with wider utilization along the successional dimension (mean position 6.5 days), and species of more generalist nature (saprophages). The utilization of carnivores along the successional gradient was more continuous, probably because of their more diversified food resources. The total niche width among the coprophages increased from early successional species to species occurring mainly later on, but was more constant in carnivores. The environmental dimensions proved to be totally independent of each other among the carnivores, while the coprophages showed a clear supplementary relationship along them. Among the carnivores, a spherical niche shape dominated, but among the coprophages elongated and disk-liked shapes were also well represented, especially in specialist species. In both groups, a significant positive correlation existed between niche width and dominance, but not between niche width and abundance. It is suggested that the patchy distribution and transient occurrence of suitable microhabitats contributes to the coexistence of many ecologically similar species.     

Variability in species richness and guild structure in two species-rich grasslands

It has been suggested that variation in the proportion of species in guilds (=guild proportionality) indicates community structuring by guilds in biotic communities. This hypothesis was tested on a subthermophilous grassland and a mesotrophic meadow at a scale of 0.09 m² based on a five-year data set. Further, variation in the total number of species, variation in the number of species belonging to a guild and non-randomness in species composition of guilds were studied. A number of criteria for guild definition were used, such as life form, Grime's C-S-R strategy, phenology, plant height, pollination and dispersal syndromes, leaf shape and anatomy and taxonomy at the family level. The observed variation in the number of guild species corresponded to the null model in which species assemblages with fixed species richness per square were randomly generated from the species pool. The observed variation in the number of guild species was often higher than the variation calculated for randomly distributed species whereas the variation in the proportion of guild species was in some cases lower than the variation calculated for randomly distributed species with fixed frequencies. Possible reasons for the discrepancy in the results based on different models are discussed. It is concluded that there is little evidence of guilds in the organization of grasslands. *** DIRECT SUPPORT *** A02DO006 00012     

Persistence of community structure: what happens when you change taxonomic scale?

It is common in community ecology to use the species as the taxonomic category of interest, yet in rich tropical assemblages containing guilds of very similar species this may not be appropriate. Such assemblages may be organized at the level of guilds rather than at the finer species level. In a ten-year study of assemblages of fish at One Tree Reef, Great Barrier Reef, we found species composition and the number of fish on a given lagoonal patch reef vary greatly across time (Sale and Douglas 1984; Sale et al. in preparation). The mean average proportional similarity of a reef's assemblage to itself at different times (censuses) is usually low at a value of around 0.5. This apparent variability may be ecologically irrelevant noise if organization is at the higher guild level. We have recast our database at the guild level to test this possibility. Thirteen guilds were defined by the diets, foraging habitats and times of the individual species comprising them. Similarity of an assemblage to itself at successive censuses was re-calculated using the number of individuals in each guild instead of the numbers in each species. This analysis yielded significantly higher levels of similarity (P<0.01) among censuses. To test whether this increase in similarity was due solely to the smaller number of categories used to calculate the similarity indices, 5 sets of randomly generated guilds were constructed using a Monte Carlo approach. No significant difference (P>0.05) was found between the average similarity among censuses when assemblages were classified into these null guilds and when they were classified according to the real guilds. These results indicate that shifting to the larger taxonomic scale of guilds does not reveal a significantly more persistent assemblage structure than that revealed when analysis is at the smaller scale of species. There is no evidence of an underlying organization of these assemblages at the guild level.     

Using fish guilds to assess community responses to temperature and flow regimes in unregulated and regulated Canadian rivers

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Secondary succession and breeding bird community structure: Patterns of resource utilization

Summary Structure of breeding bird communities was compared among four habitat types representative of stages present in most old-field successions in the eastern deciduous forest formation of North America. The successional catagories, defined by vegetational structure, were designated herbaceous (type A habitats) herb, shrub and sapling (type B), young forest (type C), and older forest (type D). Density of breeding birds was lowest in A habitats, rose sharply in B habitats and reached a maximum in D habitats. Species richness and number of feeding guild showed similar patterns. Mean number of species per guilds was highest in D habitats. Generalist feeding guilds were predominant in type A and B habitats, primarily due to the importance of the graminivore-insectivore guild. Specialist guilds increased in importance with succession due to addition of several insectivorous guilds in later successional stages. Patterns of occurrence of individual feeding guilds are analyzed and discussed with respect to changes in vegetational structure. Variance in community structure was generally highest in A habitats and lowest in D habitats; this is discussed in relation to the evolution of saturated, coadapted communities.