Plant communities and landscape diversity along a Canadian Arctic river

Abstract. We analysed the structure and diversity of the vegetation along an Arctic river to determine the relationship between species richness and plant community structure. We examined whether variation in species richness along the corridor is structured as (1) an increase in the number of communities due to increasing landscape heterogeneity, (2) an increase in the floristic distinctiveness (β-diversity) of communities, or (3) an increase in within-community richness (α-diversity) as species-poor communities are replaced by species-rich communities. We described 24 community types and analysed the relationship between site vascular species richness (γ-diversity) and β-diversity, α-diversity, site environmental heterogeneity, and the number of distinct plant communities. We also measured diversity patterns of vascular, bryophyte, and lichen species within communities and examined their relationship to community-level estimates of environmental factors. We found that an increase in site species richness correlated with an increase in the number of communities (r²= 0.323, P= 0.0173) and β-diversity (r²= 0.388, P= 0.0075), rather than an increase in the α-diversity of individual communities. Moisture and pH controlled most of the differences in composition between communities. Measures of species richness and correlations with moisture and pH within communities differed among vascular, bryophyte, and lichen species. Bryophyte richness was positively correlated with moisture (r²= 0.862, P= 0.0010) and lichen richness was negatively correlated with moisture (r²= 0.809, P= 0.0031). Vascular plants had a peak in richness at pH 6.5 (r²= 0.214, P < 0.0001). We conclude that site variation in vascular richness in this region is controlled by landscape heterogeneity, and structured as variation in the number and distinctiveness of recognizable plant communities.     

Bryophyte colonisation in experimental microcosms: the role of nutrients, defoliation and vascular vegetation

A three-year multi-factorial microcosm experiment simulating fertilisation, defoliation and the composition of vascular vegetation in a dry grassland succession was used to test four hypotheses concerning the establishment and survival of bryophytes in grassland vegetation. H₁: bryophyte cover may be used to predict bryophyte species richness. H₂: bryophyte richness is suppressed at high nutrient levels and promoted by defoliation of vascular plants. H₃: species richness of bryophytes is influenced by the species composition of the vascular vegetation. H₄: bryophyte species richness is negatively correlated with vascular plant biomass.
The relationship between bryophyte richness and bryophyte cover was found to follow the classical species-area richness curve. Bryophyte species richness responded positively to defoliation and negatively to fertilisation. The species composition of vascular vegetation had no significant effect on bryophyte richness. Bryophyte species richness was lower at high vascular plant biomass and vascular plant dry weight above 400 g m⁻² appeared fatal to bryophytes. At high nutrient levels, defoliation increased bryophyte richness, but defoliation did not fully compensate for the negative effect of fertilisation. The study reinforces the concern for short lived shuttle bryophytes in the agricultural landscape.     

Bryophyte and lichen diversity: A comparative study

Abstract We describe the regional species richness, variation in species richness and species turnover of bryophytes and lichens from 36 sites in lowland forests of southeastern Australia. The analyses subdivided the two major taxa into their constituent sub-groups: mosses, liverworts, and crustose, fruticose and foliose lichens. They also explored correlations between selected environmental variables and patterns of diversity. On a regional scale, there were 77 species of bryophytes and 69 species of lichens, giving a total of approximately one-third of the total number of vascular plant species in the region. Mean species richness was higher for lichens than bryophytes. Also, the two taxa were negatively correlated because lichens favoured dry sites and bryophytes favoured moist ones. Species turnover was greater for bryophytes than lichens, largely due to the distribution of liverwort species. Foliose lichens showed higher levels of turnover than crustose lichens. Multiple regression and canonical correspondence analysis showed that both taxa and all sub-groups responded to the same three variables: vascular plant cover, time since last fire and topographic position. Other variables, including time since logging and intensity of logging, explained little variation in bryophyte or lichen diversity. The data suggest that the strategies for the conservation of bryophyte and lichen biodiversity will be different, to reflect the different patterns of species richness and species turnover.     

Species richness of vascular plants,bryophytes, and lichens along an altitudinal gradient in western Norway

Species richness patterns of ground-dwelling vascular plants, bryophytes, and lichens were compared along an altitudinal gradient (310–1135 m a.s.l.), in western Norway. Total species richness peaked at intermediate altitudes, vascular plant species richness peaked immediately above the forest limit (at 600–700 m a.s.l.), bryophyte species richness had no statistically significant trend, whereas lichen richness increased from the lowest point and up to the forest limit, with no trend above. It is proposed that the pattern in vascular plant species richness is enhanced by an ecotone effect. Bryophyte species richness responds to local scale factors whereas the lichen species richness may be responding to the shading from the forest trees.     

Multi-scale comparisons of cliff vegetation in Colorado

We developed a nested vegetation sampling protocol to sample the plant diversity on south-facing cliffs and cliff bases in Jefferson County, Colorado. The multi-scale plots included three nested spatial scales, 1 m², 20 m², and 40 m². We compared plant species richness and species diversity among large cliffs, medium cliffs, small cliffs, and non-cliff sites using Hill's diversity numbers (N ₀, N ₁, and N ₂) for the 1-m² quadrats. Species richness (N ₀) was calculated for the 20-m² and 40-m² plots. Our results indicate that plant species diversity on the cliff faces did not increase with increasing cliff area. This pattern was consistent at all three sampling scales. A model selection was run to determine if plant species diversity values on the cliff faces were associated with cliff variables. None of the cliff variables measured were good predictors of diversity at the 1-m² scale. However, at the 20-m² scale, canyon differences and a positive relationship with increasing cliff surface roughness explained 70% of the variability in species richness. Although most plant species sampled on the cliff faces were also found in the base plots, 13 species were sampled only on the cliff faces. Additionally, dry south facing cliffs support a mix of xeric and mesic plants indicating that cliffs may provide unique microenvironments for plant establishment. This revised version was published online in August 2006 with corrections to the Cover Date.     

Species richness,alien species and plant traits in Central Argentine mountain grasslands

Abstract. Pattern of native vegetation, distribution of alien species and variation of environmental parameters were studied in mountain grasslands in a lithologically homogeneous Córdoba mountain range in Central Argentina. CCA showed that altitude was the most important factor determining the compositional variation of the vegetation, with soil nutrient status and stoniness as additional factors. Short‐grass communities, associated with the driest habitats on plateaus, showed higher small‐scale native species richness than wet‐turf communities in valleys and tall‐grass communities on slopes. Species richness was negatively correlated with soil parameters that indicate nutrient status and water availability. Also, there was a negative correlation between soil Ca‐ and Mg‐ content and richness. High native species richness coincided with high alien species richness. When smaller units – community types – were considered, it became evident that within short‐grass vegetation, the three most species‐rich community types contained significant numbers of alien species, while the other two did not. Even within one community type, the same quadrats that contained the highest number of native species, were also characterized by the highest numbers of alien species. Evidently, the same mechanism was responsible for high richness of both native and alien species. Alien species were distinguished by a greater proportion of annuals and prostrate stoloniferous plants, by lower palatability and by smaller proportion of zoochory. DCA ordination of quadrats on the basis of plant traits as attributes resulted in a clear distinction of three main vegetation types. Short‐grass vegetation was distinguished by a predomination of late flowering species, tall‐grass vegetation by the presence of high herbaceous plants and bushes, and wet‐turf vegetation by the presence of plants with storage organs, the lack of hairy leaves, and by a high proportion of cryptophytes. Quadrats with and without alien species were distinguished as well, indicating that the occurrence of aliens may be dependent on plant traits in a particular patch of a community.     

Bryophyte and Lichen Diversity Under Simulated Environmental Change Compared with Observed Variation in Unmanipulated Alpine Tundra

Effects of simulated environmental change on bryophyte and lichen species richness and diversity in alpine tundra were investigated in a 5-year experiment at Latnjajaure, northern Sweden. The experiment had a factorial design including fertilisation and temperature enhancement in one meadow and one heath plant community. Responses in species richness, biodiversity, and species composition of bryophytes and lichens to experimental treatments were compared to the observed variation in six naturally occurring plant communities. The combination of fertilisation and enhanced temperature resulted in a species impoverishment, for bryophytes in the bryophyte-dominated community, and for lichens in the lichen-dominated communities, but the species composition stayed within the observed natural variation. During the course of the study, no species new to the investigated mid-alpine landscape were recorded, but that scenario is realistic within a decade when comparing with the processes seen in vascular plants.     

Species richness of vascular plants, bryophytes and lichens in dry grasslands: The effects of environment, landscape structure and competition

We studied the relative importance of local habitat conditions and landscape structure for species richness of vascular plants, bryophytes and lichens in dry grasslands on the Baltic island of Öland (Sweden). In addition, we tested whether relationships between species richness and vegetation cover indicate that competition within and between the studied taxonomic groups is important. We recorded species numbers of vascular plants, bryophytes and lichens in 4 m² plots (n=452), distributed over dry grassland patches differing in size and degree of isolation. Structural and environmental data were collected for each plot. We tested effects of local environmental conditions, landscape structure and vegetation cover on species richness using generalized linear mixed models. Different environmental variables explained species richness of vascular plants, bryophytes and lichens. Environmental effects, particularly soil pH, were more important than landscape structure. Interaction effects of soil pH with other environmental variables were significant in vascular plants. Plot heterogeneity enhanced species richness. Size and degree of isolation of dry grassland patches significantly affected bryophyte and lichen species richness, but not that of vascular plants. We observed negative relationships between bryophyte and lichen species richness and the cover of vascular plants. To conclude, effects of single environmental variables on species richness depend both on the taxonomic group and on the combination of environmental factors on a whole. Dispersal limitation in bryophytes and lichens confined to dry grasslands may be more common than is often assumed. Our study further suggests that competition between vascular plants and cryptogams is rather asymmetric.     

Impact of Land-Use Intensity and Productivity on Bryophyte Diversity in Agricultural Grasslands

While bryophytes greatly contribute to plant diversity of semi-natural grasslands, little is known about the relationships between land-use intensity, productivity, and bryophyte diversity in these habitats. We recorded vascular plant and bryophyte vegetation in 85 agricultural used grasslands in two regions in northern and central Germany and gathered information on land-use intensity. To assess grassland productivity, we harvested aboveground vascular plant biomass and analyzed nutrient concentrations of N, P, K, Ca and Mg. Further we calculated mean Ellenberg indicator values of vascular plant vegetation. We tested for effects of land-use intensity and productivity on total bryophyte species richness and on the species richness of acrocarpous (small & erect) and pleurocarpous (creeping, including liverworts) growth forms separately. Bryophyte species were found in almost all studied grasslands, but species richness differed considerably between study regions in northern Germany (2.8 species per 16 m²) and central Germany (6.4 species per 16 m²) due environmental differences as well as land-use history. Increased fertilizer application, coinciding with high mowing frequency, reduced bryophyte species richness significantly. Accordingly, productivity estimates such as plant biomass and nitrogen concentration were strongly negatively related to bryophyte species richness, although productivity decreased only pleurocarpous species. Ellenberg indicator values for nutrients proved to be useful indicators of species richness and productivity. In conclusion, bryophyte composition was strongly dependent on productivity, with smaller bryophytes that were likely negatively affected by greater competition for light. Intensive land-use, however, can also indirectly decrease bryophyte species richness by promoting grassland productivity. Thus, increasing productivity is likely to cause a loss of bryophyte species and a decrease in species diversity.     

Goose grazing influences the fine-scale structure of a bryophyte community in arctic wetlands

Herbivores can shape plant communities, especially in the Arctic. We tested the role of geese for structuring bryophyte communities at fine spatial scales in the arctic tundra by excluding them from 4 × 4 m areas. We surveyed the presence and absence of bryophyte species in quadrats (10 × 10 cm) divided into 25 cells outside and inside these exclosures, after 5 and 11 years of treatment. Species richness per cell (4 cm²) was higher in the presence of geese, especially after 11 years of treatment, while geese had little effect on richness at larger scales (i.e. quadrat and whole exclosure). The slope of the species–area relationship within quadrats was consequently shallower outside exclosures. Our results further suggest that the community outside the exclosures was more variable in space and time than that inside the exclosures. We conclude that goose foraging activity promotes the coexistence of bryophyte species at the centimetre scale.     

Species richness in boreal swamp forests of SE Norway: The role of surface microtopography

Question: What is the relative importance of environmental gradients and surface microtopography (variation in vertical level within sampling units) for fine‐scale plant species richness in Picea abies swamp forests? Location: 11 swamp forests in SE Norway. Methods: We recorded species richness (number of species of vascular plants, mosses, Sphagnum and hepatics), depth to water table, soil base status and vertical range (microtopographic relief) in 2400 microplots, (each 1/16 m²), in 150 1‐m² plots in the 11 swamp forests. Relationships between species richness and environmental predictors were modelled by GLMM. Results: Moss and hepatic species richness increased with increasing microtopographic relief, most strongly for wet acid sites, in which similar trends were also found for Sphagnum. Relief was a poor predictor of vascular plant species richness. Conclusions: Microtopographic relief is a good predictor of local species richness in Picea abies swamp forests, partly because larger vertical variability means higher within‐plot habitat diversity with respect to the wet‐dry gradient, and partly because qualitatively new microhabitats associated with steep slopes are added in drier sites. The relationship between species richness and microtopographic relief is context dependent, differing in complex ways among species groups and among sites with different environmental conditions.     

The natural exclusion of red deer from large boulder grazing refugia and the consequences for saxicolous bryophyte and lichen ecology

Large boulder grazing refugia permitted comparison of saxicolous bryophyte and lichen assemblages with those boulder tops accessible to red deer (Cervus elaphus) on a sporting estate in northwest Scotland. Plant succession was predicted to occur unchecked by grazing on the tops of these large boulders with cascading effects on bryophytes and lichens—assuming boulders had been in place over the same time period. Fifty pairs of boulders (one ≥2 m and the other accessible to red deer) were selected at random from various locations below north-facing crags. Percentage cover of each bryophyte and lichen species was estimated from three randomly placed quadrats on each boulder top. Due consideration was given to the influence of island biogeography theory in subsequent model simplification. Mean shrub cover and height, leaf-litter, bryophyte cover and bryophyte species richness were significantly higher within quadrats on large boulder tops that naturally excluded red deer. Lichen cover and lichen species richness were significantly higher on boulder tops accessible to red deer. Lichen cover was in a significant negative relationship with bryophyte cover, shrub cover and litter cover. Bryophyte cover showed a significant positive relationship with shrub height but there was an optimum shrub cover. Natural exclusion of red deer from the tops of large boulders has facilitated plant succession. The results suggest that grazing arrests the lithosere on boulder tops accessible to red deer at an early plagioclimax favouring saxicolous lichens. The results are relevant to situations where red deer might be excluded from boulder fields that hold lichen assemblages of conservation value.     

Microtopographic heterogeneity constrains alpine plant diversity, Glacier National Park, MT

Theoretical and empirical evidence exists for a positive relationship between environmental heterogeneity and species diversity. Alpine plant communities can exhibit exceptional diversity at a fine scale, which niche theory would suggest is the result of fine scale spatial heterogeneity of the environment. To test if species diversity of alpine plants is driven by environmental heterogeneity, we sampled vascular plant species composition, microtopography, and ground cover within 1?m² plots with and without solifluction forms in Glacier National Park, MT. We analyzed the relationship between microtopographic heterogeneity and species richness at the plot and sub-plot scale with linear and quantile regression, respectively. Species richness does not differ between the plots varying in cover type. Species richness is negatively related to the fractal dimension (D) of the ground surface and non-vegetated ground cover within 1?m² plots. At a finer scale, the standard deviation of elevation and slope appear to impose a limit on species richness such that more variable sub-plots have lower species richness. Contrary to our expectations, microtopographic heterogeneity does not promote the diversity of alpine plants. The negative relationship between topographic heterogeneity and species richness is contrary to the theoretical prediction that environmental heterogeneity generally results in greater species diversity. It is possible that microtopographic variability represents a measure of soil disturbance, which would be expected to have a negative effect on species diversity in alpine tundra due to its low productivity.     

Environmental relations of the bryophytic and vascular components of a talus slope plant community

The environmental factors correlating with community structure of vegetation on talus slopes of the 785 km long Niagara Escarpment, southern Ontario, Canada, were studied using canonical correspondence and regression analysis. The bryophytes and higher vascular plants were analysed separately to see if their responses were similar or different. Both vascular plants and bryophytes responded similarly to the environmental variables that were measured. For both vegetation components, location from north to south explained most of the variance. When species richness was plotted against location for the complete vegetation and for the two components separately, the results showed that vascular plant species richness decreased with increasing latitude, while bryophyte richness increased. The magnitude of both of these trends was slight but consistent with the hypothesis that available environmental energy governs a significant amount of the variance in species richness. Since separate components of the talus vegetation were shown to respond differently to the same environmental variable, groups of taxa should not be excluded from community level studies without a consideration of the possible consequences of this bias.     

Vegetation at the Limits for Vegetation: Vascular Plants, Bryophytes and Lichens in a Geothermal Field

The effects of the chemical and physical factors associated with geothermal activity on plant community structure and composition were investigated in one of the largest geothermal fields of central Italy. The study site was located in the geothermal area of Sasso Pisano – Monte Rotondo Marittimo, Southern Tuscany. The percentage cover of all vascular plant, bryophyte and lichen species was estimated within 119 circular plots of 0.25 m². For each plot the soil pH, soil temperature, slope, aspect, incident radiation, soil nitrogen and carbon contents were also quantified. Two vascular plants, Calluna vulgaris and Agrostis castellana, were found to be the most widespread species tolerating the harshest conditions in terms of low soil pH and high soil temperature. The most widespread cryptogam species was Hypnum cupressiforme. Spatially autoregressive models showed that a proportion of about 41–51% of the variance in species richness of one group of plants (vascular or cryptogamic plants) could be modelled by using three or four uncorrelated environmental factors respectively (soil temperature, soil nitrogen and soil C/N ratio and these three plus incident radiation). For the total number of species (vascular and cryptogamic plants), the variance explained by the same three uncorrelated variables was about 57%. This study evidenced a strong environmental control of community composition and species richness, in a site subjected to extreme soil values of soil pH and temperature. The dominance of vascular over cryptogamic vegetation in this geothermal site can be explained by the combined effects of geothermal stress (low soil pH and high soil temperature) with the summer drought typical of the Mediterranean climate.     

Bryophyte diversity and distribution along an altitudinal gradient on a lava flow in La Réunion

Non‐vascular plant distribution patterns were examined in three microhabitats along an altitudinal gradient on a recent lava flow of the Piton de la Fournaise volcano (La Réunion, Mascarene archipelago). The uniform nature of the lava flow provides an excellent system to study the relationship between altitude and species diversity and distribution, and at the same time avoiding confusing multiple effects of substrate and vegetation heterogeneity. Non‐vascular plants were surveyed with quadrats within an altitudinal range from 250 m to 850 m a.s.l. Fine‐scale variations in bryophyte communities between three ecological microhabitats (the ground and on the rachises of two fern species) were investigated. Three specific questions were addressed: (1) What is the species diversity of bryophyte communities on a 19‐year‐old lava flow? (2) How does altitude influence the diversity and distribution of bryophytes on a lava flow? (3) Does microhabitat variation control bryophyte diversity? In our study, bryophyte diversity increased with altitude. Unexpectedly, species richness was very high; 70 species of bryophytes were recorded including nine new records for the island. Diversity was also controlled by ecological microhabitats. Bryophyte species were structured into six categories according to altitude and microhabitat preferences. Results suggested that the high diversity of these cryptic organisms on this lava flow is fostered in part by their host substrate and their adaptative strategies on new substrates. On a broader scale, it was concluded that lava flows as primary mineral environments are important to conserve, as they support a high diversity of pioneer organisms that constitute the early stages of the development of La Réunion's remnant lowland rainforest.     

Species richness, abundance, rarity and environmental gradients in coastal barren vegetation

Coastal barrens in Nova Scotia are heathlands characterised by short, predominantly ericaceous vegetation, sparse tree cover, exposed bedrock, pockets of Sphagnum bog, and stressful climatic conditions. Although coastal barrens are prominent in the physical and cultural landscape, they are largely unprotected. We selected six barrens along the Atlantic coast, and surveyed 20 1-m² plots at each barren for vascular plants, macrolichens, mosses and environmental factors. We recorded 173 species (105 vascular, 41 macrolichen, 27 moss), including six provincially rare vascular species found predominantly in nearshore areas with high levels of substrate salt and nutrients, variable substrate depth, and short vegetation. Although vascular plant and moss richness were similarly correlated with vegetation height, substrate depth, organic matter content, and rock exposure, there were no clear correlations between vascular plant, macrolichen and moss richness across all sites. Vascular plant rarity and species richness were not correlated, but had inverse relationships with key environmental gradients. Tailoring conservation efforts to protect areas of high richness may thus mean that rare species are missed, and vice versa. Ordination and ANOSIM show that barrens vegetation differs widely among sites; therefore, protecting any singular coastal barren will not protect the entire range of vegetation communities and species in this heathland type. Conservation planning should emphasize protecting environmental gradients correlated with richness, rarity and plant community structure, including substrate depth and moisture, and vegetation height. Additionally, protected areas should include a coastal-inland gradient and a diversity of substrate types, including exposed rock and trees.     

Bryophyte and vascular plant species richness in boreo-nemoral moist forests and mires

We compare species richness of bryophytes and vascular plants in Estonian moist forests and mires. The material was collected from two wetland nature reserves. Bryophyte and vascular plant species were recorded in 338 homogeneous stands of approximately 1 ha in nine forest and two mire types. Regional species pools for bryophytes and vascular plants were significantly correlated. The correlations between the species richnesses of bryophytes and vascular plants per stand were positive in all community types. The relative richnesses (local richness divided by the regional species pool size) were similar for bryophyte species and for vascular plant species. This shows that on larger scales, conservation of the communities rich in species of one taxonomic plant group, maintains also the species richness of the other. The minimum number of stands needed for the maintenance of the regional species pool of typical species for the every community type was calculated using the species richness accumulation curves. Less stands are needed to maintain the bryophyte species pools (300–5300 for bryophytes and 400–35 000 for vascular plants).     

Ecological Design is More Important Than Compensatory Mitigation for Successful Restoration of Alpine Spoil Heaps

Spoil heaps of surplus rock from hydropower tunnel construction negatively impact alpine landscapes unless restored. Such spoil heaps have been created for more than 100 years, but we still lack knowledge about the relative importance of compensatory mitigation (seeding and fertilization), spoil‐heap construction method, local environmental factors, and regional climatic factors for restoration success. We studied the species composition of 19 alpine spoil heaps in Western Norway and their undisturbed surroundings using ordination and statistical modeling. Substrate grain size was the principal factor explaining differences in species composition between spoil heaps and their surroundings. Soil characteristics, that is, organic matter content and pH, and reutilization of topsoil were also important. Seeding and fertilization had negligible effects on restoration success. Slow recovery was observed for total vegetation cover and species richness of vascular plants and lichens while bryophyte cover recovered rapidly. Lower bryophyte cover and bryophyte and vascular plant species richness on older than on younger spoil heaps indicated recent changes in spoil‐heap construction practices that favor plant colonization. Our results indicate that spoil‐heap design is more important for restoration success than compensatory mitigation. We therefore suggest spoil heaps designed with a fine‐grained top substrate preferably from stockpiled local topsoil, with uneven surface topography that mimics natural topographic variation, and recommend discontinuation of seeding and fertilization.     

Associations Between Weevils (Coleoptera: Curculionidea) and Plants, and Conservation Values in Two Tussock Grasslands, Otago, New Zealand

Ecosystem level processes and species interactions have become important concepts in conservation and land management. Despite being New Zealand’s greatest contributors to global diversity, native invertebrates have been largely overlooked in the assessment of land values, and their diversity has often been assumed to reflect native plant diversity without justification. Invertebrates can in fact affect plant species composition, and in ecosystems such as New Zealand’s remaining indigenous and semi-modified tussock grasslands can do so in excess of more conspicuous vertebrate grazers. An understanding of the interactions between invertebrates and their plant hosts may be informative in assessing land conservation values, increase the efficiency of rapid inventory analyses and would be applicable across the production-conservation spectrum. This study considers the Curculionoidea, vascular plant, bryophyte and lichen communities of two semi-modified tussock grasslands in the Otago region of southern New Zealand. Quantitative plant and invertebrate sampling were carried out in January 2001. Data were analysed using ANalysis Of SIMilarity (ANOSIM) and Multi-Dimensional Scaling (MDS) ordination techniques. Vascular plant, bryophyte and lichen species richness was highest in the same site and plots as native weevil species richness, however the proportion of native vegetation in these locations was lower. Associations identified between Curculionoidea and vascular plants were dismissed due to the confounding effect of species frequency across samples. This appeared to have little influence on associations with bryophytes and/or lichens and these were given more weighting. The ecological implications of associations are considered and variability in weevil composition is discussed in reference to the tussock grassland environment. The importance of plant–invertebrate relationships to conservation and the uses and limitations of the PRIMER MDS ordination technique for determining associations are discussed.