Diversity, distribution and larval habitats of North Swedish blackflies (Diptera: Simuliidae)

1. Blackfly species richness and community structure were analysed at fifty-six sites in northern Sweden in two seasons. The sites were situated in a wide range of streams and rivers from small springbrooks, bog streams and lake-outlet streams to medium-sized forest rivers and large rivers draining montane regions.
2. Thirty-nine blackfly species were found, with between two and thirteen species per site. Neither species richness nor abundance could be related to the environmental variables measured.
3. An analysis of labral fan size of blackflies indicated a clear trend for the prevalence of larvae with small fans in large rivers and larvae with larger fan size in small streams. Similarly, fan size related to current velocities so that large fans were associated with slow current velocities and small fans with high velocities.
4. A strong relationship existed between species composition and habitat, as seen in ordination by non-metric multidimensional scaling. The relationship found between fan size and habitat size-related variables, such as channel width, depth, velocity and substratum particle size, along with longitude and altitude, in partial least squares regression analysis offered an explanation of the species composition–habitat relationship.
5. In addition to testing that distributions of blackfly larvae reflect morphological traits, we tested two general hypotheses pertaining to distribution patterns: (a) that blackfly communities show bimodal distributions; and (b) that their distributions are nested. Neither of these two hypotheses was supported by our observations. However, widespread blackfly species were locally more abundant than those found at relatively few sites, thus showing a positive abundance–occupancy relationship.     

Impact of habitat diversity on the sampling effort required for the assessment of river fish communities and IBI

The spatial variation in the fish communities of four small Belgian rivers with variable habitat diversity was investigated by electric fishing to define the minimum sampling distance required for optimal fish stock assessment and determination of the Index of Biotic Integrity. This study shows that the standardised sampling distance of 100 m was not always sufficient to collect most species present. The required minimum sampling distance seems to be correlated with habitat diversity. In homogeneous streams, a mean sample distance of 282, 452 and 572 m is necessary to capture 80, 90 and 95% of all species present, respectively. In heterogeneous streams, these sample distances decrease to 217, 380 and 503 m. Hence, at least 300 m should be sampled to catch most species present with a single-pass sampling method. However, our results show that a 100 m sampling distance as presently used in the Flemish monitoring programs is sufficient to accurately describe the ecological quality since differences in IBI evaluation between adjacent stretches could at least for some rivers be explained by differences in habitat heterogeneity.     

The role of small secondary biotopes in a highly fragmented landscape as habitat and connectivity providers for dragonflies (Insecta: Odonata)

Habitat loss and degradation are considered major threats to freshwater biodiversity and to invertebrates in particular. These often irreversible processes may lead to local and regional extinctions of species, most notably of stenotopic taxa. In spite of this, a number of studies have shown that small habitat patches can sustain rich and abundant communities. The present work assesses the relevance of a group of four small man-made (secondary) wildlife ponds to Odonata species diversity and abundance. Results obtained on pond recruiting capacity, species richness, abundance and habitat use by means of exuviae collection and monitoring of adults using a capture–mark–recapture (CMR) method indicate the potential suitability of these small aquatic biotopes and the surrounding landscape as habitat providers and stepping stone connectors in the Vallès lowlands (Catalonia, Spain). This region, close to the metropolitan area of Barcelona, has severely degraded natural habitats and high landscape fragmentation due to infrastructure, urban and industrial expansion. A comparison among a greater number of sites (ponds and sections of streams and rivers) distributed across the region showed that adequately managed small waterbodies harbour richer Odonata communities than others that are unmanaged or managed specifically for other types of fauna or uses. Appropriate care of these small biotopes avoids disturbance and keeps them free from vertebrates like fish and waterfowl which, under certain conditions, may have a strong influence on the invertebrate communities because, apart from feeding on larvae, they may have a negative impact on macrophyte development and water quality. While rivers and streams, the only natural aquatic habitats in the area, may be both expensive and technically challenging to restore and manage successfully, the creation and/or restoration of small ponds and short river sections in suitable locations can be a cost-effective method for enhancing freshwater vegetation and invertebrate diversity in this impacted landscape.     

Habitat amount drives the functional diversity and nestedness of anuran communities in an Atlantic Forest fragmented landscape

Natural environments disturbed by human activities can suffer from species extinctions, but some can still harbor high taxonomic diversity. However, disturbances may have impacts beyond the species level, if the species lost represent unique functions in the ecosystem. In this study, we evaluated to what extent the amount of habitat can determine the functional diversity and nestedness of amphibian communities in an Atlantic Forest fragmented landscape in Brazil, and if there is a threshold of habitat amount beyond which there is severe loss of functional diversity. As species responses may depend on their habitat type, we performed the analyses for three different sets of species: all species, forest‐dependent species, and generalist species. We also evaluated the relative importance of turnover and nestedness components to total functional dissimilarity among sites. Habitat amount affected functional diversity of frogs, especially for forest‐dependent species where a linear reduction was detected. The functional dissimilarity among sites was mostly explained by the nestedness component. The reduction of functional diversity was mediated by an ordered loss of traits, leading to a functionally nested metacommunity. These sensitive traits were closely related to habits and reproductive modes that depend on rivers and streams. The maintenance of functional diversity of frogs in fragmented landscapes must rely on the conservation of both terrestrial and aquatic environments, as some species and their traits can disappear from remnants of native vegetation lacking some specific habitats (e.g. streams). Abstract in Portuguese is available with online material.     

Life-history traits of invasive fish in small Mediterranean streams

We compared the life-history traits of native and invasive fish species from Catalan streams in order to identify the characters of successful invasive fish species. Most of the exotic fish species were characterized by large size, long longevity, late maturity, high fecundity, few spawnings per year, and short reproductive span, whereas Iberian native species exhibited predominantly the opposite suite of traits. Species native to the southeastern Pyrenees watershed were also significantly different from species native to the rest of the Iberian Peninsula but not native to this watershed. Iberian exotic species come predominantly from large river basins, whereas Catalan streams (and other small, coastal river basins) correspond to basins and streams of a smaller size and different hydrology, with differences in species composition and life-history traits of fish. The occurrence and spread of invasive species was not significantly related to life-history traits but to introduction date. The successful prediction of future invasive species is limited due to small differences in life-history and ecological traits between native and exotic species. Fecundity, age at maturity, water quality flexibility, tolerance to pollution and habitat seem the most discriminating life-history variables. This revised version was published online in July 2006 with corrections to the Cover Date.     

Principles for linking fish habitat to fisheries management and conservation

A set of eight simple ecological and social principles is proposed that could enhance the understanding of what constitutes fish 'habitat' and, if implemented, could contribute to improved management and conservation strategies. The habitat principles are a small, interrelated sub‐set that may be coupled with additional ones to formulate comprehensive guidelines for management and conservation strategies. It is proposed that: 1) habitat can be created by keystone species and interactions among species; 2) the productivity of aquatic and riparian habitat is interlinked by reciprocal exchanges of material; 3) the riparian zone is fish habitat; 4) fishless headwater streams are inseparable from fish‐bearing rivers downstream; 5) habitats can be coupled – in rivers, lakes, estuaries and oceans, and in time; 6) habitats change over hours to centuries; 7) fish production is dynamic due to biocomplexity, in species and in habitats; 8) management and conservation strategies must evolve in response to present conditions, but especially to the anticipated future. It is contended that the long‐term resilience of native fish communities in catchments shared by humans depends on incorporating these principles into management and conservation strategies. Further, traditional strategies poorly reflect the dynamic nature of habitat, the true extent of habitat, or the intrinsic complexity in societal perspectives. Forward‐thinking fish management and conservation plans view habitat as more than water. They are multilayered, ranging from pools to catchments to ecoregions, and from hours to seasons to centuries. They embrace, as a fundamental premise, that habitat evolves through both natural and anthropogenic processes, and that patterns of change may be as important as other habitat attributes.     

Community assembly,functional traits,and phylogeny in Himalayan river birds

Heterogeneity in riverine habitats acts as a template for species evolution that influences river communities at different spatio‐temporal scales. Although birds are conspicuous elements of these communities, the roles of phylogeny, functional traits, and habitat character in their niche use or species'' assembly have seldom been investigated. We explored these themes by surveying multiple headwaters over 3000 m of elevation in the Himalayan Mountains of India where the specialist birds of montane rivers reach their greatest diversity on Earth. After ordinating community composition, species traits, and habitat character, we investigated whether river bird traits varied with elevation in ways that were constrained or independent of phylogeny, hypothesizing that trait patterns reflect environmental filtering. Community composition and trait representation varied strongly with increasing elevation and river naturalness as species that foraged in the river/riparian ecotone gave way to small insectivores with direct trophic dependence on the river or its immediate channel. These trends were influenced strongly by phylogeny as communities became more clustered by functional traits at a higher elevation. Phylogenetic signals varied among traits, however, and were reflected in body mass, bill size, and tarsus length more than in body size, tail length, and breeding strategy. These variations imply that community assembly in high‐altitude river birds reflects a blend of phylogenetic constraint and habitat filtering coupled with some proximate niche‐based moulding of trait character. We suggest that the regional co‐existence of river birds in the Himalaya is facilitated by this same array of factors that together reflect the highly heterogeneous template of river habitats provided by these mountain headwaters.     

Phylogenetic habitat filtering influences forest nucleation in grasslands

Because species–environment interactions are mediated by phenotypic tradeoffs, the maintenance of ancestral traits in some phylogenetic clades and the emergence of evolutionary novelties in others are likely to limit the types of habitats that species occupy, generating phylogenetic habitat filtering. To test for phylogenetic habitat filtering in woody sapling communities in vegetation patches scattered in southern Brazilian grasslands, I estimated if patches of different sizes encompassed species of different phylogenetic groups. I analyzed patch composition with principal coordinates of phylogenetic structure (PCPS), extracted from a matrix of phylogeny‐weighted species composition, and compared these results against net relatedness index (NRI) analyses. NRI analysis revealed that most communities were phylogenetically random, and that patches of different sizes did not differ from each other with respect to NRI. The first four PCPS contained ? 91% of total variation in phylogeny‐weighted species composition. In the first two PCPS, scores of large patches differed from those of small and medium patches, which did not differ from each other. Large patches were associated with basal plant clades, whereas small patches were mostly related to asterids, and medium patches were phylogenetically diverse. Phylogenetic habitat filtering was detected only by PCPS analysis, possibly because NRI analysis does not take into account the habitat specificity of species. Taking phylogenetic habitat filtering into account in comparative studies likely enhances our capability to understand the ways that plants interact with their environment.     

Constraints on recovery: using molecular methods to study connectivity of aquatic biota in rivers and streams

1. The ‘Field of Dreams Hypothesis’ states ‘if we build it, they will come’, referring to the assumption that if habitats are restored, species will recolonise them. However, the ability of a species to recolonise a restored site will depend not only on the appropriate habitat being present, but also on the ability to get there. This is likely to depend on both the species’ dispersal behaviour and the position of a site in the landscape. 2. Animals with good potential for dispersal are more likely to be able to disperse to newly restored sites. Similarly, sites in lowland streams with limited altitudinal differences between sites may be easier to reach than upstream sites. This is because upstream sites are connected to one another via lowland streams that have different characteristics and therefore may be difficult for animals to traverse. 3. In this paper, genetic data from a range of freshwater species that have been analysed in my laboratory are used to assess the importance of life cycle and position in the landscape (i.e. upland versus lowland streams) on connectivity patterns (and thus recolonisation potential) among populations. 4. In general, contemporary dispersal across catchment boundaries is negligible, except for aquatic insects with an adult flight stage. Dispersal among streams within catchments appears to be more limited than was predicted from knowledge on life histories, except for fish in lowland rivers and streams. 5. As predicted, dispersal of fish, crustaceans and molluscs among streams within catchments is significantly greater in lowland rivers than in upland streams. 6. Overall, these analyses suggest that, with the exception of most insects, and fishes in lowland rivers, natural recolonisation of restored sites is only likely from sites within the same stream. If a species has disappeared from the whole stream, then restoration of habitat alone may not be sufficient for its re‐establishment.     

Local habitat restoration in streams: Constraints on the effectiveness of restoration for stream biota

Summary   The restoration of physical habitat has emerged as a key activity for managers charged with reversing the damage done by humans to streams and rivers, and there has been a great expenditure of time, money and other resources on habitat restoration projects. Most restoration projects appear to assume that the creation of habitat is the key to restoring the biota ('the field of dreams hypothesis'). However, in many streams where new habitat is clearly required if populations and communities are to be restored, there may be numerous other factors that cause the expected link between habitat and biotic restoration to break down. We discuss five issues that are likely to have a direct bearing on the success, or perceived success of local habitat restoration projects in streams: (i) barriers to colonization, (ii) temporal shifts in habitat use, (iii) introduced species, (iv) long-term and large-scale processes, and (v) inappropriate scales of restoration. The purpose of the study was primarily to alert ecologists and managers involved in stream habitat restoration to the potential impacts of these issues on restoration success. Furthermore, the study highlights the opportunities provided by habitat restoration for learning how the factors we discuss affect populations, communities and ecosystems.     

Salamander occupancy in headwater stream networks

1. Stream ecosystems exhibit a highly consistent dendritic geometry in which linear habitat units intersect to create a hierarchical network of connected branches.
2. Ecological and life history traits of species living in streams, such as the potential for overland movement, may interact with this architecture to shape patterns of occupancy and response to disturbance. Specifically, large-scale habitat alteration that fragments stream networks and reduces connectivity may reduce the probability a stream is occupied by sensitive species, such as stream salamanders.
3. We collected habitat occupancy data on four species of stream salamanders in first-order (i.e. headwater) streams in undeveloped and urbanised regions of the eastern U.S.A. We then used an information–theoretic approach to test alternative models of salamander occupancy based on a priori predictions of the effects of network configuration, region and salamander life history.
4. Across all four species, we found that streams connected to other first-order streams had higher occupancy than those flowing directly into larger streams and rivers. For three of the four species, occupancy was lower in the urbanised region than in the undeveloped region.
5. These results demonstrate that the spatial configuration of stream networks within protected areas affects the occurrences of stream salamander species. We strongly encourage preservation of network connections between first-order streams in conservation planning and management decisions that may affect stream species.     

Evolutionary and ecological correlates of early seedling morphology in East African trees and shrubs

Seed size and cotyledon morphology are two key juvenile traits that have evolved in response to changes in plant species life-history strategies and habitat associations. Correlations of these traits with each other and with other juvenile traits were examined for 70 species of trees and shrubs in Kibale National Park, Uganda. Although species with photosynthetic cotyledons were more abundant than in other tropical floras, both univariate and multivariate analyses supported trait associations expected from the literature. Trait values varied continuously across species, yet mean trait values differed significantly among habitat association types. Species with large seeds, large seedlings, thick storage cotyledons, slow germination, large-stature adults, and dispersal by large animals were common in forest and gap habitats. An opposite suite of traits was common in open habitats (grassland and edge). Analyses incorporating phylogeny (independent contrasts and omnibus tests) confirmed that these suites of traits showed correlated evolution. Cotyledon functional morphology yielded a strong phylogenetic signal, while seed mass was labile. Nevertheless, contingent change tests found that evolutionary change from photosynthetic to reserve cotyledons was more likely when disperser and perhaps seed size of ancestral species were already large, suggesting a strong interdependency among these traits.     

Short‐term Impacts of Fish Removal from Small Amazonian Forest Streams

Collecting freshwater zoological material for research purposes is common and necessary, but its impact on biological communities is often overlooked. We tested the impacts of a conventional ichthyological collection in small forest streams and showed that fish assemblages recover after a few months. We highlight ecological traits and habitat characteristics that might exacerbate the impacts of repeated removal of fish in long‐term investigations, and advocate for the use of a capture‐release approach whenever possible.     

A comparison of the external morphology of the membranous inner ear in elasmobranchs

Studies on the elasmobranch inner ear have focused predominantly on a small group of sharks, particularly, carcharhinids. As a result, subsequent studies in other species have subdivided species into two main groups: those typical and those atypical of carcharhinid sharks. This study proposes a different set of inner‐ear morphology groupings to those previously suggested. The inner ears from 17 species of elasmobranchs (representing both sharks and rays) are examined in this study and based on morphometric data some groups include both rays and sharks. Four groups are now proposed based predominantly on the shape and dimensions of the membranous otoconial organs, and characteristics of the semicircular canals. Evident morphological differences between the ear types belonging to the new groups include the membranes of the semicircular canals being bound to the otoconial organs in some species, while only being connected via the canal ducts in others, as well as clear variation present in saccular organ size. Previous studies examining variation in the inner ear have attributed differences to either phylogeny or functional significance. Results from this study suggest that neither phylogeny nor feeding strategy solely accounts for the morphological diversity present in the external morphology of the elasmobranch inner ear. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

Dark diversity in Amazonian stream fish communities: What factors determine species absence along environmental gradients?

1. Species distribution models often fail to predict observed patterns of species diversity, and this is because some species within a regional pool that are tolerant of conditions at a given location may nevertheless be absent from the local community. These missing species have been termed “dark diversity”. In the present study, we investigated which factors explain dark diversity among fish assemblages in Amazonian streams.
2. We sampled 71 streams in areas with different types of land use within two river basins and estimated dark diversity from patterns of species co-occurrence, using Beals’ index, along environmental gradients. From this procedure, taxa are designated as dark diversity components when they are absent from a given stream, but often co-occur with the local species at other streams, indicating similar ecological requirements. We used generalised linear models both to determine whether environmental or landscape variables, connectivity, instream environmental heterogeneity or some combination of these factors explained dark diversity of fishes, and to evaluate whether ecomorphology is associated with the extent to which a species contributes to dark diversity and which specific traits contribute the most to explaining variation in dark diversity.
3. Mean local diversity exceeded observed dark diversity. The magnitude of dark diversity was directly associated with the proportion of secondary forest in the immediate catchment and with the index of proximity to anthropogenic impact. Species that have high affinity for environments with higher current velocity, low swimming ability and that capture food mainly on the surface contributed more to dark diversity, which suggests that swimming ability, habitat preference and aspects related to diet are key predictors of the probability that a given species will be present at locations with suitable habitat.
4. Our findings reinforce the idea that dark diversity results from interactions between species traits and environmental factors, including anthropogenic impacts. Understanding the interplay among environmental factors and species traits that contribute to dark diversity provides targets for improved ecosystem restoration and sustainability of native species assemblages.

     

Evolutionary aspects of cephalic sensory papillae of the Indo‐Pacific species of Eleotris (Teleostei: Eleotridae)

Eleotris species (Teleostei: Eleotridae) are one of the most common fish in Indo‐Pacific estuaries and insular freshwater streams. In these rivers, they are a sit‐and‐wait predator. They have an amphidromous life cycle, that is adults grow, feed and reproduce in rivers, while larvae have a marine dispersal phase. Larvae recruit back to rivers and settle in stream habitats. Primary characters used to determine Eleotris species are the presence and the disposition of cephalic sensory papillae rows on the operculum and under the eyes as well as scale row numbers. The morphology of these cephalic sensory papillae is of particular importance in this predatory genus as it is generally correlated in fish to predation and feeding. In this paper, we have established a molecular phylogeny of the genus based on the 12 mitochondrial protein‐coding genes to discuss the relationship between Indo‐Pacific Eleotris species. There is a well‐supported dichotomy in the molecular phylogeny, and this separation into two main clades is also morphologically visible, as it reveals a difference in the arrangement of cephalic sensory papillae. Indeed, the phylogeny distinguishes the species with the “open” pattern of the operculum sensory papillae and the species with the “closed” one. This phylogeny thus reflects the morphology of the opercular papillae. The evolution of this character is discussed in terms of the adaptation of the Eleotris genus to life in tropical insular river systems.     

Long-term effects of stream management on plant communities in two Danish lowland streams

Submerged macrophytes grow abundantly in most shallow streams common in the cultivated lowlands of northwestern Europe. Weed-cutting has been practised for years in many of these streams to reduce the risk of flooding of adjacent land. Our objective was to quantify long-term impacts of weed-cutting on macrophyte communities in two Danish rivers. We found that the total macrophyte coverage was similar in the weed-cut and uncut reaches in the two rivers, but species richness, diversity and patch complexity were higher in the uncut reaches. The spatial distribution of macrophytes on the stream bottom was also more heterogeneous in the uncut stream reaches. We also found evidence of a strong effect of weed-cutting on macrophyte species composition. P. natans was abundant in the uncut reaches in both streams but practically eliminated in the cut reaches, despite the fact that its basic habitat requirements were met. Also, the established phase strategy of the macrophyte community was affected by weed-cutting. Species displaying characteristically ruderal traits were more abundant in the cut reaches and species with competitive abilities were only abundant in the uncut stream reaches. We suggest that important species traits in streams, where the weed is cut regularly, are associated with rapid growth and high dispersal-capacity. Our results indicate that weed-cutting can contribute significantly to a decline in species diversity in streams. To provide optimal conditions for diverse stream macrophyte communities, we therefore suggest that weed-cutting should be minimised.     

Development of metrics based on fish body size and species traits to assess European coldwater streams

During the last decade multimetric indices (MMIs) have been greatly improved by the use of appropriate criteria to define reference conditions and by the use of statistical analysis to select a consistent set of metrics. Among the large number of MMIs developed to assess the ecological status of streams based on fish communities, the emphasis was mainly put on warmwater assemblages. When compared with warmwater fish assemblages, coldwater assemblages present depauperate faunas with a limited suit of traits. Thus, very often the number of metrics used to compute MMIs for coldwater streams is lower than for warmwaters. The objective of this study was to develop new metrics specific to European coldwater assemblages that integrate both the species traits and the body size of fish. Indeed, whereas the use of size or age classes has been highly advocated for developing MMIs, it remains largely underrepresented. Therefore, we used eight biological and ecological traits to characterize species and two size classes: small and large individuals. Among the 96 metrics tested, four were successfully related to environmental gradients and three displayed a significant response to anthropogenic pressures: the number of small rheophilous individuals, the number of small oxygen-intolerant individuals, and the number of small-habitat-intolerant individuals. These results demonstrate that metrics based on size classes could be used in the development of MMIs for coldwater streams and more generally for low-species rivers.     

Salinity-related distribution of alien amphipods in rivers provides refugia for native species

There are numerous papers upon the range extension, biology and the impact of alien amphipods upon the local fauna. However, there are no studies concerning the alien versus native species distribution patterns at the catchment scale of river systems. In total 125 sites were sampled: 41 in main rivers constituting the Polish section of the central invasion corridor (Bug, Vistula, Notec with canals, Oder) and 84 in their affluents. The conductivity of large rivers was much higher than in their small affluents. The number of alien species and their abundance was higher at sites with raised conductivity values. The reverse situation was noticed when native amphipods were taken into account. Moreover, large rivers were inhabited by alien fauna, whereas in smaller streams only native species were sampled. The absence of alien amphipods in small rivers may be explained by their ecological preference for a higher conductivity of water. Thus in smaller streams the native fauna is free from the pressure posed by alien amphipods. If not degraded by human activity, small affluents may function as refugia for native amphipod species. Otherwise, elevation of salinity related to improper catchment management or sewage treatment may lead to extinction of such local disjunct population of native amphipods.     

The stream as a habitat templet for bryophytes: species' distributions along gradients in disturbance and substratum heterogeneity

1. The structure of bryophyte communities in streams in relation to habitat characteristics, especially disturbance, is described. Disturbance in rivers is quantified as movement of the stream bed, whereas in small streams water level fluctuation is used as an indicator of disturbance frequency. 2. Canonical correspondence analysis differentiated frequently disturbed sites from more stable ones. The existence of a disturbance gradient was confirmed in a subset of the study sites using long-term records of discharge variation. A parallel change was detected in the species composition of bryophyte communities with low-statured, potentially fast colonizers dominating the disturbed end and large perennial species the stable end of the gradient. 3. A consistent pattern of zonation of bryophyte species was found along the gradient from continually submersed to persistently dry conditions in small streams and lake outlets. An abrupt increase in species richness occurred at or just above the water line, where facultatively aquatic species tolerant of both conditions formed the bulk of the community. 4. The relationship between species richness and standing crop in stream bryophyte communities was consistent with the hump-backed model of Grime (1979), especially at the within-habitat scale. Quadrats of low and very high standing crop were characterized by low species richness, while peak richness was observed at intermediate standing crops. 5. A few perennial species (e.g. Fontinalis spp. and Rhynchostegium riparioides) capable of monopolizing space dominated the most stable habitats. Species composition in low biomass sites was more variable, yet only one basic growth-form (small-statured species with high allocation to spore production) seemed possible in these highly disturbed streams. In habitats of intermediate biomass, small-scale disturbances apparently allow the formation of a more varied bryophyte community. 6. A habitat templet for stream bryophyte life strategies and community structure is presented. Disturbance is proposed to be the factor filtering out traits unsuitable for a given environment. The potential of stream bryophytes for testing and developing general ecological theory is emphasized.