Early succession on lahars spawned by Mount St. Helens

The effects of isolation on primary succession are poorly documented. I monitored vegetation recovery on two Mount St. Helens lahars (mud flows) with different degrees of isolation using contiguous plots. Seventeen years after the eruption, species richness was stable, but cover continued to increase. That isolation affects community structure was confirmed in several ways. The dominance hierarchies of the lahars differed sharply. Detrended correspondence analysis on Lahar I showed a trend related to distance from an adjacent woodland, whereas vegetation on Lahar II was relatively homogeneous. Spectra of growth forms and dispersal types also differed. Lahar I was dominated by species with modest dispersal ability, while Lahar II was dominated by species with better dispersal. Variation between plots should decline through time, a prediction confirmed on Lahar II. Lahar I remained heterogeneous despite having developed significantly higher cover. Here, the increasing distance from the forest has prevented plots from becoming more homogeneous. At this stage of early primary succession, neither lahar is converging towards the species composition of adjacent vegetation. This study shows that isolation and differential dispersal ability combine to determine initial vegetation structure. Stochastic effects resulting from dispersal limitations may resist the more deterministic effects of competition that could lead to floristic convergence.     

The impact of large herbivores on floral composition and vegetation structure in the Naukluft Mountains, Namibia

Floral composition and vegetation structure were investigated on the Naukluft Mountain Plateau to determine (i) whether or not grazing has had an impact on the flora of the Naukluft Mountains during the last two decades and (ii) if so, whether vegetation recovery is influenced by the nature of the rainy season. Vegetation subjected to grazing to large herbivores and vegetation excluded from grazing for 18 and 19 years was compared. Based on classification (TWINSPAN) and ordination (detrended and canonical correspondence analysis) of 20, 10 × 10 m stands within and outside exclosures in two seasons, a clear difference in floristic composition between grazed and not grazed areas was indicated. Although neither the height of the vegetation, not the growth form spectrum differed significantly between grazed and not grazed sites, leaf-succulents were much more abundant when grazing was excluded. This suggests that changes in the vegetation due to grazing pressure have occurred in the last two decades, but the degree of grazing impact is not severe and may be part of the natural savanna ecosystem of the Naukluft Mountains. Whether this trend also occurs in other vegetation types in the Naukluft Mountains remains to the investigated. This revised version was published online in November 2006 with corrections to the Cover Date.     

Environmental factors determining the distribution of highland plants at low-altitude algific talus sites

Algific talus is a micro-scale habitat type where highland plants (subalpine and alpine species) are found, disjunct from their typical range, in lowland forests. On algific talus, cold airflows from the interstices between talus fragments create a local microclimate colder than surrounding forests. Despite of the widely-known occurrence of unique vegetation on algific talus, critical environmental factors determining the distribution of highland species in this habitat type are unclear. In order to reveal the environmental factors enabling highland species to inhabit algific talus, we investigated the vegetation and environments of 26 algific talus sites and four reference (non-algific talus) sites in Hokkaido, northern Japan. Several algific talus sites were dominated by highland species, while some algific talus sites and all non-algific talus sites were dominated by lowland species. Community analysis based on detrended correspondence analysis (DCA) and canonical corresponding analysis (CCA) revealed that the algific talus sites dominated by highland species had lower ground temperature, more acidic soil, larger canopy openness, and less diverse vegetation than the sites dominated by lowland species. Highland plants might be maintained under conditions stressful for lowland plants, resulting in less competitive situation. Generalized linear models (GLM), used to evaluate the response of individual highland species to environmental factors, revealed that preferable environmental conditions for highland plants are highly species specific. These results indicate that the maintenance of diverse environments is crucial for the conservation of the unique vegetation and local populations of highland species in algific talus areas.     

Diversity and Abundance of Biological Soil Crust Taxa in Relation to Fine and Coarse-Scale Disturbances in a Grassy Eucalypt Woodland in Eastern Australia

Over the last 200 years the box woodlands of eastern Australia have been considerably altered by European farming practices. These changes have been accompanied by a reduction in the size and number of patches of vegetation as well as the quality of the understorey vegetation and underlying soil surface. We measured diversity and abundance of soil crust taxa in relation to habitat complexity, remnant area and width, diversity of vascular plants as well as the number, size and separation of patches of vegetation and grass butts (coarse-scale patchiness), and an index of surface stability derived from measures of seven soil surface features of small microsites (patches of bare/crusted, litter- or grass-covered soil; micro-scale) on both coarse- and fine-textured soils at 35 sites in south-eastern Australia. Fifty taxa were recorded from the 35 sites, and there were more taxa from sites with fine-textured soils (12.7) compared with coarse-textured soils (4.4). The soil crust community was dominated by a few relatively common species, with many species occurring at only a few sites. Half the number of species accounting for <1% of total abundance. Bare and crusted microsites supported more species and greater cover compared with grassy microsites. Crust diversity declined with increasing coarse-level disturbances (i.e. declines in habitat complexity, remnant area and width, and diversity of vascular plants) but the results were not consistent between soil types. No measures of fine-scale disturbance were related significantly to any of the crust diversity or abundance measures, and there was no evidence of a recent grazing effect on crust composition. The fact that few sites had many species (and visa versa) suggests to us that many sites are probably required to conserve soil crust taxa in these highly fragmented landscapes     

Succession following the catastrophic eruption of Ksudach volcano (Kamchatka, 1907)

Ksudach Volcano, southern Kamchatka Peninsula, erupted in 1907 and impacted over 2000 km² of forests with air-fall pumice deposits. We identified three impact zones. In Zone I, deposits deeper than 100 cm destroyed all vegetation. Two early successional stages occur, a lichen-dominated desert and isolated patches of a pioneer herb stage. Zone II is defined by pumice deposits 30 to 100 cm deep. Deposits of 70 to 100 cm destroyed all vegetation, but left scattered snags. Here primary succession dominates recovery, but its rate varies. Isolated trees survived in deposits of 30 to 70 cm and primary and secondary successional stages form a complex mosaic termed an intermediate succession. In Zone II, the primary stages found in Zone I are joined by a dwarf shrub-herb stage and a secondary birch forest stage. Zone III occurs where thinner deposits permitted some vegetation to survive in all locations. Secondary succession dominates in deposits of 10 to 30 cm. Trees suffered damage, but survived deposits of 20 to 30 cm, while other vegetation layers were eliminated. Deposits of 10 to 20 cm eliminated mosses and lichens and but only reduced the number of dwarf shrubs and herbs. Deposits of less than 10 cm damaged herb, moss and lichen layers but did not eliminate any species. All sampled vegetation remains in a pre-climax state, having yet to recover fully from earlier eruptions. Reconstructed vegetation maps for before 1907 and for ca. 1925 are compared to the map of vegetation in 1994. Based on degree of soil formation, vegetation recovery and colonization rates at different pumice depths, and the current vegetation, we estimate that full recovery of the soil-vegetation system will take more than 2000 years.     

汶川地震滑坡迹地植物群落与环境的关系

为了加快汶川地震滑坡迹地人工恢复植被的进程,探讨地震诱发的滑坡迹地植物群落与环境的关系。在5·12地震重灾区北川境内选取29个样地进行植被调查,采用10个环境指标刻画植物群落的地形、空间位置和土壤养分特征;利用TWINSPAN、CCA、DCA和DCCA,分析植物种、植物群落和植物生活型与环境的关系。结果显示:1)研究区的植物群落可划分为9个类型。2)研究区环境变量对植物种的解释量为21.96%,第一排序轴与pH值、海拔、土壤质地相关,反映的是植物种从次生植物群落向原生植物群落变化。通过DCCA分析得出,环境变量对植物群落的排序解释了25.7%,第一排序轴与pH值、海拔、土壤质地的相关较强,反映植物群落按照耐旱、耐贫瘠→人工或先锋植物→未受损的植被变化;第二排序轴与土壤有机质、全氮含量、坡向的相关,反映的是植物群落从草本植物→乔灌草或者灌草植物变化。3)滑坡迹地的植物群落与未受损林地的植物群落物种存在较大差异。     

Early primary succession on Mount St. Helens,Washington, USA

Abstract. The north slope of Mount St. Helens was sampled with 141 circular 100-m² plots to describe vegetation and environmental patterns 13 yr after the 1980 eruption. At least 114 vascular plant taxa were encountered. We recognized four habitat types: Refugia, Pumice barrens, Pyroclastic surfaces and Drainages. A fifth category, Lupine patches, includes samples on primary surfaces that were rapidly colonized. Refugia provided small enclaves where underground portions of several species survived the eruption. They retained an inconsistent array of forest understory species and contained 86 species (mean = 20.8 per plot). Refugia are dominated by woody species such as Penstemon cardwellii, Rubus spp., Ribes spp. and Alnus sinuata, with herbs such as Agrostis diegoensis, Luzula parviflora and Anaphalis margaritacea. Anaphalis represents a suite of species that invaded Refugia after the eruption. Diversity (N₂ and H′) is significantly greater in Refugia than in any other habitat. No plants survived on primary sites, which remain sparsely vegetated and dominated by readily dispersed taxa. Total richness ranges from 36 species (9.9 per plot) on pyroclastic surfaces, through 42 species (11.2 per plot) in drainages, to 66 (11.7 per plot) on Pumice barrens. H′ and N₂ of the three habitats do not differ significantly. Lupine-dominated vegetation occurs sporadically in Pyroclastic and drainage habitats. Lupine patches are characterized by high Lupinus cover and a suite of invaders. These sites have high cover and 52 species (12.6 per plot). H′ and N₂ scores were significantly lower than any other habitat due to strong lupine dominance. Canonical Correspondence Analysis showed that site history and slope contributed most to species composition. Geographic effects accounted for 10 25 % of the explained species-environment relationship. Forest understory species have migrated only short distances and have made negligible contributions to vegetation development. A few species common in Refugia, including Agrostis diegoensis and Carex mertensii, have invaded barren surfaces, but most have not. Refugia also have been invaded by open site species abundant on the Pumice Plain. The heterogeneity of plots within habitat types and small statistical linkage of vegetation to environmental and spatial factors suggests that stochastic events have played a leading role in early primary succession.     

Long‐term effects of drainage and initial effects of hydrological restoration on rich fen vegetation

Questions: What vegetational changes does a boreal rich fen (alkaline fen) undergo during a time period of 24 years after drainage? How is plant species richness affected, and what are the changes in composition of ecological groups of species? Is it possible to recover parts of the original flora by rewetting the rich fen? Which are the initial vegetation changes in the flora after rewetting? What are the major challenges for restoration of rich fen flora after rewetting? Location: Eastern central Sweden, southern boreal vegetational zone. Previously rich fen site, drained for forestry purposes during 1978–1979. The site was hydrologically restored (rewetted) in 2002. Method: Annual vegetation survey in permanent plots during a period of 28 years. Results: There were three successional stages in the vegetational changes. In the first stage there was a rapid (< 5 years) loss of rich fen bryophytes. The second step was an increase of sedges and early successional bryophytes, which was followed by an increase of a few emerging dominants, such as Molinia caerulea, Betula pubescens and Sphagnum spp. After rewetting, there are indications of vegetation recovery, albeit at slow rates. Depending on, for instance, initial species composition different routes of vegetation change were observed in the flora after drainage, although after 24 years, species composition became more homogenous and dominated by a few species with high cover. Conclusion: Major changes have occurred after changes in the hydrology (drainage and rewetting) with a severe impact on the biodiversity among vascular plants and bryophytes. Several rich fen bryophytes respond quickly to the changes in water level (in contrast to vascular plants). The recovery after rewetting towards the original rich fen vegetation is slow, as delayed by substrate degradation, dispersal limitation and presence of dominant species.     

Sacred sites in north‐western Morocco – naturalness of their vegetation and conservation value for vulnerable plant species

The vegetation and flora of 86 sacred sites (surroundings of tombs of local saints, cemeteries) in NW Morocco was studied to assess their conservation value for undisturbed Mediterranean evergreen forests and vulnerable plants. Habitats were mapped, based on life form, maximum height and cover value, and classified according to naturalness. Vascular plants were classified according to pre‐defined risk factors like stenochory or grazing sensitivity. Naturalness index and refugial value were calculated for each site. Only 20% of the mapped site area was closed forest. On many places, human impact had created a vegetation mosaic. Most of the vulnerable plants were mesophilous forest species or grazing sensitive taxa. Endemic species occurred only rarely. The occurrence of vulnerable plants and the naturalness of the vegetation were positively correlated. Sacred site extension had no significant effect on the number of vulnerable species. Results indicate a limited relevance of sacred sites in northern Morocco for the protection of intact forests and vulnerable plants. However, these localities can play an important role as stepping stones in agricultural landscapes. They protect plants, sensitive to opening of the tree layer, to deforestation and to intense grazing. Certain formerly widespread forest communities survived exclusively on sacred sites.     

Effects of time since fire on frog occurrence are altered by isolation,vegetation and fire frequency gradients

Aim

To quantify how frogs in terrestrial environments respond to recurrent fire, and to what extent this is mediated by isolation from breeding sites or vegetation structure.

Location

Jervis Bay, south‐eastern Australia.

Methods

We used data from 8 years of pitfall trapping, collected via a random stratified design, to quantify frog occurrence at 110 locations. We then used an information theoretic approach to compare 13 logistic generalized linear mixed models, each of which related frog occurrence to a distinct combination of additive and interactive effects of fire, vegetation structure and proximity to known breeding sites.

Results

For all four species, the effect of one or more fire variables on frog occurrence depended on both the density of breeding sites in the surrounding area, or on the vegetation structure at the trap locality. A classic “fire averse” response of initial declines followed by post‐fire recovery did occur, but only in frequently burned, low‐quality terrestrial habitats (i.e., heath vegetation for Uperoleia tyleri, or locations with few available breeding sites for the remaining species), or in some cases, when suitable habitats were infrequently burned. However, a “fire‐dependent” result of negative effects of time since fire was also evident for some species and contexts.

Main conclusions

The effect of fire on frog occurrence can be mediated by environment. Therefore, a single species could be identified as either “fire dependent” or “fire averse” depending on the combination of isolation, vegetation types and fire histories in the study region. Failure to account for the context specificity of fire response curves could lead to incomplete conclusions regarding the effect of time since fire—or the cumulative impacts of multiple fires—on faunal assemblages.     

The shrubland vegetation in western Shewa,Ethiopia and its possible recovery

Shrubland vegetation and environmental data in western Shewa, Ethiopia have been analysed. Vegetation data include cover-abundance values of vascular plant species; environmental data comprise physical and chemical properties of the soil, altitude, slope, grazing and browsing pressure. The vegetation data were subj ected to hierarchical and non-hierarchical classification and ordination with correspondence analysis. The classification resulted in seven different vegetation types, ranging from grassland with scattered shrubs to degenerated forest. Ordination of the data and biplot analysis showed that the vegetation is influenced by anthropogenic factors and altitudinal variation. Sand content is related to a low level of anthropogenic influence whereas silt content is related to a high level. This is explained by historical events rather than by the present situation. Total nitrogen, organic carbon, altitude and slope are positively correlated and these variables are negatively related to anthropogenic influences. The shrubland vegetation may have expanded from lower altitudes and drier sites as forests gradually disappeared. The recovery of an economically more rewarding vegetation type may be achieved through pro viding alternative sources of fuel and construction and through prohibiting cultivation and grazing in the shrublands on the hillsides. Regeneration can be accelerated by actively introducing seedlings of tree species that do not need a heavy canopy cover for establishment and growth.     

A new contribution to the Holocene vegetation history of the West African Sahel: pollen from Oursi,Burkina Faso and charcoal from three sites in northeast Nigeria

A pollen diagram from Oursi in Burkina Faso is compared with anthracological (charcoal analysis) results from three sites in northeast Nigeria (Konduga, Gajiganna, Lantewa). The present-day vegetation at all four sites is Sahelian or Sahelo-Sudanian and under heavy human impact. At Oursi, a closed grassland with only few trees and almost no Sudanian elements can be reconstructed for the middle Holocene. At the Nigerian sites, on the other hand, Sudanian woody plants were present during this period. We assume that the Sahel was not a uniform zone during the middle Holocene but rather a mosaic of different vegetation types according to local site conditions. In the light of these results, a simple model of latitudinally shifting vegetation zones is not applicable. Around 3000 B.P. the closed grassland at Oursi was opened by agro-pastoral activities, and at Gajiganna, plants characteristic of pasture lands can be directly related with the presence of cattle. Human impact seems to have been the dominant factor in the vegetation history of the Sahel from 3000 B.P. until today, masking possible effects of climatic change.     

Vietation changes over three decades on Santa Fe Island, Galhpagos, Ecuador

Changes in mortality and turnover of selected woody plants which dominate and characterise the arid vegetation of the island of Santa Fe, Galapagos, have been studied regularly during the period 1972–2003. After the eradication of feral goats from Santa Fe in 1971, the vegetation started to recover. The previously threatened small tree Scalesia helleri ssp. helleri recovered and some recruitment of the dominant trees Opuntia echios var. barringtonensis and Bursera graveolens took place. The dominant shrub species Cordia lutea, Encelia hispida and Lantana peduncularis increased in number, whereby the shrub layer is becoming denser and the structure of the predominant dry season deciduous steppe forest vegetation is changing. The strong impact of El Niño on the survival and recruitment of Opuntia indicates that recovery of Opuntia-dominated vegetation should be viewed on a time-scale of perhaps 150 years or more, in correspondence with the long life-expectancy of adult Opuntia individual. It is suggested that recruitment and survival of Opuntia may be hindered if future El Niño events are to occur with increased strength, and that this would alter the aspect of the Santa Fe vegetation.     

An experimental assessment of the upper elevational limit of flowering plants in the western Himalayas

Transplant experiments are a useful tool both for determining species’ ranges and understanding their cause, yet such experiments have seldom been performed in areas where plants reach extremely high elevations. We examined the position of the upper elevational limits of vascular plants in E Ladakh, India, by transplanting individuals of 14 subnival species from 5800–5850 m elevation to a control site at the same elevation, as well as to edaphically suitable sites at 5960 m (subnival belt with sparse plant cover), to 6030 m (the highest elevation reached by vascular plants in the area) and to 6160 m elevation (no vascular plants observed). Two years later, transplants of 13 species survived at the control elevation, whereas 5 species survived at 5960 m, 2 species (Waldheimia tridactylites, Poa attenuata) at 6030 m, and none at 6160 m. The highest elevation at which transplanted flowering plants survived corresponds well to the observed elevational limit. Soil temperature data at the sites suggest that the growing season, defined as the period with mean daily soil temperature above zero, lasted nearly 3 months at 5960 m, ca 1.5 month at 6030 m, but <3 weeks at 6160 m, moreover interrupted at the highest elevation by several days with temperatures remaining below zero. The experiment confirmed the observed sharp limit of vegetation, set not by any physical barrier per se (e.g. not by the top of the massif), but instead by physiological constraints of the species. The result provides support for the assumptions of mid‐domain effect models that domain limits are defined by shared organismal adaptations in relation to environmental gradients, in this case tolerance to freezing temperature.     

Species composition of coastal dune vegetation in Scotland has proved resistant to climate change over a third of a century

Climate change is expected to have an impact on plant communities as increased temperatures are expected to drive individual species' distributions polewards. The results of a revisitation study after c. 34 years of 89 coastal sites in Scotland, UK, were examined to assess the degree of shifts in species composition that could be accounted for by climate change. There was little evidence for either species retreat northwards or for plots to become more dominated by species with a more southern distribution. At a few sites where significant change occurred, the changes were accounted for by the invasion, or in one instance the removal, of woody species. Also, the vegetation types that showed the most sensitivity to change were all early successional types and changes were primarily the result of succession rather than climate‐driven changes. Dune vegetation appears resistant to climate change impacts on the vegetation, either as the vegetation is inherently resistant to change, management prevents increased dominance of more southerly species or because of dispersal limitation to geographically isolated sites.     

The ecological impact of oceanic island colonization – a palaeoecological perspective from the Azores

Aim In many cases, human colonization drastically modified the ecosystems of remote oceanic islands before scientists arrived to document the changes. Palaeoecological records before and after human colonization provide insights into the original ecosystems and an assessment of subsequent human impact. We used pollen analysis to compare the impact of 15th century colonization of the Azores with that of natural disturbances such as volcanic eruptions and climate changes. Location Azores archipelago, Atlantic Ocean. Methods Sediment records from three highland sites in the Azores (on the islands of Pico and Flores) were dated radiometrically and analysed palynologically. Pollen taxa were classified as native, endemic or introduced based on comparison with flora lists. Data were statistically zoned and temporal trends identified using detrended correspondence analysis. Results Human colonization of the Azores resulted in rapid, widespread, persistent vegetation changes on a scale unprecedented in the last 2700 years, detectable through the decline of dominant trees, the spread of grasses and fire‐tolerant species, the introduction of exotic plants, evidence for grazing and fire, and changes to soils and moisture availability. During the same period, volcanic eruptions appear to have had more localized impacts on the vegetation, lasting 500–1000 years and favouring endemic taxa. The effect of late Holocene climatic changes on the highland vegetation of the Azores seems to have been minor. Palaeoecological data indicate that at least two plant species went extinct on Pico after human colonization and that some plants regarded as introduced were almost certainly part of the original flora of the islands. Despite a consistent signal of human impact, compositional differences between Juniperus brevifolia communities on Pico and Flores remained after colonization. Main conclusions Human colonization had a greater impact on the pristine vegetation of Pico and Flores than climatic changes and volcanic activity during recent millennia. The similarity between post‐colonization changes on the Azores and other oceanic islands suggests a consistent pattern and scale to historical‐era human impact on otherwise pristine ecosystems. These characteristics could be used to further elaborate biogeographical theory and direct conservation efforts towards species that appear most susceptible to human activity.     

Remotely Monitoring Change in Vegetation Cover on the Montebello Islands,Western Australia,in Response to Introduced Rodent Eradication

The Montebello archipelago consists of 218 islands; 80 km from the north-west coast of Western Australia. Before 1912 the islands had a diverse terrestrial fauna. By 1952 several species were locally extinct. Between 1996 and 2011 rodents and cats were eradicated, and 5 mammal and 2 bird species were translocated to the islands. Monitoring of the broader terrestrial ecosystem over time has been limited. We used 20 dry-season Landsat images from 1988 to 2013 and estimation of green fraction cover in nadir photographs taken at 27 sites within the Montebello islands and six sites on Thevenard Island to assess change in vegetation density over time. Analysis of data averaged across the 26-year period suggests that 719 ha out of 2169 ha have increased in vegetation cover by up to 32%, 955 ha have remained stable and 0.6 ha have declined in vegetation cover. Over 492 ha (22%) had no vegetation cover at any time during the period analysed. Chronological clustering analysis identified two breakpoints in the average vegetation cover data occurring in 1997 and 2003, near the beginning and end of the rodent eradication activities. On many islands vegetation cover was declining prior to 1996 but increased after rodents were eradicated from the islands. Data for North West and Trimouille islands were analysed independently because of the potential confounding effect of native fauna being introduced to these islands. Mala (Lagorchestes hirsutus) and Shark Bay mice (Pseudomys fieldi) both appear to suppress native plant recruitment but not to the same degree as introduced rodents. Future research should assess whether the increase in vegetation cover on the Montebello islands is due to an increase in native or introduced plants.     

Spatial pattern, patch dynamics and successional change: chironomid assemblages in a Lake Erie coastal wetland

1. Distribution patterns in 1994 and 1995 of chironomid larvae in the sediments of wetlands on Presque Isle, Pennsylvania were used to determine whether relationships exist among species composition and wetland age, position or environmental characteristics. Canonical correspondence analysis was used to relate wetland age, degree of isolation, vegetation abundance and water chemistry to species composition.
2. Of forty-two chironomid taxa collected, Tanytarsus lugens group, Paratanytarsus sp. and Cladotanytarsus mancus group were the most widespread, but Lauterborniella agrayloides , Stictochironomus sp. and Cryptochironomus sp. were locally abundant in several sampling sites.
3. The canonical ordination based on all site characteristics was statistically significant and accounted for 54.9% of the variance in the species data and 69.5% of the variance in the species–environment relationship. The variables that contributed significantly to the ordination model were macrophyte abundance, conductivity and wetland area.
4. Partial canonical ordination based on environmental conditions with wetland age and position effects as covariables showed that environmental conditions explain a significant proportion of the variance in species composition among sites. Neither wetland age nor position variables explain a significant proportion of the variance in species composition.     

Spatial pattern, patch dynamics and successional change: chironomid assemblages in a Lake Erie coastal wetland

1. Distribution patterns in 1994 and 1995 of chironomid larvae in the sediments of wetlands on Presque Isle, Pennsylvania were used to determine whether relationships exist among species composition and wetland age, position or environmental characteristics. Canonical correspondence analysis was used to relate wetland age, degree of isolation, vegetation abundance and water chemistry to species composition.
2. Of forty-two chironomid taxa collected, Tanytarsus lugens group, Paratanytarsus sp. and Cladotanytarsus mancus group were the most widespread, but Lauterborniella agrayloides , Stictochironomus sp. and Cryptochironomus sp. were locally abundant in several sampling sites.
3. The canonical ordination based on all site characteristics was statistically significant and accounted for 54.9% of the variance in the species data and 69.5% of the variance in the species–environment relationship. The variables that contributed significantly to the ordination model were macrophyte abundance, conductivity and wetland area.
4. Partial canonical ordination based on environmental conditions with wetland age and position effects as covariables showed that environmental conditions explain a significant proportion of the variance in species composition among sites. Neither wetland age nor position variables explain a significant proportion of the variance in species composition.     

Cryptogams on decaying wood in old-growth forests of southern coastal British Columbia

Abstract. We studied the floristic composition of lignicolous cryptogams (i.e. bryophytes and lichens inhabiting decaying wood) in the old-growth coniferous forests of southern coastal British Columbia. The composition of the cryptogams was related to forest communities (described by vegetation units using forest floor cryptogams and vascular plants) and regional climates (described by biogeoclimatic zones). The study is based on a total of 247 sample plots, each of 0.04 ha in size. The plots were located in three different biogeoclimatic zones and were classified into 12 alliances and four orders in a previous study. We used indicator species analysis to determine cryptogam indicator species for each vegetation unit and biogeoclimatic zone, and used similarity analysis and multi-variate analyses (discriminant analysis and detrended correspondence analysis) to detect differences in the floristic composition of the cryptogams among sample plots, vegetation units, and biogeoclimatic zones. Most of the cryptogams in this study had a narrow distribution, and only < 5% of the species were present across all the vegetation units and biogeoclimatic zones. The overall means of Jaccard coefficients between two sample plots from the same vegetation unit (alliance or order) or biogeoclimatic zone were significantly higher (P < 0.05) than those from different vegetation units or biogeoclimatic zones. The difference in the mean Jaccard coefficients within- and between-units was highest for zones. The results of detrended correspondence analysis and discriminant analysis suggest that the composition of lignicolous cryptogams changes with the change in the floristic composition of forest floor vegetation. When the first axes of detrended correspondence analysis were compared, the Pearson's correlation coefficients between the first axes of lignicolous cryptogams and forest floor vascular plants and between the first axes of lignicolous cryptogams and forest floor cryptogams were 0.78 and 0.87, respectively. The degree of correspondence in the composition of lignicolous cryptogams and forest floor vegetation increased from alliance to order to biogeoclimatic zone. This trend suggests that the floristic composition of lignicolous cryptogams is influenced at the community level mainly by edaphic conditions and at the regional level by climatic conditions.