Diatom communities of acidic mountain streams in Poland

A comparison has been made of the species composition of diatom communities developing in acidic Polish mountain streams which flow over calcium-poor substrates: sandstones in the Silesian Beskid (section of the Western Carpathians), the witokrzyskie Mts, and over granite in the Karkonosze range (in the Sudetic Mts). The number of taxa and diversity of the diatom assemblages decreased along a decreasing pH gradient. The correlation between pH and the number of taxa was positive and significant (r ² = 0.69, p < 0.005). A small number of species (< 20) and low diversity were found in the communities developing in strongly acidic streams such as in the witokrzyskie Mts with pH 4.1–5.2, and in the Silesian Beskid with pH 3.5–4.0. In the stream of the Karkonosze Mts, with pH 5.2–6.0, the communities were characterized by their greater number of species and higher diversity.Acidobiontic and acidophilous diatoms were generally dominant. The pH-indiferent forms were less abundant, and their proportion increased above pH 5.0. Eunotia exigua, E. paludosa var. trinacria, E. tenella and Pinnularia subcapitata dominated in streams with the lowest pH, while E. exigua, E. sudetica and Achnanthes kryophila predominated in a stream with water pH above 5.2. Eunotia exigua, a common acidobiontic species was present in all the examined communities, and was a strong dominant in waters of pH 5.0. A corresponding decrease in abundance of E. exigua was observed with an increase in pH.     

Distribution of benthic diatoms in relation to environmental variables in lowland streams

The composition (% relative abundance) of diatom assemblages from soft bottom sediments was studied at 75 sites situated in 46 rivers, brooks, and ditches in the islands of Hiiumaa and Saaremaa and in the lowland of West Estonia. Although the total number of recorded taxa was 205, the studied diatom assemblages consisted of 54, 55, 48, and 50 constant species in the drainage basin of Moonsund, the Gulf of Riga, Hiiumaa, and Saaremaa, respectively. The habitats of the dominating taxa were heterogenous and the most widespread species were Achnanthidium minutissimum, Martyana martyi, Meridion circulare, Cocconeis placentula, Planothidium lanceolatum, and Amphora pediculus. The Shannon–Weaver diversity (H) index varied from 2.09 to 4.63. Multivariate analyses were used to identify the environmental variables governing the composition and structure of the benthic diatom assemblage. Principal Component Analysis (PCA) and Redundancy Analysis (RDA), based on 56 most abundant taxa, indicated differences in the composition and structure of diatom assemblages between different drainage basins as well as between the upper and lower stream courses. In the headwaters there prevailed small epipsammic diatoms (Martyana, Planothidium, Staurosira, Staurosirella). Different motile epipelic species from the genera Amphora, Navicula, Nitzschia, etc. were distributed abundantly in the lower courses of the streams. There was a positive correlation between order of the stream site and trophic level of water (R=0.35; p<0.05). Along a river system, the increasing order of the stream was accompanied by higher trophic level of water.     

Diatom communities along stream longitudinal gradients

1. Summer diatom communities on artificial substrates were sampled weekly for a month in three first- to sixth-order tributaries of the Kentucky River to determine how community structure varied with stream size. 2. Diatom cell abundances were generally higher in the headwaters. Species diversity increased in a downstream direction in two of the streams, and in an upstream direction in the third. However, diversity in general seemed more closely related to current regimes than to stream size per se, with highest species diversity at intermediate current velocities. 3. Variation in diatom accumulation rates was greater in downstream communities than in the headwater assemblages of two streams, suggesting that downstream communities may experience greater fluctuations in abundance, at least under low-flow conditions. 4. Patterns of species distributions suggested a relationship between morphological growth forms (guilds) and stream size, as well as the influence of current. Achnanthes spp., Eunotia spp., erect, and stalked taxa were more commonly associated with headwater assemblages. Filamentous and centric diatoms occurred with greater frequency downstream.     

Impact of Urban Pollution from the Hanoi Area on Benthic Diatom Communities Collected from the Red, Nhue and Tolich Rivers (Vietnam)

The effects of urban pollution from Hanoi city on the benthic diatom communities of the Nhue–Tolich river system were studied during the 2003 dry season. Benthic diatoms were allowed to grow on glass slides suspended in the water flow for 4 weeks. To reveal the relationship between water quality and diatom communities, Canonical correspondence analysis (CCA) was used on data concerning relative abundances of diatom species and environmental variables. Two diatom indices, IPS and DAIpo, were applied to evaluate water quality in the three rivers. A total of 291 diatom taxa were found in the Red, Nhue and Tolich Rivers. These were mainly cosmopolitan taxa, with some tropical, subtropical and endemic taxa. The most abundant taxa at the Red site were Aulacoseira granulata, Achnanthidium minutissimum, Encyonema minutum, Navicula recens and other halophilous taxa such as Nitzschia kurzii, Seminavis strigosa, Entomoneis paludosa, Bacillaria paradoxa. Diatom assemblages at the Tolich site consisted mainly of Nitzschia umbonata, Nitzschia palea and Eolimna minima. Diatom density ranged from 660 to 30,000 cells/cm². Environmental variables and diatom assemblage composition at all sites were significantly correlated. Two diatom indices gave similar results and indicate the Tolich River with the lowest values as a highly polluted site.     

Flood disturbance and riparian species diversity on the Colorado River Delta

We investigated the influence of channel migration and expansion on riparian plant species diversity along the lower Colorado River near the United States–Mexico border. Using repeat aerial photography in a GIS we identified and classed areas of low, moderate, and high disturbance frequency caused by channel expansion and migration. Replicate vegetation plots (12m×12m) were sampled in each of the three disturbance classes. One-way ANOVA was used to test for differences in species richness, species diversity (using the Shannon–Weiner Index) and overall percent ground cover of plants between the three disturbance classes. Regardless of disturbance class, plots were dominated by trees or shrubs, especially the non-native Tamarix ramosissima, as well as Pluchea sericea, Baccharis salicifolia and Salix goodingii. Clearly woody species constitute the great bulk of overall species richness, percent ground cover, and species diversity (H) in each disturbance group. No overall statistically significant differences were revealed among the disturbance groups for values of species richness, percent ground cover, or the Shannon–Wiener Index, though paired contrasts of means revealed that total percent ground cover on low disturbance plots was significantly higher than on moderately disturbed plots. Spatial and temporal variability in riparian diversity in the study area appears to hinge on factors other than disturbance frequency such as salt or drought stress. Alternately, our results could be interpreted as suggesting that in the presence of intensive flow regulation, disturbance plays a secondary role to ecological stresses, similar to that demonstrated by others. Intentional flood pulses are advocated as a restorative management strategy for improving plant productivity, management of exotic species (particularly T. ramosissima), and restoration of overall biodiversity.     

Effects of Canopy Cover and Ground Disturbance on Establishment of an Invasive Grass in an Australia Savanna1

Andropogon gayanus (gamba grass) is an introduced pasture grass that threatens Australia's tropical savannas by modifying fire regimes and species composition. To understand the establishment requirements of A. gayanus, we undertook a field experiment to determine the effect of canopy cover and ground layer disturbance on seedling emergence and survival. Seed was sown under three canopy treatments (undisturbed, artificial canopy gap, and natural canopy gap) and under three ground layer treatments (Control, Vegetation disturbed, and Soil disturbed). Results have shown that A. gayanus can establish and survive regardless of canopy cover or ground disturbance, although such site disturbances will increase establishment success. Disturbance of both the overstorey canopy and the ground layer increased A. gayanus emergence, whereas seedling survival to 12 mo after seed sowing was affected by ground layer disturbance alone. Disturbance of the canopy increased light transmission, which may have promoted germination. Ground layer disturbance may also have increased light transmission and suitable sites for establishment, and reduced competition for resources, such as water and nutrients. The ability of A. gayanus to spread along disturbed areas, establish in relatively undisturbed savannas, and resprout after fire within 6 mo after seedling emergence suggests that this species will become increasingly widespread in Australia's tropical savannas. Its control is urgently required.     

Canopy disturbance patterns and regeneration of Quercus species in two Ohio old-growth forests

This study was conducted to determine the abundance of Quercus species, the spatial pattern of Quercus regeneration, the current canopy disturbance pattern, and their interrelationship in two old-growth deciduous forests in Ohio (Goll Woods and Sears-Carmean Woods). Acer saccharum and Fagus grandifolia had the greatest density and basal area in both forests, yet the largest trees (by basal area) present at each site were Quercus spp. Quercus spp. appeared to be decreasing in abundance in both sites. Though Quercus seedlings were common, few Quercus saplings or subcanopy trees were present. The current disturbance regimes were dominated by small canopy gaps created by death of 1–2 trees; canopy gaps 100 m² in size were rare and only 2.5–2.8% of the forest area was covered by recognizable canopy gaps. No significant differences in the density of Quercus seedlings or saplings were found between gaps and non-gap areas at either site. Though no significant barrier to seedling establishment appeared to exist, the present disturbance regimes are not well suited for the growth of Quercus into the subcanopy size class or the recruitment of Quercus into the canopy. The most frequent gapmakers in Goll Woods were Tilia americana and Acer saccharum, and those in Sears-Carmean Woods were A. saccharum and F. grandifolia. The species most frequent as gap fillers were A. saccharum (in both sites) and F. grandifolia (in Goll Woods). These results suggest that A. saccharum will continue to increase in abundance, and Quercus decrease in abundance, in these two old-growth stands.     

Moss-inhabiting diatom communities from Heard Island,sub-Antarctic

In this paper, we list 192 diatom taxa, collected from bryophyte samples from Heard Island (52°05S, 73°30E). The Heard Island diatom flora shows a marked similarity to those of the Crozet and Kerguelen archipelagos, and is quite dissimilar to the moss-dwelling diatom flora of Macquarie and Amsterdam Island. Based on species composition, three main groups of samples could be discerned. Each group represents a different habitat. Samples dominated by Diadesmis ingeae, Pinnularia borealis and several Psammothidium species seem to prefer the driest habitats. A special form of this community was found in shaded areas. Samples from wet habitats such as flush areas and pools were dominated by species such as Adlafia bryophila, Sellaphora tumida, Fragilaria capucina and Planothidium lanceolatum. An intermediate group was observed in more acid, but terrestrial, conditions, characterised by Eunotia paludosa and Chamaepinnularia soehrensis var. muscicola. These community patterns are similar to those found on other sub-Antarctic islands, and we suggest that a group of typical sub-Antarctic species and communities exists.     

High tree alpha-diversity in Amazonian Ecuador

In a 1 ha square plot of terra firme forest at 260 m elevation in Amazonian Ecuador, all trees with diameter at breast height (dbh) 5 cm were studied. There were 1561 individuals, 473 species, 187 genera and 54 families. Of these, 693 individuals, 307 species, 138 genera and 46 families had a dbh 10 cm. This is the highest number of tree species ever recorded for a tropical rain forest sample of this size. In both dbh classes, the most species-rich families were: Fabaceae sensu lato (including Mimosaceae and Caesalpiniaceae), Lauraceae and Sapotaceae; the most species-rich genera, were Pouteria, Inga and Protium. The vertical space was partitioned among species: 166 species were found only in the 5–10 dbh cm class and were mostly sub-canopy treelets, and 307 species with dbh 10 cm were mostly large canopy trees.     

Aquatic pollution assessment based on attached diatom communities in the Pinang River Basin,Malaysia

The diatom communities collected from artificial substrates (glass slides) at 12 sampling sites along Pinang River Basin were analysed. Species composition was highly variable among sampling sites, and the highest diversity value (mean H = 3.56 bit/individual) and species richness (mean S = 17.32) was recorded at a polluted station in an estuary, which received salt-water intrusion during high tide. The downstream sampling stations were heavily organically polluted with Saprobic Index values ranging between 2.81 and 3.10, while upstream stations were relatively clean with Saprobic Index values recorded between 1.40 and 1.96. The diatom community structure and the specific sensitivity of certain diatom species can be related to the degree of water quality in Pinang River Basin. The abundance of certain diatom species could be used as biological indicators to measure impacts of river pollution.     

The composition and structure of the benthic diatom taxocene (Bacillariophyta) near Cape Fiolent (the Crimea,the Black Sea)

For the first time in the waters of the Cape Fiolent Reserve, a high species richness of benthic diatoms was revealed on natural substrates, 290 species and infraspecific taxa (IT), belonging to 281 species, 68 genera, 32 families, 19 orders, and 3 classes. A total of 68 species and 3 genera were previously registered as new for the flora of the Black Sea; 3 species and 1 taxonomic combination were previously described as new to science, as well as 4 species that did not occur in the Black sea in the past 50 and 100 years of research. Navicula (41 species and IT), Amphora (30), Nitzschia (29), Fallacia (20), Cocconeis (17) and Diploneis (16) were the most represented genera. Only one representative was identified in each of the orders Anaulales, Ardissoneales, Thalassionematales, Rhaphoneidales, and Rhopalodiales. The diversity assessment that was carried out using taxonomic distinctness indices, TaxDI, revealed that the structure of the phylogenetic diatom tree was formed by branches with various species saturation levels and different hierarchical subordination, but with the predominance of polyspecific taxa up to at genera level. The TaxDI values (Δ⁺ = 76.71; Λ⁺ = 361.52) indicate the low evenness and high variability of the structure of the benthic diatom taxocene near Cape Fiolent.     

The potential of agricultural headwater streams to retain soluble reactive phosphorus

Zooplankton may preferentially graze small, edible diatom species and therefore affect fossils relative to live assemblages by selective removal or increased sedimentation via egestion. Cladoceran zooplankton remains and diatom edibility were analyzed in sediment cores from Moon Lake and Coldwater Lake (North Dakota, USA) to assess changes in potential grazing pressure on algae and influence on diatom-inferred salinity (DIS) reconstructions. Sedimentary zooplankton in Moon Lake were dominated by littoral Cladocera, whereas Coldwater Lake assemblages were primarily small-bodied pelagic and littoral species. Relationships between cladocerans and environmental parameters over the past century varied by site and by species, with Chydorus brevilabris related most closely to drought at Moon and Bosmina sp. related to drought at Coldwater. A higher percentage of inedible diatoms occurred in the sediments of Moon Lake as compared to Coldwater Lake. DIS correlations with drought records improved in Moon Lake when only inedible diatom taxa were used to build a transfer function, but no improvement was seen for Coldwater Lake with this approach. These data suggest grazing pressure on diatoms differed between lakes and that zooplankton–phytoplankton interactions may affect the accuracy of drought reconstructions in the Great Plains.     

Assessment of Water Pollution using Diatom Community Structure and Species Distribution — A Case Study in a Tropical River Basin

A study on biological assessment of water pollution using diatom community structure and species distribution was carried out in the Linggi River Basin, Malaysia which was polluted by various urban, industrial and agricultural wastes. A total of 86 diatom taxa belonging to 21 genera were recorded from all eight sampling stations located in the basin, of which 70 species were found on artificial substrates; the remaining 16 species were recorded exclusively on natural substrates. The number of diatom species observed between the stations varied from 22-47 species. The dominant diatom species in decreasing order of abundance were Eunotia vanheurckii, Nitzschia palea, Achnanthes saxonica, Gomphonema parvulum and Achnanthes minutissima. The most common clean water species were Achnanthes minutissima, A. linearis and Synedra rumpens. The most tolerant species were Nitzschia palea followed by Gomphonema parvulum and Pinnularia braunii. Eunotia vanheurckii and Navicula cryptocephala occurred in high densities at both unpolluted and polluted stations and can be considered as the common facultative or indifferent species. Although a large number of species were recorded at the unpolluted stations, equivalent number of species were also found at the moderately polluted stations. However, the number of species was reduced at grossly polluted stations. Nevertheless, a marked variation in species association exists between the unpolluted and polluted stations, but not among the polluted stations to distinguish the type and degree of pollution.     

The radiation of spring snails of the genus Belgrandiella in Austria (Mollusca: Caenogastropoda: Hydrobiidae)

Grain density microautoradiography (MAR) was used to study uptake of dissolved amino acids (DFAA) in microalgal communities from a sandy bay on the west coast of Sweden. A mixture of fifteen ³H-labeled amino acids (final concentration 20–80 nmol l^–1) was added to sediment samples collected from two depths (0.5 and 4 m), on five occasions representing different seasons. On all sampling occasions, the microflora was dominated by diatoms (> 85% of the total biomass). Cyanobacteria occurred in the summer and autumn, but never dominated the biomass. Between 5 and 48% of all counted algal cells showed uptake (1–44% of algal biomass). Uptake was recorded for all majors microalgal groups (diatoms, cyanobacteria and autotrophic flagellates). Uptake was more frequent on the shallow site and the highest proportion of cells showing uptake occurred in May. Although uptake was common among both motile and attached growth forms, on several occasions uptake frequencies were higher for attached cells. As the attached fraction is frequently burried out of the photic zone, and not able to rapidly migrate towards light, this implies that heterotrophic capacity should be valuable. Also at species level, some taxa showed higher uptake frequencies than others, for example populations of motile diatom species that displayed a seasonal behaviour (blooming species), such as Nitzschia cf. dissipata and Cylindrotheca closterium. Other species showing frequent uptake were the coccoid colony-forming cyanobacterium Microcrocis sp., and the filamentous cyanobacterium Phormidium sp.. The results suggest that there is a high potential for DFAA uptake in the microphytobenthic community and that for some growth forms and species this could imply a competitive advantage.     

Mediterranean karst springs: diatom biodiversity hotspots under the pressure of hydrological fluctuation and nutrient enrichment

Abstract

Mediterranean karst springs are affected by strong climatic seasonality, with long, dry and hot summers and are increasingly threatened by anthropogenic pressures. In Sardinia (western Mediterranean Sea, Italy), they are largely unexplored and their biocoenoses are mostly unknown. The diatom flora from two substrates (cobbles and macrophytes) in eight springs of different areas of the island was investigated in summer 2016 and winter 2017. A total of 162 diatom taxa (58 genera) were found of which 27 (17 genera) only on cobbles and 26 (18 genera) only on macrophytes. The most abundant species from both substrates were Achnanthidium minutissimum, Amphora indistincta, Amphora pediculus, Cocconeis euglypta, Planothidium frequentissimum and P. lanceolatum. Overall, 67 taxa (40 genera) were recorded in single sites and some of these taxa showed high affinity with specific environmental conditions. Hydrological stability (water flow permanence), discharge and nutrients were the main environmental factors influencing diatom assemblages. Our results suggest that diatoms can reflect important local factors related to the vulnerability of these spring ecosystems and underline the importance of their preservation both for biodiversity and water quality maintenance.     

A functional relationship between species richness of spiders and lichens in spruce

Modern forestry has created stands with even age distribution of trees and fragmentation of the habitat. In boreal forests, the effects on biodiversity within many taxa need to be examined. We tested the hypothesis that species richness of foliose and fruticose lichens and spiders is positively related in the lower canopy of spruce (Picea abies) in forests with, or without, management in central Sweden. High species richness of lichens may increase the structural complexity of the microhabitat on spruce branches, and bring a higher species richness also in the spider community. In six areas, spruce branches were sampled in old-growth and managed boreal forest stands, respectively. Forest management did not affect the species richness of spiders or lichens, but an effect due to sampling area was found in the latter taxon. There was a significant covariation between species richness of lichens and spiders, and the hypothesised positive correlation was confirmed by separate analyses for each area and combining the probabilities. Moreover, regression analysis on mean values from each site revealed a positive relationship. We conclude that species richness of lichens and spiders covary on spruce branches for functional reasons, i.e. more lichen species promotes a more diverse spider community by increasing the structural heterogeneity. Our results might provide a shortcut for assessing biodiversity in boreal forests.     

Effects of fire on water and salinity relations of riparian woody taxa

Water and salinity relations were evaluated in recovering burned individuals of the dominant woody taxa from low-elevation riparian plant communities of the southwestern U.S. Soil elemental analyses indicated that concentrations of most nutrients increased following fire, contributing to a potential nutrient abundance but also elevated alluvium salinity. Boron, to which naturalized Tamarix ramosissima is tolerant, was also elevated in soils following fire. Lower moisture in the upper 30 cm of burned site soil profiles was attributed to shifts in evapotranspiration following fire. Higher leaf stomatal conductance occurred in all taxa on burned sites. This is apparently due to higher photosynthetic photon flux density at the midcanopy level and may be partially mitigated by reduced unit growth in resprouting burned individuals. Predawn water potentials varied little among sites, as was expected for plants exhibiting largely phreatophytic water uptake. Midday water potentials in recovering Salix gooddingii growing in the Colorado River floodplain reached levels which are considered stressful. Decreased hydraulic efficiency was also indicated for this species by examining transpiration-water potential regressions. Recovering, burned Tamarix and Tessaria sericea had enriched leaf tissue ¹³C relative to unburned controls. Higher water use efficiency following fire in these taxa may be attributed to halophytic adaptations, and to elevated foliar nitrogen in Tessaria. Consequently, mechanisms are proposed which would facilitate increased community dominance of Tamarix and Tessaria in association with fire. The theory that whole ecosystem processes are altered by invading species may thus be extended to include those processes related to disturbance.     

Testing the intermediate disturbance hypothesis on species diversity in herbaceous plant communities along a human trampling gradient using a 4-year experiment in an old-field

The intermediate disturbance hypothesis predicts unimodal relationships between species diversity and disturbance frequency/intensity. To test this hypothesis, species diversity in herbaceous plant communities along a human trampling gradient was investigated by conducting a 4-year experiment in an old-field. In general, species richness (S), the Shannon–Weiner index (H) from plant cover data and species evenness (J) showed negative linear relationships with trampling frequency, in contrast to the prediction of the intermediate disturbance hypothesis. However, the significant relationships between trampling frequency and species diversity were not observed in the fourth year without J, which showed a unimodal relationship. In all experimental years, the number of new species that colonized the plots after 1year was small under frequent trampling, and the number of species lost from the plots was large under infrequent trampling. The relative number and the relative cover of perennial species increased as trampling frequency increased in the first and second years, but this pattern was not observed in the following years because the dominance of perennials further increased at decreasing frequencies of trampling. The similarity in the species composition and the yearly changes in species dominance indicated that trampling at higher frequencies eliminated more trampling-intolerant species only in the early years of the experiment. These results suggest that trampling mediated early changes in species diversity patterns, but competitive interactions were more important in the later experimental years. The time lag in the effects of trampling and competition appears to be attributable to the infrequent occurrence of unimodal patterns of species diversity.     

Summer distribution of netphytoplankton in the Atlantic sector of the Southern Ocean

The surface distribution of netphytoplankton (>20 m) in the Atlantic sector of the Southern Ocean was investigated along two transects during early and late austral summer 1990/91. Sampling was under-taken at intervals of 60 of latitude between 34° and 70°S for the analysis of nutrients and for the identification and enumeration of netphytoplankton. Peaks in total diatom abundances were recorded at the Antarctic Polar Front (APF), in the vicinity of the South Sand wich Islands, in the marginal ice zone and in the neritic waters of the Atlantic sector of Antarctica. Cluster analysis indicates the existence of two major zones between Southern Africa and Antarctica. Diatom abundance increased south of the Antarctic Polar From along both transects, which can be partially explained by gradients of silicate concentration. Small chain-forming species (e.g. Fragilariopsis kerguelensis and Nitzschia lineata) dominated the diatom assemblages in early summer, while larger species, such as Rhizosolenia hebetata f. semispina and Corethron criophilum, dominated late summer diatom assemblages. The predominance of typically ice-associated forms in early summer suggests that the release of epontic cells during ice melt provides the initial inoculum for the netphytoplankton biomass. These small diatoms are subsequently replaced by larger species.     

Spatial distribution of canopy and subcanopy species along a sloping topography in a cool-temperate conifer-hardwood forest in the snowy region of Japan

The spatial distributions of canopy and subcanopy species (50cm stem length) were investigated within a plot extending from the top of a ridge to the valley bottom in a cool-temperate old-growth mixed forest, dominated by Cryptomeria japonica and Fagus crenata, in the snowy region of Japan. Based on the longitudinal profile of the slope, the study slope was divided into the relatively gentle upper slope position (US), the steep lower slope position (LS), the flat valley bottom (VB) and the boundary zone between the upper and lower slopes (BS). Spatial dispersal and the association patterns of species were analyzed in upperstory (10cm d.b.h) and understory (<10cm d.b.h) layers. Dominant species in the upperstory layer abruptly changed from Cryptomeria to Fagus at the BS site. In contrast, the understory trees of many species, including shade-intolerant and evergreen species, were independent of the location of conspecific upperstory trees or canopy gaps and extended their distributions on and around the BS site. Significant, diverse canopy and subcanopy species occurred at this site in both upperstory and understory layers. On the BS site, which is the lower margin of Cryptomeria-dominated vegetation, there were many medium-sized C.japonica that were killed by uprooting or breaking of the stems as a result of heavy snow pressure. It is suggested that the snow pressure gradient along a slope has a strong influence on community structure and the maintenance of diverse canopy and subcanopy species in this snowy mixed forest.