Bacterial diversity in the snow over Tibetan Plateau Glaciers

Bacterial diversity and cell abundance in the snow of the four glaciers (Guoqu, Zadang, East Rongbuk and Palong No. 4) located in different climatic zones of the Tibetan Plateau were investigated through culture-independent molecular analysis of 16S rRNA gene clone library and flow cytometry approaches. Cell abundance ranged from 0.68 × 10³ to 720 × 10³ cells mL⁻¹, with higher values in the northern glaciers than in the southern ones. Bacterial diversity was unexpectedly high in the snow habitats of the world’s highest plateau, with 15 common genera distributed widely among the glaciers. The bacterial diversity in the snow at different glaciers was related to the surrounding environments. The Guoqu Glacier, to the north near the desert zone and with the lowest temperature, preserved more bacteria closely related to a cold environment and soil than the other glaciers. However, in the Palong No. 4 Glacier located in the south warm region around vegetation, most bacteria were phylogenetically related to plant-associated bacteria.     

Prokaryotic diversity in sediments beneath two polar glaciers with contrasting organic carbon substrates

Microbial ecosystems beneath glaciers and ice sheets are thought to play an active role in regional and global carbon cycling. Subglacial sediments are assumed to be largely anoxic, and thus various pathways of organic carbon metabolism may occur here. We examine the abundance and diversity of prokaryotes in sediment beneath two glaciers (Lower Wright Glacier in Antarctica and Russell Glacier in Greenland) with different glaciation histories and thus with different organic carbon substrates. The total microbial abundance in the Lower Wright Glacier sediment, originating from young lacustrine sediment, was an order of magnitude higher (~8 × 10⁶ cells per gram of wet sediment) than in Russell Glacier sediment (~9 × 10⁵ cells g⁻¹) that is of Holocene-aged soil origin. 4% of the microbes from the Russell Glacier sediment and 0.04–0.35% from Lower Wright Glacier were culturable at 10°C. The Lower Wright Glacier subglacial community was dominated by Proteobacteria, followed by Firmicutes. The Russell Glacier library was much less diverse and also dominated by Proteobacteria. Low numbers and diversity of both Euryarchaeota and Crenarchaeota were found in both sediments. The identified clones were related to bacteria with both aerobic and anaerobic metabolisms, indicating the presence of both oxic and anoxic conditions in the sediments.     

Climate‐induced glacier and snow loss imperils alpine stream insects

Climate warming is causing rapid loss of glaciers and snowpack in mountainous regions worldwide. These changes are predicted to negatively impact the habitats of many range‐restricted species, particularly endemic, mountaintop species dependent on the unique thermal and hydrologic conditions found only in glacier‐fed and snow melt‐driven alpine streams. Although progress has been made, existing understanding of the status, distribution, and ecology of alpine aquatic species, particularly in North America, is lacking, thereby hindering conservation and management programs. Two aquatic insects – the meltwater stonefly (Lednia tumana) and the glacier stonefly (Zapada glacier) – were recently proposed for listing under the U.S. Endangered Species Act due to climate‐change‐induced habitat loss. Using a large dataset (272 streams, 482 total sites) with high‐resolution climate and habitat information, we describe the distribution, status, and key environmental features that limit L. tumana and Z. glacier across the northern Rocky Mountains. Lednia tumana was detected in 113 streams (175 sites) within Glacier National Park (GNP) and surrounding areas. The probability of L. tumana occurrence increased with cold stream temperatures and close proximity to glaciers and permanent snowfields. Similarly, densities of L. tumana declined with increasing distance from stream source. Zapada glacier was only detected in 10 streams (24 sites), six in GNP and four in mountain ranges up to ~600 km southwest. Our results show that both L. tumana and Z. glacier inhabit an extremely narrow distribution, restricted to short sections of cold, alpine streams often below glaciers predicted to disappear over the next two decades. Climate warming‐induced glacier and snow loss clearly imperils the persistence of L. tumana and Z. glacier throughout their ranges, highlighting the role of mountaintop aquatic invertebrates as sentinels of climate change in mid‐latitude regions.     

Glacial loss and its effect on riparian vegetation of alpine streams

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Comparison of Microbial Community Compositions of Two Subglacial Environments Reveals a Possible Role for Microbes in Chemical Weathering Processes

Viable microbes have been detected beneath several geographically distant glaciers underlain by different lithologies, but comparisons of their microbial communities have not previously been made. This study compared the microbial community compositions of samples from two glaciers overlying differing bedrock. Bulk meltwater chemistry indicates that sulfide oxidation and carbonate dissolution account for 90% of the solute flux from Bench Glacier, Alaska, whereas gypsum/anhydrite and carbonate dissolution accounts for the majority of the flux from John Evans Glacier, Ellesmere Island, Nunavut, Canada. The microbial communities were examined using two techniques: clone libraries and dot blot hybridization of 16S rRNA genes. Two hundred twenty-seven clones containing amplified 16S rRNA genes were prepared from subglacial samples, and the gene sequences were analyzed phylogenetically. Although some phylogenetic groups, including the Betaproteobacteria, were abundant in clone libraries from both glaciers, other well-represented groups were found at only one glacier. Group-specific oligonucleotide probes were developed for two phylogenetic clusters that were of particular interest because of their abundance or inferred biochemical capabilities. These probes were used in quantitative dot blot hybridization assays with a range of samples from the two glaciers. In addition to shared phyla at both glaciers, each glacier also harbored a subglacial microbial population that correlated with the observed aqueous geochemistry. These results are consistent with the hypothesis that microbial activity is an important contributor to the solute flux from glaciers.     

Hierarchy theory reveals multiscale predictors of Arkansas darter (Etheostoma cragini) abundance in a Great Plains riverscape

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Response of Glacier and Lake Dynamics in Four Inland Basins to Climate Change at the Transition Zone between the Karakorum And Himalayas

Inland glacier and lake dynamics on the Tibetan Plateau (TP) and its surroundings over recent decades are good indicators of climate change and have a significant impact on the local water supply and ecosystem. The glacier and lake changes in Karakoram are quite different from those of the Himalayas. The mechanisms of the complex and regionally heterogeneous behavior of the glacier and lake changes between the Karakorum and Himalayas are poorly understood. Based on satellite images and meteorological data of Shiquanhe, Hetian, and Yutian stations, we demonstrate that the overall retreat of glaciers and increase of lake area at the transition zone between the Karakoram and Himalayas (TKH) have occurred since 1968 in response to a significant global climate change. Glacial areas in the Songmuxi Co basin, Zepu Co basin, Mang Co basin and Unnamed Co decreased by -1.98 ± 0.02 km², -5.39 ± 0.02 km², -0.01 ± 0.02 km², and -0.12 ± 0.02 km² during the study period, corresponding to losses of -1.42%, -2.86%, -1.54%, and -1.57%, respectively. The lake area of the Songmuxi Co, Zepu Co, Mang Co and Unnamed Co increased by 7.57 ± 0.02 km², 8.53 ± 0.02 km², 1.35 ± 0.02 km², and 0.53±0.02 km², corresponding to growths of 30.22%, 7.55%, 11.39%, and 8.05%, respectively. Increases in temperature was the main reason for glacier retreat, whereas decreases in potential evapotranspiration of lakes, increases in precipitation, and increases in melt water from glaciers and frozen soil all contributed to lake area expansion.     

Abundance patterns of filter-feeding Caddisflies (Trichoptera : Hydropsychidae) and seston in a Montana (U.S.A.) lake outlet

The abundance patterns of four species of net-spinning caddisflies (Trichoptera : Hydropsychidae) were studied in a Montana (U.S.A.) lake outlet stream. Total Hydropsychidae reached very high densities (X > 1400 / 0.2 m²) near the lake outfall but declined precipitously downstream. Abundance patterns generally agree with a published model of filter-feeder / seston interactions in lake outlets. Larval densities were greater on moss-covered substrates and one species occupied habitats of lower water velocity compared to other species. Seston concentrations increased downstream contrary to expectation. Nutrient quality of seston may be important in predicting abundance of stream filter-feeders.     

Adult aquatic insects: Potential contributors to riparian food webs in Australia's wet–dry tropics

Abstract Changes in the abundance and biomass of aquatic and terrestrial aerial insects with distance (mid‐stream, 0, 10–15 and 160 m) from lowland streams were examined across the dry season landscape in Kakadu National Park, northern Australia. Malaise traps and sticky intercept traps were used to sample the insects at four streams, spaced over an area of 1650 km². Malaise and intercept catches were dominated by Diptera (flies and midges), both numerically and by biomass. Chironomid midges were the most abundant taxon, making up 43.4 and 51.0% of the malaise and intercept trap catches, respectively. However, most chironomids were small (less than 3 mm body length), contributing 34.9% to intercept trap biomass, but only 5.2% in malaise traps. Ceratopogonid midges and caddisflies (Trichoptera) accounted for most of the remaining adult aquatic insects. Major terrestrial components were Diptera and Hymenoptera in malaise traps and Coleoptera and Diptera in intercept traps. The total abundance and biomass of insects were much greater over streams and along the water's edge than in riparian (10–15 m) and savanna (160 m) habitats primarily because of the presence of large numbers of adult aquatic insects. The abundance and biomass of terrestrial insects in malaise traps showed no relationship with distance, but intercept trap catches suggested slightly greater abundances over the water and at the water's edge. The great abundance of aquatic insects relative to terrestrial insects close to streams suggests that they have the potential to be an important component of the diets of riparian insectivores, and predation may be an important pathway by which aquatic nutrients and energy are moved into terrestrial food webs.     

Comparison of microbial community compositions of two subglacial environments reveals a possible role for microbes in chemical weathering processes

Viable microbes have been detected beneath several geographically distant glaciers underlain by different lithologies, but comparisons of their microbial communities have not previously been made. This study compared the microbial community compositions of samples from two glaciers overlying differing bedrock. Bulk meltwater chemistry indicates that sulfide oxidation and carbonate dissolution account for 90% of the solute flux from Bench Glacier, Alaska, whereas gypsum/anhydrite and carbonate dissolution accounts for the majority of the flux from John Evans Glacier, Ellesmere Island, Nunavut, Canada. The microbial communities were examined using two techniques: clone libraries and dot blot hybridization of 16S rRNA genes. Two hundred twenty-seven clones containing amplified 16S rRNA genes were prepared from subglacial samples, and the gene sequences were analyzed phylogenetically. Although some phylogenetic groups, including the Betaproteobacteria, were abundant in clone libraries from both glaciers, other well-represented groups were found at only one glacier. Group-specific oligonucleotide probes were developed for two phylogenetic clusters that were of particular interest because of their abundance or inferred biochemical capabilities. These probes were used in quantitative dot blot hybridization assays with a range of samples from the two glaciers. In addition to shared phyla at both glaciers, each glacier also harbored a subglacial microbial population that correlated with the observed aqueous geochemistry. These results are consistent with the hypothesis that microbial activity is an important contributor to the solute flux from glaciers.     

Macrophyte beds contribute disproportionately to benthic invertebrate abundance and biomass in a sand plains stream

Macrophyte beds have been shown to influence organic matter retention and nutrient processing in streams. Less is known about the extent to which plant beds contribute to abundance, biomass, and diversity of macroinvertebrate assemblages in low-order streams. We measured aquatic invertebrate abundance, biomass, and diversity associated with plant beds and sand/gravel patches in a low-gradient second-order stream in the Central Sand Plains of Wisconsin, USA from March to October. Invertebrate abundance and biomass were higher on average in plant beds (2,552 m⁻² and 1,575 mg m⁻²) than in sand/gravel patches (893 m⁻² and 486 mg m⁻²). Although sand/gravel habitat was over three times more abundant than plant beds in the study reach, plant beds and sand/gravel patches contributed similarly to invertebrate abundance and biomass at the whole-reach scale. The abundance and biomass of invertebrates associated with plant beds decreased from spring to autumn. Non-insect invertebrates in the plant beds increased in relative abundance as the year progressed. Shannon–Weiner diversity and taxa richness of invertebrates were higher in the plant beds than in the sand/gravel habitat. Our results suggest that plant beds can represent hot spots for invertebrate abundance and production in low-gradient streams, and have implications for stream management and restoration in these types of ecosystems. Handling editor: S. I. Dodson     

Land use changes biomass and temporal patterns of insect cross-ecosystem flows

Emergent aquatic insects constitute an important food source for higher trophic levels, linking aquatic to terrestrial ecosystems. Little is known about how land use affects the biomass or composition of insect emergence. Previous studies are limited to individual time points or seasons, hampering understanding of annual biomass export patterns and detection of phenological changes. Over 1 year's primary emergence period, we continuously determined the biomass, abundance, and identity of >45,000 aquatic insects and recorded land-use-related environmental variables in 20 stream sites using a paired design with upstream forested sites and downstream agricultural sites. Total insect biomass and abundance were 2–7 mg day⁻¹ m⁻² and 7–36 ind day⁻¹ m⁻² higher in agricultural than forested sites. However, we found turnover of families between forested and agricultural sites, with more insects with shorter generation time in agriculture, indicating lower sensitivity to land-use-related stress because of higher recovery potential. Except for stoneflies, biomass and abundance of major orders were higher in agriculture, but their phenology differed. For different orders, emergence peaked 30 days earlier to 51 days later in agriculture than forest, whereas total abundance and biomass both peaked earlier in agriculture: 3–5 and 3–19 days, respectively. The most important land-use-related drivers were pesticide toxicity and electrical conductivity, which were differentially associated with different aquatic insect order abundances and biomass. Overall, we found that land use was related to changes in composition and phenology of aquatic insect emergence, which is likely to affect food-web dynamics in a cross-ecosystem context.     

Bacterial biodiversity from Roopkund Glacier, Himalayan mountain ranges, India

The bacterial diversity of two soil samples collected from the periphery of the Roopkund glacial lake and one soil sample from the surface of the Roopkund Glacier in the Himalayan ranges was determined by constructing three 16S rRNA gene clone libraries. The three clone libraries yielded a total of 798 clones belonging to 25 classes. Actinobacteria was the most predominant class (>10% of the clones) in the three libraries. In the library from the glacial soil, class Betaproteobacteria (24.2%) was the most predominant. The rarefaction analysis indicated coverage of 43.4 and 41.2% in the samples collected from the periphery of the lake thus indicating a limited bacterial diversity covered; at the same time, the coverage of 98.4% in the glacier sample indicated most of the diversity was covered. Further, the bacterial diversity in the Roopkund glacier soil was low, but was comparable with the bacterial diversity of a few other glaciers. The results of principal component analysis based on the 16S rRNA gene clone library data, percentages of OTUs and biogeochemical data revealed that the lake soil samples were different from the glacier soil sample and the biogeochemical properties affected the diversity of microbial communities in the soil samples.     

Real-time PCR assays targeting formyltetrahydrofolate synthetase gene to enumerate acetogens in natural and engineered environments

Acetogens are ubiquitous in many anaerobic habitats and play a very important role in bioconversion and biodegradation of organic compounds. Methods for rapid detection and quantification of acetogens in different environments are urgently needed to understand the in situ activities in complicated microbial communities. To overcome the limitations of culture-dependent methods and provide enhanced diagnostic tools for determination of the ecological roles of acetogens in different habitats, a quantitative real-time PCR (qrt-PCR) approach targeting functional FTHFS (fhs) gene encoding the formyltetrahydrofolate synthetase was developed. Novel primers flanking the FTHFS fragment were designed and tested. High specificity and sensitivity for estimation of the abundance of acetogens were confirmed analysis of a collection of acetogens, clone libraries and melting curves. The utility of the assay was validated and used in quantifying the FTHFS gene present in different anoxic and oxic habitats, including anoxic and oxic sludges, lake sediment, sewage sullage as well as flooded rice field soils. The abundance of FTHFS gene recovered by fhs1 assay was in the order of magnitude of 10⁵ up to 10⁷ copies per gram of dry weight sample, and the maximum calculated abundance of acetogens relative to Eubacteria was 0.6–0.9%, confirming the low proportion of acetogens to total bacteria in environments.     

Increasing donor ecosystem productivity decreases terrestrial consumer reliance on a stream resource subsidy

Because nutrient enrichment can increase ecosystem productivity, it may enhance resource flows to adjacent ecosystems as organisms cross ecosystem boundaries and subsidize predators in recipient ecosystems. Here, we quantified the biomass and abundance of aquatic emergence and terrestrial spiders in a reference and treatment stream that had been continuously enriched with nitrogen and phosphorus for 5 years. Because we previously showed that enrichment increased secondary production of stream consumers, we predicted that aquatic emergence flux would be higher in the treatment stream, subsequently increasing the biomass and abundance of terrestrial spiders. Those increases were predicted to be greatest for spiders specializing on aquatic emergence subsidies (e.g., Tetragnathidae). By adding a ¹⁵N stable isotope tracer to both streams, we also quantified nitrogen flow from the stream into the riparian community. Emergence biomass, but not abundance, was higher in the treatment stream. The average body size of emerging adult insects and the relative dominance of Trichoptera adults were also greater in the treatment stream. However, spider biomass did not differ between streams. Spiders also exhibited substantially lower reliance on aquatic emergence nitrogen in the treatment stream. This reduced reliance likely resulted from shifts in the body size distributions and community composition of insect emergence that may have altered predator consumption efficiency in the treatment stream. Despite nutrient enrichment approximately doubling stream productivity and associated cross-ecosystem resource flows, the response of terrestrial predators depended more on the resource subsidy’s characteristics that affected the predator’s ability to capitalize on such increases.     

Relationships between stream habitat and breeding territory length of the Brown Dipper (Cinclus pallasii) in Taiwan

Dippers from the genus Cinclus are highly specialised predators on aquatic invertebrates, and occupy linear territories along rivers where measurements of variations in quality are relatively straightforward. For these reasons, they are ideal model species in which to examine factors affecting territory size. In this paper, we investigated the influence of stream habitats on the territory length of the Brown Dipper (Cinclus pallasii) in Taiwan. The biomass of aquatic insects and other stream habitat variables were analyzed to determine their relationships with the territory length of Brown Dippers from November 1988 to May 1989. Compared with slow-moving waters, riffle areas contained significantly greater insect biomass (paired t test, t ₁₁ = 3.49, P < 0.01), of which trichopteran larvae contributed about 70%. Dippers spent more time foraging in riffles than in slow-moving waters. In addition, dippers preferred foraging in shallow riffles, but avoided deep, slow-moving waters (G = 62.53, df = 3, P < 0.001). Territory length (1,045 ± 165 m [SE], n = 14) was negatively related to proportion of riffles (r ² = 0.5715, P < 0.01), total aquatic insect biomass (r ² = 0.5840, P < 0.01), and altitude (r ² = 0.7176, P < 0.001). In factor analysis, four factors were extracted from the 14 stream variables. However, only factor 1 was significantly related to territory length (r ² = 0.5207, P < 0.01). Factor 1 explained 42.8% of the total variance and collectively revealed the importance of high food abundance. In other words, Brown Dipper territories were the shortest along high-altitude streams with abundant riffles, fewer pools and abundant aquatic insects. These results indicate that abundant supply of accessible invertebrate prey is the most important factor affecting the territory length of Brown Dippers. This is consistent with Cinclus species elsewhere, and reveals the importance of clean, productive river ecosystems.     

Effects of desiccation on the toxicant sensitivity of rotifers

Non-native aquatic macrophytes have invaded different types of ecosystems all over the world. The exotic submersed macrophyte Hydrilla verticillata recently invaded the Paraná basin, Brazil, being recorded by the first time in the natural habitats of this river in 2005. We investigated the effects of this species on ostracod assemblages and compared the abundance, richness, and Shannon–Wiener diversity of ostracod assemblages that colonize the invading species with those that colonize Egeria najas, a native submersed species with similar architecture and physical complexity. Fragments of these two species were left for 28 days in tanks to root and grow and then they were transferred to a floodplain lake where they remained in pairs (one plant of each species; N = 7) during 30 days for colonization by ostracods. A detrented correspondence analysis was used to summarize ostracod assemblage composition. Although there were no significant differences in ostracod abundance, richness and Shannon diversity when analyzed separately, cumulative curves, which permit to eliminate effects of abundance on richness, indicated a significantly higher number of ostracod species on H. verticillata. Assemblage composition was significantly different between both plant species, as shown by the first DCA axis. Our results show that H. verticillata might provide favorable habitats for native ostracod assemblages.     

The distribution of ciliates on Ecology Glacier (King George Island, Antarctica): relationships between species assemblages and environmental parameters

Ciliates are important consumers of pico- and nano-sized producers, are nutrient regenerators, and are an important food source for metazoans. To date, ecological research on ciliates has focused on marine ecosystems rather than on glacier habitats. This paper presents the first major study on ciliates from the Ecology Glacier (South Shetland Islands, Antarctica). The objective of the study was to investigate the structure and spatial distribution of ciliate communities and to identify the environmental factors determining the structure of the assemblages. Microbial communities were collected from three habitats: surface snow, cryoconite holes, and glacier streams. Sampling was carried out every 3–4 days from January 17 to February 24, 2012. A total of 18 ciliate taxa were identified. The species richness, abundance, and biomass of protozoa differed significantly between the stations studied with the lowest numbers in streams on the glacier surface and the highest numbers in cryoconite holes. The RDA performed to specify the direct relationships between the abundance of ciliate taxa and environmental variables showed obvious differences between studied habitats. The analysis showed that all variables together explained 62.4 % of total variance. However, variables that significantly explained the variance in ciliate communities in cryoconite holes, snow, and surface streams were temperature, conductivity, and total nitrogen. Further research is required to explain the impact of biotic factors influencing the presence of ciliates, including the abundance of bacteria, microalgae, and small Metazoa.     

Cryosphere: ice on Niwot Ridge and in the Green Lakes Valley,Colorado Front Range

Background: Subsurface ice preserved as ice lenses and within rock glaciers as well as glacial and lake ice provides sensitive indicators of climate change and serve as a late-season source of meltwater.

Aims: We synthesise the results of geomorphological, geophysical and geochemical studies during the period of 1995–2014, building on a long history of earlier work focused on ice and permafrost studies on Niwot Ridge and the adjacent Green Lakes Valley (GLV), which is part of the Niwot Ridge Long Term Ecological Research Site.

Methods: These studies are discussed in the context of how bodies of ice and rock glaciers reflect changing local climate. We review recent results from geophysical investigations (resistivity, seismic refraction and ground-penetrating radar) of the shallow subsurface, ongoing monitoring of the Arikaree Glacier, three rock glaciers and lake ice in the GLV, and interpretations of how subsurface ice melt regulates the flow and chemistry of alpine surface water after seasonal snowfields melt.

Results and conclusions: Permafrost conditions reported from Niwot Ridge in the 1970s are generally absent today, but ice lenses form and melt seasonally. Ice is present permanently within the Green Lakes 5 rock glacier and at nearby favourable sites. The Arikaree Glacier has shown a marked decline in cumulative mass balance during the past 12 years after a 30-year period when net mass balance was ca. 0. Duration of seasonal lake ice increases with elevation in GLV, but duration has decreased at all seven lakes that have been monitored during the last three decades. This decrease has been most marked at the lowest elevation where it amounted to a reduction of about 1 d year^?1 and least at Green Lake 5 where the loss has been at a rate of 0.5 d year^?1. Surface temperature measurements from rock glaciers have not shown strong trends during the past 15 years. It has been suggested that almost all of the 2.5-mm year^?1 increase in stream discharge from the upper GLV in September and October has been derived from melting of subsurface ice.     

Predicting the distribution of invasive crayfish (Pacifastacus leniusculus) in a Kusiro Moor marsh (Japan) using classification and regression trees

Summer distributions of the invasive signal crayfish (Pacifastacus leniusculus) were investigated in relation to physicochemistry in a Kusiro Moor marsh and its inflows and outflows in northern Japan. Maximum crayfish abundance and biomass were 1.04 individuals/m² and 3.56 g dry mass (DM)/m² in littoral marsh habitats, and 5.84 individuals/m² and 13.48 g DM/m² in stream habitats. Classification tree analysis was used to predict crayfish occurrence at 102 sites from all habitats (i.e. littoral marsh, pelagic marsh and stream) while regression tree analyses were used to predict crayfish abundance at littoral marsh and stream sites separately. The classification tree showed that crayfish occurrence was primarily determined by undercut bank volume regardless of habitat identity. When undercut bank volume was <0.0054 m³, crayfish were predicted to be absent at marsh sites, but expected to occur at stream sites where pH and water temperature exceeded 6.5 and 14.3°C, respectively. The regression tree using only littoral marsh sites showed that undercut bank volume, followed by dissolved oxygen level, determined the splits of the tree. Crayfish abundance was highest when undercut bank volume was >0.61 m³, and moderately high when dissolved oxygen was >9.09 mg/l and undercut bank volume was <0.61 m³. On the other hand, the regression tree using only stream sites showed that water temperature was the major predictor that determined the splits. We discuss the roles of physicochemical factors as limiting factors of the distribution pattern of the invasive crayfish.