Species composition and habitat associations of benthic algal assemblages in headwater streams of the Sierra Nevada, California

Despite their trophic importance and potential importance as bioindicators of stream condition, benthic algae have not been well studied in California. In particular there are few studies from small streams in the Sierra Nevada. The objective of this study was to determine the standing crop of chlorophyll- a and benthic algal species assemblages present in the small 1st- and 2nd-order streams of the Kings River Experimental Watersheds (KREW, watersheds of Bull, Providence, Duff, and Teakettle Creeks) and determine the associations of these measures with stream habitat. We collected samples of benthic algae from rock substrata in September 2002 (7 sites) and 2005 (the same 7 sites plus 5 additional sites). Habitat and water-quality data were collected concurrently. Chlorophyll- a values ranged from 0.2 to 3.2 mg ? m –2 . Chlorophyll- a in the Bull Creek watershed was generally lower than in the other watersheds. Benthic algal assemblages were dominated by diatoms and cyanobacteria. We collected 79 taxa of diatoms in 2002 and 126 taxa in 2005. Diatom taxa richness in individual samples ranged from 15 to 47. Nonmetric multidimensional scaling analysis of arcsine square-root transformed proportional abundances of diatoms identified 3 groups of sites. Bull Creek sites were generally different from other sites (group 1), and the sites from Bull Creek were different in 2002 (group 2) and 2005 (group 3). Five taxa appeared to be particularly important in distinguishing groups: Achnanthidium minutissimum , Cocconeis placentula , Eunotia incise , Eunotia pectinalis var. minor , and Planothidium lanceolatum . Elevation, water temperature, pH, specific conductance, and canopy were habitat variables correlated with the differences in diatom assemblages among sites. Our results provide a valuable baseline for future studies of benthic algae in Sierra Nevada headwater streams and will be particularly important in understanding the effects of different forest restoration management strategies being tested in the KREW project.     

Diatom flora of Mink Creek, Idaho, USA

Diatoms were collected from an open-canopy and closed-canopy site on Mink Creek, Bannock County, Idaho, a third-order Rocky Mountain stream. Ninety diatom taxa were identified. Achnanthes minutissima Kuetz. And Navicula lanceolata (Ag. ) Kuetz. dominated the open-canopy site, whereas Cocconeis placentula var. euglypta (Ehr.)CI. comprised greater than 40% of the diatom assemblage at the closed-canopy site. Seven of the 10 most important diatoms were present at both sites. A high degree of similarity was evident between natural and artificial substrates at both sites. Although most taxa were present at both sites, it is apparent from this study that differences in the abundance of taxa occur according to variations in light.     

Classification of the riparian vegetation along a 6-km reach of the Animas River, southwestern Colorado

Riparian ecosystems are important components of landscapes, particularly because of their role in biodiversity. A first step in using a ""coarse-filter"" approach to riparian biodiversity conservation is to determine the kinds of riparian ecosystems. These ecosystems vary substantially in plant species composition along a single river reach, as well as between rivers, and yet the river-reach scale has received little attention. We sampled the vascular plant composition of 67 contiguous patches of riparian vegetation along the reach of the Animas River, in southwestern Colorado's San Juan Mountains, that is relatively undisturbed by human land uses. Using cluster analysis and detrended correspondence analysis, we identified eight riparian community types along the reach. Using a new technique, we combined overstory size-class data and understory cover data to identify community types. The eight community types, which are in part the products of past floods, are spatially arranged along the reach in relation to variation in valley morphology, tributary location, and geomorphic landforms. These eight community types do not necessarily represent successional stages of a single potential vegetation type. This study at the river-reach scale suggests that sampling and analysis, as well as conservation, may need to be turned to the scale of patchiness produced by flood disturbances in the riverine landscape, since vegetation varies significantly at this scale.     

Assessment of cryptobiotic crust recovery

Chlorophyll a and b analyses of cryptobiotic crusts from burned and unburned sites on the Snake River plain provided an estimate of crust recovery. In less than 3 years, chlorophyll a levels were approximately 50% of the unburned reference site. Chlorophyll a/b ratios indicated that the biotic composition of the recovering cryptobiotic crust consisted largely of eukaryotic algae and bryophytes.     

Plecoptera and Trichoptera species distribution related to environmental characteristics of the metal-polluted Arkansas River, Colorado

The Upper Arkansas catchment has been polluted with heavy metals from mining for almost 140 yr. Adult Plecoptera and Trichoptera species distributions were recorded from 22 stations along 259 km of main river during 1984-85 so that these could be related to metal deposition and other environmental characteristics. Chemically or physically perturbed sites had poor species richness compared with adjacent sites. There was no sequential downstream increase in species numbers. Filter-feeders proportionally increased downstream as predators declined; these proportions were reset at a high-energy site before the trend resumed. Using canonical correspondence analysis, we found that species composition was most strongly related to changes in distance/altitude and to temperature, particularly after regulatory flows entered the river. The proportion of biological variation explained by river measurements indicated that collected adults were largely derived from the main Arkansas River. Species tolerant of high sedimentary metal concentrations were identified while some other species appeared to be sensitive. The study provides a disturbed-state reference for monitoring effects of remedial actions begun in 1991, and for comparisons with other Colorado rivers.     

Records of the yellow-nosed cotton rat ( Sigmodon ochrognathus ) in southwestern New Mexico

The yellow-nosed cotton rat ( Sigmodon ochrognathus ) previously was known only from extreme southwestern New Mexico in the Peloncillo and Animas mountains. Here I document S. ochrognathus at 3 localities from the Big Burro and Mogollon mountains in southwestern New Mexico. My northernmost capture represents an extra-limital record of 160 km north of prior records in southern New Mexico and represents the northernmost occurrence in North America. Lack of intensive mammalian surveys in the region render it difficult to interpret my recent captures, but the most parsimonious explanation is that captures likely represent overlooked populations. I captured S. ochrognathus in grassy areas ungrazed by domestic livestock along the Gila River. Understanding the use of limited grassland habitats along arid rivers by terrestrial vertebrates in the region is important because many of these grasslands have been converted to agricultural lands or are grazed by livestock.     

Parasites of the woundfin minnow, Plagopterus argentissimus , and other endemic fishes from the Virgin River, Utah

Two hundred woundfin minnows, Plagopterus argentissimus , from four sites along the Virgin River, Utah, were examined on two dates during summer 1985. The foreguts of 211 woundfin and variable numbers of other fishes from the Virgin River near Beaver Dam Wash, Arizona, and Mesquite, Nevada, were examined for cestodes on four dates throughout 1979. Seven parasites were found in P. argentissimus: Posthodiplostomum minimum (metacercariae), Diplostomum spathaceum (metacercariae), Bothriocephalus acheilognathi, Gyrodactylus sp., Lernaea cyprinacea, Trichodina sp., and Ichthyophythirius multifiliis . Fungal infections were noted on two fish during the study. Seventeen Virgin River roundtail chub, Gila robusta seminuda, were examined from two of the four sites in 1985 and 64 specimens from Beaver Dam Wash were examined in 1979. Gila robusta seminuda was infected with Posthodiplostomum minimum (metacercariae) and Bothriocephalus acheilognathi, the Asian fish tapeworm. This cestode probably gained entrance into the ichthyofauna of the Virgin River from red shiners, Notropis lutrensis, and has the potential of being very detrimental to the endemic and endangered fishes of the Virgin River. Parasite loads were correlated with water quality and habitat disturbance, with highest number and frequency occurring in "disturbed" sites. Low river flows and increased total dissolved solids appear to be associated with a higher parasite frequency and mean number in fishes of the Virgin River. These data represent the first known published records for parasites of the woundfin minnow and Virgin River roundtail chub.     

Effects of grazing exclusions on rangeland vegetation and soils, east central Idaho

Nineteen exclosures on sagebrush steppe and shadscale rangelands, varying in age from 18 to 38 years, were sampled for plant species richness, plant composition, indicators of soil erosion, ground cover, vegetative cover, and herb-low shrub layer screening cover. Features within the exclosures were compared with adjacent sites of the same size that were open to grazing by livestock and wildlife. Species richness typically was slightly greater inside exclosures than in adjacent grazed sites (median = 2 more species inside exclosures), but the difference was not significant ( P = 0.16). Similarity of plant community composition between exclosures and adjacent grazed sites ranged from 45% to 82%. Evidences of soil movement, soil pedestals, and soil flow patterns were all more pronounced outside exclosures than inside ( P ≤ 0.02), even though many sites were on flat to mild slopes (median slope = 12%). Meta-analysis of the 19 exclosure sites indicated that grazing exclusion resulted in less bare ground cover compared with adjacent grazed sites ( P ≤ 0.05). The effect of grazing exclusion on visible soil surface cryptogams was significant ( P ≤ 0.05), with generally greater cover inside exclosures. Cryptogam cover differences between grazed sites and exclosures tended to increase with the number of years of grazing exclusion ( r = 0.64, P = 0.046). Pseudoroegneria spicata , a principal livestock forage, averaged greater basal cover inside exclosures than outside on 4 of 10 sites where it occurred, although no exclosure sites had greater P. spicata cover on grazed sites. Meta-analysis of the 10 sites indicated that grazing exclusion resulted in greater P. spicata cover compared with adjacent grazed areas ( P ≤ 0.05). Poa secunda , a short-growing grass that initiates growth early in the spring and is not important livestock forage, averaged greater basal cover outside exclosures on 5 of 15 sites where it occurred. Meta-analysis of the 15 sites indicated a significant treatment effect ( P ≤ 0.05), with greater Poa secunda basal cover outside exclosures. Grazing exclusion resulted in greater screening cover in the herb-low shrub layer (0-0.5 m height; P ≤ 0.05). These results indicate that despite improved livestock grazing management over the past half century, livestock grazing still can limit the potential of native plant communities in sagebrush steppe ecosystems, and that the health of semiarid ecosystems can improve with livestock exclusion in the absence of other disturbances. A few exclosure sites were similar for the measured parameters, suggesting that these sites were ecologically stable and that exclusion of livestock grazing was not sufficient to move succession toward more pristine conditions, at least within the time periods studied. Managed disturbance such as fire or mechanical brush treatments may be necessary to restore herb productivity on these ecologically stable sites.     

Distribution, abundance, and genetic population structure of Wood River sculpin, Cottus leiopomus

The Wood River sculpin Cottus leiopomus is endemic to the Wood River Basin in central Idaho and is a nongame species of concern because of its limited distribution. However, status and genetic population structure, 2 factors often central to the conservation and management of species of concern, have not been assessed for this species. We used backpack electrofishers to survey streams that were small enough (i.e., a priori to contain, possibly contain, or not contain the sculpin, respectively. Native redband trout Oncorhynchus mykiss gairdneri were present at 21 study sites, including 18 of the 20 sites that contained Wood River sculpin. Sixty-one study sites (60%) were dry or had too little water to contain any fish. We estimated that a minimum of 1.36 million Wood River sculpin (≥20 mm total length) currently reside in the basin. The presence of Wood River sculpin was positively associated with stream width:depth ratio and percent cobble/boulder substrate and negatively associated with stream gradient. Mitochondrial DNA haplotype differences were observed between and within the 3 major river subbasins supporting sculpin, with the most striking differences observed between populations in the Camas Creek subbasin and the other 2 subbasins, among which no haplotypes were shared, suggesting relatively long-term isolation. Our results suggest that the Wood River sculpin remains relatively widespread and abundant within its endemic range, despite obvious changes in historical stream connectivity caused by irrigation diversions and other chronic habitat alterations.     

Diatoms of the Middle Fork of the Salmon River drainage, with notes on their relative abundance and distribution

A total of 145 species of diatoms was collected from 13 sites in the Middle Fork of the Salmon River drainage, Idaho, USA. Achnanthes minutissima was the prevalent species with an importance index of 19.25. Cluster analysis revealed 2 main site groupings, sites above the entrance of Loon Creek and sites below. The entrance of Loon Creek (6th order) increases the Middle Fork of the Salmon River to a 7th order stream. Shannon-Weiner diversity values were generally high.        

Home-range fidelity and use of historic habitat by adult Colorado pikeminnow ( Ptychocheilus lucius ) in the White River, Colorado and Utah

Twelve wild adult Colorado pikeminnow ( Ptychocheilus lucius ), captured in the tailwaters of Taylor Draw Dam on the White River, Colorado, were implanted with radio transmitters and their movement patterns monitored from 1992 to 1994. The spawning migration of these fish was extensive. In 1993, the only full year of the study, the fish migrated an average of 658 km from the White River to spawning sites in the Yampa or Green rivers and back to the White River. Eight of these fish were translocated in the river upstream of the dam in April 1993. These fish and the 4 others below the dam remained in the river until May 1993. All 12 had migrated down the White River to spawning sites in the Green and Yampa rivers by July 1993. The fish that were located above the dam successfully passed over the dam during their downstream migration. Seven fish migrated upstream toward the Yampa River Canyon spawning site and 5 migrated downstream toward the Green River Desolation/Gray Canyon spawning site. Five of 7 Yampa River fish were found at the spawning site. The other 2 were found 5-8 km downstream of the site. One of 5 Green River fish was found at the spawning site, the other 4 between 16 and 62 km upstream of the site. All fish migrated back to the White River by August 1993 and were found near the dam by October 1993. Two fish were recaptured and translocated above the dam in September 1993. Five fish were located below the dam and 2 above the dam in April 1994. By July 1994 seven of the same fish that had migrated toward the Yampa River in 1993 were found at the Yampa Canyon spawning site. At the same time, 3 of 5 fish that migrated toward the Green River in 1993 were found at the Desolation/Gray Canyon spawning site. This included 2 fish that had been found upstream of the site in 1993. The 12 fish traveled an average of 6 km d -1 (range: 4-10 km d -1 ) during the migration period from May through October 1993. Generally, fish moved faster to the spawning site than back from the site to the White River. These fish moved very little within their home ranges in the White River. Six fish tagged in 1992 moved only 0.1-2.3 km in the tailwater reach below Taylor Draw Dam from September 1992 through April 1993. All fish, after their spawning runs, had moved up to or near the dam by October 1993. These fish were not tracked again until April 1994. Their movement patterns in April 1994 were similar to those observed in April 1993. The greatest amount of fish movement in the White River was displayed by the 8 fish placed above Taylor Draw Dam in April 1993 and the 2 placed in Kenney Reservoir in September 1993. They moved 1.1-40.6 km in the river before and after their spawning migration in spring and autumn 1993. These spawning migrations suggest that adult Colorado pikeminnow in the White River were recruited from both Green and Yampa river spawning populations and were presumably imprinted to these respective spawning sites. Those fish placed above Taylor Draw Dam established home ranges in habitats previously occupied by Colorado pikeminnow before the dam was completed. They remained there until they migrated downstream during the spawning period. Although we did not study fish passage, our study demonstrates that adult Colorado pikeminnow will use habitat if access is provided. Translocation of wild adult fish into historic but unoccupied habitats may be a valuable recovery option.     

Structure of alpine plant communities near King's Peak, Uinta Mountains, Utah

A study was made at 18 sites with elevations between 3512 and 3768 m in the Uinta Mountains, Utah. Sites were small in extent but typified vegetation patterns found in the Uintas. Standing crop, species composition (based on dry weight), and values for several physical parameters were determined at each site. Simple linear regressions performed between the various biotic and abiotic characters revealed significant relationships between the characteristics of rocks visible at the surface (the number, size, and variation in size) and vegetation cover. This relationship was probably due to the burial of rocks as a region became vegetated. Bray and Curtis ordinations performed on the data indicated that there were several factors which influenced the species composition but that no single factor dictated the vegetational pattern.     

Aquatic vertebrate assemblages of the Upper Clear Creek Watershed, California

We sampled streams in the Upper Clear Creek Watershed in northwestern California in fall 2004 and fall 2005 to document assemblages of aquatic vertebrates and to provide resource managers with information on the importance of these assemblages in terms of regional biodiversity. We used single-pass backpack electrofishing to sample 15 sites in fall 2004 and the same 15 sites plus 4 new sites in fall 2005. We captured 10 fish taxa and 2 species of larval amphibians. Seven of the fish taxa were native species. Of the exotic species, only brook trout ( Salvelinus fontinalis ) occurred at more than 1 site. Ordinations by nonmetric multidimensional scaling indicated a gradient from sites with rainbow trout ( Oncorhynchus mykiss ), Pacific giant salamander ( Dicamptodon tenebrosus ), and tailed frog ( Ascaphus truei ) to sites dominated by riffle sculpin ( Cottus gulosus ), California roach ( Hesperoleucas symmetricus ), and Sacramento sucker ( Catostomus occidentalis ). The gradient in species composition was associated with changes in elevation, gradient, discharge, and substrate. The Upper Clear Creek Watershed represents a unique area of overlap between the North Coast California amphibian fauna and the Central Valley fish fauna with a notable paucity of exotic fishes and amphibians. Preservation of the integrity of native aquatic assemblages is an important goal for aquatic resource management in the region; our results provide a critcial baseline to gauge future management actions.     

Population density, biomass, and age-class structure of an invasive clam Corbicula fluminea in rivers of the lower San Joaquin River watershed

Corbicula fluminea is well known as an invasive filter-feeding freshwater bivalve with a variety of effects on ecosystem processes. However, C. fluminea has been relatively unstudied in the rivers of the western United States. In June 2003, we sampled C. fluminea at 16 sites in the San Joaquin River watershed of California, which was invaded by C. fluminea in the 1940s. Corbicula fluminea was common in 2 tributaries to the San Joaquin River, reaching densities of 200 clams ? m –2 , but was rare in the San Joaquin River. Biomass followed a similar pattern. Clams of the same age were shorter in the San Joaquin River than in the tributaries. Distribution of clams was different in the 2 tributaries, but the causes of the difference are unknown. The low density and biomass of clams in the San Joaquin River was likely due to stressful habitat or to water quality, because food was abundant. The success of C. fluminea invasions and subsequent effects on trophic processes likely depends on multiple factors. As C. fluminea continues to expand its range around the world, questions regarding invasion success and effects on ecosystems will become important in a wide array of environmental settings.     

Aquatic invertebrate fauna of a northern prairie stream: range extensions and water quality characteristics

The benthic macroinvertebrate fauna of southern Saskatchewan, Canada, has received little attention relative to other regions of western North America. Therefore, little is known of the related aquatic ecosystem health and biogeography of regional aquatic insects. Here we present the results of an aquatic macroinvertebrate survey for the Pipestone Creek watershed in southeastern Saskatchewan. We qualitatively sampled aquatic macroinvertebrates in 5 sites on 4 dates through spring, summer, and fall 2006. Sampling produced 294 taxa of macroinvertebrates including 25 provincial range extensions to the southeast corner of the province. Presence/absence data of taxa grouped the sites into lentic and lotic sites. However, the relative proportions of the taxa varied greatly among sites, with no 2 sites having a community similarity greater than 50%. Functional feeding group analyses separated the sites into collector-dominated and scraper/grazer-dominated sites. However, the taxonomic make-up of the feeding groups varied among sites and also among dates. A modified Hilsenhoff Biotic Index of the site communities indicated that all were influenced by organic pollution. Results of this study suggest that although the watershed is enormously diverse, its biological communities are likely influenced by organic pollution. Further, range expansions of species found here, such as the stonefly Perlesta placida , have implications for invasion pathways and post-glaciation species islands in a prairie landscape.     

Zoogeographic affinities of the stoneflies (Plecoptera) of the Raft River Mountains, Utah

We examined faunal affinities of the Raft River Mountains using stoneflies (Plecoptera) as indicators. This island-like mountain range is isolated from other major mountain ranges in the Intermountain West by low-elevation, arid regions. Thirty-seven species were recorded from collections from 19 sites in the Raft River Mountains. Cluster analysis demonstrated the Raft River Mountain stonefly assemblage to be most similar to faunas of the Sawtooth and Wasatch mountains, and quite different from that of the Sierra Nevada. An analysis of the distribution patterns of each species, on a family-by-family basis, showed that the Raft River Mountains fauna consists mostly of species widespread in western North America. Most families were represented by at least 1 species whose distribution supports faunal affinities with regions to the north and west. Logistic regression of 6 long-distance dispersal factors against stonefly presence-absence data did not support long-distance dispersal as a viable means of colonization for the Raft River Mountains. This suggest that stonefly distribution patterns may be attributed to expansion and subsequent vicariance of suitable stonefly habitats during Pleistocene climatic oscillations.     

Site characteristics and habitat requirements of the endangered dwarf bear-claw poppy ( Arctomecon humilis Coville, Papaveraceae)

Arctomecon humilis Cov. is a narrow endemic, confined to gypsiferous substrates derived from the Shnabkaib Member of the Moenkopi Formation in southwestern Utah. The characteristics of seven A. humilis sites were studied to define habitat requirements of the species. Both physical and biotic aspects of the habitat were analyzed; geology, soil chemistry, and physical properties, as well as vascular and nonvascular plant communities, were studied. Chemical and physical properties of these soils vary considerably from those supporting adjacent desert-shrub communities. The dominant vascular species on A. humilis sites are shadscale and Mormon tea. It was found that A. humilis occurred in portions of the habitat where shrubs were relatively less dense. A soil-surface, cryptogamic community contributed 84% or more of the total living cover on sites that supported A. humilis . Species composition of the cryptogamic community was highly similar among sites. Likewise, composition of the cryptogamic cover was similar when random samples were compared with samples centered on A. humilis plants. The Purgatory Flat site, which does not support the poppy, seems inseparable from A. humilis sites, in respect to soil characteristics and composition of the associated plant cover. There is reason to believe an A. humilis population could be established at the Purgatory Flat site.     

Cryptogamic soil crusts: recovery from grazing near Camp Floyd State Park, Utah, USA

The effects of grazing on the cryptogamic and vascular plant communities at two sites near Camp Floyd State Park, Utah County, Utah, were studied. The grazed site was subject to heavy grazing up until seven years prior to the study. The ungrazed site within the park boundaries had been protected from grazing for 20 years prior to the study and had a well-developed algal-lichen-moss crust. We found that the algae of the grazed site had recovered in terms of degree of crusting. There were no significant differences in the algal communities of the two sites when prevalent species were used as blocks in the ANOVAR analysis. However, when major algal groups were used as blocks, the analysis was significant, with the more recently grazed site having lower algal frequency. This difference, together with a few compositional differences, indicates that, although the algal community seven years following grazing is very similar to the algal community free of grazing for 20 years, the seven-year site is still in the process of recovery and community development. The diatom collections had a higher density in the grazed site, though the difference was not significant. Recovery of the lichen and moss community was not complete. The lichen Collema tenax and the mosses Bryum pallescens and Tortula ruralis were all significantly more abundant in the ungrazed area. Total cover of the lichen and moss components of the soil crusts was significantly lower in the more recently grazed area. Vascular cover was also lower.     

A 4000-year record of woodland vegetation from Wind River Canyon, central Wyoming

Plant macrofossil analyses of 16 radiocarbon-dated woodrat middens spanning the past 4000 years from the Wind River Canyon region in central Wyoming provide information concerning late Holocene development of juniper woodlands. The study sites are currently dominated by Juniperus osteosperma , with J. scopulorum present locally. Woodlands in the region were dominated by J. scopulorum from ca 4000 yr BP until at least 2800 yr BP. Juniperus osteosperma invaded and expanded before 2000 yr BP. This expansion fits a regional pattern of J. osteosperma colonization and expansion in north central Wyoming during a relatively dry period between 2800 and 1000 yr BP. At the time the Wind River Canyon region was colonized by J. osteosperma , the species had populations 50-100 km to both the north and south. Long-distance seed dispersal was required for establishment in the study area. Genetic studies are necessary to identify source populations and regions.     

Historical fire frequency on contrasting slope facets along the McKenzie River, western Oregon Cascades

Fire has been a critical influence on forest dynamics over much of western North America, and knowledge of historical fire regimes provides a valuable reference point for guiding ecosystem management. Spatial and temporal variations in historical (1550–1997) fire frequency were quantified for a 145-km 2 area straddling the McKenzie River in the central western Cascades of Oregon. Fire-scar and tree-origin years estimated from cut stumps in 63 clearcut sites were used to approximately date fires. The estimated natural fire rotation was 162 years for the presettlement period (1550–1849). Natural fire rotation varied considerably over the period of record, ranging from 40 years for the Euro-American settlement period (1850–1924) to 504 years for the recent (1925–1997) fire-suppression period. Such temporal variation in fire regime may be attributed to a combination of climatic and anthropogenic influences. The study area also experienced 2 periods of widespread fire recorded at a majority of sample sites: one in the mid-1500s and the other from 1850 to 1925. Historical fire frequency has also varied spatially over at least 2 distinct scales. At the slope-facet scale, fire frequency was greatest on the generally south-facing slope to the north of the McKenzie River. At the finer scale of individual 4-ha sites, local topographic influences varied according to a site’s broad geographical context north or south of the river. Results suggest that meaningful fire-regime characterization may require information derived from multiple spatial scales.