Temporal patterns of seedling emergence and early survival of Great Basin perennial plant species

We seeded Agropyron cristatum and 16 native Great Basin perennial species on Bromus tectorum -dominated sites to compare temporal patterns of seedling emergence and early survival among species. To indicate variability between sites and between years, plots were planted on 2 sites in western Utah in autumn 1994, and plantings were repeated near 1 site in autumn 1995. Two sites included burned and unburned seedbed treatments. We monitored seedling emergence and survival from early winter until July of the 1st year. Four seasonal patterns of emergence were evident, indicating considerable potential for different seasonal patterns of precipitation to favor recruitment of difference species. Emergence of all shrubs, except Ephedra nevadensis , began in February and essentially ended by April. Seedlings of Ephedra nevadensis and of the grasses Agropyron and Pseudoroegneria spicata emerged over an extended period from February through June. Most other grasses and the 2 forbs began emerging in April and continued into June. Pleuraphis jamesii emergence did not begin until May. Although many seedlings of the 3 chenopod shrubs emerged, none survived, suggesting their episodic recruitment is more a function of survival than of emergence. Survival of 2 Chrysothamnus species was also low. Pseudoroegneria and Agropyron had the highest numbers of seedlings surviving on the moister site, followed by Elymus elymoides and Stipa comata . Survival of Agropyron , but not the native Elymus , was more depressed on the drier of the 2 sites planted in 1994. Stipa, Oryzopsis hymenoides , Pleuraphis , and Ephedra survived as well on the drier site as on the moister site, or better. On the 1995 planting sites Bromus was greatly reduced by the burn treatment, and, in contrast to the effect of burn treatments on the 1994 site, survival of all species except Oryzopsis and Ephedra was higher on burned than on unburned treatments. Overall, Stipa, Oryzopsis , and Pleuraphis had low emergence but consistently higher survival.     

Seed banks of Bromus tectorum -dominated communities in the Great Basin

We seeded Agropyron cristatum and 16 native Great Basin perennial species on Bromus tectorum -dominated sites to compare temporal patterns of seedling emergence and early survival among species. To indicate variability between sites and between years, plots were planted on 2 sites in western Utah in autumn 1994, and plantings were repeated near 1 site in autumn 1995. Two sites included burned and unburned seedbed treatments. We monitored seedling emergence and survival from early winter until July of the 1st year. Four seasonal patterns of emergence were evident, indicating considerable potential for different seasonal patterns of precipitation to favor recruitment of difference species. Emergence of all shrubs, except Ephedra nevadensis , began in February and essentially ended by April. Seedlings of Ephedra nevadensis and of the grasses Agropyron and Pseudoroegneria spicata emerged over an extended period from February through June. Most other grasses and the 2 forbs began emerging in April and continued into June. Pleuraphis jamesii emergence did not begin until May. Although many seedlings of the 3 chenopod shrubs emerged, none survived, suggesting their episodic recruitment is more a function of survival than of emergence. Survival of 2 Chrysothamnus species was also low. Pseudoroegneria and Agropyron had the highest numbers of seedlings surviving on the moister site, followed by Elymus elymoides and Stipa comata . Survival of Agropyron , but not the native Elymus , was more depressed on the drier of the 2 sites planted in 1994. Stipa, Oryzopsis hymenoides , Pleuraphis , and Ephedra survived as well on the drier site as on the moister site, or better. On the 1995 planting sites Bromus was greatly reduced by the burn treatment, and, in contrast to the effect of burn treatments on the 1994 site, survival of all species except Oryzopsis and Ephedra was higher on burned than on unburned treatments. Overall, Stipa, Oryzopsis , and Pleuraphis had low emergence but consistently higher survival.     

Succession in pinyon-juniper woodlands following wildfire in the Great Basin

Twenty - one areas in pinyon ( Pinus monophylla ) - juniper ( Juniperus osteosperma ) woodlands burned by wildfire from approximately 1 to 60 years prior to sampling and adjacent unburned mature woodland stands were studied in Nevada and California to determine successional patterns and individual species responses to burning and to changing plant communities through time. One year after burning, all late successional woodland species were present in postburn plant communities except tree species. Increases in both cover and occurrence of annual and perennial forbs resulted in their dominance on early successional sites. Shrubs and annual grasses dominated midsuccessional sites, subsequently giving way to tree, shrub, and perennial grass dominance in late succession. North and east slopes generally supported high cover and occurrence of shrubs, perennial grasses, and perennial forbs, and south and west slopes generally supported high cover and occurrence of annual forbs and annual grasses. The ability to group species according to preferential occurrence on various aspects and successional stages can be used to predict plant community composition in time and space in the pinyon - juniper woodlands.         

Tree size and understory phytomass production in a western juniper woodland

Understory phytomass production in a western juniper ( Juniperus occidentalis ) woodland was examined relative to tree size in central Oregon in 1983 and 1984. Vegetation was sampled in two zones, the canopy zone (beneath the canopy) and the intercanopy zone (the space between canopies), on two adjacent sites—a lower slope site with shallow soil and an upper slope site with deeper soil. Sampling was stratified into three tree size classes. Individual species production was significantly affected by tree size and location relative to tree canopy. Production of bottlebrush squirreltail, bluebunch wheatgrass, cheatgrass, miscellaneous annual grasses, perennial forbs, and annual forbs increased with increasing tree size. Sandberg bluegrass production was greater in the intercanopy than the canopy zone, while production of bottlebrush squirreltail, bluebunch wheatgrass, miscellaneous annual grasses, and both perennial and annual forbs was greater in the canopy zone. Production of cheatgrass was determined by the interaction of tree size and zone. Phytomass relationships were expressed to a greater degree on the upper slope site, where total production exceeded that of the lower slope site by approximately 50% the second year of the study. Individual trees appear to exert a great influence on associated vegetation as western juniper woodlands progress from the seedling (tree establishment) phase to closed stands of mature trees. Original community dominants appear to be spatially segregated beneath tree canopies and associated with large trees, while formerly less common species, such as cheatgrass, come to dominate the entire site.     

Short-term effects of a prescribed burn on songbirds and vegetation in mountain big sagebrush

I measured songbird abundance and vegetation cover in and around a 420-ha prescribed burn in a mountain big sagebrush community located at 2133 m elevation. Data were collected during the 3rd growing season after the fire. Brewer's Sparrow and Sage Thrasher occurred in lower abundance on sites that were largely or completely burned relative to sites that were outside the fire perimeter or within unburned islands of shrubs. The number of Brewer's Sparrow detections was linearly related to remaining sagebrush cover. In contrast, Horned Lark occurred at higher abundances on sites where shrub cover had been removed in the prescribed burn. Cover of perennial grasses and cover of 4 of the 5 most common forbs was greater on burned sites than on unburned sites.     

Plant community patterns in unburned and burned blackbrush ( Coleogne ramosissima Torr.) shrublands in the Mojave Desert

The blackbrush vegetation type is dominated by Coleogyne ramossisima , which is thought to preclude the coexistence of many other plant species. Fire can remove blackbrush cover and possibly increase plant species richness and evenness. Fire also may increase the frequency and cover of alien annual grasses, thereby intensifying landscape flammability. We tested these predictions in unburned and burned (6-14 years postfire) blackbrush at 3 sites spanning the range of this vegetation type in the Mojave Desert. Species richness in unburned blackbrush was similar to published values for vegetation types in western North America, bur richness varied significantly among the 3 sites and 4 spatial scales (1, 10, 100, and 1000 m 2 ). Richness values declined in order from annual forbs, woody perennials, herbaceous perennials, annual grasses, cacti, to perennial grasses. Fire reduced Coleogyne cover, thus boosting species evenness. In contrast, species richness decreased after burning, although the results varied among spatial scales. Total cover was unaffected by fire because cover of wood perennials decreased, while cover of annual forbs, annual grasses, herbaceous perennials, and perennial grasses increased. Native species richness and cover decreased, whereas alien richness and cover increased after burning especially where the alien forb Erodium cicutarium was present. Fire had no effect on frequency and variable effects on cover of alien annual grasses. These results indicate that in blackbrush species richness can vary among sites and local spatial scales, and effects of fires can vary among plant life-forms and between natives and aliens.     

Community response of nontarget species to herbicide application and removal of the nonindigenous invader Potentilla recta L

Our main objective was to improve understanding of herbicide effects on community dynamics to refine the use of technology and advance the development of ecologically based weed management strategies. We hypothesized that native grasslands would exhibit reductions in culturally sensitive forb cover, biomass, and density relative to the rate of application of selective rangeland herbicides, and that hand-removal of sulfur cinquefoil ( Potentilla recta L.) would increase indigenous species cover, biomass, density, species richness, and diversity. Treatments consisted of 3 rates each of 2,4-D + clopyralid (0.28 kg ai ? ha -1 + 0.0532 kg ai ? ha -1 , 0.56 kg ai ? ha -1 + 0.1064 kg ai ? ha -1 , 0.84 kg ai ? ha -1 + 0.1596 kg ai ? ha -1 ); 2,4-D amine (0.532 kg ai ? ha -1 , 1.064 kg ai ? ha -1 , 1.596 kg ai ? ha -1 ); metsulfuron (0.0042 kg ai ? ha -1 , 0.021 kg ai ? ha -1 , 0.032 kg ai ? ha -1 ); picloram (0.14 kg ai ? ha -1 , 0.28 kg ai ? ha -1 , 0.56 kg ai ? ha -1 ); and clopyralid (0.05025 kg ai ? ha -1 , 0.21 kg ai ? ha -1 , 0.42 kg ai ? ha -1 ). This experiment was replicated 3 times at 2 lateseral, noninfested sites in southeastern Montana. In a companion study, sulfur cinquefoil was removed adjacent to paired nonremoved controls in 5 replicates at 2 sites in 1-m 2 plots for 2 growing seasons. Canopy cover, density, and biomass were collected 24 months after initial treatment at all sites. Indigenous perennial grass cover and biomass increased with herbicide application; however, picloram, metsulfuron, and clopyralid reduced native forb density at 1 site, and picloram reduced forb cover at both sites regardless of rate. Effects of herbicides on species richness or diversity were not detected. Hand-removing sulfur cinquefoil increased total plant richness, especially that of native forbs. Restoring species richness and diversity may be difficult using selective broadleaf herbicides because key functional groups, such as forbs, appear to be at risk.     

Native plant community composition and genetic diversity associated with long-term weed invasions

Many studies have assessed genetic changes in exotic plant species in their native and introduced ranges, but none have focused on genetic variation in native plant species in response to exotic invasion. We examine characteristics of native plant communities within and outside old (> 25 year) invasions of Acroptilon repens (Russian knapweed) and Cardaria draba (hoary cress). We also document genetic variability of 4 native grass populations ( Hesperostipa comata [needle and thread], Achnatherum hymenoides [Indian ricegrass], Sporobolus airoides [alkali sacaton], and Poa secunda [Sandberg bluegrass]) from 2 areas: adjacent to and within weed invasions. Native plant species richness and diversity did not differ between invaded and noninvaded areas. Inter-simple sequence repeat (ISSR) analysis of individual native perennial grasses of each of the 4 species suggests that populations exposed to long-term coexistence with exotics may differ from adjacent noninvaded populations. We suggest that future research efforts should focus on intraspecific diversity of native plant species to identify possible candidates for restoration following weed control.     

Vegetation response to fire and postburn seeding treatments in juniper woodlands of the Grand Staircase-Escalante National Monument, Utah

We compared 3 naturally ignited burns with unburned sites in the Grand Staircase-Escalante National Monument. Each burn site was restored with native and nonnative seed mixes, restored with native seeds only, or regenerated naturally. In general, burned sites had significantly lower native species richness (1.8 vs. 2.9 species), native species cover (11% vs. 22.5%), and soil crust cover (4.1% vs. 15%) than unburned sites. Most burned plots, seeded or not, had significantly higher average nonnative species richness and cover and lower average native species richness and cover than unburned sites. Regression tree analyses suggest site variation was equally important to rehabilitation results as seeding treatments. Low native species richness and cover, high soil C, and low cover of biological soil crusts may facilitate increased nonnative species richness and cover. Our study also found that unburned sites in the region had equally high cover of nonnative species compared with the rest of the Monument. Cheatgrass ( Bromus tectorum ) dominated both burned and unburned sites. Despite the invasion of cheatgrass, unburned sites still maintain higher native species richness; however, the high cover of cheatgrass may increase fire frequency, further reduce native species richness and cover, and ultimately change vegetation composition in juniper woodlands.     

Species richness and California voles in an annual and a perennial grassland

Populations of a common burrowing rodent, Microtus californicus (the California vole), thrive in ungrazed or lightly grazed grasslands in coastal California. Two sites ungrazed by livestock, one dominated by native perennial grasses and another dominated by invasive annuals, were evaluated over 2 consecutive years for the relationship between plant species richness and location of M. californicus burrow entrances (burrows). Plant species and burrows were sampled as present or absent in contiguous 1-m 2 quadrats on a 100-m 2 grid. Quadrats with burrows averaged significantly more plant species than quadrats without them (11.3 vs. 9.9 species, P < 0.001). Burrows found in 1996 were not correlated with species richness in 1995, suggesting that voles affect richness rather than seek it out. Vole burrow locations showed significant clumping on the annual site and trended toward clumping on the perennial site in both 1995 and 1996. Because voles seem to create a clumped pattern with their burrow entrances, the associated increase in plant species richness may have a strong effect on the overall structure of the plant community. A quantitative comparison of the 2 sites showed that the plant matrix of the perennial site contained flora of the annual site. This similarity in plant species composition may allow for similar treatment of our 2 types of sites and potentially other California grasslands. Undetected increases in vole populations with livestock grazing reduction may account for the erratic results from grasslands management research and the inconsistent success of derived management practices.     

Livestock and vegetation responses to rotational grazing in short-grass steppe

There is a paucity of information concerning livestock and vegetation responses to rotational grazing in short-grass steppe. We compared effects between 1995 and 2003 of time-controlled, short-duration rotational grazing against season-long continuous grazing, at a moderate stocking rate (1.95 ha ? animal unit month –1 ), on livestock gains and on foliar and basal cover in short-grass steppe. Livestock average daily gains, grazing-season gains, and beef production did not differ between grazing systems. We observed no relationship between precipitation (annual or growing-season) and average daily gain. In contrast, both grazing-season gains and beef production exhibited a significant curvilinear response to both growing-season and annual precipitation. Basal and foliar cover of all plant functional groups (C 3 annual grasses, C 3 perennial grasses and grass-likes, C 4 perennial grasses, cactus, annual forbs, perennial forbs, and shrubs/subshrubs) did not differ between grazing systems. Litter and bare ground were also unaffected by grazing system, with litter increasing and bare ground decreasing over the duration of this experiment. Land managers in shortgrass steppe who are implementing rotational grazing at moderate stocking rates should not expect increased livestock gains or changes in basal and foliar cover of the plant community. Rotational grazing may be a useful management tool for achieving conservation goals through modifications of seasonality and intensity of use in paddocks within larger management units to accommodate spatial variability in vegetation and temporal variation in precipitation.     

Differential effects of cattle and sheep grazing on high mountain meadows in the Strawberry Valley of central Utah

Species diversity, niche metrics, cover, frequency, and soil relationships were studied on high mountain meadows on adjacent cattle and sheep allotments in Strawberry Valley, Wasatch County, Utah. The cattle allotment vegetation was predominantly Mountain bluebell (Mertensia ciliata) , and the sheep allotment vegetation was predominantly Smallwing sedge (Carex microptera) . Other species of importance on both areas included Letterman needlegrass (Stipa lettermanii) , Mountain bromegrass (Bromus carinatus) , and Yarrow (Achillea millefolium) . Tall forbs were most abundant on the cattle allotment, and low forbs, perennial grasses, and sedges were most abundant on the sheep allotment. Vegetation composition on the two allotments was significantly different. &nbsp;&nbsp;&nbsp;     

Long-term effects of tebuthiuron on Bromus tectorum

Use of herbicides to thin dense stands of Artemisia spp. (sagebrush) can free up resources for herbaceous plants and increase forage production, but may also facilitate weed invasion. We revisited a sagebrush thinning experiment in a north central Wyoming big sagebrush–grassland 11 years after application of tebuthiuron (N-[5-(1,1- dimethylethyl)-1,3,4-thiadiazol-2-yl]-N-N′-dimethylurea) to determine the long-term responses of shrubs, available soil resources, perennial grasses, and Bromus tectorum L. (downy brome). Tebuthiuron reduced shrub cover by more than half, from 31% in untreated plots to 15% in treated plots ( P = 0.002), and increased downy brome cover approximately 4-fold, from 0.9% in untreated plots to 3.5% in treated plots ( P = 0.02). Treatment with tebuthiuron also resulted in marginally significant increases in cover of perennial grasses (from 9% to 12.3%; P = 0.07) and bare ground (from 39.1% to 43.9%; P = 0.08). In comparisons of resource availability among microsites, available NO 3 was higher under dead sagebrush than under live sagebrush ( P = 0.03). No significant differences in soil water content were detected. The relatively recent expansion of downy brome populations at this site and the high NO 3 –N levels observed under dead sagebrush suggest that conditions facilitating downy brome invasion may persist for many years following sagebrush thinning. We demonstrate that sagebrush thinning can cause increases in downy brome populations years after initial treatment and suggest that managers should use caution when considering thinning sagebrush if downy brome is present, even if initial populations are small.     

Potential for controlling the spread of Centaurea maculosa with grass competition

Spotted knapweed ( Centaurea maculosa Lam.) is a major rangeland and roadside weed of the northern Rocky Mountains. It is often found in plant communities dominated by Pseudoroegneria spicatum or Festuca idahoensis , but it rarely invades roadsides dominated by Bromus tnermis Leyss. Aboveground biomass of the 3 grass species grown in mixture with Centaurea was compared to growth in monoculture at a range of nitrogen input levels. The results suggest that Bromus is capable of suppressing the growth of Centaurea with the degree of suppression increasing with increasing nitrogen levels. The 2 native grasses had no impact on Centaurea under the controlled environment conditions of this study.     

C 3 and C 4 Species Changes Identified by δ 13 C values of soil organic matter in a Colorado prairie

We measured carbon isotope signatures (δ 13 C) from 0-10 cm and 10-20 cm soil depth intervals for grassland soils near Boulder, Colorado. These grasslands included tall-, short-, and mixed-grass prairies that were grazed, ungrazed, or hayed. Soils exhibited δ 13 C signatures consistent with observations that current sites are a mix of C 3 and C 4 species, with C 3 plants more abundant in mixed-grass than in native tall- or shortgrass prairies. The δ 13 C signatures were not significantly different for grassland types; however, management treatments (grazing, no grazing, haying) significantly influenced changes in soil δ 13 C signatures from the 0-10 cm to 10-20 cm soil depth intervals. We observed a correlation ( r = 0.63) between isotopic values of surface soils and percent native species in total vegetation cover. Overall, the community type with the lowest percentage of nonindigenous species cover had the most enriched δ 13 C signature. Sites currently grazed by prairie dogs, cattle, or both herbivores had stronger C 3 signatures, indicating that grazing may have increased C 3 plant productivity in these communities at the expense of C 4 grasses. This finding differs from studies of native shortgrass steppe where grazing has the opposite effect on the relative abundance of these 2 functional groups of plants. This result, along with the correlation between C 3 isotopic values and nonnative vegetation abundance, provides evidence that management practices that maintain dominance of C 4 grasses should be encouraged.     

Allelopathic effects of bur buttercup tissue on germination and growth of various grasses and forbs in vitro and in soil

The allelopathic effects of bur buttercup ( Ranunculus testiculatus ) tissue on selected grasses and forbs varied according to the substratum for germination and growth. The in vitro effects of an aqueous extract of buttercup tissue on germination and root development of five grasses were strongly inhibitory in all cases. &nbsp; However, in soil the effects of buttercup tissue on germination and growth of seven grasses and two dicotyledonous herbs were small to nonsignificant. Deleterious effects were less severe in fine - as opposed to coarse-textured soils. Under field conditions, the ability of seedlings of the grasses to compete with buttercup varied with the species.     

Long-term vegetation dynamics in a cut western juniper woodland

Western juniper ( Juniperus occidentalis spp. occidentalis Hook.) expansion in the northern Great Basin has reduced shrubsteppe productivity and diversity. Chainsaw cutting of western juniper woodlands is commonly applied to remove tree interference and restore sagebrush plant communities. Studies assessing understory response following cutting have been limited to early successional stages and have not evaluated the effects of western juniper debris on plant succession. Cutting western juniper produces a large amount of debris which is commonly left on site, occupying a significant portion of treated areas. This study evaluated successional dynamics spanning 13 years after western juniper cutting. Four 0.45-ha blocks were selected on Steens Mountain in southeastern Oregon. Western juniper cover averaged 26% and mature tree density averaged 250 trees ? ha –1 . Blocks were cut in late summer 1991. Understory standing crop, cover, and density were compared among 3 locations: old canopy litter mats (canopy), interspace, and area underneath cut western juniper (debris). In the interspace, perennial grasses increased in cover and in standing crop relative to other functional groups. In canopy and debris locations, species composition shifted in the 6th year after cutting as annual grass cover, density, and standing crop increased. However, by 2003, perennial grass biomass was 2 times greater than annual grass biomass in canopy and debris locations. Because annual grasses increased in areas of debris accumulation, managers need to be cognizant of western juniper treatments that create safe sites that are favorable to the establishment of weedy species. Retaining western juniper debris on this site did not increase establishment and growth of perennial grasses compared to the interspace. A shift in perennial grass dominance from Thurber's needlegrass ( Achnatherum thurberianum [Piper] Barkworth) to bottlebrush squirreltail ( Elymus hystrix [Nutt.] Smith) occurred in areas of debris accumulation. Our results demonstrated that long-term vegetation evaluations are necessary to properly assess management activities and disturbance.     

Relationships between Centaurea maculosa and indigenous plant assemblages

Ecological impacts of invasive plants include displacement of indigenous species and declines in species richness and diversity. The objective of this study was to characterize the functional relationship between plant community composition and Centaurea maculosa Lam. (spotted knapweed) within a Festuca idahoensis / Pseudoroegneria spicatum habitat type in Montana. Density, cover, and biomass of all species were collected along a gradient of spotted knapweed cover ranging from 0% to about 100%. Step-down regression was used to determine the relationship among C. maculosa , indigenous species, species richness, and Shannon-Weavers diversity index. Regressions showed that indigenous perennial grass cover, species richness, and species diversity were inversely related to C. maculosa cover. There was no relationship between C. maculosa and indigenous forbs. While this study does not imply a causal relationship, the literature suggests that C. maculosa displaces indigenous species and/or invades areas of reduced indigenous plant cover, low diversity, or low species richness. Knowing levels of indigenous perennial grass cover will help managers predict the outcome of weed management on rangelands that are vulnerable to weed infestation.     

Density, biomass, and diversity of grasshoppers (Orthoptera: Acrididae) in a California native grassland

A native California perennial grassland was sampled for grasshopper populations. The grassland is managed for the preservation of the native perennial bunchgrass, Nassella pulchra Hitch. Grasshopper density, biomass, diversity, and richness were measured from July 1993 to October 1994. Average density of all grasshoppers was 2.30 hoppers/m 2 (0.66 s ) for 1994 (June through August). Overall forage consumed for 1994 was 140 kg/ha, suggesting that grasshopper populations exist at economically damaging levels. Grasshoppers do not appear in the grasslands until late spring, after annual grasses have set seed. Biomass of grasshoppers peaks in July when adults are predominant. Both grasshopper density and biomass were higher in 1993 than in 1994, and a total of 5 species were found throughout the study. Melanoplus sanguinipes Fabricus dominated the acridid communities and accounted for more than 95% of the individuals.     

Arthropod and plant communities as indicators of land rehabilitation effectiveness in a semiarid shrubsteppe

We describe a case study evaluating the ecological impact of Bromus tectorum L. (cheatgrass) invasion following fire disturbance and the effectiveness of revegetation as a means of rehabilitation in a degraded semiarid shrubsteppe system. The effectiveness of rehabilitation efforts was assessed relative to arthropod richness, vegetation and arthropod community composition, and ground-cover characteristics in 3 habitats: undisturbed, burned and weed infested ( B. tectorum ), and burned then rehabilitated with native and nonnative vegetation. Arthropods were collected in each habitat using pitfall traps. Differences in arthropod richness were compared using rarefaction curves. Nonmetric multidimensional scaling and nonparametric multivariate statistical procedures, including analysis of similarity and similarity percentage routines, were used to compare arthropod and vegetation community composition and ground-cover characteristics between habitats. Arthropod communities in the rehabilitated habitat were distinct from those observed in the undisturbed and weed-infested habitats. Rehabilitation in this study resulted in a shift toward conditions observed in an undisturbed habitat and perhaps is an intermediate step to complete restoration. Arthropod richness, arthropod and vegetation community composition, and ground-cover characteristics were all useful indicators but returned slightly different results. Assessing multiple variables yielded the most complete understanding of the habitats studied.