Effective control measures are required for the invasive forage grass smooth brome in native prairie to maintain native prairie diversity and function. The objective of this study was to assess the long-term effectiveness of glyphosate as a control method for smooth brome and to evaluate the subsequent recovery of native prairie species at Kernen Prairie near Saskatoon, SK, Canada. In 1999 and 2000, a total of forty 6- to 8-m-diameter patches of smooth brome were spot sprayed with glyphosate; community composition in each patch was monitored for 17 yr. Following glyphosate application, the abundance of smooth brome decreased, and recovery of native species richness and the abundance of important native species, including plains rough fescue, was observed. In the long term however, the elimination of smooth brome created empty niche space ultimately occupied by other invasive species, particularly Kentucky bluegrass. The spot application of glyphosate is thus an effective control method for reducing smooth brome in native prairie; however, maintaining desirable native species composition in this system posttreatment depends on other factors, including the presence of additional invasive species that may move in after the elimination of smooth brome.

Understanding competition for soil nitrate between common shrub-steppe, potential reclamation species, and common invasive species is necessary to identify mechanisms associated with ecosystem invasion and can assist with developing weed management scenarios. We designed a field experiment to evaluate the differential competitive effects of the invasive annual grass downy brome, the invasive biennial forb dyer's woad, and the reclamation shrub prostrate kochia, on nitrate acquisition of the perennial grass crested wheatgrass, the native forb western yarrow, and the native shrub big sagebrush. Individual plants were grown in two-plant neighborhoods, and a K15NO3 tracer was injected into the soil between plants and recovered from leaf material after 5 to 11 d. We also evaluated neighbor effects on shoot and root growth, leaf carbon : nitrogen ratio, and leaf nitrogen concentrations to better understand how these traits are associated with differences in nitrate acquisition and nitrogen allocation among the three growth forms. Nitrate acquisitions by crested wheatgrass and western yarrow were significantly lower when competing with downy brome than with dyer's woad and prostrate kochia; however, competitors had similar, negative effects on nitrate acquisition by big sagebrush. Nitrate acquisition ratios between competing neighbors revealed that: (1) the grasses always acquired more nitrate than neighbors of a different growth form, (2) western yarrow was equally competitive with dyer's woad and prostrate kochia, and (3) all neighbors acquired more nitrate than big sagebrush. More successful competition for nitrate in the grasses was associated with greater specific root length. Compared to species of the same respective growth form, the two invasive weeds (downy brome and dyer's woad) and prostrate kochia always had significantly lower leaf carbon : nitrogen ratio, and greater leaf nitrogen concentration, which have been broadly correlated with leaf lifespan and nutrient use efficiency, and indicate differing strategies to persist in the semiarid shrub-steppe ecosystems.

The introduced exotic vines pale and black swallowwort rapidly have become invasive throughout regions of the northeastern United States and adjoining areas of Canada. Preliminary studies have reported that the species are allelopathic, possibly contributing to their competitive ability and invasiveness. Results from our laboratory assays indicated that swallowwort root exudates caused significant root length reductions (e.g., 40% for butterfly milkweed and 20% for large crabgrass) and reduced germination (e.g., 25% for lettuce) of indicator species. Additional bioassays with dried swallowwort tissues demonstrated that tissue leachates caused varied responses in indicators, with both significant stimulatory and inhibitory effects. In particular, significant congeneric interactions were noted between the two swallowwort species. Evidence from this study of swallowwort tissue phytotoxicity has important implications for developing effective management and habitat restoration strategies for the two invasive species.

Invasion of closed canopy forests by shade-tolerant alien plants has the potential to modify species composition, stand structure, ecosystem function, and long-term forest development patterns. Ligustrum sinense is a shade-tolerant alien shrub that has invaded bottomland forests throughout the southeastern United States. This species has received comparatively little attention in the literature despite its potential to drastically alter invaded sites. The overarching goal of our study was to document the relationships between Ligustrum sinense invasion and woody plant biodiversity and development patterns in an intact southeastern U.S. bottomland forest. The forest was dominated by Quercus nigra and Liquidambar styraciflua. Ligustrum sinense ranked fifth in basal area contribution, occurred in 97% of our plots, and represented 95% of all understory stems. Spearman's rho for dominance (based on basal area of stems > 5 cm diameter at breast height [dbh]) of L. sinense and woody plant species richness for each plot revealed a significant negative relationship (rs = −0.69, P < 0.01). A similar relationship was revealed between L. sinense density and woody plant species diversity (rs = −0.78, P < 0.01) and evenness (rs = −0.82, P < 0.01). Spearman's rho for L. sinense density and native understory stem density (individuals ≥ 1 m height, < 5 cm dbh) also revealed a significant negative association (rs = −0.48, P < 0.01). Under the current disturbance regime and without active management, we projected the forest would shift to support a stronger component of L. sinense and that structure would transition from tree to shrub dominance for sites within the forest.

The European vines pale swallowwort and black swallowwort are invading various habitats in northeastern North America. It is unclear how these plants might respond to potential biological control agents, as they experience little herbivore damage in North America, or longer durations of mowing given the reported lack of efficacy of mechanical control. We evaluated the effect of six seasons of artificial defoliation (50 or 100% defoliation once or twice per season) and clipping (once, twice, or four times at 8 cm above the soil level) on the survival, growth, and reproduction of mature plants of the two species grown in a common garden field experiment. No plants died from damage after 6 yr. Black swallowwort produced more aboveground biomass, whereas pale swallowwort produced more root biomass and root crown buds, compared with its congener species. For most damage treatments, root biomass and the number of crown buds and stems increased over time, whereas aboveground biomass and viable seeds per plant generally did not change. Substantial overlap in plant size and seed production occurred among damage treatments and species. The most severe defoliation treatment did not substantially limit growth and reproduction compared with undamaged plants. While two clippings per season sometimes prevented seed production, four clippings per season was the only type of damage that consistently prevented plant growth and eliminated seed production. Pale and black swallowwort display a high tolerance to aboveground tissue loss in high-light environments without plant competition. The annual increase in plant size calls into question the potential efficacy of a defoliating insect against field populations of swallowworts, and it seems likely the only benefits of a long-term mowing regime will be to eliminate seed production.

Aminopyralid efficacy on Canada thistle and changes in species composition in both Canada thistle–infested and native plant communities were evaluated in Theodore Roosevelt National Park, ND. Aminopyralid at 120 g ae/ha was applied in September 2004 to one-half of 30 native or 30 Canada thistle–infested sites. Canada thistle density 10 mo after treatment (MAT) averaged 2 stems/m2 compared to 31 stems/m2 in the control and 22 MAT averaged 16 stems/m2 compared to 42 stems/m2. Although Canada thistle was reduced in treated sites, species richness was similar to the untreated control. Species evenness increased in treated subplots compared to the control as foliar cover of low- and high-seral forbs and high-seral monocot species increased. Species richness was reduced in native plant communities treated with aminopyralid from approximately 12 to 9 species but evenness was not affected. Plant species diversity was similar in Canada thistle–infested treated and nontreated subplots but was reduced by aminopyralid in native subplots. Despite the decrease in Canada thistle and increased foliar cover of native perennial plants, composition and structure 22 MAT in the Canada thistle–infested plant community did not resemble native plant communities. Native plant communities were composed of many perennial monocots and high-seral forbs, which had increased but were in lower relative abundance in the treated Canada thistle–infested subplots.

Invasive plants continue to spread in riparian ecosystems, causing both ecological and economic damage. This research investigated the impacts of common reed, purple loosestrife, riparian shrubland, and riparian woodlands on the quality and quantity of sandhill crane roosting habitat in the central Platte River, Nebraska, using a discrete choice model. A more detailed investigation of the impacts of common reed on sandhill crane roosting habitat was performed by forecasting a spread or contraction of this invasive plant. The discrete choice model indicates that riparian woodlands had the largest negative impact on sandhill crane roosting habitat. The forecasting results predict that a contraction of common reed could increase sandhill crane habitat availability by 50%, whereas an expansion could reduce the availability by as much as 250%. This suggests that if the distribution of common reed continues to expand in the central Platte River the availability of sandhill crane roosting habitat would likely be greatly reduced.

Understanding seed characteristics and seedling establishment patterns is essential for the development of effective management strategies for invasive annual species. Dittrichia graveolens (stinkwort) has increased its range rapidly within California since 1995, yet its biology is not well understood, which has led to poorly timed management. In this study, seed viability, germination, longevity, and dormancy, as well as seedling emergence characteristics of D. graveolens were evaluated in field, greenhouse, and laboratory experiments in Davis, CA, over a 2-yr period (fall 2010 to summer 2012). In the laboratory, seed germination of D. graveolens occurred at a wide range of constant temperatures (12 to 34 C). Cumulative germination was comparable to total seed viability (80 to 95%) at optimal germination temperatures, indicating that primary (innate) dormancy is likely absent. The base temperature for germination was identified using a thermal time model: 6.5 C and 4 C for 2010 and 2011 seed populations, respectively. In the field, seedlings emerged from fall through spring following precipitation events. A very low percentage of seedlings (2.5%) emerged in the second year after planting. Equivalent seedling emergence was observed over a wide range of light conditions (100, 50, 27, and 9% of available sunlight) in a greenhouse experiment, indicating that seed germination is not limited by high or low light. Results from these seed experiments improve our understanding of the reproductive biology of this rapidly expanding exotic annual and provide valuable information for developing effective timing and longevity of management programs.

Cogongrass, an invasive grass native to Asia, has infested thousands of hectares in the southeastern United States. Although numerous studies have examined cogongrass control, no published studies, to our knowledge, have tested strategies for cogongrass eradication. Cogongrass has a persistent, thick rhizome mat but an ephemeral seedbank; therefore, successful eradication methods must largely focus on the rhizomes. A field study to evaluate specific herbicide treatments and application timings for cogongrass patch eradication was conducted at two locations in southwestern Alabama. Herbicide treatments included glyphosate at 4.48 kg ai ha−1, imazapyr at 0.84 kg ai ha−1, and a tank mix of glyphosate and imazapyr at the same rates. Treatments were applied in May, August, or October for 3 consecutive yr, and the May glyphosate treatment included a second annual application each October. Cogongrass visual control, shoot biomass, rhizome biomass, rhizome depth, and total nonstructural carbohydrate (TNC) content were sampled during the course of the study. Cogongrass response to treatments varied by location but by 36 mo after initial treatment (MAIT), complete elimination of cogongrass shoot and rhizome biomass and 100% visual control was achieved in several herbicide treatment–timing combinations at both locations. These included glyphosate plus imazapyr at any application timing, imazapyr in August or October, and glyphosate applied in May and October each year. TNC levels of surviving healthy rhizomes were not affected by herbicide treatments, but a seasonal pattern was observed. The maximum live-rhizome depth was not influenced by any treatment, indicating that herbicides were not preferentially leaving deeper, surviving rhizomes. These results demonstrate, for the first time, that the entire rhizome layer of cogongrass can be eliminated within 3 yr with multiple treatment options and that cogongrass patch eradication is possible for many land managers.

The invasive annual grass downy brome (Bromus tectorum L.) is a critical threat to the semiarid shrublands that characterize western North America. More abundant fine fuel after invasion typically increases fire frequency in plant communities adapted to relatively infrequent burning, reducing the likelihood of native plant persistence. Currently, imazapic is most often used to manage B. tectorum, but reinvasion from the seedbank after treatment is common. Indaziflam is a newer herbicide recently labeled for use in rangelands grazed by livestock, and many research trials have demonstrated its ability to deplete invasive annual grass seedbanks. We evaluated the effectiveness of indaziflam and imazapic for reducing B. tectorum density and cover over a period of approximately 5 yr (57 mo after treatment [MAT]) at two invaded sagebrush-grassland sites near Pinedale, WY. Treatments included three different indaziflam rates (51, 73, and 102 g ai ha−1) and one imazapic rate (123 g ai ha−1), and these treatments were reapplied to half of each plot at 45 MAT to evaluate the effects of two sequential applications. We also measured perennial grass cover, because positive perennial grass responses were observed after release from B. tectorum competition in other studies, and perennial grasses may provide resistance to B. tectorum reinvasion. Intermediate and high indaziflam rates (73 and 102 g ha−1, respectively) reduced B. tectorum cover and density at 45 MAT, and perennial grass cover responded positively to some treatments, mostly early in the study (≤33 MAT). Imazapic reduced B. tectorum initially, but did not affect density or cover at either site beyond 21 MAT. Reapplication did not substantially improve B. tectorum control at 57 MAT in plots treated with intermediate and high indaziflam rates, suggesting that long-term control with a single indaziflam treatment may be possible in some cases.

Broom snakeweed (snakeweed) is an aggressive native range-weed found throughout arid and semiarid areas of the western United States, that increases following disturbances such as overgrazing, drought, or wildfire. Ecologically based strategies that include controlling snakeweed and reestablishing desirable herbaceous species are needed to restore productivity and diversity to invaded areas. The objective of this study was to compare the ability of selected introduced and native grass species and prostrate kochia (kochia) to prevent reinvasion of snakeweed, downy brome, and annual forbs following control. This field study was replicated at two sites (Howell and Nephi, Utah) within the sagebrush-steppe biome. Snakeweed and downy brome were controlled by picloram (0.25 kg/ha [0.22 lb/ac]) and glyphosate (1.5 kg/ha [1.3 lb/ac]). The seeding treatments were comprised of three introduced grasses and a mix of these species, three native grasses and a mix of these species, and kochia. The treatments were seeded into 3 by 15-m (10 by 50 ft) plots in October 2003. Frequency and biomass of seeded species, snakeweed, downy brome, other grasses, and annual forbs were measured in 2004, 2005, and 2008. Seeded species were evaluated for success of establishment and persistence, and their ability to restrict reinvasion of snakeweed, downy brome, and annual forbs. Crested wheatgrass and big squirreltail had the best initial establishment at both locations (> 75%). In contrast, kochia and Russian wildrye did not establish well at either site, and western wheatgrass did not establish well at Nephi. Snakeweed reestablishment was restricted in all treatments except the kochia treatments, in which kochia did not establish well. Frequency of downy brome increased at both sites, and annual forb frequency increased at Nephi to near 100%, but the better established grasses suppressed biomass production of these weedy species. Crested wheatgrass established best, had the greatest production, and provided greatest suppression of downy brome and annual weeds.

The invasive annual grass downy brome is the most ubiquitous weed in sagebrush systems of western North America. The center of invasion has largely been the Great Basin region, but there is an increasing abundance and distribution in the Rocky Mountain States. We evaluated postfire vegetation change using very large–scale aerial (VLSA) and near-earth imagery in an area where six different fires occurred over a 4-yr period at elevations ranging from 1,900 to over 2,700 m. The frequency of downy brome increased from 8% in 2003 to 44% in 2008 and downy brome canopy cover increased from < 1% in 2003 to 6% in 2008 across the entire study area. Principal component analyses of vegetation cover indicate a shift from plant communities characterized by high bare soil and forbs immediately postfire to communities with increasing downy brome cover with time after fire. The highest-elevation sampling area exhibited the least downy brome cover, but cover at some midelevation locations approached 100%. We postulate that the loss of ground-level shade beneath shrubs and conifers, accompanied by diminished perennial vegetative cover, created conditions suitable for downy brome establishment and dominance. Without a cost-effective means of landscape-scale downy brome control, and with infestation levels and climate warming increasing, we predict there will be continued encroachment of downy brome at higher elevations and latitudes where disturbance creates suitable conditions.

We investigated the single and combined effects of two biological control agents, the stem-mining weevil Hadroplontus litura and the pathogen Pseudomonas syringae pv. tagetis, with a herbicide (reduced or full application of glyphosate: 0.63 kg ae ha−1, or 3.78 kg ae ha−1, respectively) on the growth of Canada thistle, Cirsium arvense. We hypothesized that first, although each control method would have a negative effect on Canada thistle shoot biomass, root biomass, and shoot number, the integration of more than one control method would have greater impact than individual control methods. Second, we hypothesized that the order in which control methods are applied affects the outcome of the management program, with a pathogen application following weevil infestation being more effective than one prior to it. Although control methods impacted Canada thistle growth (P < 0.001, expect for a nonsignificant impact of glyphosate on shoot number), the combined effect of the three control methods behaved, generally, in an additive manner. A marginal interaction between the pathogen and the herbicide (P = 0.052) indicated a slight antagonistic interaction between these control methods. An interaction between the two biological control agents tested (P < 0.001) indicated that application of a pathogen prior to the release of weevil larvae could be more deleterious to Canada thistle than a late application. The observed, mostly additive, relationship between biological control agents and herbicides implies that integrating control methods rather than using a single approach could lead to greater Canada thistle control.

Introduced plants threaten biodiversity and ecosystem processes, including carbon (C) and nitrogen (N) cycles, but little is known about the threshold at which such effects occur. We examined the impact of the invasive shrub Amur honeysuckle on soil organic carbon (SOC) and N density at study sites that varied in invasion history. In plots with and without honeysuckle, we measured honeysuckle abundance and size (basal area) and extracted soil cores. SOC and N densities were highest at the site with the longest invasion history and highest invasion intensity (i.e., greatest abundance and basal area of honeysuckle). Basal area of honeysuckle positively affected SOC and N densities likely because of increased litter decomposition and altered microbial communities. Because honeysuckle increases forest net primary productivity (NPP) and SOC, it also may play a role in C sequestration. Our results demonstrate the need to consider the influence of invasion history and intensity when evaluating the potential impact of invasive species.

The giant reed, Arundo donax is one of the worst invasive alien species globally, including South Africa, where it invades riparian areas across the country. Biological control is being considered to address the invasive potential and negative impacts of the weed. This study investigated the phylogeography of A. donax to guide the biological control program. To determine plant haplotype and genetic diversity, three regions of the chloroplast were sequenced and three microsatellite markers were analyzed in 40 samples from across the plant’s distribution in South Africa. It was determined that all populations of A. donax in South Africa were haplotype M1, which is the most widely distributed haplotype worldwide, believed to originate from the Indus Valley, Asia. In addition, no genetic diversity was found, indicating that all the A. donax populations in South Africa are essentially one clone. The results indicate that suitable biological control agents are likely to be found in the ancient native range of haplotype M1. This research has contributed to the global understanding of the phylogeography of A. donax and will guide the biological control program in South Africa.

Since its introduction to the United States in 1852, Chinese privet (Ligustrum sinense Lour.) has spread throughout the Southeast, invading many natural areas. Manual control by cutting or shredding is one of the most common strategies many land managers employ. However, rapid sprouting from the root collar and lateral roots commonly results in poor control. Cutting followed by either glyphosate or triclopyr application to the stumps is generally effective, but the efficacy of these herbicides in relation to treatment timing and L. sinense root collar diameter has not been evaluated. The objective of this experiment was to determine the effectiveness of glyphosate and triclopyr cut stump treatments compared with cutting alone at spring and fall timings across a range of L. sinense size classes. Studies were conducted at two locations in Auburn, AL. Treatments included cut stump+no herbicide, cut stump+ glyphosate (120 g L−1), or cut stump+triclopyr (90 g L−1). Treatments were applied to at least 50 experimental units each at April and November timings. Root collar diameter was recorded for each stem, stems were cut 2.5 cm above the ground, and herbicide treatments were applied within 30 s. Ligustrum sinense mortality and sprouting were quantified 6, 12, and 18 mo after treatment. Both glyphosate and triclopyr amine were very effective in controlling L. sinense at both spring and fall timings. However, glyphosate provided slightly better results than triclopyr when lateral sprouting was included. Application timing also was significant, with a lower percentage of sprouting following November treatments than April treatments. Stem size influenced treatment success, as larger stumps tended to sprout more than smaller stumps. These results indicate L. sinense can be controlled with cut stump herbicide treatment using either glyphosate or triclopyr with spring or fall timings at concentrations much lower than typically used.

Nonnative, invasive plants are becoming increasingly widespread and abundant throughout the southwestern United States, leading to altered fire regimes and negative effects on native plant communities. Models of potential invasion are pertinent tools for informing regional management. However, most modeling studies have relied on occurrence data, which predict the potential for nonnative establishment only and can overestimate potential risk. We compiled locations of presence and high abundance for two problematic, invasive plants across the southwestern United States: red brome (Bromus rubens L.) and African mustard (Brassica tournefortii Gouan). Using an ensemble of five climate projections and two types of distribution model (MaxEnt and Bioclim), we modeled current and future climatic suitability for establishment of both species. We also used point locations of abundant infestations to model current and future climatic suitability for abundance (i.e., impact niche) of both species. Because interpretations of future ensemble models depend on the threshold used to delineate climatically suitable from unsuitable areas, we applied a low threshold (1 model of 10) and a high threshold (6 or more models of 10). Using the more-conservative high threshold, suitability for Bromus rubens presence expands by 12%, but high abundance contracts by 42%, whereas suitability for Brassica tournefortii presence and high abundance contract by 34% and 56%, respectively. Based on the low threshold (worst-case scenario), suitability for Bromus rubens presence and high abundance are projected to expand by 65% and 64%, respectively, whereas suitability for Brassica tournefortii presence and high abundance expand by 29% and 28%, respectively. The difference between results obtained from the high and low thresholds is indicative of the variability in climate models for this region but can serve as indicators of best- and worst-case scenarios.

Invasive species are widely recognized as a major threat to global diversity and an important factor associated with global change. Species distribution models (SDMs) have been widely applied to determine the range that invasive species could potentially occupy, but most examples focus on predictive variables at a single spatial scale. In this study, we simultaneously considered a broad range of variables related to climate, topography, land cover, land use, and propagule pressure to predict what areas in the southeastern United States are more susceptible to invasion by 45 invasive terrestrial plant species. Using expert-verified occurrence points from EDDMapS, we modeled invasion susceptibility at 30-m resolution for each species using a maximum entropy (MaxEnt) modeling approach. We then analyzed how environmental predictors affected susceptibility to invasion at different spatial scales. Climatic and land-use variables, especially minimum temperature of coldest month and distance to developed areas, were good predictors of landscape susceptibility to invasion. For most of the species tested, human-disturbed systems such as developed areas and barren lands were more prone to be invaded than areas that experienced minimal human interference. As expected, we found that landscape heterogeneity and the presence of corridors for propagule dispersal significantly increased landscape susceptibility to invasion for most species. However, we also found a number of species for which the susceptibility to invasion increased in landscapes with large core areas and/or less-aggregated patches. These exceptions suggest that even though we found the expected general patterns for susceptibility to invasion among most species, the influence of landscape composition and configuration on invasion risk is species specific.

Some plants being considered as bioenergy crops share traits with invasive species and have histories of spreading outside of their native ranges, highlighting the importance of evaluating the invasive potential before the establishment of large-scale plantings. The Asian grass Miscanthus × giganteus is currently being planted as a bioenergy crop in the north central region of the United States. Our goal was to understand the demographic rates and vegetative spread of this species in unmanaged arable lands in Illinois to compare with those of large-statured invasive grasses (LSIGs). We collected data from 13 M. × giganteus plantings in Illinois, ranging in age from 1 to 7 yr, recording tiller number, plant spatial extent, spikelet production, and plant survival over 4 yr. Additionally, to understand recruitment potential, we conducted a greenhouse germination experiment, and, to estimate establishment from rhizome fragments, field trials were performed. Miscanthus × giganteus demographic rates were age dependent. Spikelet production was high, with 1- and 4-yr plants producing an annual average of more than 10,000 and 180,000 spikelets plant−1, respectively; however, data from our germination trial suggested that none of these spikelets had the potential to yield seedlings. On average, plants expanded vegetatively 0.15 m yr−1. Tiller density within the center of a clone decreased with age, possibly leading to a “dead center” found among some LSIGs. Rhizome establishment increased with weight, ranging from 0 to 42%. Survival was low, 24%, for first-year plants but quickly climbed to an asymptote of 98% survival for 4-yr-old plants. Our results suggest that efforts should be made to eradicate plants that escape biomass production fields within a year of establishment, before the onset of high survival. Future work is needed to determine what types of natural and anthropogenic disturbances can fragment rhizomes, leading to regeneration.

Early detection of an invading nonindigenous plant species (NIS) may be critical for efficient and effective management. Adaptive survey sampling methods may provide unbiased sampling for best estimates of distribution of rare and spatially clustered populations of plants in the early stages of invasion. However, there are few examples of these methods being used for nonnative plant surveys in which travelling distances away from an initial or source patch, or away from a road or trail, can be time consuming due to the topography and vegetation. Nor is there guidance as to which of the many adaptive methods would be most appropriate as a basis for invasive plant mapping and subsequent management. Here we used an empirical complete census of four invader species in early to middle stages of invasion in a management area to assess the effectiveness and efficiency of three nonadaptive methods, four adaptive cluster methods, and four adaptive web sampling methods that all originated from transects. The adaptive methods generally sampled more NIS-occupied cells and patches than standard transect approaches. Sampling along roads only was time-efficient and effective, but only for species with restricted distribution along the roads. When populations were more patchy and dispersed over the landscape the adaptive cluster starting at the road generally proved to be the most time-efficient and effective NIS detection method.