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Improving national-scale invasion maps: Tamarisk in the western United States

New invasions, better field data, and novel spatial-modeling techniques often drive the need to revisit previous maps and models of invasive species. Such is the case with the at least 10 species of Tamarix , which are invading riparian systems in the western United States and expanding their range throughout North America. In 2006, we developed a National Tamarisk Map by using a compilation of presence and absence locations with remotely sensed data and statistical modeling techniques. Since the publication of that work, our database of Tamarix distributions has grown significantly. Using the updated database of species occurrence, new predictor variables, and the maximum entropy (Maxent) model, we have revised our potential Tamarix distribution map for the western United States. Distance-to-water was the strongest predictor in the model (58.1%), while mean temperature of the warmest quarter was the second best predictor (18.4%). Model validation, averaged from 25 model iterations, indicated that our analysis had strong predictive performance (AUC = 0.93) and that the extent of Tamarix distributions is much greater than previously thought. The southwestern United States had the greatest suitable habitat, and this result differed from the 2006 model. Our work highlights the utility of iterative modeling for invasive species habitat modeling as new information becomes available.

Western United States

Information on black-footed ferret biology collected within the framework of ferret conservation

Once feared to be extinct, black-footed ferrets ( Mustela nigripes ) were rediscovered near Meeteetse, Wyoming, in 1981, resulting in renewed conservation and research efforts for this highly endangered species. A need for information directly useful to recovery has motivated much monitoring of ferrets since that time, but field activities have enabled collection of data relevant to broader biological themes. This special feature is placed in a context of similar books and proceedings devoted to ferret biology and conservation. Articles include general observations on ferrets, modeling of potential impacts of ferrets on prairie dogs ( Cynomys spp.), discussions on relationships of ferrets to prairie dog habitats at several spatial scales (from individual burrows to patches of burrow systems) and a general treatise on the status of black-footed ferret recovery.

Wyoming

Use of multi-opening burrow systems by black-footed ferrets

Multi-opening burrow systems constructed by prairie dogs (Cynomys) ostensibly provide escape routes when prairie dogs are pursued by predators capable of entering the burrows, such as black-footed ferrets (Mustela nigripes), or by predators that can rapidly dig into the tunnels, such as American badgers (Taxidea taxus). Because badgers also prey on ferrets, ferrets might similarly benefit from multi-opening burrow systems. Using an air blower, white-tailed prairie dog (Cynomys leucurus) burrow openings were tested for connectivity on plots occupied by black-footed ferrets and on randomly selected plots in Wyoming. Significantly more connected openings were found on ferret-occupied plots than on random plots. Connected openings might be due to modifications by ferrets in response to plugging by prairie dogs, due to selection by ferrets for complex systems with multiple openings that are already unobstructed, or simply due to ferrets lingering at kill sites that were multi-opening systems selected by their prairie dog prey.

Wyoming

Black-footed ferret digging activity in summer

Black-footed ferrets (Mustela nigripes) excavate soil from prairie dog (Cynomys spp.) burrows, thereby creating characteristic soil deposits at burrow openings. These soil deposits have been observed only rarely in summer. We monitored adult ferrets during June–October of the years 2007 and 2008 on a 452-ha colony of black-tailed prairie dogs (Cynomys ludovicianus) in the Conata Basin, South Dakota. We located and identified ferret excavations during nighttime spotlight surveys for ferrets and daytime sampling of prairie dog burrow openings around locations where ferrets were located via spotlight. We accumulated 48 observations of in-process or recently completed ferret excavations during spotlight surveys (21 in 2007, 27 in 2008) and located 51 diggings during daytime burrow sampling (25 in 2007, 26 in 2008). We located diggings during 5.5% of spotlight observations, most frequently in July–August. These results collectively suggest ferrets may frequently excavate soil in summer, because prairie dogs frequently use soil to plug burrow openings and tunnels in defense against ferrets. Prairie dogs might frequently destroy soil deposits left by ferrets during summer, thereby reducing detection of diggings by biologists.

South Dakota

Digging behaviors of radio-tagged black-footed ferrets near Meeteetse, Wyoming, 1981-1984

Intensive radio-tracking during August–December enabled us to collect detailed information on digging behaviors of a small sample of black-footed ferrets ( Mustela nigripes ) occupying colonies of white-tailed prairie dogs ( Cynomys leucurus ). A sample of 33 prairie dogs, also radio-tagged, progressively ceased aboveground activity during late summer and fall, presumably as they descended into burrows to hibernate. Most of the time ferrets spent digging was in November–December when >95% of the radio-tagged prairie dogs were inactive, suggesting that digging was primarily to excavate hibernating prey. Although 43.9% of the burrow openings were estimated to be in large mounds, which are common on colonies of white-tailed prairie dogs, all of a sample of 17 deposits of soil (diggings) made by ferrets were excavated at small mounds or nonmounded openings. The average duration of 23 nocturnal sessions of digging by ferrets was 112.2 minutes. A digging session consisted of multiple bouts of soil movement typically lasting about 5 min, and sessions were separated by pauses above- or belowground lasting several minutes. Bouts of moving soil from a burrow involved round-trips of 12.5–30.3 s to remove an average of 35 cm 3 of soil per trip. These digging bouts are energetically costly for ferrets. One female moved 16.8 kg of soil an estimated 3.3 m during bouts having a cumulative duration of 178 minutes, removing a soil plug estimated to be 178 cm long. Increasing evidence suggests that some behaviors of ferrets and prairie dogs are coevolutionary responses between this highly specialized predator and its prairie dog prey.

Wyoming

Black-footed ferrets and recreational shooting influence the attributes of black-tailed prairie dog burrows

Black-tailed prairie dogs (Cynomys ludovicianus) plug burrows occupied by black-footed ferrets (Mustela nigripes), and they also plug burrows to entomb dead prairie dogs. We further evaluated these phenomena by sampling connectivity and plugging of burrow openings on prairie dog colonies occupied by ferrets, colonies where recreational shooting was allowed, and colonies with neither shooting nor ferrets. We counted burrow openings on line surveys and within plots, classified surface plugging, and used an air blower to examine subsurface connectivity. Colonies with ferrets had lower densities of openings, fewer connected openings (suggesting increased subsurface plugging), and more surface plugs compared to colonies with no known ferrets. Colonies with recreational shooting had the lowest densities of burrow openings, and line-survey data suggested colonies with shooting had intermediate rates of surface plugging. The extent of surface and subsurface plugging could have consequences for the prairie dog community by changing air circulation and escape routes of burrow systems and by altering energetic relationships. Burrow plugging might reduce prairie dogs' risk of predation by ferrets while increasing risk of predation by American badgers (Taxidea taxus); however, the complexity of the trade-off is increased if plugging increases the risk of predation on ferrets by badgers. Prairie dogs expend more energy plugging and digging when ferrets or shooting are present, and ferrets increase their energy expenditures when they dig to remove those plugs. Microclimatic differences in plugged burrow systems may play a role in flea ecology and persistence of the flea-borne bacterium that causes plague (Yersinia pestis).

Western North American Naturalist

Patterns of surface burrow plugging in a colony of black-tailed prairie dogs occupied by black-footed ferrets

Black-tailed prairie dogs ( Cynomys ludovicianus ) can surface-plug openings to a burrow occupied by a black-footed ferret ( Mustela nigripes ). At a coarse scale, surface plugs are more common in colonies of prairie dogs occupied by ferrets than in colonies without ferrets. However, little is known about spatial and temporal patterns of surface plugging in a colony occupied by ferrets. In a 452-ha colony of black-tailed prairie dogs in South Dakota, we sampled burrow openings for surface plugs and related those data to locations of ferrets observed during spotlight surveys. Of 67,574 burrow openings in the colony between June and September 2007, 3.7% were plugged. In a colony-wide grid of 80 m × 80 m cells, the occurrence of surface plugging (≥1 opening plugged) was greater in cells used by ferrets (93.3% of cells) than in cells not observably used by ferrets (70.6%). Rates of surface plugging (percentages of openings plugged) were significantly higher in cells used by ferrets (median = 3.7%) than in cells without known ferret use (median = 3.2%). Also, numbers of ferret locations in cells correlated positively with numbers of mapped surface plugs in the cells. To investigate surface plugging at finer temporal and spatial scales, we compared rates of surface plugging in 20-m-radius circle-plots centered on ferret locations and in random plots 1–4 days after observing a ferret (Jun–Oct 2007 and 2008). Rates of surface plugging were greater in ferret-plots (median = 12.0%) than in random plots (median = 0%). For prairie dogs and their associates, the implications of surface plugging could be numerous. For instance, ferrets must dig to exit or enter plugged burrows (suggesting energetic costs), and surface plugs might influence microclimates in burrows and consequently influence species that cannot excavate soil (e.g., fleas that transmit the plague bacterium Yersinia pestis ).

South Dakota

Importance of lunar and temporal conditions for spotlight surveys of adult black-footed ferrets

Black-footed ferrets (Mustela nigripes) spend most daylight hours underground in prairie dog (Cynomys) burrows and exhibit aboveground movements primarily at night. Moonlight can influence the activity patterns of ferrets and, consequently, might influence the efficiency of spotlight surveys used by biologists to monitor ferret populations. We related detection of adult ferrets during postbreeding spotlight surveys to lunar and temporal conditions. We most frequently located ferrets during surveys in which the moon breached the horizon. The data suggested intersexual differences in response to moonlight. We located male ferrets most frequently during nights with greater moon illumination, but we did not detect a correlation between moon illumination and spotlight detection of female ferrets. In general, moonlight could facilitate aboveground navigation by ferrets. However, it seems activity under bright moonlight could be costly for female ferrets while they raise young. Detection of ferrets also varied among months. We detected female ferrets most frequently in August–September, when mothers increase hunting efforts to acquire prey for growing offspring (kits). Detection of adult female ferrets declined in October, when kits were likely independent of their mother. We located male ferrets most frequently in September–October, when males might increase activity to monitor female ferrets and male competitors. Consideration of lunar and temporal influences and standardization of postbreeding surveys could enhance site-specific assessment of reintroduction success and across-site assessment of species recoveiy progress. We suggest that postbreeding surveys for ferrets should be enhanced by concentrating efforts in August–September during moonlit nights when the moon is above the horizon.

Western North American Naturalist

Resource selection models are useful in predicting fine-scale distributions of black-footed ferrets in prairie dog colonies

Wildlife-habitat relationships are often conceptualized as resource selection functions (RSFs)—models increasingly used to estimate species distributions and prioritize habitat conservation. We evaluated the predictive capabilities of 2 black-footed ferret ( Mustela nigripes ) RSFs developed on a 452-ha colony of black-tailed prairie dogs ( Cynomys ludovicianus ) in the Conata Basin, South Dakota. We used the RSFs to project the relative probability of occurrence of ferrets throughout an adjacent 227-ha colony. We evaluated performance of the RSFs using ferret space use data collected via postbreeding spotlight surveys June–October 2005–2006. In home ranges and core areas, ferrets selected the predicted "very high" and "high" occurrence categories of both RSFs. Count metrics also suggested selection of these categories; for each model in each year, approximately 81% of ferret locations occurred in areas of very high or high predicted occurrence. These results suggest usefulness of the RSFs in estimating the distribution of ferrets throughout a black-tailed prairie dog colony. The RSFs provide a fine-scale habitat assessment for ferrets that can be used to prioritize releases of ferrets and habitat restoration for prairie dogs and ferrets. A method to quickly inventory the distribution of prairie dog burrow openings would greatly facilitate application of the RSFs.

South Dakota

Fine-scale habitat use of reintroduced black-footed ferrets on prairie dog colonies in New Mexico

Black-footed ferrets ( Mustela nigripes ) are among the most endangered animals in North America. Reintroductions of captive-born ferrets onto prairie dog ( Cynomys spp.) colonies are crucial to the conservation of the species. In September 2007, captive-born ferrets were released on a black-tailed prairie dog ( Cynomys ludovicianus ) colony at the Vermejo Park Ranch, New Mexico. Ferret kits experimentally released in areas of comparatively low and high prairie dog burrow densities were located via spotlight surveys. Some maturing ferret kits were subsequently translocated to areas of low and high burrow densities on nearby prairie dog colonies. For 2 months, fine-scale habitat use was quantified by mapping all burrow openings within a 30-m radius of each ferret location. Spatial statistics accounted for autocorrelation in the burrow densities in areas used by ferrets. It was hypothesized that ferrets would select areas of high burrow densities within colonies; however, burrow densities in areas used by ferrets were generally similar to the available burrow densities. Because ferrets used areas with burrow densities similar to densities available at the colony level and because of the potential energetic benefits for ferrets using areas with high burrow densities, releasing ferrets on colonies with high burrow densities might increase reintroduction success.

New Mexico

Salinity of the Little Colorado River in Grand Canyon confers anti-parasitic properties on a native fish

Water in the Little Colorado River within Grand Canyon is naturally high in salt (NaCl), which is known to prohibit development of external fish parasites such as Ich ( Ichthyophthirius multifiliis ). The naturally high salinity (>0.3%) of the Little Colorado River at baseflow may be one factor allowing survival and persistence of larval and juvenile humpback chub ( Gila cypha ) and other native fishes in Grand Canyon. We compared salinity readings from the Little Colorado River to those reported in the literature as being effective at removing protozoan parasites from fish. In laboratory tests, 10 juvenile roundtail chub ( Gila robusta ; 61–90 mm TL) were randomly placed into each of 12, 37-L aquaria filled with freshwater, water obtained from the Little Colorado River (0.3% salinity), or freshwater with table salt added until the salinity reached 0.3%. Roundtail chub was used as a surrogate for humpback chub in this study because the species is not listed as endangered but is morphologically and ecologically similar to humpback chub. All roundtail chub infected with Ich recovered and survived when placed in water from the Little Colorado River or water with 0.3% salinity, but all experimental fish placed in freshwater died because of Ich infection. The naturally high salinity of the Little Colorado River at baseflow (0.22%–0.36%), appears sufficiently high to interrupt the life cycle of Ich and may allow increased survival of larval and juvenile humpback chub relative to other areas within Grand Canyon.

Arizona

Associations of benthic macroinvertebrate assemblages with environmental variables in the upper Clear Creek watershed, California

Benthic macroinvertebrates are integral components of stream ecosystems and are often used to assess the ecological integrity of streams. We sampled streams in the upper Clear Creek drainage in the Klamath—Siskiyou Ecoregion of northwestern California in fall 2004 (17 sites) and 2005 (original 17 plus 4 new sites) with the objectives of documenting the benthic macroinvertebrate assemblages supported by the streams in the area, determining how those assemblages respond to environmental variables, assessing the biological condition of the streams using a benthic index of biotic integrity (IBI), and understanding the assemblages in the context of biodiversity of the ecoregion. We collected both reach-wide (RW) and targeted-riffle (TR) macroinvertebrate samples at each site. The macroinvertebrate assemblages were diverse, with over 150 genera collected for each sampling protocol. The macroinvertebrate assemblages appeared to be most responsive to a general habitat gradient based on stream size, gradient, flow, and dominance of riffles. A second important habitat gradient was based on elevation and dominance of riffles. A gradient in water quality based on concentrations of dissolved ions and metals was also important. Models based on these 3 gradients had Spearman's rank correlations with macroinvertebrate taxonomic composition of 0.60 and 0.50 for the TR and RW samples, respectively. The majority (>50%) of the sites were in good or very good biological condition based on IBI scores. The diversity of macroinvertebrate assemblages is associated with the diversity of habitats available in the Klamath—Siskiyou Ecoregion. Maintaining the aquatic habitats in good condition is important in itself but is also vital to maintaining biodiversity in this diverse and unique ecoregion.

California

Recreational trails as corridors for alien plants in the Rocky Mountains, USA

Alien plant species often use areas of heavy human activity for habitat and dispersal. Roads and utility corridors have been shown to harbor more alien species than the surrounding vegetation and are therefore believed to contribute to alien plant persistence and spread. Recreational trails represent another corridor that could harbor alien species and aid their spread. Effective management of invasive species requires understanding how alien plants are distributed at trailheads and trails and how their dispersal may be influenced by native vegetation. Our overall goal was to investigate the distribution of alien plants at trailheads and trails in the Rocky Mountains of Colorado. At trailheads, we found that although the number of alien species was less than the number of native species, alien plant cover ( x̄=50%) did not differ from native plant cover, and we observed a large number of alien seedlings in the soil seed bank, suggesting that alien plants are a large component of trailhead communities and will continue to be so in the future. Along trails, we found higher alien species richness and cover on trail (as opposed to 4 m from the trail) in 3 out of 4 vegetation types, and we observed higher alien richness and cover in meadows than in other vegetation types. Plant communities at both trailheads and trails, as well as seed banks at trailheads, contain substantial diversity and abundance of alien plants. These results suggest that recreational trails in the Rocky Mountains of Colorado may function as corridors that facilitate the spread of alien species into wildlands. Our results suggest that control of alien plants should begin at trailheads where there are large numbers of aliens and that control efforts on trails should be prioritized by vegetation type.

Colorado

Habitat interaction between two species of chipmunk in the Basin and Range Province of Nevada

Interspecies interactions can affect how species are distributed, put constraints on habitat expansion, and reduce the fundamental niche of the affected species. Using logistic regression, we analyzed and compared 174 Tamias palmeri and 94 Tamias panamintinus within an isolated mountain range of the Basin and Range Province of southern Nevada. Tamias panamintinus was more likely to use pinyon/ponderosa/fir mixed forests than pinyon alone, compared to random sites. In the presence of T palmeri , however, interaction analyses indicated T. panamintinus was less likely to occupy the mixed forests and more likely near large rocks on southern aspects. This specie s-by-habitat interaction data suggest that T. palmeri excludes T panamintinus from areas of potentially suitable habitat. Climate change may adversely affect species of restricted distribution. Habitat isolation and species interactions in this region may thus increase survival risks as climate temperatures rise.

Nevada

Distribution and movement of Big Spring spinedace ( Lepidomeda mollispinis pratensis ) in Condor Canyon, Meadow Valley Wash, Nevada

Big Spring spinedace (Lepidomeda mollispinis pratensis) is a cyprinid whose entire population occurs within a section of Meadow Valley Wash, Nevada. Other spinedace species have suffered population and range declines (one species is extinct). Managers, concerned about the vulnerability of Big Spring spinedace, have considered habitat restoration actions or translocation, but they have lacked data on distribution or habitat use. Our study occurred in an 8.2-km section of Meadow Valley Wash, including about 7.2 km in Condor Canyon and 0.8 km upstream of the canyon. Big Spring spinedace were present upstream of the currently listed critical habitat, including in the tributary Kill Wash. We found no Big Spring spinedace in the lower 3.3 km of Condor Canyon. We tagged Big Spring spinedace ≥70 mm fork length (range 70–103 mm) with passive integrated transponder tags during October 2008 (n = 100) and March 2009 (n = 103) to document movement. At least 47 of these individuals moved from their release location (up to 2 km). Thirty-nine individuals moved to Kill Wash or the confluence area with Meadow Valley Wash. Ninety-three percent of movement occurred in spring 2009. Fish moved both upstream and downstream. We found no movement downstream over a small waterfall at river km 7.9 and recorded only one fish that moved downstream over Delmue Falls (a 12-m drop) at river km 6.1. At the time of tagging, there was no significant difference in fork length or condition between Big Spring Spinedace that were later detected moving and those not detected moving. We found no significant difference in fork length or condition at time of tagging of Big Spring spinedace ≥70 mm fork length that were detected moving and those not detected moving. Kill Wash and its confluence area appeared important to Big Spring spinedace; connectivity with these areas may be key to species persistence. These areas may provide a habitat template for restoration or translocation. The lower 3.3 km of Meadow Valley Wash in Condor Canyon may be a good candidate section for habitat restoration actions.

Nevada

Insect prey eaten by Hoary Bats ( Lasiurus cinereus ) prior to fatal collisions with wind turbines

Wind turbines are being deployed all across the world to meet the growing demand for energy, and in many areas, these turbines are causing the deaths of insectivorous migratory bats. One of the hypothesized causes of bat susceptibility is that bats are attracted to insects on or near the turbines. We examined insect remains in the stomachs and intestines of hoary bats ( Lasiurus cinereus ) found dead beneath wind turbines in New York and Texas to evaluate the hypothesis that bats die while feeding at turbines. Most of the bats we examined had full stomachs, indicating that they fed in the minutes to hours leading up to their deaths. However, we did not find prey in the mouths or throats of any bats that would indicate the bats died while capturing prey. Hoary bats fed mostly on moths, but we also detected the regular presence of beetles, true bugs, and crickets. Presence of terrestrial insects in stomachs indicates that bats may have gleaned them from the ground or the turbine surfaces, yet aerial capture of winged insect stages cannot be ruled out. Our findings confirm earlier studies that indicate hoary bats feed during migration and eat mostly moths. Future studies on bat behaviors and insect presence at wind turbines could help determine whether feeding at turbines is a major fatality risk for bats.

Western North American Naturalist

Net primary productivity of subalpine meadows in Yosemite National Park in relation to climate variability

Subalpine meadows are some of the most ecologically important components of mountain landscapes, and primary productivity is important to the maintenance of meadow functions. Understanding how changes in primary productivity are associated with variability in moisture and temperature will become increasingly important with current and anticipated changes in climate. Our objective was to describe patterns and variability in aboveground live vascular plant biomass in relation to climatic factors. We harvested aboveground biomass at peak growth from four 64-m 2 plots each in xeric, mesic, and hydric meadows annually from 1994 to 2000. Data from nearby weather stations provided independent variables of spring snow water content, snow-free date, and thawing degree days for a cumulative index of available energy. We assembled these climatic variables into a set of mixed effects analysis of covariance models to evaluate their relationships with annual aboveground net primary productivity (ANPP), and we used an information theoretic approach to compare the quality of fit among candidate models. ANPP in the xeric meadow was negatively related to snow water content and thawing degree days and in the mesic meadow was negatively related to snow water content. Relationships between ANPP and these 2 covariates in the hydric meadow were not significant. Increasing snow water content may limit ANPP in these meadows if anaerobic conditions delay microbial activity and nutrient availability. Increased thawing degree days may limit ANPP in xeric meadows by prematurely depleting soil moisture. Large within-year variation of ANPP in the hydric meadow limited sensitivity to the climatic variables. These relationships suggest that, under projected warmer and drier conditions, ANPP will increase in mesic meadows but remain unchanged in xeric meadows because declines associated with increased temperatures would offset the increases from decreased snow water content.

Western North American Naturalist

Rodent-Mediated Interactions Among Seed Species of Differing Quality in a Shrubsteppe Ecosystem

Interactions among seeds, mediated by granivorous rodents, are likely to play a strong role in shrubsteppe ecosystem restoration. Past studies typically consider only pairwise interactions between preferred and less preferred seed species, whereas rangeland seedings are likely to contain more than 2 seed species, potentially leading to complex interactions. We examined how the relative proportion of seeds in a 3-species polyculture changes rodent seed selectivity (i.e., removal) and indirect interactions among seeds. We presented 2 rodent species, Peromyscus maniculatus (deer mice) and Perognathus parvus (pocket mice), in arenas with 3-species seed mixtures that varied in the proportion of a highly preferred, moderately preferred, and least preferred seed species, based on preferences determined in this study. We then conducted a field experiment in a pocket mouse—dominated ecosystem with the same 3-species seed mixtures in both “treated” (reduced shrub and increased forb cover) and “untreated” shrubsteppe. In the arena experiment, we found that rodents removed more of the highly preferred seed when the proportions of all 3 seeds were equal. Moderately preferred seeds experienced increased removal when the least preferred seed was in highest proportion. Removal of the least preferred seed increased when the highly preferred seed was in highest proportion. In the field experiment, results were similar to those from the arena experiment and did not differ between treated and untreated shrubsteppe areas. Though our results suggest that 3-species mixtures induce complex interactions among seeds, managers applying these results to restoration efforts should carefully consider the rodent community present and the potential fate of removed seeds.

Utah