Effects of exotic grasses on soil seed banks in southeastern Arizona grasslands

Species richness in Madrean mixed-grass prairies dominated by native or exotic species in southeastern Arizona was characterized at the community and point scales using ten 1-m 2 quadrats nested within each of eight 1000-m 2 plots. In the 1000-m 2 plots average richness was significantly higher in oak savanna (OS, 121.0 species) than in exotic grassland on mesa tops (EMT, 52.0 species), whereas native grassland on mesa slopes (NMS, 92.5 species) and native grassland on mesa tops (NMT, 77.0 species) did not differ significantly in richness from OS or EMT. When richness was partitioned by life form, EMT was notably poorer than other community types in species of perennial grasses, perennial herbs, and summer annuals. In the 1-m 2 quadrats, OS (21.2 species), NMS (20.9 species), and NMT (20.7 species) were significantly richer than EMT (5.9 species). Cover in 1-m 2 plots was significantly higher in EMT than in NMT, NMS, or OS. Species richness at the point scale showed a unimodal relation to canopy cover, with cover accounting for 30% of the variation in number of species in 1-m2 quadrats. Competitive exclusion and allelopathy have perhaps limited species richness at the point scale in exotic grassland. There was no evidence of a species-pool effect between point and community scales, but such an effect between community and landscape scales was supported. Madrean mixed-grass prairies are landscapes with high species richness in comparison to other grassland types in North America, providing a large pool of potential colonizing species at the community scale. Beta-diversity (between communities) within the landscape of the Appleton-Whittell Research Ranch was consequently high despite a relative lack of habitat diversity.     

New approaches for sampling and modeling native and exotic plant species richness

We demonstrate new multi-phase, multi-scale approaches for sampling and modeling native and exotic plant species to predict the spread of invasive species and aid in control efforts. Our test site is a 54,000-ha portion of Rocky Mountain National Park, Colorado, USA. This work is based on previous research wherein we developed vegetation sampling techniques to identify hot spots of diversity, important rare habitats, and locations of invasive plant species. Here we demonstrate statistical modeling tools to rapidly assess current patterns of native and exotic plant species to determine which habitats are most vulnerable to invasion by exotic species. We use stepwise multiple regression and modified residual kriging to estimate numbers of native species and exotic species, as well as probability of observing an exotic species in 30 × 30-m cells. Final models accounted for 62% of the variability observed in number of native species, 51% of the variability observed in number of exotic species, and 47% of the variability associated with observing an exotic species. Important independent variables used in developing the models include geographical location, elevation, slope, aspect, and Landsat TM bands 1-7. These models can direct resource managers to areas in need of further inventory, monitoring, and exotic species control efforts.     

Leafhopper assemblages on native and reseeded grasslands in southwestern Montana

Using sweep samples, we surveyed leafhoppers (Homoptera: Cicadellidae) on grassland sites in the Gallatin Valley of Montana during 1988 and 1991. We sampled 12 sites representing 2 habitat types defined by their dominant plant species in an undisturbed state ( Stipa comata / Bouteloua gracilis and Festuca idahoensis / Agropyron spicatum ). At half of the sites the native plant communities were present, whereas the remainder had been reseeded with either Agropyron spicatum (to replace the S. comata / B. gracilis assemblage ) or Bromus inermis (to replace the F. idahoensis / A. spicatum assemblage). We found at least 66 species of leafhoppers among 44,428 adults collected. Seven taxa comprised 83% of all individuals collected: Doratura stylata (26%), Ceratagallia spp. (18%), Endria inimica (17%), Orocastus perpusillus (7%), Sorhoanus spp. (6%), Athysanella spp. (5%), and Psammotettix lividellus (4%). Sites with similar vegetation had broadly similar leafhopper assemblages, and assemblages differed most between the relatively xeric Stipa comata / Bouteloua gracilis sites and the more mesic sites dominated by Bromus inermis . The composition of a leafhopper assemblage at a site tended to be more similar to those on noncontiguous sites with the same overall vegetation than to those on contiguous sites with different vegetation. These patterns are likely related to the fact that many Cicadellidae are host specialists. In fact, variation in abundance of some of the most common leafhopper taxa on our sites was correlated with the percent cover of their known host plants. Our analyses of the leafhopper assemblages generally support the contention that terrestrial plant associations are among the more useful indicators of insect community composition.     

Natural enemy assemblages on native and reseeded grasslands in southwestern Montana: a family-level analysis

We conducted a 2-year survey, using sweep sampling and family-level taxon identification, of the predatory and parasitoid insects on grassland sites in the Gallatin Valley of southwestern Montana. The 25 sites were divided into 4 habitat classes: 2 native habitat types ( Stipa comata/Bouteloua gracilis and Festuca idahoensis/Agropyron spicatum ) and 2 that had been reseeded with either crested wheatgrass ( Agropyron cristatum ) or smooth brome ( Bromus inermis ). Our major goal was to make quantitative comparisons of the abundance of insects among native and reseeded habitats. Of 51 families in 5 insect orders identified, 7 Hymenoptera (Encyrtidae, Braconidae, Ichneumonidae, Pteromalidae, Eulophidae, Scelionidae, and Torymidae), 3 Hemiptera (Lygaeidae, Nabidae, and Reduviidae), 1 Coleoptera (Coccinellidae), and 1 Diptera (Asilidae) comprised 90% of the natural enemies sampled. Ordination analyses provided no strong evidence that the 4 habitat classes contained distinct overall natural enemy communities. However, contiguous native and reseeded sites usually had relatively different overall natural enemy assemblages, suggesting that vegetation was often a more important correlate of community composition than was close spatial proximity of sites. Furthermore, several common families exhibited differential abundances across habitat classes in one or both years. For example, in 1989, Eulophidae, Pteromalidae, and Torymidae were more abundant on native Festuca/Agropyron sites, whereas Encyrtidae and Nabidae were more abundant on Festuca/Agropyron sites reseeded with Bromus inermis . Although analyses of insect assemblages classified to the family level provide somewhat limited information on functional ecological differences among habitats, they allow one to survey a broad array of taxa to identify focal groups for future conservation and land management studies.     

Defining and evaluating exotic species: issues for Yellowstone park policy

Exotics are species that are foreign to an ecological assemblage in the sense that they have not significantly adapted to resident biota or to local abiotic conditions, and resident species have not significantly adapted to them. Although they need not be human introduced nor damaging, when they are, a negative appraisal of such exotic species can be justified. Human introduction of exotics into natural systems typically increases human influence over those systems, thus diminishing their wildness. Valuing nature for its wildness is a rationale for the national parks policy of letting nature take its course. Thus, Yellowstone Park has a strong reason for removing human-introduced exotics and for welcoming naturally migrating exotics. Disvaluing exotics that are neither human introduced nor damaging simply because they are foreign smacks of xenophobia. But given that wanton human mixing of species threatens to homogenize the earths biological communities, biological nativism is justified as a way to preserve the diversity between such communities.     

Plant invasions in protected areas at multiple scales : Linaria vulgaris (Scrophulariaceae) in the West Yellowstone area

Invasive alien plants have long been recognized as a threat to low-elevation, disturbed environments, but the case of Linaria vulgaris Mill. in Yellowstone National Park and Gallatin National Forest shows that invasions can also spread to high-elevation natural reserves. Because invasions in protected areas are a product of complex processes occurring over a broad range of scales, we argue that a multi-scale research approach is needed to capture both patterns and potential mechanisms of the invasion process. Mapping L. vulgaris at the landscape scale, we found the species occupying a broad range of sites, apparently originating from just 2 historical sources, colonizing both human-caused and natural disturbances. Analyzed at the stand scale, patches tend to aggregate in newly invaded areas and disperse in heavily infested areas. The data suggest that patches grow in size by clonal growth and in number by creation of new satellite patches. Radial patch growth rates are related to site characteristics. Clonal patch scale analysis shows that ramet densities and Linaria 's effects on native plants are highest in patch centers. Both mean ramet height and reproductive vs. vegetative ramet height ratio are higher in patch cores. These results suggest that L. vulgaris may displace natural vegetation by maintaining vigor even in large and old clonal patches. Our results confirm that L. vulgaris is a significant threat to native biodiversity in open, human- or naturally disturbed environments in protected areas of the Rocky Mountains. A multi-scale method can allow managers to better understand patterns of invasion and prioritize management activities to control invasive alien plants, especially in heterogeneous protected area landscapes.     

Horizontal and vertical diameter of burrows of five small mammal species in southeastern Idaho

Burrow diameters of five small mammal species, Townsend's ground squirrel ( Spermophilus townsendii ), Wyoming ground squirrel ( S. elegans ), Ord's kangaroo rat ( Dipodomys ordii ), montane vole ( Microtus montanus ), and deer mouse ( Peromyscus maniculatus ), were examined. Burrow cross sections were noncircular for all species with horizontal diameters 1.2-1.6 times wider than vertical diameters. Montane vole and deer mouse burrows were the smallest diameter, burrows of Wyoming and Townsend's ground squirrels were the largest, and kangaroo rat burrows were intermediate. Soil bulk density and texture significantly affected burrow diameters of montane voles and deer mice but not the other three species.     

Effects of habitat type and degradation on avian species richness in Great Basin riparian habitats

The overwhelming majority of bird species in the Great Basin region are found in riparian habitats. However, most previous research on the impact of change in habitat condition through degradation on these bird communities failed to account for the large intersite differences, in both habitat type and extent of degradation. We examined songbird communities in 4 riparian habitat types (meadows, willow-birch-, and aspen-dominated forest stands) during summers 1994 (last year of a 7-yr drought) and 1995 (following the 6th wettest winter recorded) in the Toiyabe Mountain Range of central Nevada. Habitat degradation significantly influenced bird species richness in riparian areas, but the impact was dependent upon habitat type. While meadow bird communities were affected adversely by habitat degradation, with significant drops in species richness on degraded sites, bird species richness in forested riparian habitats was consistently greater on degraded sites. Data for the 6 most common species seen during our study indicated that degradation may have influenced distribution of American Robins ( Turdus migratorius ) and Yellow Warblers ( Dendroica petechia ), but habitat type was the best predictor of abundance for House Wrens ( Troglodytes aedon ), Red-naped Sapsuckers ( Sphyrapicus nuchalis ), Warbling Vircos ( Vireo gilvus ), and Brewer's Blackbirds ( Euphagus cyanocephalus ). Avian species diversity in meadow habitats may be linked to moisture levels during specific times of the year. Diversity increased during the pre-migratory period of the dry year (1994) when compared with that of the breeding season, but was unchanged in the wet year (1995).     

Asian tapeworm ( Bothriocephalus acheilognathi ) in native fishes from the Little Colorado River, Grand Canyon, Arizona

Examination of gastrointestinal tracts of native cyprinids from the Little Colorado River (LCR) in Grand Canyon, Arizona, 1990-1994, revealed varying rates of prevalence and infrapopulation levels of Asian tapeworm ( Bothriocephalus acheilognathi ). Mean prevalence was 28% (range 0-78%) in humpback chub ( Gila cypha ) and 8% (range 0-46%) in speckled dace ( Rhinichthys osculus ), with infrapopulations as high as 46 and 28, respectively. We also note Asian tapeworm infection of nonnatives common carp ( Cyprinus carpio ), fathead minnow ( Pimephales promelas ), and plains killifish ( Fundulus zebrinus ) from the LCR. Reported pathogenic and chronic effects of this cestode to its definitive hosts add concern for the status of the Grand Canyon population of the federally endangered humpback chub. The rapidity which Asian tapeworm has spread to different drainages of the Colorado River Basin likely portends an eventual cosmopolitan basin distribution in lower elevations suitable to the parasite's thermophilic life history. Such biotic changes must be considered among the most serious threats to conservation and recovery of native fish populations.     

Genetic analysis of the rare species Salix arizonica (Salicacea) and associated willows in Arizona and Utah

Management decisions affecting the rare plant Arizona willow ( Salix arizonica ) will be aided by understanding genetic similarities among populations of this species. Random Amplified Polymorphic DNA (RAPD) analysis was conducted on 20 populations of S. arizonica , 12 populations of 5 congeners, and 2 samples of outgroup, Populus tremuloides . A phenogram based on DNA markers shows clear separation of populations of S. arizonica from those of co-occurring willow species, but similarity is low (～ 37%) between Utah and Arizona populations of S. arizonica . Evaluation of the relationship of habitat characteristics and geographic distance to genetic similarity reveals that environment and genetic similarity are poorly correlated. Considering Arizona versus Utah populations, we found a significant negative relationship between geographic distance and genetic similarity ( r = 0.936), but no significant relationship between interpopulation distance and genetic similarity within Arizona or Utah. The wide geographic disjunction of S. arizonica populations in Utah and Arizona appears to have existed for a long period during which genetic drift, random mutations, and selection for somewhat different habitats have pushed the 2 regional complexes along separate evolutionary trajectories. Preservation of genetic variation within S. arizonica will require protection of multiple populations in Arizona and Utah.     

Trichoptera and other macroinvertebrates in springs of the Great Basin: species composition, richness, and distribution

Detailed surveys of aquatic invertebrates in 28 springs along the western border of the Great Basin were made using pan traps, emergence traps, black lights, and benthic sampling devices from 1994 to 1998. Although all macroinvertebrate groups were collected, caddisflies (Insecta: Trichoptera) were sampled most intensively. Fifty-eight species of caddisflies were collected from the 28 springs, with up to 18 species from a single spring. Although several springs had very similar physicochemical characteristics, none had identical trichopteran species composition. Using Wards minimum variance clustering technique, and with all macroinvertebrates identified to the lowest possible level, we found 3 taxa assemblages. These assemblages occurred in springs with the following physical conditions: (1) warm water (15.9° C), low elevation (1794 m); (2) cold water (8.5° C), mid-elevation (2334 m); and (3) cold water (6.6° C), high elevation (1794 m). Although the warm water group had a distinct assemblage of invertebrates (amphipods, gastropods, Trichoptera), distinct assemblages did not separate the 2 cold water groups. Discriminant analysis indicated that temperature, conductivity, alkalinity, and elevation were most responsible for discriminating group 1 from groups 2 and 3; temperature and elevation distinguished the latter 2 groups. Spring permanence, lack of disturbance, and cold water temperatures were the factors most responsible for explaining higher species richness of Trichoptera.     

Selection of dwarf mistletoe-infected ponderosa pines by Ips species (Coleoptera, Scolytidae) in northern Arizona

Following an Ips bark beetle outbreak in 2002, mortality of ponderosa pine ( Pinus ponderosa Douglas ex C. Lawson) was evaluated in 2 study areas infested with southwestern dwarf mistletoe ( Arceuthobium vaginatum [Willd.] Presl subsp. cryptopodum [Engelm.] Hawksw. & Wiens) in the Coconino and Tonto National Forests, Arizona. A pairwise comparison of dwarf mistletoe ratings for live and dead ponderosa pines was conducted to determine whether dead ponderosa pines had higher dwarf mistletoe ratings than pines that were not attacked. In both study areas, dead ponderosa pines had significantly higher dwarf mistletoe ratings, indicating an association between the severity of dwarf mistletoe infection and susceptibility to attack by Ips spp. We suggest that the probability of ponderosa pine mortality is greater in stands severely infested with southwestern dwarf mistletoe in northern Arizona.     

Long-term interactions of climate, productivity, species richness, and growth form in relictual sagebrush steppe plant communities

Vegetation trends due to climatic changes are difficult to separate from disturbances caused by varying land uses. To separate climatic influences from livestock grazing and fire disturbances within sagebrush steppe, we compared vegetation structure and productivity during 2 periods (10-year sequences of data from the late 1950s to the late 1960s and 3 years in the early 1990s) at 12 stands within 3 relict areas in or near the Great Rift of southern Idaho. Yearto- year fluctuations in annual net aboveground phytomass accumulation (ANAPA) were considerable in response to varying climate during both periods. More importantly, an apparently significant increase in ANAPA was observed during the latter study period. However, this apparent increase may have been due to the unusually wet conditions of 1993. Throughout both study periods, ANAPA was even more positively correlated with species richness than with precipitation, although both influences were statistically significant. Aggregation of these data by plant growth forms showed that shrubs, particularly sagebrushes, have progressively increased their proportion relative to herbaceous understory. This directional (successional) change in the proportional contributions of growth forms to ANAPA was probably due more to long-term absence of fire at these isolated locations than to any detectable climate change or lack of grazing. This finding reduces our confidence in the use of such areas as reference sites for monitoring or as living examples for restoration efforts, but it does increase our understanding of vegetation dynamics within sagebrush steppe. ACRONYMS. ANAPA—annual net aboveground phytomass accumulations, ANOVA—analysis of variance, ANPP—aboveground net primary production, COV—coefficient of variation, CY—crop year, CYP—crop-year precipitation, ES—ecological site, MANOVA—multivariate analysis of variance, MAT—mean annual temperature, NRCS—Natural Resources Conservation Service, PDSI—Palmer Drought Severity Index, RUE—rainfall use efficiency.     

Spatial niche variation in two sympatric species of Bokermannohyla (Anura: Hylidae) in southeastern Brazil

We studied patterns of microhabitat use by adults of two sympatric Bokermannohyla species at the Reserva Particular do Patrimônio Natural Santuário do Caraça, southeastern Brazil. We selected three streams, one where both species occurred in syntopy and the other two where each one occurred alone. We sampled 150-m transects in each stream throughout 1 year, recording microhabitat features for each frog located (substrate type, height and distance from water). Microhabitat availability varied between dry and wet seasons in all streams, and overall microhabitat diversity changed in two streams. Bokermannohyla nanuzae seemed to have a niche contraction in the presence of Bokermannohyla martinsi, but this only happened during the dry season. Microhabitat requirements during the wet season may be closely linked to similar reproductive needs that probably represent a strong selective pressure, forcing niche overlap.     

Microvelia rasilis Drake in Arizona: a species new to the United States (Heteroptera: Veliidae)

Microvelia rasilis Drake was taken in Montezuma Well, Yavapai Co., Arizona, along with Microvelia hinei Drake and Hydrometra aemula Drake (Heteroptera: Hydrometridae). These are all new to this locality; however, the latter two species are previously known from Arizona.     

Recurrence of Mexican long-tongued bats ( Choeronycteris mexicana ) at historical sites in Arizona and New Mexico

The Mexican long-tongued bat ( Choeronycteris mexicana ) is a nectar-eating species that seasonally inhabits the southwestern United States. Since 1906, fewer than 1500 individuals of C. mexicana have been documented throughout the range of the species. We conducted a field survey in Arizona and New Mexico during summer 1999 to check historically occupied areas for recurrence of C. mexicana . We observed C. mexicana occupying a majority (75%, n = 18) of visited sites. Multiple individuals were observed at many sites, including young-of-year. Choeronycteris mexicana roosted in lighted areas close to entrances within mine adits, abandoned buildings, wide rock crevices, and caves. All occupied sites in Madrean evergreen woodlands or semidesert grasslands where species of Agave were present. Most sites were located near a water source and, with the exception of a single site, near areas of riparian vegetation. Sites at which we did not encounter C. mexicana were frequently disturbed, difficult to search, or historically occupied by single individuals. Based on the relatively high rate of bat recurrence, we do not believe that populations of C. mexicana in the region have declined dramatically over the past several decades.     

You are what you eat: native versus exotic Crotalaria species (Fabaceae) as host plants of the Ornate Bella Moth,Utetheisa ornatrix (Lepidoptera: Erebidae: Arctiinae)

Crotalaria plants and Utetheisa ornatrix are closely linked to each other: the larvae destroy the seeds, while the moth depends on hostplants for alkaloids. To better understand the ongoing co-evolution, the present study examines how native hostplants compare to exotic ones. Leaf-feeding on Crotalaria pumila, C. rotundifolia, and C. incana, native to the moth’s range, led to faster larval development than on the exotic C. lanceolata, C. spectabilis, and C. pallida. Seed-feeding on all species of Crotalaria led to accelerated larval development and a resultant larger adult moth, and correlates with a higher nitrogen content in the plant tissues. These results add a novel dimension to the previous studies of reproductive biology of this model organism. In controlled settings, mature larvae showed preference for leaves of C. spectabilis over those of other species, perhaps due to the higher alkaloid content. Differences in morphology and phenology of Crotalaria determine the ecology of U. ornatrix populations in Florida. The introduction of novel hostplants, on which U. ornatrix can have a significant negative effect and which are of concern to humans as invasive toxic weeds, has greatly expanded the niche occupied by this moth. Possible co-evolution of Utetheisa sensu lato and plant defences in the genus Crotalaria is discussed.     

On Cowania and its intergeneric hybrids in Arizona

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A new genus and two new species of Parastenocarididae (Copepoda: Harpacticoida) from southeastern India

Indocaris gen. nov. with two new species, Indocaris imbricata sp. nov. and Indocaris inopinata sp. nov., and also for the already known Indocaris tirupatiensis (Ranga Reddy 2011a) comb. nov. – all from the groundwaters in peninsular India. The highly diagnostic synapomorphy of the new genus is a composite character associated with the male leg 4 basis: five or six prominent, imbricate, enlarged, petal-like spinules, arranged as a semi-whorl at the insertion of the endopod and increasing in size from internal to external. Another distinctive feature of the same appendage is that its one-segmented endopod is dilated or bulbous in the proximal half, produced distally into an incurved spiniform or horn-like structure about as long as the corresponding first exopodal segment, and ornamented with three or four fine spinules on the subproximal outer margin. The three species also share a unique constellation of other salient morphologic features, which along with the phylogenetic position of Indocaris gen. nov. within the family Parastenocarididae are discussed. Indocaris gen. nov. has closest phylogenetic affinity with the Neotropical Remaneicari Jakobi, 1972. A short note on the ecology and biogeography of the parastenocaridid species of the Indian subcontinent is provided besides a key for their identification.