Demographic and environmental relations of two rare Astragalus species endemic to Washington County, Utah: Astragalus holmgreniorum and A. ampullarioides

Two endemic locoweeds of Washington County, Utah, Astragalus holmgreniorum and A. ampullarioides , were recently listed as federally endangered plant species (Federal Register 2001). They both occur in few, small populations surrounded by increasing human activity in the St. George, Utah, area. Demographic and habitat studies have been ongoing since 1992 and results of those studies are presented here. Astragalus holmgreniorum (Holmgren locoweed) is presently limited to 1 larger population and 2 smaller, isolated populations on the eastern and western edges of its range. It is a short-lived perennial that grows primarily on the Virgin limestone member of the Moenkopi Formation. Density of A. holmgreniorum over the 10-year study averaged about 2 plants per m 2 . The number of living plants of this species perhaps never exceeds 10,000. In drought years A. holmgreniorum populations are as much as 95% smaller than in years with adequate water, and few plants produce flowers that successfully contribute to the seed bank. Astragalus ampullarioides (Shivwits locoweed) occurs in 4 populations where plant density fluctuated between 0.5 and 4.8 per m 2 during our study period (average = 2.6). This locoweed is restricted to clay soils of the Chinle Formation. Direct threats to both Astragalus species include widespread urbanization, road construction, population fragmentation, and off-highway vehicle use. Indirect threats include competition from aggressive, introduced annual species such as Bromus rubens, B. tectorum, Malcolmia africana , and Erodium cicutarium . Our data show that 3 of 4 species most closely associated with these Astragalus species are introduced. Studies of seed banks and reproduction biology, as well as continued monitoring of these Astragalus species, are critically needed. Studies of the consequences of competition from alien and native species on seedling establishment of these locoweeds are especially needed.     

Ecology and conservation of the endangered quillwort Isoetes sinensis in China

Isoetes sinensis Palmer is an East Asia endemic fern that faces impending extirpation from the expansive area of Mainland China. Presently, only two natural populations are known to exist in this vast region. Both populations have decreased by about 50% within the last 4 years. This reduction is most probably related to the decline in the size of the wetlands since reproductive activity, spore production, and recruitment were noticeably lower at the microsites occurring in smaller ponds. The decline in population size seems to be associated with succession from marsh to dry land at the habitats of both populations. The study revealed that individuals in both populations produced copious numbers of spores suggesting adequate fecundity, furthermore high levels of spore germination were obtained in laboratory experiments. However, inefficient spore dispersal appears to have contributed to the limited spread of the species. The sex ratio in both populations was male‐biased. Several factors are identified here as posing a threat to Isoetes sinensis populations in China, chiefly the loss of habitat due to infrastructure construction and competition with adventive weed species for establishment on available sites. Disturbance by introduced mammals during browsing and trampling by livestock farm workers, as well as tourists, further contribute to the diminishing numbers of Isoetes sinensis at the study sites.     

High foal mortality limits growth of a desert feral horse population in Nevada

A population of feral horses ( Equus caballus ) in the southern Great Basin Desert of Nevada was monitored from 1989 to 1998 to determine size, distribution, and population trends. All individual horses observed in the population were identified by unique markings during the first 2 yr of study, and most animals could be observed annually. During this study no new horses were identified in the population, indicating that no immigration occurred from outside populations. The population reached a high of 65 horses yearling or older in 1992 and declined each year thereafter, reaching a low of 36 horses in 1998. Estimated foal survival averaged < 12% over 8 yr. Only 11 horses were recruited into the population as yearlings or older animals during the study. Mountain lion predation is hypothesized as major factor limiting growth of this feral horse population.     

Vegetal change on a northern Utah foothill range in the absence of livestock grazing between 1948 and 1982

Reexamination of a semiarid foothill rangeland, first evaluated in 1948, indicated that secondary succession continues to shift toward a perennial grass-forb community formerly dominated by xeric shrubs, particularly big sagebrush ( Artemisia tridentata spp. vaseyana ). The direct role in livestock grazing in establishment and maintenance of shrub-dominant plant communities appears confirmed in the decline of shrubs upon cessation of livestock grazing in summer and continued browsing by mule deer in winter. The reduction of shrub forages on mule deer winter ranges is a major factor in population declines.     

Abundance and condition indices of coyotes on Hart Mountain National Antelope Refuge, Oregon

U.S. Fish and Wildlife Service staff at Hart Mountain National Antelope Refuge (HMNAR), southeastern Oregon, documented high pronghorn ( Antilocapra americana ) fawn mortality, subsequent low fawn recruitment, and declining pronghorn numbers from 1996 to 1999. Coyote ( Canis latrans ) predation was the primary cause, accounting for 60-85% of fawn mortalities each year, and fawns were not physiologically predisposed to predation. Therefore, we investigated certain coyote population parameters (age structure, survival, density, physiology) to evaluate how or if these factors influence coyote predation rates on pronghorn fawns. We captured 11 coyotes (5 male and 6 female) in December 1998. Age of captured animals ranged from 1.7 to 10.7 yrs (￣x = 5.0 years), and all coyotes appeared healthy upon capture. There were no known mortalities through December 1999. We estimated pre-whelping (December through February 1997-1999) density from howling surveys conducted within HMNAR to be 0.40-0.53 km -2 . Compared to other published studies, we found significant ( P ≤ 0.05) differences in selected blood parameters (e.g., blood urea nitrogen, total protein, white blood cell counts), indicating coyote nutrition may be marginal to deficient during winter at HMNAR. A high percentage of coyotes (91%) tested positive for serum-neutralizing antibodies to canine parvovirus. We judged that parasite ( Toxascaris spp., Alaria spp., Sarcocystis spp., and Isospora spp.) prevalence and intensity were not high enough to influence coyote condition. Based on our data, the coyote population at HMNAR is old aged, at a relatively high density, and stable, but their nutrition may be marginal to deficient during winter. Presently, we are unable to draw direct conclusions relating the parameters we sampled with predation rates by this unexploited coyote population.     

Morphological and genetic variation among populations of the rare Kachina daisy ( Erigeron kachinensis ) from southeastern Utah

Erigeron kachinensis , the Kachina daisy, is a rare species restricted to canyons in southeastern Utah. The species is known to exhibit low fecundity due to low percent fertilization of ovules and high percent abortion of fertilized ovules. Previous reproductive studies suggest that low fecundity is a consequence of small population size and inbreeding depression. This study examines genetic diversity within and among populations of E. kachinensis in Natural Bridges National Monument using enzyme electrophoresis. Field populations are found to have significantly different morphologies. However, morphological differences were less pronounced among populations grown in the greenhouse. The Kachina daisy exhibits levels of genetic variability in its populations similar to that of other outcrossed species. Genetic diversity statistics demonstrate that only 22.8% of genetic variation is distributed among populations. Genetic distance could not be correlated with geographic distance. Most of the populations showed significant deviation of fixation indices from zero for multiple loci. Observation of genotype frequencies demonstrates that populations are fixing on different genotypes and may be experiencing initial stages of genetic drift. Mean observed heterozygosity was 0.166 and was found to increase with increasing size and/or age in populations.     

Growth and demography of one population of the lizard Sceloporus mucronatus mucronatus

We report on growth and demography of Sceloporus mucronatus mucronatus , a lizard subspecies endemic to central Mexico. We characterize the life history of this subspecies, provide quantitative information relevant to conservation, and add to the growing literature on the diversity of life histories in the genus Sceloporus . We calculated body growth rates and fitted them to the Von Bertalanffy, the logistic-by-length, and the logistic-by-weight growth models. The Von Bertalanffy model provided the best fit, and we used it to analyze the growth pattern. Growth rates were similar during the 1st year of life in both sexes, but after that point males grew faster and reached maturity earlier (20 months) than females (31 months). We used a population projection matrix to model population dynamics during 2003–2004 and found a positive population growth rate (λ = 1.769). However, based on the projected stable size-class vector ( w ), this population does not appear to have reached stability, and it might be currently experiencing considerable interannual fluctuations. Elasticity values showed that the transition from the juvenile stage to the 1st adult stage was the vital rate that contributes the most to population growth rate, followed by fecundity and stasis of the 1st reproductive category. While total elasticities for demographic processes were similar, elasticities per size class showed the relatively high importance of small adults in comparison to juveniles and large adults. The restriction of this endemic subspecies to central Mexico, where human activities and consequent habitat destruction are increasing, demands further quantitative evaluation and monitoring of populations, even though our results indicate a potential for population growth.     

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.     

Predator-specific mortality of pronghorn on Yellowstone’s northern range

Yellowstone National Park supports a small population ( Antilocapra americana ). Some individuals migrate during summer to areas characterized by reduced visibility, mixed habitat types, and a diverse predator community. Across areas selected by migratory and nonmigratory pronghorn, we documented cause-specific mortality of adults and fawns and assessed relative risk of predation by various predators. Coyotes ( Canis latrans ) accounted for 56% of adult predation and up to 79% of fawn predation. Cougars ( Puma concolor ) and wolves ( Canis lupus ) accounted for additional predation of adults, while cougars, black bears ( Ursus americanus ), and Golden Eagles ( Aquila chrysaetos ) were documented killing fawns on rare occasions. Our results suggest that even when pronghorn are in the presence of multiple predator species, coyote predation on adults and fawns may predominate for populations inhabiting shrubsteppe habitat. However, the risk of predation by sympatric predators, particularly cougars, may be high for female pronghorn selecting mixed cover types during migration or for birthing purposes. While the direct effect of wolves on overall mortality was low, wolves may indirectly influence survival rates of adult females and fawns by altering the behavior and space use of sympatric predators, particularly coyotes.     

The reproductive ecology of the Tahoe sucker, Catostomus tahoensis, in Pyramid Lake, Nevada

The Tahoe sucker spawns in Pyramid Lake from April to August at lake temperatures of 11.7 to 22.7 C. The spawning population is comprised of a large lake spawning group and a numerically smaller river running group. The river running group is smaller in length and was not considered during this study. The sex ratio of sampled suckers significantly favored the females. This is the result of the longer life of females and greater mortality of males during spawning. &nbsp; Pyramid Lake Tahoe suckers reach sexual maturity at two to three years of age; however, those in Lake Tahoe do not mature until four or five years of age. The size at sexual maturity is different in both populations, which suggests that size or rate of growth rather than age determines sexual maturity. &nbsp; The fecundity of Tahoe suckers is positively correlated with fork length, weight, and age. Additional analysis showed that a better correlation occurred between fish size (either length or net weight) and total ovary weight. We believe that fish size is primarily correlated with total reproductive tissue produced and secondarily with fecundity. A comparison of the Pyramid Lake population and the Lake Tahoe population demonstrated that size, not age, is the most important determinant of Tahoe sucker fecundity.     

Age and population structure of Joshua trees ( Yucca brevifolia ) in the northwestern Mojave Desert

Many desert perennials are long-lived, but there are few data on ages or population structures of desert plants. We used 2 methods to estimate ages of a population of Yucca brevifolia (Joshua tree) in southwestern Utah from a 14-year census of plant sizes. Plant height at the 1st census ranged from 0.08 m to 6.0 m, and trees grew in height at a mean rate of 3.75 cm ? yr -1 . Plants also increased slowly in basal diameter (0.142 cm ? yr -1 ) and branch length (0.024 m ? yr -1 ), but basal diameter varied greatly from year to year, with many plants decreasing in diameter between sequential censuses. Forms of a nonlinear growth equation (the Richards function) did not fit the data on growth in height, but a simple linear relationship fit the data well. Using height as a linear predictor of age, we estimated that ages of a sample of 69 Joshua trees ranged from 300 years old, with 75% of the trees between 20 and 100 years and 11% over 150 years old. These data support the assumption that Joshua trees are indeed long-lived.     

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.     

Effects of convict cichlids on growth and recruitment of White River springfish

Observations of changes in population density of native White River springfish ( Crenichthys baileyi ) in Pahranagat Valley led to the following hypothesis: introduced convict cichlids ( Cichlasoma nigrofasciatum ) cause reduced growth and recruitment; cover reduces the magnitude of the effect. The hypothesis was tested by establishing sympatric and allopatric groups of the two species in experimental aquaria with and without cover. Change in volume (= mass) and length of the two species over a three-month period in spring 1986 and 1987 was measured and analyzed using 2 × 2 factorial analyses. Convict cichlids caused reduced growth and eliminated recruitment of springfish under the experimental conditions. Cover did not influence growth but positively affected recruitment of springfish in allopatry. It is likely that a portion of the reduced springfish population densities in nature can be attributed to adverse effects from introduced cichlids.     

Predation of artificial Sage Grouse nests in treated and untreated sagebrush

We measured predation on 120 artificial Sage Grouse ( Centrarcus urophasianus ) nests in montane sagebrush grassland in northern Utah. We examined nests in areas that had been chained and seeded 25 years previously (treated areas) and in areas that were untreated. Predation rates of artificial nests were higher in areas of untreated sagebrush, even though these areas had greater sagebrush cover, taller shrubs, and greater horizontal plant cover. These results differ from those previously hypothesized for treated sagebrush habitat and may reflect a greater abundance of other potential prey species, especially lagomorphs, in untreated areas that attracted greater densities of predators. In addition, over 80% of nests were depredated by mammals, which hunt using olfaction and are less likely than avian predators to be affected by nest cover. We conclude that, after treated sagebrush has recovered to some degree, predation rates of Sage Grouse nests may be lower in treated sagebrush. Consequently, factors other than nest predation (e.g., winter food, thermal cover, insects, perennial forb abundance) may be more important reasons for preserving mature sagebrush stands for Sage Grouse.     

Reproductive ecology of Dusky Flycatchers in montane meadows of the central Sierra Nevada

Avian species with expansive ranges or those that occupy more than one vegetative association may vary in aspects of their life histories across their ranges. The distribution of Dusky Flycatchers encompasses a variety of vegetative associations, including riparian communities. However, much of the literature on this species details studies conducted in upland areas. Our objectives were to describe the breeding ecology and fecundity of Dusky Flycatchers nesting in montane meadows of the central Sierra Nevada, California. We monitored 36 territories and located 37 Dusky Flycatcher nests in 8 meadows. Average clutch size was 3.9 eggs. Egg laying, incubation, and nestling stages were 4, 15.4, and 16.4 days, respectively. Eighteen nests successfully fledged young, with an average of 3.3 fledglings per successful nest. Nest success was 43% and nest predation was the leading cause of nest failure. Estimated annual fecundity was 1.62 fledglings per pair; however, because all renesting attempts were not located, this should be viewed as the minimum annual fecundity. Dusky Flycatchers we monitored may have had higher fecundity than those nesting in upland areas because riparian areas often have higher arthropod abundances. While the importance of riparian conservation to riparian-obligate bird species is obvious, our study indicates that these areas also may be of value to Dusky Flycatchers that breed in riparian areas and upland areas.     

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.     

Comparative population biology of Uca rapax (Smith, 1870) (Brachyura,Ocypodidae) from two subtropical mangrove habitats on the Brazilian coast

The structure of two populations of the fiddler crab Uca rapax in two subtropical mangrove habitats near Ubatuba, State of São Paulo, Brazil were compared. The size–frequency distribution, sex ratio, and recruitment were evaluated. Sampling was performed monthly from April 2001 to March 2002 in the Itamambuca and Ubatumirim habitats. Crabs were caught manually for 15?min by two collectors during low tide. The carapace width of each crab was measured with a digital caliper, and the sex and ovigerous state were recorded. The median size of the carapace width of males was greater than that of females at both sites (P<0.05). The median size of the crabs from Itamambuca was larger than at Ubatumirim (P<0.05). Only 28 ovigerous females were obtained from both mangroves, which suggested that females might remain in their burrows during the incubation period. The highest recruitment pulse occurred in winter for both populations, probably as a consequence of high reproductive activity during summer. The sex ratio in the size classes showed an anomalous pattern, with a higher frequency of females in the intermediate size classes. This may be related to a greater energy requirement for reproduction in females, thus delaying growth. The variable environmental conditions to which Uca rapax populations are subject appear to act directly or indirectly on the population, causing variations in growth and reproductive processes in the different populations investigated here.     

The occurrence of facultative paedomorphosis in a lacustrine population of the Pyrenean newt (Calotriton asper): morphology and age structure

Facultative paedomorphosis is a common polyphenism in newts where two alternative phenotypes can coexist: metamorphs and paedomorphs (mature with larval traits, i.e. retaining gills). This phenomenon has been reported in some lacustrine populations of the Pyrenean newt (Calotriton asper). The morphology and life-history traits were studied in a lacustrine population of this species with the occurrence of facultative paedomorphosis. The results showed that 24.3% of adults retained gills at different levels of reabsorption and had a smoother skin compared with the other populations, which were fully metamorphic. The body size and shape showed significant sexual differences, with males being larger than females. The presence of paedomorphic traits also affected the body size and shape, revealing a complex pattern of growth in which metamorphic adults were larger than paedomorphs. The age structure was different between sexes, with a median age of 10.5 and 7 years and longevity of 19 and 14 years in males and females, respectively. Considering each sex separately, the age structure was not significantly different between metamorphic and paedomorphic newts. In addition, metamorphosed and branchiate juveniles were found with a maximum age of 13 and 20 years, respectively. Remarkably, long-lived larvae with an exceptional individual of 18 years were found, which has not previously been reported in any other newt species. The mechanisms that can explain the evolution of paedomorphosis are complex and comprise the interaction of multiple factors related to the cost of reproduction and growth.     

Effects of livestock grazing exlosure on aquatic macroinvertebrates in a montane stream, New Mexico

Aquatic macroinvertebrate populations inhabiting reaches of a stream within areas excluded from livestock grazing for a decade were markedly different from those in grazed areas when density, biomass, biotic condition indices, and mean chi square indices of the two populations were compared. Increased densities and biomasses of more tolerant forms of macroinvertebrates were observed in grazed reaches. Because pretreatment data were not available, differences in macroinvertebrate populations and relative tolerances of taxa in grazed and ungrazed areas could be as easily attributed to linear changes in stream habitat as to removal of domestic livestock. Results of this study have implications for the design of future research on the effects of livestock grazing on stream environments and biota: (1) baseline/pretreatment information is prerequisite, and (2) the study should take a watershed (ecosystem) approach.