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At least 919 records · Page 51Linked to original sources

Natural and human dimensions of a quasi-wild species: The case of kudzu

The human dimensions of biotic invasion are generally poorly understood, even among the most familiar invasive species. Kudzu ( Pueraria montana (Lour.) Merr.) is a prominent invasive plant and an example of quasi-wild species, which has experienced repeated introduction, cultivation, and escape back to the wild. Here, we review a large body of primary scientific and historic records spanning thousands of years to characterize the complex relationships among kudzu, its natural enemies, and humans, and provide a synthesis and conceptual model relevant to the ecology and management of quasi-wild invasive species. We documented over 350, mostly insect, natural enemy species and their impacts on kudzu in its native East Asian range. These natural enemies play a minor role in limiting kudzu in its native range, rarely generating severe impacts on populations of wild kudzu. We identified a number of significant influences of humans including dispersal, diverse cultural selection, and facilitation through disturbances, which catalyzed the expansion and exuberance of kudzu. On the other hand, harvest by humans appears to be the major control mechanism in its native areas. Humans thus have a complex relationship with kudzu. They have acted as both friend and foe, affecting the distribution and abundance of kudzu in ways that vary across its range and over time. Our conceptual model of kudzu emphasizes the importance of multiple human dimensions in shaping the biogeography of a species and illustrates how kudzu and other quasi-wild species are more likely to be successful invaders.

Biological Invasions↗

Insular ecosystems of the southeastern United States—A regional synthesis to support biodiversity conservation in a changing climate

In the southeastern United States, insular ecosystems—such as rock outcrops, depression wetlands, high-elevation balds, flood-scoured riparian corridors, and insular prairies and barrens—occupy a small fraction of land area but constitute an important source of regional and global biodiversity, including concentrations of rare and endemic plant taxa. Maintenance of this biodiversity depends upon regimes of abiotic stress and disturbance, incorporating factors such as soil surface temperature, widely fluctuating hydrologic conditions, fires, flood scouring, and episodic droughts that may be subject to alteration by climate change. Over several decades, numerous localized, site-level investigations have yielded important information about the floristics, physical environments, and ecological dynamics of these insular ecosystems; however, the literature from these investigations has generally remained fragmented. This report consists of literature syntheses for eight categories of insular ecosystems of the southeastern United States, concerning (1) physical geography, (2) ecological determinants of community structures including vegetation dynamics and regimes of abiotic stress and disturbance, (3) contributions to regional and global biodiversity, (4) historical and current anthropogenic threats and conservation approaches, and (5) key knowledge gaps relevant to conservation, particularly in terms of climate-change effects on biodiversity. This regional synthesis was undertaken to discern patterns across ecosystems, identify knowledge gaps, and lay the groundwork for future analyses of climate-change vulnerability. Findings from this synthesis indicate that, despite their importance to regional and global biodiversity, insular ecosystems of the southeastern United States have been subjected to a variety of direct and indirect human alterations. In many cases, important questions remain concerning key determinants of ecosystem function. In particular, few empirical investigations in these ecosystems have focused on possible climate-change effects, despite the well-documented ecological effects of climate change at a global level. Long-term management of these ecosystems could benefit from increased scientific effort to characterize and quantify the linkages between changing environmental conditions and the ecological processes that sustain biodiversity.

Alabama, Arkansas, Florida, Georgia, Kentucky, Lou↗

NASA and USGS invest in invasive species modeling to evaluate habitat for Africanized Honey Bees

Invasive non-native species, such as plants, animals, and pathogens, have long been an interest to the U.S. Geological Survey (USGS) and NASA. Invasive species cause harm to our economy (around $120 B/year), the environment (e.g., replacing native biodiversity, forest pathogens negatively affecting carbon storage), and human health (e.g., plague, West Nile virus). Five years ago, the USGS and NASA formed a partnership to improve ecological forecasting capabilities for the early detection and containment of the highest priority invasive species. Scientists from NASA Goddard Space Flight Center (GSFC) and the Fort Collins Science Center developed a longterm strategy to integrate remote sensing capabilities, high-performance computing capabilities and new spatial modeling techniques to advance the science of ecological invasions [Schnase et al., 2002].

The Earth Observer↗

Deer do not affect short-term rates of vegetation recovery in overwash fans on Fire Island after Hurricane Sandy

1. Coastal resilience is threatened as storm-induced disturbances become more frequent and intense with anticipated changes in weather patterns. After severe storms, rapid recovery of vegetation, especially that of dune-stabilizing plants, is a fundamental property of coastal resilience. Herbivores may affect resilience by selectively foraging palatable plant species in disturbed areas. Knowledge of herbivore impacts to dune recovery after storm events is crucial to management of coastal properties. 2. We combined imagery classification, wildlife monitoring, and trend analysis to investigate effects of white-tailed deer on recovery rates of vegetation over four years post-storm in nine areas overwashed. We estimated local deer density with trail cameras, coupled deer density with primary productivity, and assessed the relationship between deer density and rates of vegetation recovery in overwash fans. 3. Pre-storm vegetation cover consisted of shrubs and sporadic patches of beachgrass. Post-storm cover was dominated by beachgrass. At current recovery rates, vegetation coverage will return to pre-storm conditions within the decade, though community transition from grasses to shrubs will take longer and will vary by site with the rate of dune formation. 4. The effect of deer on rates of vegetation recovery was negative, but not statistically significant nor biologically compelling. Although effects of trampling are evident in classified imagery, deer foraging on beachgrass had little effect on rates of spread throughout overwash fans. 5. Our research provides perhaps the first assessment of the effects of deer on post-storm vegetation recovery. While the rate of spread of the primary dune-building plant species was not deleteriously affected by deer, locally high deer densities will likely affect the establishment and development of more palatable herbs and shrubs.

New York↗

A floodplain continuum for Atlantic coast rivers of the Southeastern US: Predictable changes in floodplain biota along a river's length

Floodplains are among the world’s economically-most-valuable, environmentally-most-threatened, and yet conceptually-least-understood ecosystems. Drawing on concepts from existing riverine and wetland models, and empirical data from floodplains of Atlantic Coast rivers in the Southeastern US (and elsewhere when possible), we introduce a conceptual model to explain a continuum of longitudinal variation in floodplain ecosystem functions with a particular focus on biotic change. Our hypothesis maintains that major controls on floodplain ecology are either external (ecotonal interactions with uplands or stream/river channels) or internal (wetland-specific functions), and the relative importance of these controls changes progressively from headwater to mid-river to lower-river floodplains. Inputs of water, sediments, nutrients, flora, and fauna from uplands-to-floodplains decrease, while the impacts of wetland biogeochemistry and obligate wetland plants and animals within-floodplains increase, along the length of a river floodplain. Inputs of water, sediment, nutrients, and fauna from river/stream channels to floodplains are greatest mid-river, and lower either up- or down-stream. While the floodplain continuum we develop is regional in scope, we review how aspects may apply more broadly. Management of coupled floodplain-river ecosystems would be improved by accounting for how factors controlling the floodplain ecosystem progressively change along longitudinal riverine gradients.

Wetlands↗

Fungal endophyte effects on invasive Phragmites australis performance in field and growth chamber environments

Manipulating plant microbiomes may provide control of invasive species. Invasive Phragmites australis has spread rapidly in North American wetlands, causing significant declines in native biodiversity. To test microbiome effects on host growth, we inoculated four common fungal endophytes into replicated Phragmites genotypes and monitored their growth in field and growth chamber environments. Inoculations were highly successful in the growth chamber but inoculated plants in the field were rapidly colonized by diverse endophytes from the local environment. There were significant genotype effects and minimal inoculation effects in both experiments with a significant inoculation × genotype interaction on tiller height in the field. Our results demonstrate that endophyte inoculation treatments are feasible, but repeated inoculations may be required to maintain high titer in plants subject to endophyte colonization from the local environment. Future studies should investigate a wider range of fungal endophytes to identify taxa that inhibit Phragmites and other invaders.

Fungal Ecology↗

Fine-scale plant defence variability increases top-down control of an herbivore

Herbivore populations are regulated by a combination of plant defences and natural enemies. While plant defence can suppress herbivore populations, these defences can also adversely affect natural enemies, thereby releasing herbivores from top-down control. Over their life spans, herbivores and their natural enemies may experience substantial variation in plant defence. Recent studies have demonstrated that individual herbivores feeding on diets containing variable concentrations of plant toxins suffer substantially reduced fitness compared to herbivores feeding on a constant toxin concentration, even when both groups of herbivores experience equivalent means. However, the impacts of defence variability on natural enemies and top-down control of herbivores are unknown. Using artificial diets, we independently manipulated the mean concentration and variation of a plant toxin experienced by individual Trichoplusia ni caterpillars and its parasitoid Copidosoma floridanum . Additionally, by combining the performance of individual caterpillars on different constant diet concentrations of toxin, we were able to estimate the effect of toxin variability between herbivores using nonlinear averaging. Increases in the mean toxin concentration in the diet of parasitized T. ni hosts decreased the fitness of C. floridanum , while variance in individual diets did not impact parasitoid fitness, even though both mean and variance decreased the fitness of T. ni caterpillars. Increased variability in encountered plant defences suppressed individual herbivore fitness with no perceptible cost to top-down control. At the population level, however, increased variability between individual herbivore diets decreased the success of parasitoids relative to herbivores, thus reducing the strength of top-down control. Our study highlights the importance of defence variability at different scales in regulating herbivore performance. Variability in plant defence has the potential to reduce herbivore populations through a combination of bottom-up and top-down effects, but only at small spatial scales experienced by individual herbivores.

Functional Ecology↗

Patterning emergent marsh vegetation assemblages in coastal Louisiana, USA, with unsupervised artificial neural networks

Question Are self‐organizing maps (SOMs) useful for patterning coastal wetland vegetation communities? Do SOMs provide robust alternatives to traditional classification methods, particularly when underlying species response functions are unknown or difficult to approximate, or when a need exists to continuously classify new samples obtained under ongoing long‐term ecosystem monitoring programs as they become available?. Location Coastal Louisiana, USA. Methods A SOM was trained from in situ observations of 559 vegetation species relative cover data from 2526 samples collected over eight years at 343 locations across coastal Louisiana. Hierarchical cluster analysis was applied to the SOM output to delineate vegetation community types, and indicator species analysis was conducted. Salinity and flood duration were compared across the delineated community types. Results The SOM patterned the 2526 training samples into 260 output neurons, which were further clustered into eleven community types. Clear gradients in salinity and flood duration existed among the community types, and geographic zonation of the communities was evident across the landscape. At some locations assemblages were temporally stable; at other locations they varied considerably. Samples not used in training the network were effectively projected onto the SOM and assigned to one of the delineated community types. Conclusions The SOM was effective in delineating plant communities in the region that were qualitatively similar to those obtained in previous investigations. Being robust to skewed distributions and the presence of outliers, SOMs provide an alternative to traditional distribution‐based statistical approaches. Their ability to efficiently classify new data into existing community types makes their use an ideal approach to classifying samples obtained from ongoing, long‐term ecological monitoring programs.

Louisiana↗

Managing effects of drought and other water resource challenges in Alaska and the Pacific Northwest

This is a Cooperator Report. As such, there is no specific abstract. The physical, ecological, and social environments of Alaska and the Pacific Northwest (PNW) region of the United States are extremely diverse. Alaska ranges from the Arctic Ocean and the very cold, dry environments of the North Slope to the cool and very rainy coastal North Pacific region of Southeast Alaska. Most precipitation falls as snow at higher elevations. In Arctic Alaska, average annual temperature is 14.6 F, and average annual precipitation is 11 inches. By contrast, in Southeast Alaska, average annual temperature is 35.8 F, and annual average precipitation is 143 inches. The PNW, defined here as Idaho, Oregon, and Washington, ranges from the Pacific Coast (annual precipitation of 200 inches) to interior semi-arid regions (annual precipitation of 8 inches). Precipitation patterns in the PNW are strongly governed by orographic phenomena, with high, persistent snowpack in the higher mountains (e.g., record annual snowfall of 1,130 in at Mount Baker, Washington in 1999-2000). Ecosystems in the PNW include productive temperate coniferous forests near the Pacific coast and along the (wet) west slope of the Cascade Range, less productive mixed-conifer forest along the (dry) east slope of the Cascades and in interior mountain ranges, and sagebrush-steppe and shrublands at lower elevations in much of the interior and mountain valleys. Large rivers and thousands of smaller tributaries form an extensive network of riparian, wetland, and estuarine systems that provide both critical hydrologic function and biological diversity at broad and fine spatial scales. Although Alaska and the Pacific Northwest differ in important physical, ecological, and social features, the importance of natural resources is evident in both regions. Water is important for wildlife and people. Water provides critical habitat for salmon, which are culturally and economically valuable species. Timber production has declined in recent decades. Recreation has emerged as a major revenue source.

Alaska, Oregon, Washington↗

Site-scale disturbance best predicts moss, vascular plant, and amphibian indices in Ohio wetlands

Loss of wetland habitats and their associated biological communities is a major environmental concern. Quality assessment indices (QAIs) and indices of biological integrity (IBIs) are useful for assessing the responses of taxa to wetland habitat quality and land use in the surrounding landscape. We synthesized the results of our previous predictive modeling studies of five IBIs and QAIs for communities of mosses, vascular plants, and amphibians in forested and emergent wetlands in Ohio (USA). Overall, the single best predictor of these indices was a metric that estimated site-scale (i.e., within the wetland boundaries) substrate and habitat development, alteration, and disturbance. The second most important predictor was a metric that assessed site-scale wetland plant community types and quality, degree of interspersion, and microtopography. Landscape-scale variables better predicted moss and amphibian indices than either vascular plant index. Our results indicate that applying management practices that reduce the effects of site-scale anthropogenic disturbances and increase habitat complexity, such as creating forested buffers surrounding wetlands, increasing wetland contiguity, and creating hummocks and tussocks may simultaneously enhance amphibian, vascular plant, and moss communities in forested and emergent wetlands. Such a focused strategy may enable management agencies to more effectively apportion resources for wetland restoration and construction projects.

Ecological Restoration↗

Spatial and temporal patterns across an ecological boundary: Allochthonous effects of a young saltwater lake on a desert ecosystem

We documented changes in the abundance and composition of terrestrial flora and fauna with respect to distance from the sea edge and timing of large allochthonous inputs from the Salton Sea, California. We found significant effects that were most pronounced within 300 m of the shore, but extended 3 km inland via coyote scat deposition. The zone within 300 m of the sea had a higher density of vegetation with a distinctly different plant composition. The denser vegetation supported higher abundances of birds and reptiles. Coyotes exhibited spatial and temporal responses to marine subsidies of fish, while birds were likely subsidized by aquatic aerial insects. Top-down control, as well as dietary and habitat preferences, may have resulted in reduced number of ants, beetles, and small mammals near the sea. Species responses to the habitat edge appeared to be associated with life history, as the near shore habitat favored habitat generalists and shore specialists, while inland desert habitat favored many sand and open desert specialists. Ecosystem responses support current theories of allochthonous spatial subsidies and consumer-resource dynamics but were limited in scope, magnitude, and distance.

Journal of Arid Environments↗

Comparative performance of invasive and native Celastrus species across environmental gradients

The ability to understand and predict the success of invasive plant species in their new ranges is increased when there is a sympatric native congener available for comparison. Celastrus orbiculatus (oriental bittersweet) is a liana introduced into the United States in the mid-1800s from East Asia as an ornamental plant. Its native congener, Celastrus scandens (American bittersweet), ranges from the east coast of the United States as far west as Wyoming. In the Northeastern United States, C. orbiculatus is continuing to expand its range while C. scandens appears to be in serious decline. One hypothesis for this decline is that C. scandens does not have such a wide range of ecological tolerances in the current landscape as C. orbiculatus, which seems to tolerate a greater range of resource conditions. To investigate this hypothesis, we transplanted these two species into ten sites that spanned a full range of light and soil moisture conditions to compare their establishment and performance in terms of aboveground growth (biomass and height) and mortality. After two years, C. orbiculatus showed significantly lower mortality and greater biomass across all resource conditions compared to C. scandens. In addition, C. orbiculatus preferred more mesic soil moisture conditions, while C. scandens performed better in drier soil moisture conditions. Since much of the Northeastern United States is now forested, this preference for mesic soil conditions could make it more successful than C. scandens in the region. This study shows the utility of manipulative experiments, particularly those using congeneric native species as benchmarks, for assessing the causes and predicting the course of invasions. ?? 2007 Springer-Verlag.

Oecologia↗

Vegetation of semi-stable rangeland dunes of the Navajo Nation, Southwestern USA

Dune destabilization and increased mobility is a worldwide issue causing ecological, economic, and health problems for the inhabitants of areas with extensive dune fields. Dunes cover nearly a third of the Navajo Nation within the Colorado Plateau of southwestern USA. There, higher temperatures and prolonged drought beginning in 1996 have produced significant increases in dune mobility. Vegetation plays an important role in dune stabilization, but there are few studies of the plants of the aeolian surfaces of this region. We examined plant species and their attributes within a moderately vegetated dune field of the Navajo Nation to understand the types and characteristics of plants that stabilize rangeland dunes. These dunes supported a low cover of mixed grass-scrubland with fifty-two perennial and annual species including extensive occurrence of non-native annual Salsola spp. Perennial grass richness and shrub cover were positively associated with increased soil sand composition. Taprooted shrubs were more common on sandier substrates. Most dominant grasses had C4 photosynthesis, suggestive of higher water-use efficiencies and growth advantage in warm arid environments. Plant cover was commonly below the threshold of dune stabilization. Increasing sand movement with continued aridity will select for plants adapted to burial, deflation, and abrasion. The study indicates plants tolerant of increased sand mobility and burial but more investigation is needed to identify the plants adapted to establish and regenerate under these conditions. In addition, the role of Salsola spp. in promoting decline of perennial grasses and shrubs needs clarification.

Arizona, Colorado, New Mexico, Utah↗

The role of microtopography and resident species in post-disturbance recovery of arid habitats in Hawaiʻi

Habitat-suitability indices (HSI) have been employed in restoration to identify optimal sites for planting native species. Often, HSI are based on abiotic variables and do not include biotic interactions, even though similar abiotic conditions can favor both native and nonnative species. Biotic interactions such as competition may be especially important in invader-dominated habitats because invasive species often have fast growth rates and can exploit resources quickly. In this study, we test the utility of an HSI of microtopography derived from airborne LiDAR to predict post-disturbance recovery and native planting success in native shrub-dominated and nonnative, invasive grass-dominated dryland habitats in Hawaiʻi. The HSI uses high-resolution digital terrain models to classify sites' microtopography as high, medium, or low suitability, based on wind exposure and topographic position. We used a split-plot before-after-control-impact design to implement a disturbance experiment within native shrub ( Dodonaea viscosa ) and nonnative, invasive grass ( Cenchrus clandestinus )-dominated ecosystems across three microtopography categories. In contrast to previous studies using the same HSI, we found that microtopography was a poor predictor of pre-disturbance conditions for soil nutrients, organic matter content, or foliar C:N, within both Dodonaea and Cenchrus vegetation types. In invader-dominated Cenchrus plots, microtopography helped predict cover, but not as expected (i.e., highest cover would be in high-suitability plots): D. viscosa had the greatest cover in low-suitability and C. clandestinus had the greatest cover in medium-suitability plots. Similarly, in native-dominated Dodonaea plots, microtopography was a poor predictor of D. viscosa , C. clandestinus , and total plant cover. Although we found some evidence that microtopography helped inform post-disturbance plant recovery of D. viscosa and total plant cover, vegetation type was a more important predictor. Important for considering the success of plantings, percent cover of D. viscosa decreased while percent cover of C. clandestinus increased within both vegetation types 20 months after disturbance. Our results are evidence that HSIs based on topographic features may prove most useful for choosing planting sites in harsh habitats or those already dominated by native species. In more productive habitats, competition from resident species may offset any benefits gained from “better” suitability sites.

Hawaii↗

Greater sage-grouse respond positively to intensive post-fire restoration treatments

Habitat loss is the most prevalent threat to biodiversity in North America. One of the most threatened landscapes in the United States is the sagebrush ( Artemisia spp.) ecosystem, much of which has been fragmented or converted to non-native grasslands via the cheatgrass-fire cycle. Like many sagebrush obligates, greater sage-grouse ( Centrocercus urophasianus ) depend upon sagebrush for food and cover and are affected by changes to this ecosystem. We investigated habitat selection by 28 male greater sage-grouse during each of 3 years after a 113,000-ha wildfire in a sagebrush steppe ecosystem in Idaho and Oregon. During the study period, seeding and herbicide treatments were applied for habitat restoration. We evaluated sage-grouse responses to vegetation and post-fire restoration treatments. Throughout the 3 years post-fire, sage-grouse avoided areas with high exotic annual grass cover but selected strongly for recovering sagebrush and moderately strongly for perennial grasses. By the third year post-fire, they preferred high-density sagebrush, especially in winter when sagebrush is the primary component of the sage-grouse diet. Sage-grouse preferred forb habitat immediately post-fire, especially in summer, but this selection preference was less strong in later years. They also selected areas that were intensively treated with herbicide and seeded with sagebrush, grasses, and forbs, although these responses varied with time since treatment. Wildfire can have severe consequences for sagebrush-obligate species due to loss of large sagebrush plants used for food and for protection from predators and thermal extremes. Our results show that management efforts, including herbicide application and seeding of plants, directed at controlling exotic annual grasses after a wildfire can positively affect habitat selection by sage-grouse.

Ecology and Evolution↗

Passive restoration following ungulate removal in a highly disturbed tropical wet forest devoid of native seed dispersers

Overabundant ungulate populations can alter forests. Concurrently, global declines of seed dispersers may threaten native forest structure and function. On an island largely devoid of native vertebrate seed dispersers, we monitored forest succession for 7 years following ungulate exclusion from a 5-ha area and adjacent plots with ungulates still present. We observed succession from open scrub to forest and understory cover by non-native plants declined. Two trees, native Hibiscus tiliaceus and non-native Leucaena leucocephala , accounted for most forest regeneration, with the latter dominant. Neither species is dependent on animal dispersers nor was there strong evidence that plants dependent on dispersers migrated into the 5-ha study area. Passive restoration following ungulate removal may facilitate restoration, but did not show promise for fully restoring native forest on Guam. Restoration of native forest plants in bird depopulated areas will likely require active outplanting of native seedlings, control of factors resulting in bird loss, and reintroduction of seed dispersers.

Guam↗

The transformative impact of genomics on sage-grouse conservation and management

For over two decades, genetic studies have been used to assist in the conservation and management of both Greater Sage-grouse ( Centrocercus urophasianus ) and Gunnison Sage-grouse ( C. minimus ), addressing a wide variety of topics including taxonomy, parentage, population connectivity, and demography. The field of conservation genetics has been transformed by dramatic improvements in sequencing technology, facilitating genomic studies in many wildlife species. The quality and amount of data generated by genomic methods vastly exceed that of traditional genetic studies, allowing for increased precision in estimating genetic parameters of interest. Perhaps more importantly, genomic methods can provide insight into non-neutral evolution such as adaptive divergence. Here we recount the shift from genetic to genomic methods using two wildlife species of substantial conservation interest, focusing on the improved capabilities and advantages of genomic methods. For instance, reassessment of divergence in sage-grouse using genomic methods confirmed strong differentiation between the two species and revealed that a small population in the state of Washington was more genetically distinct than previously recognized. Further, new genomic resources and approaches have been used to identify a family of genes linked to local dietary adaptation suggesting that sage-grouse may possess digestive and metabolic adaptations that mitigate the effects of consuming plant secondary metabolites like those found in sagebrush. Genetic variation among populations in these gene regions is thought to be involved with local dietary adaptations, and therefore maintaining the tie between sage-grouse and the chemistry of local sagebrush may be an important management consideration. We posit that the integration of newly developed genomic resources combined with the vast wealth of ecological and behavioral data for sage-grouse has the potential to shed light on mechanistic relationships that ultimately are vital to the conservation and management of these species.

Book chapter↗

Notes on status and ecology of the endangered Hawaiian annual Awiwi, Centaurium sebaeoides (Gentianaceae)

The annual, endemic, coastal herb Centaurium sebaeoides is the only native Hawaiian species in the gentian family. The U.S. Fish and Wildlife Service listed it as Endangered under the Endangered Species Act on 29 October 1991. Before surveys reported here, the total population of this species statewide was estimated at 80-110 individuals in eight populations. During counts made in April and May 1997, following ample winter rains, 12 populations of C. sebaeoides with a total of 6300-6600 plants were noted on five islands (Kaua'i, O'ahu, Lana'i, Moloka'i, and Maui). Five populations were mapped with a global positioning system and counted; in the remaining seven populations, the numbers of individuals were estimated. More recent surveys in 1998-1999 estimated a total of only 60-80 individuals at all sites. Such dramatic population fluctuations are believed to be related to the sporadic occurrence of winter rains. Threats that further contribute to the rarity of the species include (1) displacement and overtopping by salt-tolerant nonnative woody species, especially Casuarina spp ., (2) trampling and erosion of habitat by ungulates, and (3) damage caused by off-road vehicles.

Hawaii↗