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Predicting performance for ecological restoration: A case study using Spartina altemiflora

The success of population-based ecological restoration relies on the growth and reproductive performance of selected donor materials, whether consisting of whole plants or seed. Accurately predicting performance requires an understanding of a variety of underlying processes, particularly gene flow and selection, which can be measured, at least in part, using surrogates such as neutral marker genetic distances and simple latitudinal effects. Here we apply a structural equation modeling approach to understanding and predicting performance in a widespread salt marsh grass, Spartina alterniflora, commonly used for ecological restoration throughout its native range in North America. We collected source materials from throughout this range, consisting of eight clones each from 23 populations, for transplantation to a common garden site in coastal Louisiana and monitored their performance. We modeled performance as a latent process described by multiple indicator variables (e.g., clone diameter, stem number) and estimated direct and indirect influences of geographic and genetic distances on performance. Genetic distances were determined by comparison of neutral molecular markers with those from a local population at the common garden site. Geographic distance metrics included dispersal distance (the minimum distance over water between donor and experimental sites) and latitude. Model results indicate direct effects of genetic distance and latitude on performance variation among the donor sites. Standardized effect strengths indicate that performance was roughly twice as sensitive to variation in genetic distance as to latitudinal variation. Dispersal distance had an indirect influence on performance through effects on genetic distance, indicating a typical pattern of genetic isolation by distance. Latitude also had an indirect effect on genetic distance through its linear relationship with dispersal distance. Three performance indicators had significant loadings on performance alone (mean clone diameter, mean number of stems, mean number of inflorescences), while the performance indicators mean stem height and mean stem width were also influenced by latitude. We suggest that dispersal distance and latitude should provide an adequate means of predicting performance in future S. alterniflora restorations and propose a maximum sampling distance of 300 km (holding latitude constant) to avoid the sampling of inappropriate ecotypes. ?? 2010 by the Ecological Society of America.

Ecological Applications

Large, high-intensity fire events in Southern California shrublands: Debunking the fine-grain age patch model

We evaluate the fine-grain age patch model of fire regimes in southern California shrublands. Proponents contend that the historical condition was characterized by frequent small to moderate size, slow-moving smoldering fires, and that this regime has been disrupted by fire suppression activities that have caused unnatural fuel accumulation and anomalously large and catastrophic wildfires. A review of more than 100 19th-century newspaper reports reveals that large, high-intensity wildfires predate modern fire suppression policy, and extensive newspaper coverage plus first-hand accounts support the conclusion that the 1889 Santiago Canyon Fire was the largest fire in California history. Proponents of the fine-grain age patch model contend that even the very earliest 20th-century fires were the result of fire suppression disrupting natural fuel structure. We tested that hypothesis and found that, within the fire perimeters of two of the largest early fire events in 1919 and 1932, prior fire suppression activities were insufficient to have altered the natural fuel structure. Over the last 130 years there has been no significant change in the incidence of large fires greater than 10000 ha, consistent with the conclusion that fire suppression activities are not the cause of these fire events. Eight megafires (???50 000 ha) are recorded for the region, and half have occurred in the last five years. These burned through a mosaic of age classes, which raises doubts that accumulation of old age classes explains these events. Extreme drought is a plausible explanation for this recent rash of such events, and it is hypothesized that these are due to droughts that led to increased dead fine fuels that promoted the incidence of firebrands and spot fires. A major shortcoming of the fine-grain age patch model is that it requires age-dependent flammability of shrubland fuels, but seral stage chaparral is dominated by short-lived species that create a dense surface layer of fine fuels. Results from the Behave Plus fire model with a custom fuel module for young chaparral shows that there is sufficient dead fuel to spread fire even under relatively little winds. Empirical studies of fuel ages burned in recent fires illustrate that young fuels often comprise a major portion of burned vegetation, and there is no difference between evergreen chaparral and semi-deciduous sage scrub. It has also been argued that the present-day fire size distribution in northern Baja California is a model of the historical patterns that were present on southern California landscapes. Applying this model with historical fire frequencies shows that the Baja model is inadequate to maintain these fire-prone ecosystems and further demonstrates that fire managers in southern California are not likely to learn much from studying modern Baja California fire regimes. Further supporting this conclusion are theoretical cellular automata models of fire spread, which show that, even in systems with age dependent flammability, landscapes evolve toward a complex age mosaic with a plausible age structure only when there is a severe stopping rule that constrains fire size, and only if ignitions are saturating. ?? 2009 by the Ecological Society of America.

Ecological Applications

A less field-intensive robust design for estimating demographic parameters with Mark-resight data

The robust design has become popular among animal ecologists as a means for estimating population abundance and related demographic parameters with mark-recapture data. However, two drawbacks of traditional mark-recapture are financial cost and repeated disturbance to animals. Mark-resight methodology may in many circumstances be a less expensive and less invasive alternative to mark-recapture, but the models developed to date for these data have overwhelmingly concentrated only on the estimation of abundance. Here we introduce a mark-resight model analogous to that used in mark-recapture for the simultaneous estimation of abundance, apparent survival, and transition probabilities between observable and unobservable states. The model may be implemented using standard statistical computing software, but it has also been incorporated into the freeware package Program MARK. We illustrate the use of our model with mainland New Zealand Robin (Petroica australis) data collected to ascertain whether this methodology may be a reliable alternative for monitoring endangered populations of a closely related species inhabiting the Chatham Islands. We found this method to be a viable alternative to traditional mark-recapture when cost or disturbance to species is of particular concern in long-term population monitoring programs. ?? 2009 by the Ecological Society of America.

Ecology

Predicting community responses to perturbations in the face of imperfect knowledge and network complexity

How best to predict the effects of perturbations to ecological communities has been a long-standing goal for both applied and basic ecology. This quest has recently been revived by new empirical data, new analysis methods, and increased computing speed, with the promise that ecologically important insights may be obtainable from a limited knowledge of community interactions. We use empirically based and simulated networks of varying size and connectance to assess two limitations to predicting perturbation responses in multispecies communities: (1) the inaccuracy by which species interaction strengths are empirically quantified and (2) the indeterminacy of species responses due to indirect effects associated with network size and structure. We find that even modest levels of species richness and connectance (∼25 pairwise interactions) impose high requirements for interaction strength estimates because system indeterminacy rapidly overwhelms predictive insights. Nevertheless, even poorly estimated interaction strengths provide greater average predictive certainty than an approach that uses only the sign of each interaction. Our simulations provide guidance in dealing with the trade-offs involved in maximizing the utility of network approaches for predicting dynamics in multispecies communities.

Ecology

Proximate and landscape factors influence grassland bird distributions

Ecologists increasingly recognize that birds can respond to features well beyond their normal areas of activity, but little is known about the relative importance of landscapes and proximate factors or about the scales of landscapes that influence bird distributions. We examined the influences of tree cover at both proximate and landscape scales on grassland birds, a group of birds of high conservation concern, in the Sheyenne National Grassland in North Dakota, USA. The Grassland contains a diverse array of grassland and woodland habitats. We surveyed breeding birds on 2015 100 m long transect segments during 2002 and 2003. We modeled the occurrence of 19 species in relation to habitat features (percentages of grassland, woodland, shrubland, and wetland) within each 100-m segment and to tree cover within 200-1600 m of the segment. We used information-theoretic statistical methods to compare models and variables. At the proximate scales, tree cover was the most important variable, having negative influences on 13 species and positive influences on two species. In a comparison of multiple scales, models with only proximate variables were adequate for some species, but models combining proximate with landscape information were best for 17 of 19 species. Landscape-only models were rarely competitive. Combined models at the largest scales (800-1600 m) were best for 12 of 19 species. Seven species had best models including 1600-m landscapes plus proximate factors in at least one year. These were Wilson's Phalarope (Phalaropus tricolor), Sedge Wren (Cistothorus platensis), Field Sparrow (Spizella pusilla), Grasshopper Sparrow (Ammodramus savannarum), Bobolink (Dolychonix oryzivorus), Red-winged Blackbird (Agelaius phoeniceus), and Brown-headed Cowbird (Molothrus ater). These seven are small-bodied species; thus larger-bodied species do not necessarily respond most to the largest landscapes. Our findings suggest that birds respond to habitat features at a variety of scales. Models with only landscape-scale tree cover were rarely competitive, indicating that broad-scale modeling alone, such as that based solely on remotely sensed data, is likely to be inadequate in explaining species distributions. ?? 2006 by the Ecological Society of America.

Ecological Applications

Habitat structure mediates predation risk for sedentary prey: Experimental tests of alternative hypotheses

Predation is an important and ubiquitous selective force that can shape habitat preferences of prey species, but tests of alternative mechanistic hypotheses of habitat influences on predation risk are lacking. 2. We studied predation risk at nest sites of a passerine bird and tested two hypotheses based on theories of predator foraging behaviour. The total-foliage hypothesis predicts that predation will decline in areas of greater overall vegetation density by impeding cues for detection by predators. The potential-prey-site hypothesis predicts that predation decreases where predators must search more unoccupied potential nest sites. 3. Both observational data and results from a habitat manipulation provided clear support for the potential-prey-site hypothesis and rejection of the total-foliage hypothesis. Birds chose nest patches containing both greater total foliage and potential nest site density (which were correlated in their abundance) than at random sites, yet only potential nest site density significantly influenced nest predation risk. 4. Our results therefore provided a clear and rare example of adaptive nest site selection that would have been missed had structural complexity or total vegetation density been considered alone. 5. Our results also demonstrated that interactions between predator foraging success and habitat structure can be more complex than simple impedance or occlusion by vegetation. ?? 2008 British Ecological Society.

Journal of Animal Ecology

Linking occurrence and fitness to persistence: Habitat-based approach for endangered Greater Sage-Grouse

Detailed empirical models predicting both species occurrence and fitness across a landscape are necessary to understand processes related to population persistence. Failure to consider both occurrence and fitness may result in incorrect assessments of habitat importance leading to inappropriate management strategies. We took a two-stage approach to identifying critical nesting and brood-rearing habitat for the endangered Greater Sage-Grouse (Centrocercus urophasianus) in Alberta at a landscape scale. First, we used logistic regression to develop spatial models predicting the relative probability of use (occurrence) for Sage-Grouse nests and broods. Secondly, we used Cox proportional hazards survival models to identify the most risky habitats across the landscape. We combined these two approaches to identify Sage-Grouse habitats that pose minimal risk of failure (source habitats) and attractive sink habitats that pose increased risk (ecological traps). Our models showed that Sage-Grouse select for heterogeneous patches of moderate sagebrush cover (quadratic relationship) and avoid anthropogenic edge habitat for nesting. Nests were more successful in heterogeneous habitats, but nest success was independent of anthropogenic features. Similarly, broods selected heterogeneous high-productivity habitats with sagebrush while avoiding human developments, cultivated cropland, and high densities of oil wells. Chick mortalities tended to occur in proximity to oil and gas developments and along riparian habitats. For nests and broods, respectively, approximately 10% and 5% of the study area was considered source habitat, whereas 19% and 15% of habitat was attractive sink habitat. Limited source habitats appear to be the main reason for poor nest success (39%) and low chick survival (12%). Our habitat models identify areas of protection priority and areas that require immediate management attention to enhance recruitment to secure the viability of this population. This novel approach to habitat-based population viability modeling has merit for many species of concern. ?? 2007 by the Ecological Society of America.

Ecological Applications

Avifaunal responses to fire in southwestern montane forests along a burn severity gradient

The effects of burn severity on avian communities are poorly understood, yet this information is crucial to fire management programs. To quantify avian response patterns along a burn severity gradient, we sampled 49 random plots (2001-2002) at the 17 351-ha Cerro Grande Fire (2000) in New Mexico, USA. Additionally, pre-fire avian surveys (1986-1988, 1990) created a unique opportunity to quantify avifaunal changes in 13 pre-fire transects (resampled in 2002) and to compare two designs for analyzing the effects of unplanned disturbances: after-only analysis and before-after comparisons. Distance analysis was used to calculate densities. We analyzed after-only densities for 21 species using gradient analysis, which detected a broad range of responses to increasing burn severity: (I) large significant declines, (II) weak, but significant declines, (III) no significant density changes, (IV) peak densities in low- or moderate-severity patches, (V) weak, but significant increases, and (VI) large significant increases. Overall, 71% of the species included in the after-only gradient analysis exhibited either positive or neutral density responses to fire effects across all or portions of the severity gradient (responses III-VI). We used pre/post pairs analysis to quantify density changes for 15 species using before-after comparisons; spatiotemporal variation in densities was large and confounded fire effects for most species. Only four species demonstrated significant effects of burn severity, and their densities were all higher in burned compared to unburned forests. Pre- and post-fire community similarity was high except in high-severity areas. Species richness was similar pre- and post-fire across all burn severities. Thus, ecosystem restoration programs based on the assumption that recent severe fires in Southwestern ponderosa pine forests have overriding negative ecological effects are not supported by our study of post-fire avian communities. This study illustrates the importance of quantifying burn severity and controlling confounding sources of spatiotemporal variation in studies of fire effects. After-only gradient analysis can be an efficient tool for quantifying fire effects. This analysis can also augment historical data sets that have small samples sizes coupled with high non-process variation, which limits the power of before-after comparisons. ?? 2007 by the Ecological Society of America.

Ecological Applications

Thermal and energetic constraints on ectotherm abundance: A global test using lizards

Population densities of birds and mammals have been shown to decrease with body mass at approximately the same rate as metabolic rates increase, indicating that energetic needs constrain endotherm population densities. In ectotherms, the exponential increase of metabolic rate with body temperature suggests that environmental temperature may additionally constrain population densities. Here we test simple bioenergetic models for an ecologically important group of ectothermic vertebrates by examining 483 lizard populations. We find that lizard population densities decrease as a power law of body mass with a slope approximately inverse to the slope of the relationship between metabolic rates and body mass. Energy availability should limit population densities. As predicted, environmental productivity has a positive effect on lizard density, strengthening the relationship between lizard density and body mass. In contrast, the effect of environmental temperature is at most weak due to behavioral thermoregulation, thermal evolution, or the temperature dependence of ectotherm performance. Our results provide initial insights into how energy needs and availability differentially constrain ectotherm and endotherm density across broad spatial scales. ?? 2008 by the Ecological Society of America.

Ecology

The relative importance of disturbance and exotic-plant abundance in California coastal sage scrub

Many ecosystems of conservation concern require some level of disturbance to sustain their species composition and ecological function. However, inappropriate disturbance regimes could favor invasion or expansion of exotic species. In southern California coastal sage scrub (CSS) fire is a natural disturbance, but because of human influence, frequencies may now be unnaturally high. Other anthropogenic disturbances such as grazing also occur in reserve areas. Managers charged with imposing or tolerating fire or other disturbance within their reserves are concerned that habitat quality may be degraded by an increasing abundance of exotic plants. We used vegetation monitoring data from Camp Pendleton, California, USA, to assess the correlation between past disturbances (frequent fire, agriculture, or grazing and mechanical disturbances) and current exotic species abundance in CSS. We found that disturbance history was only modestly related to exotic abundance overall, but fire frequency showed the strongest association. We also examined whether cover and richness of various native plant life forms (woody species, perennial herbs, and annual herbs) were more strongly influenced by disturbance history or by exotic-plant abundance. Native plant responses varied among life forms, but woody species and annual herbs were generally more strongly and negatively associated with exotic abundance than with disturbance. Effective CSS conservation will require developing means to curb the negative impacts of exotic plants, which may abound with or without severe or recent disturbance. Additionally, more focus should be given to understory herbs showing sensitivity to invasion. Though understudied, native herbs comprise the greatest portion of plant diversity in CSS and are critical to preservation of the community as a whole. ?? 2009 by the Ecological Society of America.

Ecological Applications

Disturbance alters local-regional richness relationships in appalachian forests

Whether biological diversity within communities is limited by local interactions or regional species pools remains an important question in ecology. In this paper, we investigate how an experimentally applied tree-harvesting disturbance gradient influenced local-regional richness relationships. Plant species richness was measured at three spatial scales (2 ha = regional; 576 m 2 and 1 m 2 = local) on three occasions (one year pre-disturbance, one year post-disturbance, and 10 years post-disturbance) across five disturbance treatments (uncut control through clearcut) replicated throughout the southern Appalachian Mountains, USA. We investigated whether species richness in 576-m 2 plots and 1-m 2 subplots depended on species richness in 2-ha experimental units and whether this relationship changed through time before and after canopy disturbance. We found that, before disturbance, the relationship between local and regional richness was weak or nonexistent. One year after disturbance local richness was a positive function of regional richness, because local sites were colonized from the regional species pool. Ten years after disturbance, the positive relationship persisted, but the slope had decreased by half. These results suggest that disturbance can set the stage for strong influences of regional species pools on local community assembly in temperate forests. However, as time since disturbance increases, local controls on community assembly decouple the relationships between regional and local diversity. ?? 2009 by the Ecological Society of America.

Ecology

Fire helps restore natural disturbance regime to benefit rare and endangered marsh birds endemic to the Colorado River

Large flood events were part of the historical disturbance regime within the lower basin of most large river systems around the world. Large flood events are now rare in the lower basins of most large river systems due to flood control structures. Endemic organisms that are adapted to this historical disturbance regime have become less abundant due to these dramatic changes in the hydrology and the resultant changes in vegetation structure. The Yuma Clapper Rail is a federally endangered bird that breeds in emergent marshes within the lower Colorado River basin in the southwestern United States and northwestern Mexico. We evaluated whether prescribed fire could be used as a surrogate disturbance event to help restore historical conditions for the benefit of Yuma Clapper Rails and four sympatric marsh-dependent birds. We conducted call-broadcast surveys for marsh birds within burned and unburned (control) plots both pre-and post-burn. Fire increased the numbers of Yuma Clapper Rails and Virginia Rails, and did not affect the numbers of Black Rails, Soras, and Least Bitterns. We found no evidence that detection probability of any of the five species differed between burn and control plots. Our results suggest that prescribed fire can be used to set back succession of emergent marshlands and help mimic the natural disturbance regime in the lower Colorado River basin. Hence, prescribed fire can be used to help increase Yuma Clapper Rail populations without adversely affecting sympatric species. Implementing a coordinated long-term fire management plan within marshes of the lower Colorado River may allow regulatory agencies to remove the Yuma Clapper Rail from the endangered species list. ?? 2010 by the Ecological Society of America.

Ecological Applications

The accuracy of matrix population model projections for coniferous trees in the Sierra Nevada, California

1 We assess the use of simple, size-based matrix population models for projecting population trends for six coniferous tree species in the Sierra Nevada, California. We used demographic data from 16 673 trees in 15 permanent plots to create 17 separate time-invariant, density-independent population projection models, and determined differences between trends projected from initial surveys with a 5-year interval and observed data during two subsequent 5-year time steps. 2 We detected departures from the assumptions of the matrix modelling approach in terms of strong growth autocorrelations. We also found evidence of observation errors for measurements of tree growth and, to a more limited degree, recruitment. Loglinear analysis provided evidence of significant temporal variation in demographic rates for only two of the 17 populations. 3 Total population sizes were strongly predicted by model projections, although population dynamics were dominated by carryover from the previous 5-year time step (i.e. there were few cases of recruitment or death). Fractional changes to overall population sizes were less well predicted. Compared with a null model and a simple demographic model lacking size structure, matrix model projections were better able to predict total population sizes, although the differences were not statistically significant. Matrix model projections were also able to predict short-term rates of survival, growth and recruitment. Mortality frequencies were not well predicted. 4 Our results suggest that simple size-structured models can accurately project future short-term changes for some tree populations. However, not all populations were well predicted and these simple models would probably become more inaccurate over longer projection intervals. The predictive ability of these models would also be limited by disturbance or other events that destabilize demographic rates. ?? 2005 British Ecological Society.

Journal of Ecology

Habitat classification modeling with incomplete data: Pushing the habitat envelope

Habitat classification models (HCMs) are invaluable tools for species conservation, land-use planning, reserve design, and metapopulation assessments, particularly at broad spatial scales. However, species occurrence data are often lacking and typically limited to presence points at broad scales. This lack of absence data precludes the use of many statistical techniques for HCMs. One option is to generate pseudo-absence points so that the many available statistical modeling tools can be used. Traditional techniques generate pseudoabsence points at random across broadly defined species ranges, often failing to include biological knowledge concerning the species-habitat relationship. We incorporated biological knowledge of the species-habitat relationship into pseudo-absence points by creating habitat envelopes that constrain the region from which points were randomly selected. We define a habitat envelope as an ecological representation of a species, or species feature's (e.g., nest) observed distribution (i.e., realized niche) based on a single attribute, or the spatial intersection of multiple attributes. We created HCMs for Northern Goshawk (Accipiter gentilis atricapillus) nest habitat during the breeding season across Utah forests with extant nest presence points and ecologically based pseudo-absence points using logistic regression. Predictor variables were derived from 30-m USDA Landfire and 250-m Forest Inventory and Analysis (FIA) map products. These habitat-envelope-based models were then compared to null envelope models which use traditional practices for generating pseudo-absences. Models were assessed for fit and predictive capability using metrics such as kappa, thresholdindependent receiver operating characteristic (ROC) plots, adjusted deviance (Dadj2), and cross-validation, and were also assessed for ecological relevance. For all cases, habitat envelope-based models outperformed null envelope models and were more ecologically relevant, suggesting that incorporating biological knowledge into pseudo-absence point generation is a powerful tool for species habitat assessments. Furthermore, given some a priori knowledge of the species-habitat relationship, ecologically based pseudo-absence points can be applied to any species, ecosystem, data resolution, and spatial extent. ?? 2007 by the Ecological Society of America.

Ecological Applications

Mangrove recruitment after forest disturbance is facilitated by herbaceous species in the Caribbean

Plant communities along tropical coastlines are often affected by natural and human disturbances, but little is known about factors influencing recovery. We focused on mangrove forests, which are among the most threatened ecosystems globally, to examine how facilitation by herbaceous vegetation might improve forest restoration after disturbance. We specifically investigated whether recovery of mangrove forests in harsh environments is accelerated by nurse plants and whether the beneficial effects are species-specific. Quantification of standardized effects allowed comparisons across performance parameters and over time for: (1) net effect of each herbaceous species on mangrove survival and growth, (2) effects of pre- and post-establishment factors associated with each herbaceous species, and (3) need for artificial planting to enhance growth or survival of mangrove seedlings. Mangrove recruitment in a clear-cut forest in Belize was accelerated by the presence of Sesuvium portulacastrum (succulent forb) and Distichlis spicata (grass), two coastal species common throughout the Caribbean region. The net effect of herbaceous vegetation was positive, but the magnitude of effects on mangrove survival and growth differed by species. Because of differences in their vegetative structure and other features, species effects on mangroves also varied by mechanism: (1) trapping of dispersing propagules (both species), (2) structural support of the seedling (Distichlis), and/or (3) promotion of survival (Sesuvium) or growth (Distichlis) through amelioration of soil conditions (temperature, aeration). Artificial planting had a stronger positive effect on mangrove survival than did edaphic conditions, but planting enhanced mangrove growth more in Sesuvium than in Distichlis patches. Our study indicates that beneficial species might be selected based on features that provide multiple positive effects and that species comparisons may be improved using standardized effects. Our findings are not only relevant to the coastal environments found in the Caribbean region, but our assessment methods may be useful for developing site-specific information to restore disturbed mangrove forests worldwide, especially given the large pool of mangrove associates (>45 genera) available for screening. ?? 2007 by the Ecological Society of America.

Ecological Applications

Climate alters response of an endemic island plant to removal of invasive herbivores

Islands experience higher rates of species extinction than mainland ecosystems, with biological invasions among the leading causes; they also serve as important model systems for testing ideas in basic and applied ecology. Invasive removal programs on islands are conservation efforts that can also be viewed as powerful manipulative experiments, but few data are available to evaluate their effects. We collected demographic and herbivore damage data for Castilleja mollis Pennell, an endangered plant endemic to Santa Rosa Island, California, over a 12-year period before, during, and after the implementation of control for introduced cattle, deer, and elk. We used these long-term data to explore mechanisms underlying herbivore effects, assess the results of herbivore reduction at the scales of both individual plants and populations, and determine how temporal variability in herbivory and plant demography influenced responses to herbivore removals. For individual plants, herbivore effects mediated by disturbance were greater than those of grazing. Deer and elk scraping of the ground substantially increased plant mortality and dormancy and reduced flowering and growth. Stem damage from browsing did not affect survivorship but significantly reduced plant growth and flower production. Herbivore control successfully lowered damage rates, which declined steeply between 1997 and 2000 and have remained relatively low. Castilleja mollis abundances rose sharply after 1997, suggesting a positive effect of herbivore control, but then began to decline steadily again after 2003. The recent decline appears to be driven by higher mean growing season temperatures; interestingly, not only reductions in scraping damage but a period of cooler conditions were significant in explaining increases in C. mollis populations between 1997 and 2002. Our results demonstrate strong effects of introduced herbivores on both plant demography and population dynamics and show that climate-driven variation may counteract and mask positive responses to herbivore removal. Regional mean temperatures already have risen significantly over the last 50 years, suggesting that climate change could increasingly swamp the effects of management targeted at other environmental problems. Similar interactions between climate and invasive species will play an even greater role in future management, with long-term data sets like this critical to quantifying such effects. ?? 2009 by the Ecological Society of America.

Ecological Applications

Do competitors modulate rare plant response to precipitation change?

Ecologists increasingly suspect that climate change will directly impact species physiology, demography, and phenology, but also indirectly affect these measures via changes to the surrounding community. Unfortunately, few studies examine both the direct and indirect pathways of impact. Doing so is important because altered competitive pressures can reduce or magnify the direct responses of a focal species to climate change. Here, we examine the effects of changing rainfall on three rare annual plant species in the presence and absence of competition on the California Channel Islands. We used rain-out shelters and hand watering to exclude and augment early, late, and season-long rainfall, spanning the wide range of precipitation change forecast for the region. In the absence of competition, droughts reduced the population growth rates of two of three focal annuals, while increased rainfall was only sometimes beneficial, As compared to the focal species, the dominant competitors were more sensitive to the precipitation treatments, benefiting from increased season-long precipitation and harmed by droughts. Importantly, the response of two of three competitors to the precipitation treatments tended to be positively correlated with those of the focal annuals. Although this leads to the expectation that increased competition will counter the direct benefits of favorable conditions, such indirect effects of precipitation change proved weak to nonexistent in our experiment. Competitors had little influence on the precipitation response of two focal species, due to their low sensitivity to competition and highly variable precipitation responses. Competition did affect how our third focal species responded to precipitation change, but this effect only approached significance, and whether it truly resulted from competitor response to precipitation change was unclear. Our work suggests that even when competitors respond to climate change, these responses may have little effect on the focal species. Ultimately, the strength of the indirect effect depends on how strongly climate change alters competition, and how sensitive focal species are to changes in competition. ?? 2010 by the Ecological Society of America.

Ecology

Two-species occupancy models: A new parameterization applied to co-occurrence of secretive rails

Two-species occupancy models that account for false absences provide a robust method for testing for evidence of competitive exclusion, but previous model parameteriza-tions were inadequate for incorporating covariates. We present a new parameterization that is stable when covariates are included: the conditional two-species occupancy model, which can be used to examine alternative hypotheses for species' distribution patterns. This new model estimates the probability of occupancy for a subordinate species conditional upon the presence of a dominant species. It can also be used to test if the detection of either species differs when one or both species are present, and if detection of the subordinate species depends on the detection of the dominant species when both are present. We apply the model to test if the presence of the larger Virginia Rail (Rallus limicola) affects probabilities of detection or occupancy of the smaller California Black Rail (Laterallus jamaicensis coturniculus) in small freshwater marshes that range in size from 0.013 to 13.99 ha. We hypothesized that Black Rail occupancy should be lower in small marshes when Virginia Rails are present than when they are absent, because resources are presumably more limited and interference competition should increase. We found that Black Rail detection probability was unaffected by the detection of Virginia Rails, while, surprisingly, Black and Virginia Rail occupancy were positively associated even in small marshes. The average probability of Black Rail occupancy was higher when Virginia Rails were present (0.74 ?? 0.053, mean ?? SE) than when they were absent (0.36 ?? 0.069), and for both species occupancy increased with marsh size. Our results contrast with recent findings from patchy forest systems, where small birds were presumed to be excluded from small habitat patches by larger competitors. ?? 2010 by the Ecological Society of America.

Ecological Applications