Rapid genetic restoration of a keystone species exhibiting delayed demographic response

Genetic founder effects are often expected when animals colonize restored habitat in fragmented landscapes, but empirical data on genetic responses to restoration are limited. We examined the genetic response of banner‐tailed kangaroo rats (Dipodomys spectabilis) to landscape‐scale grassland restoration in the Chihuahuan Desert of New Mexico, USA. Dipodomys spectabilis is a grassland specialist and keystone species. At sites treated with herbicide to remove shrubs, colonization by D. spectabilis is slow and populations persist at low density for ≥10 years (≥6 generations). Persistence at low density and low gene flow may cause strong founder effects. We compared genetic structure of D. spectabilis populations between treated sites and remnant grasslands, and we examined how the genetic response to restoration depended on treatment age, area, and connectivity to source populations. Allelic richness and heterozygosity were similar between treated sites and remnant grasslands. Allelic richness at treated sites was greatest early in the restoration trajectory, and genetic divergence did not differ between recently colonized and established populations. These results indicated that founder effects during colonization of treated sites were weak or absent. Moreover, our results suggested founder effects were not mitigated by treatment area or connectivity. Dispersal is negatively density‐dependent in D. spectabilis, and we hypothesize that high gene flow may occur early in the restoration trajectory when density is low. Our study shows genetic diversity can be recovered more rapidly than demographic components of populations after habitat restoration and that founder effects are not inevitable for animals colonizing restored habitat in fragmented landscapes.     

Response of Grassland Birds in Sand Shinnery Oak Communities Restored Using Tebuthiuron and Grazing in Eastern New Mexico

Sand shinnery oak (Quercus havardii) communities are a unique component of grassland bird habitat in eastern New Mexico and have been impacted by human activities for decades. These communities are frequently managed with livestock grazing and herbicide application for shrub control, strategies that potentially can be used to restore the historical shrub–grass composition of this plant community. During spring migration and the breeding seasons of 2004 and 2005, we compared density and community structure of grassland bird species among four combinations of tebuthiuron application and grazing treatments that were being evaluated for restoration of shinnery oak communities. We performed biweekly point transects on sixteen 65‐ha study plots in these communities. Density of all avian species combined did not differ between grazed and ungrazed plots. Tebuthiuron‐treated plots had a 40% higher average density for combined species than untreated plots. There was a 41% higher average density of all species during spring 2005 than 2004, but density was similar during the breeding season of both years. These trends were predominantly influenced by densities of migratory Cassin’s Sparrow (Aimophila cassinii), which were greater in tebuthiuron‐treated plots in both years. Densities of resident Meadowlarks (Sturnella spp.) exhibited little response to tebuthiuron or grazing treatments. Avian species richness, evenness, and diversity were only minimally affected by the tebuthiuron and grazing treatments. This study occurred over a period of highly variable precipitation, so future assessments, spanning longer wet–dry cycles and maturing plant communities, may be necessary to completely determine avian response to these restoration efforts.     

栖息地荒漠化对蜥蜴群落组成的影响

栖息地改变往往对动物群落及其物种多样性有重要影响,但是目前关于栖息地荒漠化如何影响蜥蜴的群落组成鲜有研究。在内蒙古自治区鄂尔多斯市准格尔旗和达拉特旗地区通过样线法调查了三类不同荒漠化程度样地上的蜥蜴群落组成差异,用典范对应分析探究了关键的影响因素。结果表明,栖息地荒漠化使蜥蜴的群落组成发生显著变化,从固定沙丘、半固定沙丘到流动沙丘,草原沙蜥均是蜥蜴群落的优势物种,密点麻蜥逐渐减少,丽斑麻蜥则消失于荒漠化严重的半固定沙丘和流动沙丘中。随栖息地荒漠化程度的增加,蜥蜴的数量显著减少,密度显著降低,蜥蜴物种的丰富度、香农-威纳(Shannon-Weaver)多样性指数和皮洛(Pielou)均匀性指数均逐次显著下降,而辛普森(Simpson)优势度指数则逐次显著升高。这说明荒漠化不仅使蜥蜴群落组成变得简单,同时造成了蜥蜴物种多样性的下降甚至丧失。荒漠化也使蜥蜴的栖息条件发生显著变化。植被高度、油蒿比例、裸地比例、隐蔽度、表层土壤含水量和表层土壤孔隙度在三类栖息样地之间均存在极显著差异。典范对应分析结果表明,这些环境因子与蜥蜴群落组成的变化密切相关。总体而言,草原沙蜥偏好裸地,密点麻蜥偏好隐蔽性好且土壤湿润又疏松的区域,而丽斑麻蜥则偏好植被高的区域。     

Multivariate analysis of the response of overgrown semi-natural calcareous grasslands to restorative shrub cutting

Monitoring and evaluation of restoration management often suffers from poor sampling design and a lack of statistical rigour, seldom considering the spatial and temporal variability of habitat. We tested the effectiveness of shrub cutting on the restoration of calcareous grasslands, using a 4-year Before–After/Reference-Control-Impact (BARCI) project design with replicated patches of habitat. Departure from the control sites and convergence with the reference sites were analysed using the Principal Response Curves (PRC) method.The structure of communities was compared 1 year before and 3 years after management on control scrubland, restored ex-arable land, restored and reference grassland. Results show that shrub cutting did not suffice to restore the community composition of the semi-natural calcareous grasslands. The restored ex-arable areas maintained a community structure extremely similar to the control scrubland.The restored grassland, on the other hand, deviated from the control and slightly converged to the reference grassland, though not significantly. The dominant scrub species that was cut, Prunus spinosa, showed higher cover values compared to the reference grassland in any of the treatments, even the first year after cutting. Species typical of xeric and/or calcareous grassland were more abundant in the reference than in the restored grassland, while arable and ruderal species were more frequent in the restored sites.This study has demonstrated that the BARCI approach is a powerful tool for the evaluation of restoration management, as it was possible to evaluate not only departure from the unmanaged control, but also convergence with the reference community. The PRC method provided a comprehensive overview of the response of the 153 species involved in the study.Our results also indicate that the second PRC should be considered, when significant, to not exclude important information from the assessment.     

Non-trophic interactions in deserts: Facilitation,interference, and an endangered lizard species

Research on plant–animal interactions has been focused on direct consumer interactions (i.e. plants as resources), but non-trophic interactions including providing shelter or interference with movement can also affect the fine-scale distribution of animals. In particular, non-trophic interactions that are positive could support threatened animal populations. Positive interactions have been used in the restoration of plant communities, but have not yet been extended to the management of animal habitat. In this study, we tested the hypothesis that non-trophic interactions influence the occurrence of an endangered lizard species in an arid shrub-annual system. At a location known to have a population of blunt-nosed leopard lizards (Gambelia sila), we geotagged 700 shrubs, measured shrub morphometric traits, collected biomass samples, and surveyed for lizard presence using scat detection dogs over two years. Relative to 2014, in 2013 plant productivity was high and lizard scats were found more frequently in areas with low invasive grass cover (i.e. residual dry matter, RDM). In 2014, plant productivity was low because of an extreme drought year, and lizard scats were more frequently observed under shrub canopies, particularly those with relatively dense cover. These findings support the novel theory that positive non-trophic interactions are a critical form of plant–animal interactions in addition to consumption. Dominant shrubs can act as a foundation species by functioning as a basal node in structuring both plant and animal communities through a network of interactions. Managing dominant plants, in addition to habitat, is therefore important for conserving animal species in arid ecosystems.     

Impact of Non-Native Plant Removal on Lizards in Riparian Habitats in the Southwestern United States

Many natural processes in the riparian cottonwood (Populus deltoides) forest of the Middle Rio Grande (MRG) in the southwestern United States have been disrupted or altered, allowing non‐native plants such as saltcedar (Tamarix spp.) and Russian olive (Elaeagnus angustifolia) to establish. We investigated reptilian responses to restoration efforts by sampling communities of lizards at 12 study sites invaded by non‐native plants along the MRG in New Mexico for 7 years (2000–2006). Sites within three regions were randomly assigned to one of the three treatments to remove non‐native plants and woody debris, or as untreated controls. We used pitfall and funnel traps to capture, mark, and release lizards from June to September. Principal components analysis of 15 vegetation variables identified five factors that best explained variation among sites before and after removal of non‐native plants. Relative abundances for four of six common species of lizards were associated with vegetation characteristics that significantly changed after plant removal. Species were either positively associated with the more open, park‐like understory found in treated sites or negatively associated with debris heaps and thickets of non‐native plants found in untreated sites. Eastern fence lizards (Sceloporus consobrinus) and New Mexico whiptails (Aspidoscelis neomexicana) increased in relative abundance after non‐native plants were removed. Overall, removal of non‐native plants seems beneficial, or at least is non‐damaging, to lizard communities of the MRG forest. Providing information on habitat associations of lizard communities will help land managers balance management objectives with other considerations, such as providing important wildlife habitat.     

Effects of long‐term fox baiting on species composition and abundance in an Australian lizard community

Abstract We report on the effects of almost a decade of 1080‐fox baiting on a lizard community in a mosaic Australian habitat. Replicated comparisons of baited versus non‐baited control areas with near‐identical histories of bush fires, grazing and climate showed a higher density of red fox tracks (Canis vulpes) in the non‐baited areas. Furthermore, the fox‐baited areas showed a more than five times higher density of sand goannas (Varanus gouldii), a species that strongly overlaps the red fox in food niche breadth and is itself a direct target of fox predation, in particular its eggs and young. Exclusion of predators from a natural habitat led to significant increases in the density of small lizards, suggesting that predation can drive lizard population dynamics in this ecosystem. Replicated pitfall‐trapping in three habitats in the control areas (with high fox and low goanna density) versus the baited areas (with low fox and high goanna density) showed that fox baiting had positive effects on the density of diurnal scincid lizards in open grassland, whereas the control areas showed higher density of nocturnal gecko lizards. Our interpretation is that fox removal may result in a shift in the top predator towards the sand goanna. Historically, this indigenous, endemic species was the natural top predator. It has co‐evolved with its prey and that may have moulded it into a more efficient lizard predator per encounter than the introduced fox.     

Spider community responses to grassland restoration: balancing trade‐offs between abundance and diversity

Spiders (Araneae) play key roles in ecosystems, not only as common and abundant generalist predators, but also as major contributors to biodiversity in many areas. In addition, due to their short generation times and high mobility, spiders respond rapidly to small changes in their environment, potentially making them useful indicators for restoration monitoring. However, few studies have focused on spider responses to grassland restoration in the United States. We compared degraded, native, and restored grassland sites to examine how spider communities and habitat respond to arid grassland restoration. We also examined how responses varied with the age of the restoration project. Spider communities in native sites differed from those in restored and degraded sites in several ways: native sites had fewer spiders and a different community composition than degraded and restored sites. However, native and restored sites had more species than degraded sites. Chronosequence data showed trends for lower abundance, higher species richness, and changing community composition as restoration projects mature. Several habitat variables were closely linked to variation in spider communities including cover of invasive annual grasses, litter, and biological soil crusts. Our data suggest that spider and vegetation responses to grassland restoration efforts can be successful in the long term—with resulting communities becoming more similar to native ones—and that spiders are useful indictors of grassland restoration. Our results also suggest that restoration may involve balancing trade‐offs between ecosystem services, with potential losses in predatory control offset by increases in biodiversity with restoration effort.     

Restoration Practices Have Positive Effects on Breeding Bird Species of Concern in the Chihuahuan Desert

Woody plant encroachment into grasslands is a global concern. Efforts to restore grasslands often assume that removal of woody plants benefits biodiversity but assumptions are rarely tested. In the Chihuahuan Desert of the Southwestern United States, we tested whether abundances of grassland specialist bird species would be greater in plant communities resulting from treatment with herbicides to remove encroaching shrubs compared with untreated shrub‐dominated areas that represented pre‐treatment conditions. In 2010, we surveyed breeding birds and vegetation at 16 treated–untreated pairs. In 2011, we expanded the survey effort to 21 treated–untreated pairs, seven unpaired treatment areas, and five reference grassland areas. Vegetation in treatment areas had higher perennial grass foliar and basal cover and lower shrub foliar cover compared with untreated areas. Several regionally declining grassland specialists exhibited higher occurrence and relative abundance in treated areas. A shrubland specialist, however, was associated with untreated areas and may be negatively impacted by shrub removal. Bird community composition differed between treated and untreated areas in both years. Our results indicate that shrub removal can have positive effects on grassland specialist bird species, but that a mosaic of treated and untreated areas might be most beneficial for regional biodiversity .     

Predation by an exotic lizard,Anolis sagrei,alters the ant community structure in betelnut palm plantations in southern Taiwan

Abstract 1. Predators can affect prey directly by reducing prey abundance and indirectly by altering behavioural patterns of prey. From previous studies, there is little evidence that ant community structure is affected by vertebrate predation. 2. Researchers tend to consider the interactions between vertebrate predators and ants to be weak. The present study examined the impact of the exotic invasive lizard, Anolis sagrei, on the ant community structure by manipulating the density of lizards within enclosures. The natural density of A. sagrei in the field was surveyed and used as the stocking density rate in the lizard‐present sub‐enclosures. 3. Before the lizard density was manipulated, there was no difference in the ant diversity between sub‐enclosures. After the lizard density manipulation, the ant diversity in sub‐enclosures with A. sagrei present was significantly different from that of enclosures where the lizards were absent, although the overall ant abundance did not differ significantly. 4. The ant diversity difference was generated by a significant reduction of the ant species Pheidole fervens in sub‐enclosures with A. sagrei present. Such an abundance change might be the result of direct predation by the lizards, or it might be generated by a foraging site shift by this ant. 5. The results of this study thus demonstrated that the invasion of an exotic vertebrate can significantly alter the community structure of ants, perhaps through the combined direct and indirect effects of lizards on ants.     

Negative Effects of an Exotic Grass Invasion on Small-Mammal Communities

Exotic invasive species can directly and indirectly influence natural ecological communities. Cheatgrass (Bromus tectorum) is non-native to the western United States and has invaded large areas of the Great Basin. Changes to the structure and composition of plant communities invaded by cheatgrass likely have effects at higher trophic levels. As a keystone guild in North American deserts, granivorous small mammals drive and maintain plant diversity. Our objective was to assess potential effects of invasion by cheatgrass on small-mammal communities. We sampled small-mammal and plant communities at 70 sites (Great Basin, Utah). We assessed abundance and diversity of the small-mammal community, diversity of the plant community, and the percentage of cheatgrass cover and shrub species. Abundance and diversity of the small-mammal community decreased with increasing abundance of cheatgrass. Similarly, cover of cheatgrass remained a significant predictor of small-mammal abundance even after accounting for the loss of the shrub layer and plant diversity, suggesting that there are direct and indirect effects of cheatgrass. The change in the small-mammal communities associated with invasion of cheatgrass likely has effects through higher and lower trophic levels and has the potential to cause major changes in ecosystem structure and function.     

Implications of Food Web Interactions for Restoration of Missouri Ozark Glade Habitats

Ozark glades are gaps in forested areas that are dominated by grasses and forbs growing in rocky, nutrient‐poor soil. Historically, these open, patchy habitats were maintained by natural and anthropogenic fire cycles that prohibited tree encroachment. However, because of decades of fire suppression, glades have become overgrown by fire‐intolerant species such as Eastern red cedar (Juniperus virginiana). Current restoration practices include cutting down invasive cedars and burning brush piles, which represent habitat for Northern fence lizards (Sceloporus undulatus). Because Sceloporus actively consumes herbivores, we hypothesized that the presence of these lizards in and around brush piles might result in a trophic cascade, whereby damage on native plants is reduced. Field surveys across six Missouri glades indicated that lizard activity was minimal beyond 1 m from habitat structures. This activity pattern reduced grasshopper abundance by 75% and plant damage by over 66% on Echinacea paradoxa and Rudbeckia missouriensis near structures with lizards. A field transplant experiment demonstrated similar reductions in grasshopper abundance and damage on two other glade endemic species, Aster oblongifolius and Schizachyrium scoparium. These results demonstrate that future glade restoration efforts might benefit from considering top‐down effects of predators in facilitating native plant establishment.     

Reestablishment of a Rodent Community in Restored Desert Scrub

I conducted small-mammal trapping surveys on a desert scrub restoration site in Palm Springs, California, to document concomitant recovery of the rodent community. These surveys were conducted following quantitative vegetation sampling efforts that indicated that a predefined successful restoration criterion of 15% total shrub cover had been met throughout most of the area. But shrub cover, native shrub cover, herb cover, native herb cover, total cover, and total native cover remained significantly lower in the restoration area than in undeveloped desert scrub immediately surrounding the site. Native herb species richness was also generally lower in the restoration area. Despite these vegetation differences, rodent diversity, evenness, and abundance were very similar between the restoration and natural areas (they were consistently slightly higher in the restoration area). More diverse microhabitats, proximity to water, and reduced competition with harvester ants may have contributed to this outcome. If ecosystem restoration is the goal, reestablishment of a faunal community in restored habitat, rather than surpassing a predefined percent cover of vegetation, may be a better indicator of success, because plant cover proved to be a poor predictor of mammal success.     

Effects of grazing regime on the relative body mass of the endangered pygmy bluetongue lizard (Tiliqua adelaidensis)

Many animals rely on fat reserves, to keep them alive through extended periods of food shortage, such as the winter, and to provide additional energy for reproduction. Fat reserves, measured relative to an animal's size, are often referred to as the animal's body condition. The present study investigated how different levels of grazing by domestic stock in native grassland habitat affect the body condition of the pygmy bluetongue lizard (Tiliqua adelaidensis), and if these effects are related to changes in the abundance and size of grasshoppers which are the lizards primary source of food. The initial hypothesis was that lizards would have the highest body condition in moderately grazed paddocks, because those paddocks would have more grasshoppers than heavily grazed paddocks, and better visual conditions for lizards to catch those grasshoppers than in ungrazed paddocks. The results, however, showed that both lizard body condition and the abundance of grasshoppers increased with decreasing grazing intensity. The connection between lizard body condition and abundance of grasshoppers was complex. Within an activity season, lizard body condition generally declined from spring to summer, while the number of grasshoppers grew. The mean size of grasshoppers seemed to be more important, as lizard body condition was higher in spring, the time of year with the largest grasshoppers. These results show that the intensity of grazing by domestic stock influences the body condition of pygmy bluetongue lizards, but that this effect is not entirely due to the reduction in the number of grasshoppers resulting from grazing.     

Soil disturbance favours threatened beetle species in sandy grasslands

Soil disturbance is recognised as an important restoration measure for conserving biodiversity in sandy soils. We used a soil disturbance (ploughing) experiment in a sandy grassland as well as a semi-natural disturbance (slope erosion enhanced by cattle trampling) gradient on a sandy slope to test the soil disturbance effects on the ground-living beetle community. Both experimental disturbance and semi-natural disturbance favoured sandy grassland specialists, but there was no overall effect on beetle richness and abundance. Amara lucida and Harpalus spp. were favoured by disturbance while Calathus melanocephalus was disfavoured. Experimental ploughing significantly increased the proportion of red-listed species in disturbed plots compared to non-disturbed controls. In the semi-natural disturbance gradient we found that the beetle community on the disturbed slope differed from that of the flat areas, and there were tendencies for a higher proportion of red-listed species on the slope. We conclude that increasing the area of bare sand in sandy grasslands can have positive effects on many threatened species. Soil disturbance should thus be included as a regular measure in sandy grasslands under conservation management and as a measure to restore high biodiversity in areas where bare sand is rare.     

Effects on the thermoregulatory efficiency of two native lizards as a consequence of the habitat modification by the introduction of the exotic tree Acacia longifolia

Habitat modification alters several aspects of the original fauna, among them the opportunity for thermoregulation. Here, we studied the thermal biology of sympatric populations of two lizard species (Liolaemus multimaculatus and Liolaemus wiegmannii) in two different situations; a grassland without trees (natural habitat) and in a grassland plus the exotic tree Acacia longifolia (modified habitat), aiming to assess whether the structural alteration of native Pampean coastal grasslands of Argentina affects the thermal biology of these lizards. Field body temperatures, laboratory preferred temperatures, micro-environmental temperatures, operative temperatures, thermoregulatory efficiency and spatial distribution of each species were analyzed in both habitats. Environmental operative temperature was 0.64 °C lower in the modified habitat (Te=38.39 °C) than in the natural (Te=39.03 °C). Thermoregulatory efficiency (E) of L. wiegmannii was lower in modified sites (E=0.58) than in natural sites (E=0.70). This difference may be because this lizard occupied shaded microhabitats under acacias, with suboptimal thermal features. In contrast, L. multimaculatus in the modified habitat restricted its activity to open microenvironments that retained a similar structure to that of the native habitat, while maintaining high thermoregulatory efficiency in both habitat types (E_modified=0.92; E_natural=0.96). Although these two lizard species are phylogenetically close, they respond differently to human-induced changes in their thermal environments. The introduction of A. longifolia into coastal grasslands for L. wiegmannii in particular, this introduction converts its native habitat into a suboptimal thermal environment.     

Frugivore behaviour determines plant distribution: a spatially‐explicit analysis of a plant‐disperser interaction

The spatial distribution of plants (and other primarily sessile organisms) depends on the interplay between their ecological requirements and the spatial template set before, during, and after the dispersal process. In the case of animal‐dispersed plants, the spatial characteristics of animal behaviour during the seed dispersal process are likely to leave a lasting imprint on plant distribution. Here, we hypothesize that the activity patterns of the frugivorous lizard Podarcis lilfordi directly influence the spatial distribution of the fleshy‐fruited shrub Daphne rodriguezii. To evaluate this hypothesis, we first analysed lizard activity, following radio‐tracked lizards during the plant's fruiting period, and identified its main determinants at several spatial scales of habitat aggregation (from 12.5 to 150 m). We hypothesised that lizard activity depends on differential habitat features explaining its territory use plus habitat preferences associated with each movement bout. In a second step, the most important determinants of lizard activity plus the variables describing habitat structure were used to predict the presence of adult and juvenile plants. Predictability of lizard activity (based on AUC and Pearson regression coefficients) was higher at broad spatial‐scales of habitat aggregation (75 m). The two best predictors of lizard activity were the habitat features of and the distance to the core area (defined as the area enclosing the 0.50 cumulative probability of lizard locations). Plant presence was best predicted by models based on a combination of lizard activity and habitat features at local spatial scales (1.5 m). Best models included habitat features and lizard activity for adult plants, and local‐scale habitat features, the proximity of adult plants and lizard activity for juveniles. In both cases, most plants (50–60%) were located at ‘optimal sites’ (both favourable for lizards and with adequate habitat features), whereas a small fraction of them (3–10%) were located at dispersal‐limited sites (i.e. with adequate habitat features but suboptimal for lizards). Our results thus suggest that the interplay between lizard activity and local habitat features determines the spatial patterns of juvenile‐plant presence and leaves a lasting signature on adult‐plant distribution.     

长江三峡库区蝶类群落的等级多样性指数

在三峡库区蝶类物种多样性调查基础上,从等级多样性角度进行分析,结果表明,蝴蝶属级,科级和等级多样性指数在灌丛生境较高,森林、农田和草地生境多样性较低;非参数检验和方差分析结果表明4种生境在属级和等级多样性上差异显著,灌丛生境和其他生境差异显著。海拔500-1500m高度,蝴蝶的生物多样性相对较高,是库区蝴蝶最重要的生存地带。属级和科级多样性指数的变异系数在森林植被型样本间最大,前者有4个小生境在100％以上,后者有6个小生境超过100%;农田科的变异系数有1个小生境超过100%;草地的属和科变异系数均是1个小生境超过100%;灌丛则全部低于100%。这些数值,反映了三峡库区的形成对蝴蝶的生存及分布的产生了很大的影响,进一步证明库区蝶类生存环境的多样性和破碎化;其中,森林对蝴蝶生存的影响是最复杂的;通过蝴蝶做环境质量评估时,灌丛可作为首选考察对象。