Developing landscape habitat models for rare amphibians with small geographic ranges: a case study of Siskiyou Mountains salamanders in the western USA

To advance the development of conservation planning for rare species with small geographic ranges, we determined habitat associations of Siskiyou Mountains salamanders (Plethodon stormi) and developed habitat suitability models at fine (10 ha), medium (40 ha), and broad (202 ha) spatial scales using available Geographic Information Systems data and logistic regression analysis with an information theoretic approach. Across spatial scales, there was very little support for models with structural habitat features, such as tree canopy cover and conifer diameter. Model-averaged 95% confidence intervals for regression coefficients and associated odds ratios indicated that the occurrence of Siskiyou Mountains salamanders was positively associated with rocky soils and Pacific madrone (Abutus menziesii) and negatively associated with elevation and white fir (Abies concolor); these associations were consistent across 3 spatial scales. The occurrence of this species also was positively associated with hardwood density at the medium spatial scale. Odds ratios projected that a 10% decrease in white fir abundance would increase the odds of salamander occurrence 3.02–4.47 times, depending on spatial scale. We selected the model with rocky soils, white fir, and Oregon white oak (Quercus garryana) as the best model across 3 spatial scales and created habitat suitability maps for Siskiyou Mountains salamanders by projecting habitat suitability scores across the landscape. Our habitat suitability models and maps are applicable to selection of priority conservation areas for Siskiyou Mountains salamanders, and our approach can be easily adapted to conservation of other rare species in any geographical location.     

Evidence for ecological release over a fine spatial scale in a lizard from the White Sands formation

When a population experiences relaxation of selective pressures due to reduced predation or competition, it may undergo ecological release. Ecological release often manifests as increased niche breadth and trait variation, as individuals have the opportunity to exploit a wider variety of resources. At the recently‐colonized White Sands dune formation in New Mexico, lizards from white sand habitat have fewer predators and competitors than their dark soil counterparts, and show evidence of ecological release compared to dark soil lizards. To determine whether the dynamics of ecological release also play out over an even finer habitat gradient, we studied southwestern fence lizards in the center of the White Sands dune formation and the narrow ecotone between White Sands and the surrounding dark soil habitats. We predicted that lizards from the central dunes would exhibit ecological release in terms of broadened resource use compared to lizards from the ecotone. We first conducted avian surveys in both the central dunes and ecotone habitats to measure abundance, richness and diversity of avian species that could act as lizard predators. Next, we measured microhabitat‐scale resource use by comparing perch selection of lizards in both habitats. Finally, we measured landscape‐scale resource use by quantifying home range sizes and daily distances traveled by lizards in both habitats. We found that central dunes lizards used a greater diversity of perch types, and had greater variation in both home range size and daily distances traveled, than ecotone lizards. There were fewer predatory bird species in the central dunes than on the ecotone, and there was a tendency for fewer predation events in the central dunes than on the ecotone. Our results demonstrate that ecological release is detectable over a fine habitat gradient, such as between the center of a recently‐colonized habitat and its ecotone with the surrounding ecosystem.     

An integrated method to create habitat suitability models for fragmented landscapes

Given the pervasive influence of human induced habitat fragmentation in ecological processes, landscape models are a welcome advance. The development of GIS software has allowed a greater use of these models and of analyses of the relationship between species and habitat variables. Habitat suitability models are thus theoretical concepts that can be used for planning in fragmented landscapes and habitat conservation. The most commonly used models are based on single species and on the assignment of suitability values for some environmental variables. Generally the cartographic basis for modeling suitability are thematic maps produced by a Boolean logic. In this paper we propose a model based on a set of focal species and on maps produced by a fuzzy classification method. Focal species, selected by an expert-based approach, provide a practical way of extending the scope of habitat suitability models to the conservation of biodiversity at landscape scale. The utilisation of a classification method that applies a continuity criterion may allow more consideration of the connectivity of an area because it allows a better detection of ecological gradients within a landscape. We applied this methodology to the Tuscany region focusing on terrestrial mammals. Performing a fuzzy classification we produced five land cover maps and through image processing operations we obtained a suitability model which applies a continuity criterion. The resulting suitability fuzzy model seems better for the study of connectivity and fragmentation, especially in areas with high spatial complexity.     

Biodiversity potential of <Emphasis Type="Italic">Nothofagus</Emphasis> forests in Tierra del Fuego (Argentina): tool proposal for regional conservation planning

It is difficult to map and quantify biodiversity at landscape level in areas with low data availability, despite demand from decision-makers. We propose a methodology to determine potential biodiversity pattern using habitat suitability maps of the understory plant species with highest cover and occurrence frequency in the three different forests types of Tierra del Fuego (Argentina). We used a database of vascular plants from 535 surveys from which we identified 35 indicative species. We explored more than 50 potential explanatory variables to develop habitat suitability maps of the indicative species, which were combined to develop a map of the potential biodiversity. Correlation among environmental, topographic and forest landscape variables were discussed, as well as the marginality and the specialization of the indicative species. We detected differences in the niches of the species prevailing in the three forest types. The developed map of potential biodiversity uncovered hotspots of biodiversity in the ecotone of Nothofagus pumilio and N. antarctica as well as in the wettest part of the mixed N. pumilio–N. betuloides forests. It allowed thus to identify forest areas with different conservation potential and can be readily used as a decision support system for conservation and management strategies at different scales including the identification of land-use conflicts (e.g. of biodiversity with timber production and livestock) and the development of a network of protected areas, which currently does not cover the forests of highest conservation value.     

Monitoring sandy desertification of Otindag Sandy Land based on multi-date remote sensing images

Sandy desertification is now the main ecological problem in the Otindag Sandy Land. In order to reveal the process of land degradation, especially the latest situation of sandy desertification, a method integrating remote sensing, Geographic Information System (GIS) and field survey was employed to build a sandy desertification dataset for analysis. Remote sensing images included the Landsat Thematic Mapper (TM) image in 1987, the Enhanced Thematic Mapper Plus (ETM⁺) image in 2000, and the image with the Charge-Coupled Device Camera (CCD) on the China-Brazil Earth Resources Satellite (CBERS) in 2006. Five land-cover classes, including active sand dunes, fixed sand dunes, semi-fixed sand dunes, inter-dune grassland and wetlands, were identified. Results showed that the Otindag Sandy Land has been suffering sandy desertification since 1987 with 2 different desertified stages. The first stage from 1987 to 2000 was a severe sandy desertification period, characterized by the fixed sand dunes decreasing at a high speed, and the semi-fixed and active sand dunes increasing remarkably. The second stage spanned from 2000 to 2006 and the sandy desertification was weakened greatly. Although a large area of fixed sand dunes were transformed to other types, fixed sand dunes were still the dominant type in the Ointdag region at 2006. Spatial change detection based on active sand dunes showed that the expansion area was much larger than the reversion area in the past two decades, and that several active sand belts had been formed, suggesting that sandy desertification controlling of the Otindag Sandy Land will be a long-term task.     

Monitoring sandy desertification of Otindag Sandy Land based on multi-date remote sensing images

Sandy desertification is now the main ecological problem in the Otindag Sandy Land. In order to reveal the process of land degradation, especially the latest situation of sandy desertification, a method integrating remote sensing, Geographic Information System (GIS) and field survey was employed to build a sandy desertification dataset for analysis. Remote sensing images included the Landsat Thematic Mapper (TM) image in 1987, the Enhanced Thematic Mapper Plus (ETM⁺) image in 2000, and the image with the Charge-Coupled Device Camera (CCD) on the China-Brazil Earth Resources Satellite (CBERS) in 2006. Five land-cover classes, including active sand dunes, fixed sand dunes, semi-fixed sand dunes, inter-dune grassland and wetlands, were identified. Results showed that the Otindag Sandy Land has been suffering sandy desertification since 1987 with 2 different desertified stages. The first stage from 1987 to 2000 was a severe sandy desertification period, characterized by the fixed sand dunes decreasing at a high speed, and the semi-fixed and active sand dunes increasing remarkably. The second stage spanned from 2000 to 2006 and the sandy desertification was weakened greatly. Although a large area of fixed sand dunes were transformed to other types, fixed sand dunes were still the dominant type in the Ointdag region at 2006. Spatial change detection based on active sand dunes showed that the expansion area was much larger than the reversion area in the past two decades, and that several active sand belts had been formed, suggesting that sandy desertification controlling of the Otindag Sandy Land will be a long-term task.     

不同沙地生境下黄柳(Salix gordejevii)的根系分布和冠层结构特征

对分布于3种不同沙地生境（半固定沙丘、固定沙丘、丘间低地）黄柳灌丛的根系分布（根的数目与深度分布）和冠层结构特征（叶面积指数、平均叶倾角以及当年枝长度分布）进行了研究。半固定沙丘上的黄柳根系最发达（根系分布深,数量大）,固定沙丘的黄柳明显不及前者,而丘间低地的黄柳根系最不发达。粗根（Φ≥5mm）的分布与土壤水分和土壤容重呈显著相关性,细根（Φ<5mm）的分布与土壤容重和紧实度呈极显著相关。不同程度的根系发育导致地上部分的繁茂程度不同分布于半固定沙丘上的黄柳灌丛叶面积指数较大,当年枝平均长度较大,枝长频度分布趋于正态分布;丘间低地的黄柳灌丛叶面积指数较小,当年枝平均长度较小,枝长频度分布趋于对数正态分布;固定沙丘上灌丛的上述特征大致介于二者之间,而其当年枝长度频度符合Weibull分布。     

不同沙地生境下黄柳(Salix gordejevii)种群若干数量特征的比较研究

对分布于３种不同沙地生境（半固定沙丘、固定沙丘、丘间低地）黄柳灌丛的个体形态、生长和群体数量特征进行了比较研究,同时测定了灌丛下草本层植物种类组成及频率。结果显示：分布于丘间低地的黄柳灌丛当年枝平均长度增加缓慢,短枝条的比例较高,枝长频度分布（当年枝）峰值偏左,分枝也较多,叶片短而狭,长宽比大;与之相比,分布于半固定沙丘上的灌丛当年枝平均长度增加较快,长枝条比例较高,枝长频度分布趋于正态分布,分枝     

基于多时相遥感影像的浑善达克沙地沙漠化监测

沙漠化是浑善达克沙地面临的主要环境问题.为了揭示该地区沙漠化过程,特别是其最新状态,利用1987年和2000年的Landsat TM/ETM 及2006年的中巴资源卫星CCD数据,在地面考察资料及地理信息系统(GIS)的支持下,提取了固定沙地、半固定沙地、流动沙地、丘间草原及湿地5种地表覆被类型,建立了浑善达克沙地沙漠化数据库.结果表明浑善达克沙地在过去20a里发生了沙漠化,但是可分为两个阶段.第一阶段从1987年到2000年,为沙漠化快速发展期,固定沙地面积显著减少,而半固定沙地和流动沙地则明显增加;第二阶段从2000年到2006年,为沙漠化缓减期.尽管发生了沙漠化,但固定沙地仍然是浑善达克沙地面积最大的类型.针对流动沙地的空间变化检测表明在过去20a里沙地恶化的面积远远大于逆转的面积,并且已经形成了几条流动沙带,这意味着浑善达克沙地沙漠化的遏制是一个长期的过程.     

Identifying Core Areas in a Species’ Range Using Temporal Suitability Analysis: an Example Using Little Bustards <Emphasis Type="Italic">Tetrax Tetrax</Emphasis> L. in Spain

Variations in habitat quality impact on breeding success, leading to strong selection pressure for the best sites to be occupied first during a population increase and last during a decline. Coupled with dispersal and metapopulation processes, the result is that snapshot surveys of wildlife distributions may fail to reveal core areas that conservation seeks to protect. At a local scale, territory occupancy is a good indicator of quality but data are not readily available to assess occupancy for rarer species, in remote areas, and over large spatial extents. We introduce temporal suitability analysis as a way to generate an analogue of occupancy from a single survey and illustrate it using data on the little bustard in Spain. We first used Generalised Additive Modelling (GAM) to build a predictive distribution model using Geographic Information System (GIS) coverages and satellite imagery, and then applied the model retrospectively to a time series of satellite images to produce one distribution map for each year. These annual maps differed in the extent of Spain predicted as suitable for little bustards. By overlaying the maps, we identified areas predicted as suitable in one to n years. We show that this temporal suitability map correlates with a conventional habitat suitability map based on a single year but contains extra information on hierarchical use of habitats and the lag between suitability and use. The technique may be applied at a variety of spatial scales to reveal changes in expected occupancy as land use or external factors determining land cover types vary over time.     

Coastal dune vegetation and pollen representation in south Buenos Aires Province, Argentina

Aim To establish the relationship between coastal dune vegetation and its pollen representation as an aid to interpret Holocene vegetation dynamics and environmental changes from pollen assemblages. Location The study area is situated on the temperate Atlantic coast of south Buenos Aires Province, Argentina (c. 39° S and 61°20′ W). Methods The vegetation of the active dune area adjacent to the beach was described on the basis of its floristic composition from 25 plots. Classification of the vegetation into distinct zones was carried out by cluster analysis. Surface samples were collected from each vegetation stand and analysed for their pollen composition. Pollen percentage data were analysed using principal components analysis in order to investigate the degree to which the different vegetation units can be distinguished by their pollen spectra. Pollen–vegetation relationships for selected taxa were explored using simple scatter plots and indices of association, under‐ and over‐representation. Indices of floristic diversity and palynological richness were used to assess the representation of the vegetation in the pollen spectra. Results and conclusions Five vegetation zones are defined on the basis of species composition and their quantitative variation: back shore, mobile dunes, slacks, semi‐fixed and fixed dunes. Pollen assemblages from back shore, mobile dunes and slacks are clearly differentiated from semi‐fixed and fixed dunes. Pollen assemblages differ considerably from the associated vegetation composition. Major discrepancies are caused by large differences in pollen and vegetation proportion of Hyalis argentea and Discaria americana. There is a considerable proportion of non‐local pollen in every spectrum. Pollen representation in the coastal dunes at Monte Hermoso is influenced by differences in pollen production, dispersal and preservation of individual taxa as well as by the spatial distribution of the vegetation, the topography of the dune system and the wind pattern. The pollen–vegetation relationship established in this study has important implication for understanding and interpreting fossil pollen records from coastal dune environments.     

Contrasting habitat use of morphologically similar bat species with differing conservation status in south‐eastern Australia

The east‐coast free‐tailed bat Mormopterus norfolkensis Gray, 1839 is a threatened insectivorous bat that is poorly known and as such conservation management strategies are only broadly prescribed. Insectivorous bats that use human‐modified landscapes are often adapted to foraging in open microhabitats. However, few studies have explored whether open‐adapted bats select landscapes with more of these microhabitat features. We compared three morphologically similar and sympatric, molossid bats (genus Mormopterus) with different conservation status in terms of their association with vegetation, climate, landform and land‐use attributes at landscape and local habitat element scales. We predicted that these species would use similar landscape types, with semi‐cleared and low density urban landscapes used more than forested and heavily cleared landscapes. Additionally, we explored which environmental variables best explained the occurrence of each species by constructing post‐hoc models and habitat suitability maps. Contrary to predictions, we found that the three species varied in their habitat use with no one landscape type used more extensively than other types. Overall, M. norfolkensis was more likely to occur in low‐lying, non‐urban, riparian habitats with little vegetation cover. Mormopterus species 2 occupied similar habitats, but was more tolerant of urban landscapes. In contrast, Mormopterus species 4 occurred more often in cleared than forested landscapes, particularly dry landscapes with little vegetation cover. The extensive use of coastal floodplains by the threatened M. norfolkensis is significant because these habitats are under increasing pressure from human land‐uses and the predicted increase in urbanization is likely to further reduce the amount of suitable habitat.     

On‐land habitat preferences of female New Zealand sea lions at Sandy Bay,Auckland Islands

Terrestrial habitat is important for breeding in most pinnipeds. On land, most species remain near the shore, but New Zealand (NZ) sea lions, Phocarctos hookeri, often rest inland up to 1.5 km from the sea. Only three breeding areas of NZ sea lions exist today after the species was extirpated from its historical range (NZ mainland). The study was conducted at the Sandy Bay breeding colony, Auckland Islands, between December 2002 and March 2003. We used daily Global Positioning System locations of breeding females with pups and mapping in a Geographic Information System to determine terrestrial habitat use and preferences. Slopes less than 20° were preferred throughout the study. Females chose nursing sites with a seasonal change, preferentially based on the distance from the sea and habitat type. Comparisons with the other breeding colonies of NZ sea lions are presented and data are discussed in the context of the recolonization of the NZ mainland. Overall, the most suitable terrestrial habitat configuration for a breeding aggregation of NZ sea lions appears to be a sandy beach, with a wide area above high tide and moderate intertidal zone (for breeding), backed with vegetated sand dunes and forest on primarily flat terrain (for later dispersion).     

卡拉麦里山有蹄类自然保护区鹅喉羚生境适宜性评价

在卡拉麦里山保护区内选取具有代表性的研究样区约1 447 km² 作为评价区,基于2005 年以来对卡拉麦里山有蹄类自然保护区鹅喉羚野外调查和样方采集,利用模糊赋值法建立生境评价模型,结合GIS 技术的空间分析功能,以植被类型、坡度、坡向和人类干扰活动为评价因子进行生境适宜性研究。结果表明:春、夏、秋、冬四季实际生境面积分别为 1146.9km² 、1137.1 km² 、991.6 km² 和499.8 km² ,分别占研究区域总面积的79.3% 、78.6% 、68.5% 和34.5% ;其中适宜生境面积分别为304.1 km² 、599.4 km² 、303.6 km² 和56.2 km² ,分别占研究区域总面积的21.0% 、41.4% 、21.0% 和3.9% 。人类活动影响下,一些潜在生境转变为不适宜生境,春、夏、秋、冬四季生境分别丧失4.0% 、3.1% 、4.3% 和48.5%。在各种人类活动中,居民点、国道、采矿点对鹅喉羚生境具有一定的干扰作用,使其质量有所降低,但未造成严重影响;而冬季牧民的放牧行为,导致鹅喉羚的适宜生境被家畜占据,鹅喉羚生境质量大面积下降。本文基于评价结果提出了保护措施。     

Landscape heterogeneity metrics as indicators of bird diversity: Determining the optimal spatial scales in different landscapes

Species distribution models are often used to study the biodiversity of ecosystems. The modelling process uses a number of parameters to predict others, such as the occurrence of determinate species, population size, habitat suitability or biodiversity. It is well known that the heterogeneity of landscapes can lead to changes in species’ abundance and biodiversity. However, landscape metrics depend on maps and spatial scales when it comes to undertaking a GIS analysis.We explored the goodness of fit of several models using the metrics of landscape heterogeneity and altitude as predictors of bird diversity in different landscapes and spatial scales. Two variables were used to describe biodiversity: bird richness and trophic level diversity, both of which were obtained from a breeding bird survey by means of point counts. The relationships between biodiversity and landscape metrics were compared using multiple linear regressions. All of the analyses were repeated for 14 different spatial scales and for cultivated, forest and grassland environments to determine the optimal spatial scale for each landscape typology.Our results revealed that the relationships between species’ richness and landscape heterogeneity using 1:10,000 land cover maps were strongest when working on a spatial scale up to a radius of 125–250 m around the sampled point (circa 4.9–19.6 ha). Furthermore, the correlation between measures of landscape heterogeneity and bird diversity was greater in grasslands than in cultivated or forested areas. The multi-spatial scale approach is useful for (a) assessing the accuracy of surrogates of bird diversity in different landscapes and (b) optimizing spatial model procedures for biodiversity mapping, mainly over extensive areas.     

Novel insights into habitat suitability for Amazonian freshwater mussels linked with hydraulic and landscape drivers

Novel insights into habitat suitability for two Unionida freshwater mussels, Castalia ambigua Lamarck, 1819 (Hyriidae) and Anodontites elongatus (Swainson, 1823) (Mycetopodidae), are presented on the basis of hydraulic variables linked with the riverbed in six 500‐m reaches in an eastern Amazonian river basin. Within the reaches, there was strong habitat heterogeneity in hydrodynamics and substrate composition. In addition, we investigated stressors based on landscape modification that are associated with declines in mussel density. We measured hydraulic variables for each 500‐m reach, and landscape stressors at two spatial scales (subcatchment and riparian buffer forest). We used the Random Forest algorithm, a tree‐based model, to predict the hydraulic variables linked with habitat suitability for mussels, and to predict which landscape stressors were most associated with mussel density declines. Both mussel species were linked with low substrate heterogeneity and greater riverbed stability (low Froude and Reynolds numbers), especially at high flow (low stream power). Different sediment grain size preferences were observed between mussel species: Castalia ambigua was associated with medium sand and Anodontites elongatus with medium and fine sand. Declines in mussel density were associated with modifications linked to urbanization at small scales (riparian buffer forest), especially with percent of and distance from rural settlements, distance to the nearest street, and road density. In summary, the high variance explained in both hydraulic and landscape models indicated high predictive power, suggesting that our findings may be extrapolated and used as a baseline to test hypotheses of habitat suitability in other Amazonian rivers for Castalia ambigua and Anodontites elongatus and also for other freshwater mussel species. Our results highlight the urgent need for aquatic habitat conservation to maintain sheltered habitats during high flow as well as mitigate the effects of landscape modifications at the riparian buffer scale, both of which are important for maintaining dense mussel populations and habitat quality.     

Assessment of habitat quality for four small mammal species of the Monte Desert, Argentina

In the temperate desert of Argentina, the combined action of climatic and anthropogenic factors has contributed to the formation of a highly heterogeneous landscape. In the central region of the Monte desert, four small mammal species (Eligmodontia typus, Calomys musculinus, Akodon molinae and Graomys griseoflavus) coexist and show different habitat uses in response to spatial variability. Three main habitat types are present in the region: mesquite forest, the creosotebush community and sand dunes. These habitat types are present also in the surrounding grazing area.The objective of this study was to determine habitat quality for these species in a protected area (Reserve MaB Ñacuñán) and in the adjacent grazed area. For each species we estimated demographic parameters that are highly correlated to fitness in each habitat, and for both treatments (protected and grazed).We found that the protected area offered a higher quality habitat than the grazed area for all species, but principally A. molinae and G. griseoflavus. At a local scale, we found that A. molinae and C. musculinus clearly showed higher fitness in the more complex habitats as the creosotebush community and the mesquite forest. In contrast, for E. typus, open and simplest patches, such as sand dunes, were optimal for its survival and reproduction.     

科尔沁沙地植物群落圆环状分布成因地统计学分析

调查发现,在科尔沁沙地中部部分梁窝状沙丘的丘顶、丘坡和丘间地,植物群落依生境梯度而分布,呈典型圆环状分布格局．沙层水分状况分析表明,同一沙丘不同部位的沙层含水量为丘顶低于丘坡地,且二者均低于丘间地．地统计学分析表明,无论是流动沙丘还是固定沙丘,从丘顶、丘坡到丘间地,沙层含水量的空间异质性呈规律性的变化,块金值与基台值之比逐渐减小,变程逐渐减小,分维数逐渐增大,表明随机部分引起的空间异质性逐渐降低,而由空间自相关部分引起的空间异质性逐渐增高．因此,沙丘不同部位沙层含水量及其空间异质性的有规律变化是科尔沁沙地中部部分梁窝状沙丘植物群落呈圆环状分布格局的一个重要原因．     

Relationships between survival and habitat suitability of semi‐aquatic mammals

Spatial distribution and habitat selection are integral to the study of animal ecology. Habitat selection may optimize the fitness of individuals. Hutchinsonian niche theory posits the fundamental niche of species would support the persistence or growth of populations. Although niche‐based species distribution models (SDMs) and habitat suitability models (HSMs) such as maximum entropy (Maxent) have demonstrated fair to excellent predictive power, few studies have linked the prediction of HSMs to demographic rates. We aimed to test the prediction of Hutchinsonian niche theory that habitat suitability (i.e., likelihood of occurrence) would be positively related to survival of American beaver (Castor canadensis), a North American semi‐aquatic, herbivorous, habitat generalist. We also tested the prediction of ideal free distribution that animal fitness, or its surrogate, is independent of habitat suitability at the equilibrium. We estimated beaver monthly survival probability using the Barker model and radio telemetry data collected in northern Alabama, United States from January 2011 to April 2012. A habitat suitability map was generated with Maxent for the entire study site using landscape variables derived from the 2011 National Land Cover Database (30‐m resolution). We found an inverse relationship between habitat suitability index and beaver survival, contradicting the predictions of niche theory and ideal free distribution. Furthermore, four landscape variables selected by American beaver did not predict survival. The beaver population on our study site has been established for 20 or more years and, subsequently, may be approaching or have reached the carrying capacity. Maxent‐predicted increases in habitat use and subsequent intraspecific competition may have reduced beaver survival. Habitat suitability‐fitness relationships may be complex and, in part, contingent upon local animal abundance. Future studies of mechanistic SDMs incorporating local abundance and demographic rates are needed.