Climate Change,Marine Environments,and the U.S. Endangered Species Act

Climate change is expected to be a top driver of global biodiversity loss in the 21st century. It poses new challenges to conserving and managing imperiled species, particularly in marine and estuarine ecosystems. The use of climate‐related science in statutorily driven species management, such as under the U.S. Endangered Species Act (ESA), is in its early stages. This article provides an overview of ESA processes, with emphasis on the mandate to the National Marine Fisheries Service (NMFS) to manage listed marine, estuarine, and anadromous species. Although the ESA is specific to the United States, its requirements are broadly relevant to conservation planning. Under the ESA, species, subspecies, and “distinct population segments” may be listed as either endangered or threatened, and taking of most listed species (harassing, harming, pursuing, wounding, killing, or capturing) is prohibited unless specifically authorized via a case‐by‐case permit process. Government agencies, in addition to avoiding take, must ensure that actions they fund, authorize, or conduct are not likely to jeopardize a listed species’ continued existence or adversely affect designated critical habitat. Decisions for which climate change is likely to be a key factor include: determining whether a species should be listed under the ESA, designating critical habitat areas, developing species recovery plans, and predicting whether effects of proposed human activities will be compatible with ESA‐listed species’ survival and recovery. Scientific analyses that underlie these critical conservation decisions include risk assessment, long‐term recovery planning, defining environmental baselines, predicting distribution, and defining appropriate temporal and spatial scales. Although specific guidance is still evolving, it is clear that the unprecedented changes in global ecosystems brought about by climate change necessitate new information and approaches to conservation of imperiled species. El Cambio Climático, los Ecosistemas Marinos y el Acta Estadunidense de Especies en Peligro     

Latent Extinction and Invasion Risk of Crayfishes in the Southeastern United States

Abstract: Crayfishes are both a highly imperiled invertebrate group as well as one that has produced many invasive species, which have negatively affected freshwater ecosystems throughout the world. We performed a trait analysis for 77 crayfishes from the southeastern United States in an attempt to understand which biological and ecological traits make these species prone to imperilment or invasion, and to predict which species may face extinction or become invasive in the future. We evaluated biological and ecological traits with principal coordinate analysis and classification trees. Invasive and imperiled crayfishes occupied different positions in multivariate trait space, although crayfishes invasive at different scales (extraregional vs. extralimital) were also distinct. Extraregional crayfishes (large, high fecundity, habitat generalists) were most distinct from imperiled crayfishes (small, low fecundity, habitat specialists), thus supporting the “two sides of the same coin” hypothesis. Correct classification rates for assignment of crayfishes as invasive or imperiled were high (70–80%), even when excluding the highly predictive but potentially confounding trait of range size (75–90%). We identified a number of species that, although not currently listed as imperiled or found outside their native range, possess many of the life‐history and ecological traits characteristic of currently invasive or imperiled taxa. Such species exhibit a high latent risk of extinction or invasion and consequently should be the focus of proactive conservation or management strategies. Our results illustrate the utility of trait‐based approaches for taxonomic groups such as invertebrates, for which detailed species‐specific data are rare and conservation resources are chronically limited.     

Estimation of Core Terrestrial Habitat for Stream-Breeding Salamanders and Delineation of Riparian Buffers for Protection of Biodiversity

Abstract:  Many species of wildlife depend on riparian habitats for various life-history functions (e.g., breeding, foraging, overwintering). Although this unique habitat is critical for many species, delineations of riparian zones and buffers for various taxa are lacking. Typically when buffer zones are determined to mitigate edge effects, they are based on criteria that protect aquatic resources alone and do not consider impacts to wildlife and other terrestrial resources. Using two different survey methods (area-constrained daytime searches and nighttime visual encounter searches), we estimated core terrestrial habitat and buffer widths for stream-breeding salamanders in southern Appalachian streams from May to August 2004. A core terrestrial habitat of 27.0 m encompassed 95% of the salamander assemblage (four species of stream plethodontids), and an additional 50 m (to buffer edge effects) yielded a total buffer of 77.0 m. When each species of the assemblage was analyzed separately, the maximum core terrestrial habitat needed for the Blue Ridge two-lined salamander ( Eurycea wilderae ), a dominant member and the farthest-ranging species from the stream, was 42.6 m. Thus, we recommend an overall buffer width of 92.6 m in southern Appalachian streams. To protect stream amphibians and other wildlife dependent on riparian areas, land managers and policy makers must consider conserving more than aquatic resources alone. Developing core terrestrial habitat estimates and buffer zone widths for wildlife populations is a critical first step in the conservation of many semiaquatic organisms and protecting biodiversity.     

Mitigation of Habitat "Take": Application to Habitat Conservation Planning

One of the most important provisions of the U.S. Endangered Species Act precludes the "taking" of listed species on both public and private land. In past Endangered Species Act litigation, take has been broadly interpreted to include the destruction or modification of habitats as well as the direct killing of animals. This requirement created an extensive burden on private landowners to provide habitats for listed species. This burden was substantially lessened when the ESA was modified in 1982 to allow incidental takings conditioned on preparation of a satisfactory "habitat conservation plan." Because the majority of listed species are imperiled due to habitat modification, most habitat conservation plans must demonstrate defensible methods to mitigate against incidental habitat loss. A review of HCPs for the Northern Spotted Owl ( Strix occidentalis), and other species, indicates that mitigation solutions are often arbitrary, lacking an empirical foundation in the species' life history requirements. Based on data from the Spotted Owl, we illustrate a biologically based method for estimating the areal requirements necessary to mitigate against the take of essential habitats. Toward this goal we adopt the concept of "core area," that portion of an animal's home range that receives disproportionate use. We estimated core areas by means of the adaptive kernel density function and tested against a null distribution of animal use that assumes a bivariate, uniform distribution of locations within the home range. The method we illustrate, which is defensible, repeatable, and empirical, is a clear improvement over the ad hoc methods used in many habitat conservation plans. Further, the methods we propose should be applicable to a large number of terrestrial species for which home range is a meaningful concept.     

Quantitative Determination of Rarity of Freshwater Fishes and Implications for Imperiled‐Species Designations

Abstract: Conserving rare species and protecting biodiversity and ecosystem functioning depends on sound information on the nature of rarity. Rarity is multidimensional and has a variety of definitions, which presents the need for a quantitative classification scheme with which to categorize species as rare or common. We constructed such a classification for North American freshwater fishes to better describe rarity in fishes and provide researchers and managers with a tool to streamline conservation efforts. We used data on range extents, habitat specificities, and local population sizes of North American freshwater fishes and a variety of quantitative methods and statistical decision criteria, including quantile regression and a cost‐function algorithm to determine thresholds for categorizing a species as rare or common. Species fell into eight groups that conform to an established framework for rarity. Fishes listed by the American Fisheries Society (AFS) as endangered, threatened, or vulnerable were most often rare because their local population sizes were low, ranges were small, and they had specific habitat needs, in that order, whereas unlisted species were most often considered common on the basis of these three factors. Species with large ranges generally had few specific habitat needs, whereas those with small ranges tended to have narrow habitat specificities. We identified 30 species not designated as imperiled by AFS that were rare along all dimensions of rarity and may warrant further study or protection, and we found three designated species that were common along all dimensions and may require a review of their imperilment status. Our approach could be applied to other taxa to aid conservation decisions and serve as a useful tool for future revisions of listings of fish species.     

Extinction Rates of North American Freshwater Fauna

Abstract: Since 1900, 123 freshwater animal species have been recorded as extinct in North America. Hundreds of additional species of fishes, mollusks, crayfishes, and amphibians are considered imperiled. Using an exponential decay model, we derived recent and future extinction rates for North American freshwater fauna that are five times higher than those for terrestrial fauna. Assuming that imperiled freshwater species will not survive throughout the next century, our model projects a future extinction rate of 4% per decade, which suggests that North America's temperate freshwater ecosystems are being depleted of species as rapidly as tropical forests.     

Using expert knowledge to support Endangered Species Act decision-making for data-deficient species

Many questions relevant to conservation decision-making are characterized by extreme uncertainty due to lack of empirical data and complexity of the underlying ecologic processes, leading to a rapid increase in the use of structured protocols to elicit expert knowledge. Published ecologic applications often employ a modified Delphi method, where experts provide judgments anonymously and mathematical aggregation techniques are used to combine judgments. The Sheffield elicitation framework (SHELF) differs in its behavioral approach to synthesizing individual judgments into a fully specified probability distribution for an unknown quantity. We used the SHELF protocol remotely to assess extinction risk of three subterranean aquatic species that are being considered for listing under the U.S. Endangered Species Act. We provided experts an empirical threat assessment for each known locality over a video conference and recorded judgments on the probability of population persistence over four generations with online submission forms and R-shiny apps available through the SHELF package. Despite large uncertainty for all populations, there were key differences between species’ risk of extirpation based on spatial variation in dominant threats, local land use and management practices, and species’ microhabitat. The resulting probability distributions provided decision makers with a full picture of uncertainty that was consistent with the probabilistic nature of risk assessments. Discussion among experts during SHELF's behavioral aggregation stage clearly documented dominant threats (e.g., development, timber harvest, animal agriculture, and cave visitation) and their interactions with local cave geology and species’ habitat. Our virtual implementation of the SHELF protocol demonstrated the flexibility of the approach for conservation applications operating on budgets and time lines that can limit in-person meetings of geographically dispersed experts.     

A Diverse and Endangered Aquatic Ecosystem of the Southeast United States

We document the biodiversity and conservation status of an extraordinarily diverse and endangered ecosystem in the United States that has failed to attract the same attention as tropical ecosystems—the rivers and streams of Alabama and adjoining states. Relative to North America as a whole, Alabama is a highlight of aquatic diversity supporting 38% of native fresh water fishes, 43% of native freshwater gill-breathing snails, 60% of native mussels, and 52% of native freshwater turtles. Of these, 41%, 77%, 34%, and 22% of the fishes, snails, mussels, and turtles, respectively, are endemic to Alabama and an adjacent state. Like many tropical systems of developing nations, this fauna is in an imperiled state, with 10%, 65%, 69%, and 43% of Alabama's fishes, gill-breathing snails, mussels, and turtles, respectively, considered either extinct, endangered, threatened, or of special concern. Unlike tropical systems, however, little effort has been made to protect the taxa and their habitats. Only 40% of fishes, 1% of gill-breathing snails, 32% of mussels, and 20% of freshwater turtles are formally listed as either threatened or endangered via the U.S. Endangered Species Act of 1973; no critical habitat has been protected. Clearly, the biodiversity crisis in not limited to tropical systems of developing nations. Although the Endangered Species Act of 1973 helps to ensure a future of sustainable diversity, efforts must be made to hasten recognition, protection, and recovery of critical habitat, particularly for hotspots such as the aquatic systems of Alabama.     

Terrestrial Reserve Networks Do Not Adequately Represent Aquatic Ecosystems

Abstract: Protected areas are a cornerstone of conservation and have been designed largely around terrestrial features. Freshwater species and ecosystems are highly imperiled, but the effectiveness of existing protected areas in representing freshwater features is poorly known. Using the inland waters of Michigan as a test case, we quantified the coverage of four key freshwater features (wetlands, riparian zones, groundwater recharge, rare species) within conservation lands and compared these with representation of terrestrial features. Wetlands were included within protected areas more often than expected by chance, but riparian zones were underrepresented across all (GAP 1–3) protected lands, particularly for headwater streams and large rivers. Nevertheless, within strictly protected lands (GAP 1–2), riparian zones were highly represented because of the contribution of the national Wild and Scenic Rivers Program. Representation of areas of groundwater recharge was generally proportional to area of the reserve network within watersheds, although a recharge hotspot associated with some of Michigan's most valued rivers is almost entirely unprotected. Species representation in protected areas differed significantly among obligate aquatic, wetland, and terrestrial species, with representation generally highest for terrestrial species and lowest for aquatic species. Our results illustrate the need to further evaluate and address the representation of freshwater features within protected areas and the value of broadening gap analysis and other protected‐areas assessments to include key ecosystem processes that are requisite to long‐term conservation of species and ecosystems. We conclude that terrestrially oriented protected‐area networks provide a weak safety net for aquatic features, which means complementary planning and management for both freshwater and terrestrial conservation targets is needed.     

An unusual trophic subsidy and species dominance in a tropical stream

In classical theory, species are assumed to achieve dominance through competitive exclusion, but if food resources are limiting, cross-habitat trophic subsidies could also underpin dominance. The impact of dominant species on community dynamics may depend on the energy base of population size. We report on an unusual, spatially subsidized population of a tropical, stream-dwelling crab that dominates the benthic fauna of a Kenyan stream. Diet and stable isotope analyses indicated that this crab is a true omnivore, with terrestrial subsidies dominating both plant and animal resources. Unusually, the animal prey included almost no aquatic invertebrates. Instead, a single species of ant constituted approximately 35% of the annual diet (stomach contents analysis) and up to 90% of assimilated nitrogen (estimates from stable isotope analysis). Ants may be pivotal to enabling crab dominance, and this crab may be largely disconnected from the local trophic network for its dietary needs. The paucity of other invertebrates in the stream community suggests that this super-dominant crab is a strong interactor that suppresses aquatic invertebrate populations. Common stabilizing attributes of spatially subsidized food webs (e.g., asynchronous prey availability, wide feeding niche, consumer migration) were absent from this system, and although apparently stable, it may be vulnerable to disturbance in the donor habitat.     

Going with the flow: using species-discharge relationships to forecast losses in fish biodiversity

In response to the scarcity of tools to make quantitative forecasts of the loss of aquatic species from anthropogenic effects, we present a statistical model that relates fish species richness to river discharge. Fish richness increases logarithmically with discharge, an index of habitat space, similar to a species-area curve in terrestrial systems. We apply the species-discharge model as a forecasting tool to build scenarios of changes in riverine fish richness from climate change, water consumption, and other anthropogenic drivers that reduce river discharge. Using hypothetical reductions in discharges (of magnitudes that have been observed in other rivers), we predict that reductions of 20-90% in discharge would result in losses of 2-38% of the fish species in two biogeographical regions in the United States (Lower Ohio-Upper Mississippi and Southeastern). Additional data on the occurrence of specific species relative to specific discharge regimes suggests that fishes found exclusively in high discharge environments (e.g., Shovelnose sturgeon) would be most vulnerable to reductions in discharge. Lag times in species extinctions after discharge reduction provide a window of opportunity for conservation efforts. Applications of the species-discharge model can help prioritize such management efforts among species and rivers.     

Diverging response patterns of terrestrial and aquatic species to hydromorphological restoration

Although experiences with ecological restoration continue to accumulate, the effectiveness of restoration for biota remains debated. We complemented a traditional taxonomic analysis approach with information on 56 species traits to uncover the responses of 3 aquatic (fish, macroinvertebrates, macrophytes) and 2 terrestrial (carabid beetles, floodplain vegetation) biotic groups to 43 hydromorphological river restoration projects in Germany. All taxonomic groups responded positively to restoration, as shown by increased taxonomic richness (10–164%) and trait diversity (habitat, dispersal and mobility, size, form, life history, and feeding groups) (15–120%). Responses, however, were stronger for terrestrial than aquatic biota, and, contrary to our expectation, taxonomic responses were stronger than those of traits. Nevertheless, trait analysis provided mechanistic insights into the drivers of community change following restoration. Trait analysis for terrestrial biota indicated restoration success was likely enhanced by lateral connectivity and reestablishment of dynamic processes in the floodplain. The weaker response of aquatic biota suggests recovery was hindered by the persistence of stressors in the aquatic environment, such as degraded water quality, dispersal constraints, and insufficient hydromorphological change. Therefore, river restoration requires combined local- and regional-scale approaches to maximize the response of both aquatic and terrestrial organisms. Due to the contrasting responses of aquatic and terrestrial biota, the planning and assessment of river restoration outcomes should consider effects on both components of riverine landscapes.     

Effect of scale on trait predictors of species responses to agriculture

Species persistence in human‐altered landscapes can depend on factors operating at multiple spatial scales. To understand anthropogenic impacts on biodiversity, it is useful to examine relationships between species traits and their responses to land‐use change. A key knowledge gap concerns whether these relationships vary depending on the scale of response under consideration. We examined how local‐ and large‐scale habitat variables influence the occupancy dynamics of a bird community in cloud forest zones in the Colombian Chocó‐Andes. Using data collected across a continuum of forest and agriculture, we examined which traits best predict species responses to local variation in farmland and which traits best predict species responses to isolation from contiguous forest. Global range size was a strong predictor of species responses to agriculture at both scales; widespread species were less likely to decline as local habitat cover decreased and as distance from forest increased. Habitat specialization was a strong predictor of species responses only at the local scale. Open‐habitat species were particularly likely to increase as pasture increased, but they were relatively insensitive to variation in distance to forest. Foraging plasticity and flocking behavior were strong predictors of species responses to distance from forest, but not their responses to local habitat. Species with lower plasticity in foraging behaviors and obligate flock‐following species were more likely to decline as distance from contiguous forest increased. For species exhibiting these latter traits, persistence in tropical landscapes may depend on the protection of larger contiguous blocks of forest, rather than the integration of smaller‐scale woodland areas within farmland. Species listed as threatened or near threatened on the International Union for Conservation of Nature Red List were also more likely to decline in response to both local habitat quality and isolation from forest relative to least‐concern species, underlining the importance of contiguous forests for threatened taxa.     

Topology of Extinction and Endangerment of Native Fishes in the Pacific Northwest and California (U.S.A.)

Recent studies have provided a broad data base on extinction and endangerment of species, subspecies, and distinct populations of inland fishes in western North America. Development of a synoptic, regional-scale image of extinction and risk of extinction is complicated by the small size and linear distribution of fluvial aquatic habitats and by interspecific variation in areal extent of populations. I developed a regional map of extinction-risk isopleths based on the number of extinct and persistently declining species in drainage basins of the Pacific Northwest and California. This topological synthesis is useful for delineating and monitoring areas of historic and ongoing loss of aquatic biodiversity, and for relating losses to patterns of land use and habitat modification, climate hydrology, and geomorphology. From an ecological perspective, endangerment of numerous indigenous populations of seven widely distributed species of anadromous salmonids in this region is as important as the more local, diffuse effects of declines in more than 60 endemic, nonanadromous species and subspecies. The simultaneous decline of numerous taxa in basins not afflicted with dams or diversions suggests that cumulative damage to aquatic habitats caused by logging, grazing, urbanization, and other land uses plays a major role in icthyofaunal impoverishment.     

Demographic Consequences of Terrestrial Habitat Loss for Pool‐Breeding Amphibians: Predicting Extinction Risks Associated with Inadequate Size of Buffer Zones

Abstract: Much of the biodiversity associated with isolated wetlands requires aquatic and terrestrial habitat to maintain viable populations. Current federal wetland regulations in the United States do not protect isolated wetlands or extend protection to surrounding terrestrial habitat. Consequently, some land managers, city planners, and policy makers at the state and local levels are making an effort to protect these wetland and neighboring upland habitats. Balancing human land‐use and habitat conservation is challenging, and well‐informed land‐use policy is hindered by a lack of knowledge of the specific risks of varying amounts of habitat loss. Using projections of wood frog (Rana sylvatica) and spotted salamander (Ambystoma maculatum) populations, we related the amount of high‐quality terrestrial habitat surrounding isolated wetlands to the decline and risk of extinction of local amphibian populations. These simulations showed that current state‐level wetland regulations protecting 30 m or less of surrounding terrestrial habitat are inadequate to support viable populations of pool‐breeding amphibians. We also found that species with different life‐history strategies responded differently to the loss and degradation of terrestrial habitat. The wood frog, with a short life span and high fecundity, was most sensitive to habitat loss and isolation, whereas the longer‐lived spotted salamander with lower fecundity was most sensitive to habitat degradation that lowered adult survival rates. Our model results demonstrate that a high probability of local amphibian population persistence requires sufficient terrestrial habitat, the maintenance of habitat quality, and connectivity among local populations. Our results emphasize the essential role of adequate terrestrial habitat to the maintenance of wetland biodiversity and ecosystem function and offer a means of quantifying the risks associated with terrestrial habitat loss and degradation.     

Not Knowing, Not Recording, Not Listing: Numerous Unnoticed Mollusk Extinctions

Abstract:  Mollusks are the group most affected by extinction according to the 2007 International Union for Conservation of Nature (IUCN) Red List, despite the group having not been evaluated since 2000 and the quality of information for invertebrates being far lower than for vertebrates. Altogether 302 species and 11 subspecies are listed as extinct on the IUCN Red List. We reevaluated mollusk species listed as extinct through bibliographic research and consultation with experts. We found that the number of known mollusk extinctions is almost double that of the IUCN Red List. Marine habitats seem to have experienced few extinctions, which suggests that marine species may be less extinction prone than terrestrial and freshwater species. Some geographic and ecologic biases appeared. For instance, the majority of extinctions in freshwater occurred in the United States. More than 70% of known mollusk extinctions took place on oceanic islands, and a one-third of these extinctions may have been caused precipitously by introduction of the predatory snail Euglandina rosea. We suggest that assessment of the conservation status of invertebrate species is neglected in the IUCN Red List and not managed in the same way as for vertebrate species .     

Ecological Attributes of Extinction-Prone Species: Loss of Freshwater Fishes of Virginia

I examined patterns of extirpation among Virginia's 197 historically native freshwater fish species to address the following questions: (1) Are extinction-prone species ecologically distinct? and (2) Are distinctive features similar to those identified for extinction-prone species in terrestrial systems? All species were assigned to categories for a series of attributes reflecting geographical distribution, habitat use, trophic habits, life history, size, and reproductive behavior. Associations between species that had been extirpated and those that had not were examined for each attribute. Univariate associations were observed between extirpation and three ecological attributes: diadromy, limited physiographic range, and limited range of water sizes. Species specialized with respect to multiple ecological attributes also were especially likely to be extirpated. These associations reflected the effects of reduced habitat area and increased isolation (insularization), which are also important determinants of extinction in terrestrial systems. Multivariate analyses suggested that extirpated species were ecologically similar to each other, but were not completely distinct from the nonextirpated fauna. My results suggest that ecological knowledge of species can help identify extinction-prone species and provide a basis for proactive conservation. Current approaches to conservation, which are largely reactive and piecemeal, are inadequate to protect biodiversity. Because aquatic degradation is complex and pervasive, conservation of aquatic biodiversity requires proactive comprehensive approaches to water resource management, including emphasis on protecting the ecological integrity of entire systems.     

Scaling up private land conservation to meet recovery goals for grassland birds

Long-term population declines have elevated recovery of grassland avifauna to among the highest conservation priorities in North America. Because most of the Great Plains is privately owned, recovery of grassland bird populations depends on voluntary conservation with strong partnerships between private landowners and resource professionals. Despite large areas enrolled in voluntary practices through U.S. Department of Agriculture's Lesser Prairie-chicken (Tympanuchus pallidicinctus) Initiative (LPCI), the effectiveness of Farm Bill investments for meeting wildlife conservation goals remains an open question. Our objectives were to evaluate extents to which Conservation Reserve Program (CRP) and LPCI-grazing practices influence population densities of grassland birds; estimate relative contributions of practices to regional bird populations; and evaluate percentages of populations conserved relative to vulnerability of species. We designed a large-scale impact-reference study and used the Integrated Monitoring in Bird Conservation Regions program to evaluate bird population targets of the Playa Lakes Joint Venture. We used point transect distance sampling to estimate density and population size for 35 species of grassland birds on private lands enrolled in native or introduced CRP plantings and LPCI-prescribed grazing. Treatment effects indicated CRP plantings increased densities of three grassland obligates vulnerable to habitat loss, and LPCI grazing increased densities of four species requiring heterogeneity in dense, tall-grass structure (α = 0.1). Population estimates in 2016 indicated the practices conserved breeding habitat for 4.5 million birds (90% CI: 4.0–5.1), and increased population sizes of 16 species , totaling 1.8 million birds (CI: 1.4–2.4). Conservation practices on private land benefited the most vulnerable grassland obligate species (AIC_c weight = 0.53). By addressing habitat loss and degradation in agricultural landscapes, conservation on private land provides a solution to declining avifauna of North America and scales up to meet population recovery goals for the most imperiled grassland birds.     

Life-history adaptations to arboreality in snakes

If selective forces on locomotor ability and reproductive biology differ among habitats, we expect to see relationships between habitat, morphology, and life-history traits. Comparative (phylogenetically based) analysis of data from 12 pythonid and 12 boid snake species reveals multiple evolutionary shifts in habitat use, notably in the evolution of arboreal habits. Compared to terrestrial and aquatic taxa of the same overall body size, arboreal species have narrower and more laterally compressed bodies and relatively longer tails. Offspring sizes are not affected by arboreality, but presumably reflecting space constraints within their narrow bodies, arboreal species (1) produce smaller clutch sizes relative to maternal body length and (2) have left and right ovaries that overlap little if at all along the length of the body (i.e., the right ovary is positioned anterior to the left ovary) whereas in terrestrial snakes the two ovaries overlap along much of their length. This modification of ovarian morphology in arboreal snakes presumably reduces the degree of bodily distension during vitellogenesis and pregnancy, thus enhancing climbing ability and camouflage among the branches.     

异质生境空心莲子草植物群落组成及物种多样性研究

研究草海湿地空心莲子草（Alternanthera philoxeroides）入侵对异质生境植物群落生活型、物种组成、重要值及物种多样性的影响,结果表明,①异质生境中群落生活型组成比例有差异,水生生境地面芽植物（H）∶隐芽植物（Cr）∶一年生植物（Th）为56∶38∶6;湿生生境为43∶36∶21。②异质生境中物种组成差异明显,水生生境试验样方包含9科13属16种,对照样方（无空心莲子草入侵）包含10科19属23种;湿生生境试验样方包含8科14属14种,对照样方有10科19属22种。空心莲子草入侵不同生境后群落内物种重要值变化及差异均较大,部分乡土物种在试验样方中减少甚至消失,水生生境和湿生生境群落中分别有30.44%、36.36%物种消失。③异质生境中空心莲子草重要值与物种多样性指数、生态优势度指数、均匀度指数均呈二项式函数关系;两种生境的多样性指数和均匀度指数均表现为随空心莲子草重要值先增加后减少,生态优势度指数均为先减少后增加,空心莲子草入侵对乡土植物群落物种多样性产生正向影响和负向影响,在群落中其重要值达25.8（水生生境）、18.9（湿生生境）时,是乡土植物多样性下降的临界点。