The Effectiveness of Surrogate Taxa for the Representation of Biodiversity

Abstract: Biodiversity is too complex to measure directly, so conservation planning must rely on surrogates to estimate the biodiversity of sites. The species richness of selected taxa is often used as a surrogate for the richness of other taxa. Surrogacy values of taxa have been evaluated in diverse contexts, yet broad trends in their effectiveness remain unclear. We reviewed published studies testing the ability of species richness of surrogate taxa to capture the richness of other (target) taxa. We stratified studies into two groups based on whether a complementarity approach (surrogates used to select a combination of sites that together maximize total species richness for the taxon) or a richness‐hotspot approach (surrogates used to select sites containing the highest species richness for the taxon) was used. For each comparison of one surrogate taxon with one target, we used the following predictor variables: biome, spatial extent of study area, surrogate taxon, and target taxon. We developed a binary response variable based on whether the surrogate taxon provided better than random representation of the target taxon. For studies that used an evaluation approach that was not based on better than random representation of target taxa, we based the response variable on the interpretation of results in the original study. We performed a categorical regression to elucidate trends in the effectiveness of surrogate taxa with regard to each of the predictor variables. A surrogate was 25% more likely to be effective with a complementarity approach than with a hotspot approach. For hotspot‐based approaches, biome, extent of study, surrogate taxon, and target taxon significantly influenced effectiveness of the surrogate. For complementarity‐based approaches, biome, extent, and surrogate taxon significantly influenced effectiveness of the surrogate. For all surrogate evaluations, biome explained the greatest amount of variation in surrogate effectiveness. From most to least, extent, surrogate taxon, and target taxon explained the most variation after biome. Surrogate taxa were most effective in grasslands and in some cases boreal zones, deserts, and tropical forests; surrogate taxa also were more effective in studies examining larger areas. Herpetofauna were the most effective taxon as both surrogate and target when a richness‐hotspot approach was used; however, herpetofauna were analyzed in fewer studies, so this result is tentative. For complementarity approaches, taxa that are easy to measure and tend to have a large number of habitat specialists distributed collectively across broad environmental gradients (e.g., plants, birds, and mammals) were the most effective surrogates.     

Factors influencing incidental representation of previously unknown conservation features in marine protected areas

Spatially explicit information on species distributions for conservation planning is invariably incomplete; therefore, the use of surrogates is required to represent broad‐scale patterns of biodiversity. Despite significant interest in the effectiveness of surrogates for predicting spatial distributions of biodiversity, few researchers have explored questions involving the ability of surrogates to incidentally represent unknown features of conservation interest. We used the Great Barrier Reef marine reserve network to examine factors affecting incidental representation of conservation features that were unknown at the time the reserve network was established. We used spatially explicit information on the distribution of 39 seabed habitats and biological assemblages and the conservation planning software Marxan to examine how incidental representation was affected by the spatial characteristics of the features; the conservation objectives (the minimum proportion of each feature included in no‐take areas); the spatial configuration of no‐take areas; and the opportunity cost of conservation. Cost was closely and inversely correlated to incidental representation. However, incidental representation was achieved, even in a region with only coarse‐scale environmental data, by adopting a precautionary approach that explicitly considered the potential for unknown features. Our results indicate that incidental representation is enhanced by partitioning selection units along biophysical gradients to account for unknown within‐feature variability and ensuring that no‐take areas are well distributed throughout the region; by setting high conservation objectives that (in this case >33%) maximize the chances of capturing unknown features incidentally; and by carefully considering the designation of cost to planning units when using decision‐support tools for reserve design. The lessons learned from incidental representation in the Great Barrier Reef have implications for conservation planning in other regions, particularly those that lack detailed environmental and ecological data.     

The effectiveness of surrogate taxa to conserve freshwater biodiversity

Establishing protected areas has long been an effective conservation strategy and is often based on readily surveyed species. The potential of any freshwater taxa to be a surrogate for other aquatic groups has not been explored fully. We compiled occurrence data on 72 species of freshwater fishes, amphibians, mussels, and aquatic reptiles for the Great Plains, Wyoming (U.S.A.). We used hierarchical Bayesian multispecies mixture models and MaxEnt models to describe species’ distributions and the program Zonation to identify areas of conservation priority for each aquatic group. The landscape‐scale factors that best characterized aquatic species’ distributions differed among groups. There was low agreement and congruence among taxa‐specific conservation priorities (<20%), meaning no surrogate priority areas would include or protect the best habitats of other aquatic taxa. Common, wideranging aquatic species were included in taxa‐specific priority areas, but rare freshwater species were not included. Thus, the development of conservation priorities based on a single freshwater aquatic group would not protect all species in the other aquatic groups.     

Anthropogenic areas as incidental substitutes for original habitat

One speaks of ecological substitutes when an introduced species performs, to some extent, the ecosystem function of an extirpated native species. We suggest that a similar case exists for habitats. Species evolve within ecosystems, but habitats can be destroyed or modified by natural and human‐made causes. Sometimes habitat alteration forces animals to move to or remain in a suboptimal habitat type. In that case, the habitat is considered a refuge, and the species is called a refugee. Typically refugee species have lower population growth rates than in their original habitats. Human action may lead to the unintended generation of artificial or semiartificial habitat types that functionally resemble the essential features of the original habitat and thus allow a population growth rate of the same magnitude or higher than in the original habitat. We call such areas substitution habitats and define them as human‐made habitats within the focal species range that by chance are partial substitutes for the species’ original habitat. We call species occupying a substitution habitat adopted species. These are 2 new terms in conservation biology. Examples of substitution habitats are dams for European otters, wheat and rice fields for many steppeland and aquatic birds, and urban areas for storks, falcons, and swifts. Although substitution habitats can bring about increased resilience against the agents of global change, the conservation of original habitat types remains a conservation priority.     

Effectiveness of vegetation‐based biodiversity offset metrics as surrogates for ants

Biodiversity offset schemes are globally popular policy tools for balancing the competing demands of conservation and development. Trading currencies for losses and gains in biodiversity value at development and credit sites are usually based on several vegetation attributes combined to yield a simple score (multimetric), but the score is rarely validated prior to implementation. Inaccurate biodiversity trading currencies are likely to accelerate global biodiversity loss through unrepresentative trades of losses and gains. We tested a model vegetation multimetric (i.e., vegetation structural and compositional attributes) typical of offset trading currencies to determine whether it represented measurable components of compositional and functional biodiversity. Study sites were located in remnant patches of a critically endangered ecological community in western Sydney, Australia, an area representative of global conflicts between conservation and expanding urban development. We sampled ant fauna composition with pitfall traps and enumerated removal by ants of native plant seeds from artificial seed containers (seed depots). Ants are an excellent model taxon because they are strongly associated with habitat complexity, respond rapidly to environmental change, and are functionally important at many trophic levels. The vegetation multimetric did not predict differences in ant community composition or seed removal, despite underlying assumptions that biodiversity trading currencies used in offset schemes represent all components of a site's biodiversity value. This suggests that vegetation multimetrics are inadequate surrogates for total biodiversity value. These findings highlight the urgent need to refine existing offsetting multimetrics to ensure they meet underlying assumptions of surrogacy. Despite the best intentions, offset schemes will never achieve their goal of no net loss of biodiversity values if trades are based on metrics unrepresentative of total biodiversity.     

Surrogates for Macrofungi and Mosses in Reservation Planning

Abstract: Our knowledge of cryptogam taxonomy and species distributions is currently too poor to directly plan for their conservation. We used inventory data from four distinct vegetation types, near Hobart Tasmania, to address the proposition that vegetation type, vascular plant taxon composition, and environmental variables can act as surrogates for mosses and macrofungi in reservation planning. The four vegetation types proved distinct in their taxon composition for all macrofungi, mosses, and vascular plants. We tested the strength of the relationships between the composition of cryptogam taxonomic groups and vascular plant composition and between the environmental variables and canopy cover. Taxon composition of woody vascular plants and vascular plants was the best predictor of the taxon composition of mosses and macrofungi. Combinations of environmental variables and canopy cover were also strong predictors of the taxon composition of mosses and macrofungi. We used an optimization routine for vascular plant taxa and woody plant species and determined the representation of cryptogam taxa in these selections. We identified sites with approximately 10% and 30% of the greatest proportions of vascular plants and woody vascular plants and calculated representation of mosses and macrofungi at these sites. We compared the results of these site selections with random site selections and random selections stratified by vegetation type. Random selection of sites by vegetation type generally captured more cryptogams than site selection by vascular plants at the 10% level. Vascular plant and woody plant taxon composition, vegetation type, and environmental and structural characteristics, all showed promise as surrogates for capturing common cryptogams in reserve systems.     

An Empirical Assessment of the Focal Species Hypothesis

Biodiversity surrogates and indicators are commonly used in conservation management. The focal species approach (FSA) is one method for identifying biodiversity surrogates, and it is underpinned by the hypothesis that management aimed at a particular focal species will confer protection on co‐occurring species. This concept has been the subject of much debate, in part because the validity of the FSA has not been subject to detailed empirical assessment of the extent to which a given focal species actually co‐occurs with other species in an assemblage. To address this knowledge gap, we used large‐scale, long‐term data sets of temperate woodland birds to select focal species associated with threatening processes such as habitat isolation and loss of key vegetation attributes. We quantified co‐occurrence patterns among focal species, species in the wider bird assemblage, and species of conservation concern. Some, but not all, focal species were associated with high levels of species richness. One of our selected focal species was negatively associated with the occurrence of other species (i.e., it was an antisurrogate)—a previously undescribed property of nominated focal species. Furthermore, combinations of focal species were not associated with substantially elevated levels of bird species richness, relative to levels associated with individual species. Our results suggest that although there is some merit to the underpinning concept of the FSA, there is also a need to ensure that actions are sufficiently flexible because management tightly focused on a given focal species may not benefit some other species, including species of conservation concern, such of which might not occur in species‐rich assemblages. Una Evaluación Empírica de la Hipótesis de Especie Focal     

Environmental diversity as a surrogate for species representation

Because many species have not been described and most species ranges have not been mapped, conservation planners often use surrogates for conservation planning, but evidence for surrogate effectiveness is weak. Surrogates are well‐mapped features such as soil types, landforms, occurrences of an easily observed taxon (discrete surrogates), and well‐mapped environmental conditions (continuous surrogate). In the context of reserve selection, the idea is that a set of sites selected to span diversity in the surrogate will efficiently represent most species. Environmental diversity (ED) is a rarely used surrogate that selects sites to efficiently span multivariate ordination space. Because it selects across continuous environmental space, ED should perform better than discrete surrogates (which necessarily ignore within‐bin and between‐bin heterogeneity). Despite this theoretical advantage, ED appears to have performed poorly in previous tests of its ability to identify 50 × 50 km cells that represented vertebrates in Western Europe. Using an improved implementation of ED, we retested ED on Western European birds, mammals, reptiles, amphibians, and combined terrestrial vertebrates. We also tested ED on data sets for plants of Zimbabwe, birds of Spain, and birds of Arizona (United States). Sites selected using ED represented European mammals no better than randomly selected cells, but they represented species in the other 7 data sets with 20% to 84% effectiveness. This far exceeds the performance in previous tests of ED, and exceeds the performance of most discrete surrogates. We believe ED performed poorly in previous tests because those tests considered only a few candidate explanatory variables and used suboptimal forms of ED's selection algorithm. We suggest future work on ED focus on analyses at finer grain sizes more relevant to conservation decisions, explore the effect of selecting the explanatory variables most associated with species turnover, and investigate whether nonclimate abiotic variables can provide useful surrogates in an ED framework.     

Text analysis tools for identification of emerging topics and research gaps in conservation science

Keeping track of conceptual and methodological developments is a critical skill for research scientists, but this task is increasingly difficult due to the high rate of academic publication. As a crisis discipline, conservation science is particularly in need of tools that facilitate rapid yet insightful synthesis. We show how a common text‐mining method (latent Dirichlet allocation, or topic modeling) and statistical tests familiar to ecologists (cluster analysis, regression, and network analysis) can be used to investigate trends and identify potential research gaps in the scientific literature. We tested these methods on the literature on ecological surrogates and indicators. Analysis of topic popularity within this corpus showed a strong emphasis on monitoring and management of fragmented ecosystems, while analysis of research gaps suggested a greater role for genetic surrogates and indicators. Our results show that automated text analysis methods need to be used with care, but can provide information that is complementary to that given by systematic reviews and meta‐analyses, increasing scientists’ capacity for research synthesis.