The Silent Mass Extinction of Insect Herbivores in Biodiversity Hotspots

Abstract: Habitat loss is silently leading numerous insects to extinction. Conservation efforts, however, have not been designed specifically to protect these organisms, despite their ecological and evolutionary significance. On the basis of species–host area equations, parameterized with data from the literature and interviews with botanical experts, I estimated the number of specialized plant‐feeding insects (i.e., monophages) that live in 34 biodiversity hotspots and the number committed to extinction because of habitat loss. I estimated that 795,971–1,602,423 monophagous insect species live in biodiversity hotspots on 150,371 endemic plant species, which is 5.3–10.6 monophages per plant species. I calculated that 213,830–547,500 monophagous species are committed to extinction in biodiversity hotspots because of reduction of the geographic range size of their endemic hosts. I provided rankings of biodiversity hotspots on the basis of estimated richness of monophagous insects and on estimated number of extinctions of monophagous species. Extinction rates were predicted to be higher in biodiversity hotspots located along strong environmental gradients and on archipelagos, where high spatial turnover of monophagous species along the geographic distribution of their endemic plants is likely. The results strongly support the overall strategy of selecting priority conservation areas worldwide primarily on the basis of richness of endemic plants. To face the global decline of insect herbivores, one must expand the coverage of the network of protected areas and improve the richness of native plants on private lands.     

Conservation of Insect Diversity: a Habitat Approach

Abstract: Neither time nor resources exist to design conservation plans for every species, particularly for little-studied, noncharismatic, but ecologically important taxa that make up most of biodiversity. To explore the feasibility of basing conservation action on community-level biogeography, we sampled a montane insect community. We addressed three issues: (1) the appropriate scale for sampling insect communities; (2) the association of habitat specialization—perhaps a measure of extinction vulnerability—with other ecological or physical traits; and (3) the correlation of diversity across major insect groups. Using malaise traps in Gunnison County, Colorado, we captured 8847 Diptera (identified to family and morphospecies), 1822 Hymenoptera (identified to morphospecies), and 2107 other insects (identified to order). We sampled in three habitat types—meadow, aspen, and conifer—defined on the basis of the dominant vegetation at the scale of hundreds of meters. Dipteran communities were clearly differentiated by habitat type rather than geographic proximity. This result also holds true for hymenopteran communities. Body size and feeding habits were associated with habitat specialization at the family level. In particular, habitat generalists at the family level—taxa perhaps more likely to survive anthropogenic habitat alteration—tended to be trophic generalists. Dipteran species richness was marginally correlated with hymenopteran species richness and was significantly correlated with the total number of insect orders sampled by site. Because these correlations result from differences in richness among habitat types, insect taxa may be reasonable surrogates for one another when sampling is done across habitat types. In sum, community-wide studies appear to offer a practical way to gather information about the diversity and distribution of little-known taxa.     

Risk of Local Extinction of Odonata Freshwater Habitat Generalists and Specialists

Understanding the risk of a local extinction in a single population relative to the habitat requirements of a species is important in both theoretical and applied ecology. Local extinction risk depends on several factors, such as habitat requirements, range size of species, and habitat quality. We studied the local extinctions among 31 dragonfly and damselfly species from 1930 to 1975 and from 1995 to 2003 in Central Finland. We tested whether habitat specialists had a higher local extinction rate than generalist species. Approximately 30% of the local dragonfly and damselfly populations were extirpated during the 2 study periods. The size of the geographical range of the species was negatively related to extinction rate of the local populations. In contrast to our prediction, the specialist species had lower local extinction rates than the generalist species, probably because generalist species occurred in both low‐ and high‐quality habitat. Our results are consistent with source–sink theory. Riesgo de Extinción Local de Odonatos de Agua Dulce Generalistas y Especialistas de Hábitat     

The contribution of policy,law, management,research, and advocacy failings to the recent extinctions of three Australian vertebrate species

Extinctions typically have ecological drivers, such as habitat loss. However, extinction events are also influenced by policy and management settings that may be antithetical to biodiversity conservation, inadequate to prevent extinction, insufficiently resourced, or poorly implemented. Three endemic Australian vertebrate species—the Christmas Island pipistrelle (Pipistrellus murrayi), Bramble Cay melomys (Melomys rubicola), and Christmas Island forest skink (Emoia nativitatis)—became extinct from 2009 to 2014. All 3 extinctions were predictable and probably preventable. We sought to identify the policy, management, research, and other shortcomings that contributed to their extinctions or failed to prevent them. These included a lack within national environmental legislation and policy of explicit commitment to the prevention of avoidable extinctions, lack of explicit accountability, inadequate resources for conservation (particularly for species not considered charismatic or not of high taxonomic distinctiveness), inadequate biosecurity, a slow and inadequate process for listing species as threatened, recovery planning that failed to consider the need for emergency response, inability of researchers to identify major threatening factors, lack of public engagement and involvement in conservation decisions, and limited advocacy. From these 3 cases, we recommend: environmental policy explicitly seeks to prevent extinction of any species and provides a clear chain of accountability and an explicit requirement for public inquiry following any extinction; implementation of a timely and comprehensive process for listing species as threatened and for recovery planning; reservation alone not be assumed sufficient to maintain species; enhancement of biosecurity measures; allocation of sufficient resources to undertake actions necessary to prevent extinction; monitoring be considered a pivotal component of the conservation response; research provides timely identification of factors responsible for decline and of the risk of extinction; effective dissemination of research results; advocacy by an informed public for the recovery of threatened species; and public involvement in governance of the recovery process. These recommendations should be applicable broadly to reduce the likelihood and incidence of extinctions.     

Identifying and Managing Threatened Invertebrates through Assessment of Coextinction Risk

Abstract: Invertebrates with specific host species may have a high probability of extinction when their hosts have a high probability of extinction. Some of these invertebrates are more likely to go extinct than their hosts, and under some circumstances, specific actions to conserve the host may be detrimental to the invertebrate. A critical constraint to identifying such invertebrates is uncertainty about their level of host specificity. We used two host‐breadth models that explicitly incorporated uncertainty in the host specificity of an invertebrate species. We devised a decision protocol to identify actions that may increase the probability of persistence of a given dependent species. The protocol included estimates from the host‐breadth models and decision nodes to identify cothreatened species. We applied the models and protocol to data on 1055 insects (186 species) associated with 2 threatened (as designated by the Australian Government) plant species and 19 plant species that are not threatened to determine whether any insect herbivores have the potential to become extinct if the plant becomes extinct. According to the host‐breadth models, 18 species of insect had high host specificity to the threatened plant species. From these 18 insects, the decision protocol highlighted 6 species that had a high probability of extinction if their hosts were to become extinct (3% of all insects examined). The models and decision protocol have added objectivity and rigor to the process of deciding which dependent invertebrates require conservation action, particularly when dealing with largely unknown and speciose faunas.     

Vascular plant extinction in the continental United States and Canada

Extinction rates are expected to increase during the Anthropocene. Current extinction rates of plants and many animals remain unknown. We quantified extinctions among the vascular flora of the continental United States and Canada since European settlement. We compiled data on apparently extinct species by querying plant conservation databases, searching the literature, and vetting the resulting list with botanical experts. Because taxonomic opinion varies widely, we developed an index of taxonomic uncertainty (ITU). The ITU ranges from A to F, with A indicating unanimous taxonomic recognition and F indicating taxonomic recognition by only a single author. The ITU allowed us to rigorously evaluate extinction rates. Our data suggest that 51 species and 14 infraspecific taxa, representing 33 families and 49 genera of vascular plants, have become extinct in our study area since European settlement. Seven of these taxa exist in cultivation but are extinct in the wild. Most extinctions occurred in the west, but this outcome may reflect the timing of botanical exploration relative to settlement. Sixty-four percent of extinct plants were single-site endemics, and many occurred outside recognized biodiversity hotspots. Given the paucity of plant surveys in many areas, particularly prior to European settlement, the actual extinction rate of vascular plants is undoubtedly much higher than indicated here.     

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 .     

Conservation Corridors and Contagious Disease: A Cautionary Note

Recent conservation proposals frequently include the establishment of corridors to connect isolated patches of wildlife habitat. Much attention has been focused on the potential benefits of corridors with little note given to potentially adverse consequences. A simulation model is developed here to study the effect of corridors on the survival of a metapopulation in the presence of a fatal disease that is spread by direct contact between susceptible and infected individuals. For the disease modeled here, a landscape of patches connected by corridors generally suffers fewer metapopulation extinctions than a landscape of isolated patches. However, under a narrow range of conditions, results suggest that corridors may dramatically increase the probability of metapopulation extinction. This occurs when disease-induced mortality is low enough to allow infected individuals to spread the disease, but high enough to reduce population levels to the point that random demographic and environmental events cause frequent metapopulation extinctions. This has important implications for the design and management of conservation reserve networks. Although discussion focuses primarily on conservation corridors, the model results apply to any management techniques that increase the movement of individuals among populations.     

Major Conservation Policy Issues for Biodiversity in Oceania

Abstract:  Oceania is a diverse region encompassing Australia, Melanesia, Micronesia, New Zealand, and Polynesia, and it contains six of the world's 39 hotspots of diversity. It has a poor record for extinctions, particularly for birds on islands and mammals. Major causes include habitat loss and degradation, invasive species, and overexploitation. We identified six major threatening processes (habitat loss and degradation, invasive species, climate change, overexploitation, pollution, and disease) based on a comprehensive review of the literature and for each developed a set of conservation policies. Many policies reflect the urgent need to deal with the effects of burgeoning human populations (expected to increase significantly in the region) on biodiversity. There is considerable difference in resources for conservation, including people and available scientific information, which are heavily biased toward more developed countries in Oceania. Most scientific publications analyzed for four threats (habitat loss, invasive species, overexploitation, and pollution) are from developed countries: 88.6% of Web of Science publications were from Australia (53.7%), New Zealand (24.3%), and Hawaiian Islands (10.5%). Many island states have limited resources or expertise. Even countries that do (e.g., Australia, New Zealand) have ongoing and emerging significant challenges, particularly with the interactive effects of climate change. Oceania will require the implementation of effective policies for conservation if the region's poor record on extinctions is not to continue .     

Extinction, Colonization, and Metapopulations: Environmental Tracking by Rare Species

Extinction and metapopulation theories emphasize that stochastic fluctuations in local populations cause extinction and that local extinctions generate empty habitat patches that are then available for recolonization. Metapopulation persistence depends on the balance of extinction and colonization in a static environment. For many rare and declining species, I argue (1) that extinction is usually the deterministic consequence of the local environment becoming unsuitable (through habitat loss or modification, introduction of a predator, etc.); (2) that the local environment usually remains unsuitable following local extinction, so extinctions only rarely generate empty patches of suitable habitat; and (3) that colonization usually follows improvement of the local environment for a particular species (or long-distance transfer by humans). Thus, persistence depends predominantly on whether organisms are able to track the shifting spatial mosaic of suitable environmental conditions or on maintainance of good conditions locally.     

Future habitat loss and extinctions driven by land‐use change in biodiversity hotspots under four scenarios of climate‐change mitigation

Numerous species have been pushed into extinction as an increasing portion of Earth's land surface has been appropriated for human enterprise. In the future, global biodiversity will be affected by both climate change and land‐use change, the latter of which is currently the primary driver of species extinctions. How societies address climate change will critically affect biodiversity because climate‐change mitigation policies will reduce direct climate‐change impacts; however, these policies will influence land‐use decisions, which could have negative impacts on habitat for a substantial number of species. We assessed the potential impact future climate policy could have on the loss of habitable area in biodiversity hotspots due to associated land‐use changes. We estimated past extinctions from historical land‐use changes (1500–2005) based on the global gridded land‐use data used for the Intergovernmental Panel on Climate Change Fifth Assessment Report and habitat extent and species data for each hotspot. We then estimated potential extinctions due to future land‐use changes under alternative climate‐change scenarios (2005–2100). Future land‐use changes are projected to reduce natural vegetative cover by 26‐58% in the hotspots. As a consequence, the number of additional species extinctions, relative to those already incurred between 1500 and 2005, due to land‐use change by 2100 across all hotspots ranged from about 220 to 21000 (0.2% to 16%), depending on the climate‐change mitigation scenario and biological factors such as the slope of the species–area relationship and the contribution of wood harvest to extinctions. These estimates of potential future extinctions were driven by land‐use change only and likely would have been higher if the direct effects of climate change had been considered. Future extinctions could potentially be reduced by incorporating habitat preservation into scenario development to reduce projected future land‐use changes in hotspots or by lessening the impact of future land‐use activities on biodiversity within hotspots.     

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.     

Objectives for Multiple-Species Conservation Planning

Abstract:  The first step in conservation planning is to identify objectives. Most stated objectives for conservation, such as to maximize biodiversity outcomes, are too vague to be useful within a decision-making framework. One way to clarify the issue is to define objectives in terms of the risk of extinction for multiple species. Although the assessment of extinction risk for single species is common, few researchers have formulated an objective function that combines the extinction risks of multiple species. We sought to translate the broad goal of maximizing the viability of species into explicit objectives for use in a decision-theoretic approach to conservation planning. We formulated several objective functions based on extinction risk across many species and illustrated the differences between these objectives with simple examples. Each objective function was the mathematical representation of an approach to conservation and emphasized different levels of threat. Our objectives included minimizing the joint probability of one or more extinctions, minimizing the expected number of extinctions, and minimizing the increase in risk of extinction from the best-case scenario. With objective functions based on joint probabilities of extinction across species, any correlations in extinction probabilities had to be known or the resultant decisions were potentially misleading. Additive objectives, such as the expected number of extinctions, did not produce the same anomalies. We demonstrated that the choice of objective function is central to the decision-making process because alternative objective functions can lead to a different ranking of management options. Therefore, decision makers need to think carefully in selecting and defining their conservation goals.     

The difference conservation makes to extinction risk of the world's ungulates

Previous studies show that conservation actions have prevented extinctions, recovered populations, and reduced declining trends in global biodiversity. However, all studies to date have substantially underestimated the difference conservation action makes because they failed to account fully for what would have happened in the absence thereof. We undertook a scenario‐based thought experiment to better quantify the effect conservation actions have had on the extinction risk of the world's 235 recognized ungulate species. We did so by comparing species’ observed conservation status in 2008 with their estimated status under counterfactual scenarios in which conservation efforts ceased in 1996. We estimated that without conservation at least 148 species would have deteriorated by one International Union for Conservation of Nature (IUCN) Red List category, including 6 species that now would be listed as extinct or extinct in the wild. The overall decline in the conservation status of ungulates would have been nearly 8 times worse than observed. This trend would have been greater still if not for conservation on private lands. While some species have benefited from highly targeted interventions, such as reintroduction, most benefited collaterally from conservation such as habitat protection. We found that the difference conservation action makes to the conservation status of the world's ungulate species is likely to be higher than previously estimated. Increased, and sustained, investment could help achieve further improvements.     

Understanding the drivers of expert opinion when classifying species as extinct

The criteria as laid out by the International Union for the Conservation of Nature (IUCN) Red List are the gold standard by which the extinction risk of a species is assessed and where appropriate biological extinctions are declared. However, unlike all other categories, the category of extinct lacks a quantitative framework for assigning this category. Given its subjective nature, we surveyed expert assessors working on a diversity of taxa to explore the attributes they used to declare a species extinct. Using a choice experiment approach, we surveyed 674 experts from the IUCN Species Survival Commission specialist groups and taskforces. Data availability, time from the last sighting, detectability, habitat availability, and population decline were all important attributes favored by assessors when inferring extinction. Respondents with red-listing experience assigned more importance to the attributes data availability, time from the last sighting, and detectability when considering a species extinction, whereas those respondents working with well-known taxa gave more importance to the time from the last sighting. Respondents with no red-listing experience and those working with more well-known taxa (i.e., mammals and birds) were overall less likely to consider species extinct. Our findings on the importance assessors place on attributes used to declare a species extinct provide a basis for informing the development of specific criteria for more accurately assessing species extinctions.     

A Matrix‐Calibrated Species‐Area Model for Predicting Biodiversity Losses Due to Land‐Use Change

Abstract: Application of island biogeography theory to prediction of species extinctions resulting from habitat loss is based on the assumption that the transformed landscape matrix is completely inhospitable to the taxa considered, despite evidence demonstrating the nontrivial influence of matrix on populations within habitat remnants. The island biogeography paradigm therefore needs refining to account for specific responses of taxa to the area of habitat “islands” and to the quality of the surrounding matrix. We incorporated matrix effects into island theory by partitioning the slope (z value) of species–area relationships into two components: γ, a constant, and σ, a measure of taxon‐specific responses to each component of a heterogeneous matrix. We used our matrix‐calibrated model to predict extinction and endangerment of bird species resulting from land‐use change in 20 biodiversity hotspots and compared these predictions with observed numbers of extinct and threatened bird species. We repeated this analysis with the conventional species–area model and the countryside species–area model, considering alternative z values of 0.35 (island) or 0.22 (continental). We evaluated the relative strength of support for each of the five candidate models with Akaike's information criterion (AIC). The matrix‐calibrated model had the highest AIC weight (w_i = 89.21%), which means the weight of evidence in support of this model was the optimal model given the set of candidate models and the data. In addition to being a valuable heuristic tool for assessing extinction risk, our matrix‐calibrated model also allows quantitative assessment of biodiversity benefits (and trade‐offs) of land‐management options in human‐dominated landscapes. Given that processes of secondary regeneration have become more widespread across tropical regions and are predicted to increase, our matrix‐calibrated model will be increasingly appropriate for practical conservation in tropical landscapes.     

The Role of Behavior in Recent Avian Extinctions and Endangerments

Abstract: Understanding patterns of differential extinction and predicting the relative risks of extinction among extant species are among the most important problems in conservation biology. Although recent studies reveal that behavior can be a critical component in many species' extinctions or endangerments, current approaches to the problem of predicting extinction patterns largely ignore behavior. I reviewed how behavior can affect population persistence and then used recent avian extinctions and endangerments to illustrate behaviors relevant to extinction risk. Behaviors that affect population persistence can be grouped as aggregation, interspecific responses, dispersal, habitat selection, intraspecific behavior, and maladaptive behavior. Behavior that can affect extinction risk is not limited to birds; for example, in many taxonomic groups (vertebrate and invertebrate) there is evidence of socially facilitated reproduction in colonial species, Allee effects on reproductive success and survival, behavioral regulation of population size, and conspecific attraction to breeding sites. Incorporating specific behaviors into models predicting extinction probabilities and patterns should improve their predictions.     

Examining the relationship between local extinction risk and position in range

Over half of globally threatened animal species have experienced rapid geographic range loss. Identifying the parts of species’ distributions most vulnerable to local extinction would benefit conservation planning. However, previous studies give little consensus on whether ranges decline to the core or edge. We built on previous work by using empirical data to examine the position of recent local extinctions within species’ geographic ranges, address range position as a continuum, and explore the influence of environmental factors. We aggregated point‐locality data for 125 Galliform species from across the Palearctic and Indo‐Malaya into equal‐area half‐degree grid cells and used a multispecies dynamic Bayesian occupancy model to estimate rates of local extinctions. Our model provides a novel approach to identify loss of populations from within species ranges. We investigated the relationship between extinction rates and distance from range edge by examining whether patterns were consistent across biogeographic realm and different categories of land use. In the Palearctic, local extinctions occurred closer to the range edge than range core in both unconverted and human‐dominated landscapes. In Indo‐Malaya, no pattern was found for unconverted landscapes, but in human‐dominated landscapes extinctions tended to occur closer to the core than the edge. Our results suggest that local and regional factors override general spatial patterns of recent local extinction within species’ ranges and highlight the difficulty of predicting the parts of a species’ distribution most vulnerable to threat.     

Habitat Loss and Changes in the Species-Area Relationship

Abstract: The species-area relationship (SAR) has been used successfully to predict extinction from extent of habitat reduction. These extinction estimates assume that species have uniformly distributed range requirements and a minimum abundance level required for persistence; how many species are lost depends solely on how much habitat is removed, not on where it is removed. We consider another limiting case in which range requirements, rather than abundances, determine extinctions. We used a new method for constructing SARs based on assumptions about geographic ranges of species. Our results show that habitat destruction can change the SAR and consequently the number of species predicted to be lost due to habitat destruction. Our method generates SARs that vary in shape according to the specific distributions of geographic range and occupancy but that have the common feature of being described by a power law with an exponent of <1. When the geographic range of species was included in the SAR, the way habitat was lost became important. Although the SAR before habitat destruction is often used to predict species loss after habitat destruction, assumptions must be clearly stated. To predict the damage caused by habitat loss with our model, it is necessary to know the fraction of aggregated species, the distribution of geographic ranges, the form of habitat destruction, and the sampling protocol. The remaining theoretical challenge is to develop a full theory that links abundance and range.