Threatened Vertebrate Species Research Articles

Identifying protected areas in biodiversity hotspots at risk from climate and human-induced compound events for conserving threatened species

Anthropogenic pressure in areas of biodiversity importance erodes the integrity of the ecosystems they harbour, making features of biodiversity less buffered against extreme climatic events. We define the combination of these disturbances as compound events. We assessed compound event risk in protected areas (PAs) applying a spatial framework guided by criteria and quantitative thresholds associated with exposure to cyclones, drought, and intense human pressure. This assessment was used in a relational matrix to classify PAs with different risk of compound event occurrence. We identified PAs of higher conservation concern by quantifying the extent of human pressure in their surrounding landscape while harbouring large numbers of threatened vertebrate species. Of the 39,694 PAs assessed, very high risk of compound events was determined for 6965 PAs (17.5 %) related to cyclones and human pressure (mainly island hotspots), 6367 PAs (16 %) related to droughts and human pressure (island and continental hotspots), and 2031 PAs (5 %) to cyclones, drought and human pressure (mainly in island hotspots). From the subset of 2031 PAs assessed at very high risk, we identified 239 PAs of higher conservation concern distributed predominantly in the Caribbean Islands, Japan, North America Coastal Plain, Philippines, and Southwest Australia. Our work highlights PAs in the biodiversity hotspots where high risk of compound event occurrence poses a greater threat to species. We encourage researchers to adapt and apply this framework across other globally significant sites for conserving biodiversity to identify high risk-prone areas, and prevent further biodiversity decline.

The Ethiopian wolf can act as a flagship and umbrella species to protect the Afroalpine ecosystem and foster sustainable development

SummaryHabitat alteration and climate change are important threats to terrestrial biodiversity in the tropics. Endorsing flagship or umbrella species can help conserve sympatric biodiversity, restore degraded ecosystems and achieve United Nations Sustainable Development Goals (UN SDGs). The Ethiopian wolf (Canis simensis) is a rare and endemic Ethiopian canid. It is Africa’s most endangered canid species and is restricted to several isolated patches of Afroalpine habitats. While its behavioural ecology and conservation biology have been well studied, studies of the Ethiopian wolf’s significance for the conservation of its habitat and sympatric species are lacking. Here we use geographical range overlap and geospatial modelling to evaluate the importance of the Ethiopian wolf as a flagship and/or umbrella species. We assess whether conservation interventions targeting the Ethiopian wolf could help to restore and protect Afroalpine habitat and conserve sympatric species whilst simultaneously providing a wide range of socioeconomic and environmental benefits. We found that Ethiopian wolves share their range with 73 endemic and/or threatened vertebrate species, 68 of which are Afroalpine ecosystem species, and at least 121 endemic and/or threatened plant species. Ethiopian wolves are taxonomically distinctive and charismatic species classified as Endangered on the International Union for Conservation of Nature (IUCN) Red List. Thus, they meet both the flagship and umbrella species criteria to restore Afroalpine habitats and conserve threatened sympatric species. A conservation strategy protecting and restoring Afroalpine habitat has the potential to contribute to achieving at least five of the 17 UN SDGs. The protection of flagship and umbrella species should be integrated into broader regional biodiversity and habitat conservation.

Difficult times for amphibians: Effects of land-use change at the local and landscape scales in the Iberá Wetlands

Land-use change and management practices have led to habitat loss, one of the greatest factors in biodiversity decline. Particularly, amphibians comprise the highest number of threatened vertebrate species at the global scale. In this work, amphibian communities were analysed in three differing landscapes: a protected wetland, a livestock-impaired rangeland and a pine afforestation. In each landscape, amphibian species were sampled. Land-cover type and composition, as well as primary vegetation types were characterised at the landscape and local scale, respectively. The relationship between these environmental variables and the amphibian communities was analysed. Twenty-one amphibian species were identified; the protected and afforested landscapes were the richest, whereas the rangeland showed the lowest species richness and diversity estimates. At the local scale, vegetation, water coverage and land-use category explained the higher presence of amphibian species and their abundance. These results show how different land-uses, especially livestock farming, modify the composition of amphibian communities. This work constitutes a foundation for the development of sustainable management practices for conserving amphibians in landscape-level altered habitats.

The value of private properties for the conservation of biodiversity in the Brazilian Cerrado

Areas set aside for conservation within private lands may be key to enhancing biodiversity-friendly landscapes. This conservation strategy should be especially effective in highly threatened regions that are poorly protected by public lands, such as the Brazilian Cerrado. Brazil's Native Vegetation Protection Law has included set-aside areas within private properties, but their relevance to conservation has not been evaluated. We assess whether private lands are contributing to biodiversity in the Cerrado, a global biodiversity conservation priority and major region for food production, where land use conflicts are often at odds with conservation objectives. We determined that private protected areas accommodate up to 14.5% of threatened vertebrate species ranges, which increases to 25% when considering the distribution of remaining native habitat. Moreover, the spatial spread of private protected areas benefits a large number of species. Ecological restoration of private protected lands would improve the benefits of this protection system, especially in the Southeastern Cerrado, where a large economic hub meets a threat hotspot.

The socio-environmental impacts of tropical crop expansion on a global scale: A case study in cashew

Tropical crops like cashew (Anacardium occidentale) receive limited socio-ecological research attention despite expansion in regions of high biodiversity. We investigated how cashew plantations affected biodiversity and livelihoods globally by conducting a systematic literature review and modelling the potential overlaps between cashew land suitability and threatened vertebrates. Nine studies on the effects of cashew on biodiversity from India, Guinea- Bissau and Nigeria showed four to 84 % species richness reductions for fungi, plants, butterflies, birds, and terrestrial mammals across cashew and ‘reference habitats’ such as forests, woodlands, coffee plantations and sand dunes. Three studies reported that cashew harboured a subset of generalist butterfly and bird species when compared to woodlands, coffee, and forests with no evidence of forest-dependent species in cashew. Preliminary studies (n = 3) showed that the potential for facilitative coexistence among humans and wildlife in cashew land use, although further research is necessary to evaluate this. Livelihood effects of cashew farms from 29 studies across the tropics showed that cashew farms are dominated by smallholders who rely increasingly on cultivar cashew. Latin American and Asian countries showed spatial overlaps between land suitable for cashew and areas of threatened vertebrate species richness. FAO data showed that land area under cashew is expanding in African and Asian countries, but not in Latin American countries. Our study highlights the importance of studying tropical crops that lack large, apparent land-use footprints but affect biodiverse regions through steady, small-scale expansion.

The behavioural ecology toolkit for fish management and conservation

AbstractFishes include some of the most threatened vertebrate species globally. As such, efforts to effectively conserve and manage fish populations and their habitats are vast. Here, we present conceptual tools from behavioural ecology to establish a framework for studies of fish conservation and management, connecting questions relevant to managers and practitioners with behavioural theories and methodologies. We apply predictions developed from a theory to diet choice, patch use, habitat selection, movement, and social behaviours. We present questions and issues in fisheries management and conservation for which theory, hypotheses, and methodologies would be both novel and complementary to current assessment strategies or conservation efficacy studies. In each case, theory approaches the ecological trade‐offs associated with a given behavioural trait through the lens of adaptations and fitness implications—the foundational principles of behavioural ecology. We show key methodologies used to effectively apply behavioural theory to specific hypotheses relevant to a given management question. We then compile the conceptual and methodological approaches to assemble a toolkit through which fisheries managers may assess, for example, habitat selection behaviours via novel study designs and/or new ways of interpreting commonly collected data (e.g., distribution and abundance relative to habitat type). Finally, we propose training of aquatic and marine natural resource specialists and conservation agency fish biologists be complemented with behavioural ecology theories and methodologies.

The value of flagship and umbrella species for restoration and sustainable development: Bale monkeys and bamboo forest in Ethiopia

Forest loss and degradation are the most significant threats to terrestrial biodiversity in the tropics. Promoting flagship or umbrella species is a strategy that can be used to conserve intact forests and restore degraded ecosystems, conserve biodiversity, and achieve sustainable development goals. The Bale monkey (Chlorocebus djamdjamensis) is an arboreal, forest-dwelling, threatened primate restricted to a small range in the southern Ethiopian Highlands, which relies mostly on a single species of bamboo (Arundinaria alpina) and prefers bamboo forest habitat. Most of the Bale monkey’s range lies outside protected areas and most of its historical bamboo forest habitat is degraded or destroyed. The conservation of Bale monkeys and bamboo is highly inter-dependent; however, the value of using the Bale monkey as a flagship or umbrella species for forest restoration has not been evaluated. Here we use geographic range overlap and geospatial modeling to evaluate Bale monkeys as a flagship and/or umbrella species. We also assess if conservation intervention on behalf of Bale monkeys can help restore bamboo forest, while simultaneously providing a wide range of socioeconomic and environmental benefits. We found that Bale monkeys share their range with 52 endemic and/or threatened vertebrate species and at least 9 endemic and/or threatened plant species. Our results show that Bale monkeys meet both the flagship and umbrella species criteria to restore bamboo forest and conserve threatened co-occurring species. Since bamboo is fast-growing and can be harvested every year, we suggest that a science-based sustainable harvest and management regime for bamboo would help to improve the livelihood of both the local community and Ethiopians in general without significantly affecting the long-term survival of Bale monkeys and regional biodiversity. Further, a conservation management strategy protecting and restoring bamboo forest has the potential to achieve at least six of the 17 United Nations Sustainable Development Goals.

A global ecological signal of extinction risk in terrestrial vertebrates.

To determine the distribution and causes of extinction threat across functional groups of terrestrial vertebrates, we assembled an ecological trait data set for 18,016 species of terrestrial vertebrates and utilized phylogenetic comparative methods to test which categories of habitat association, mode of locomotion, and feeding mode best predicted extinction risk. We also examined the individual categories of the International Union for Conservation of Nature Red List extinction drivers (e.g., agriculture and logging) threatening each species and determined the greatest threats for each of the four terrestrial vertebrate groups. We then quantified the sum of extinction drivers threatening each species to provide a multistressor perspective on threat. Cave dwelling amphibians (p < 0.01), arboreal quadrupedal mammals (all of which are primates) (p < 0.01), aerial and scavenging birds (p < 0.01), and pedal (i.e., walking) squamates (p < 0.01) were all disproportionately threatened with extinction in comparison with the other assessed ecological traits. Across all threatened vertebrate species in the study, the most common risk factors were agriculture, threatening 4491 species, followed by logging, threatening 3187 species, and then invasive species and disease, threatening 2053 species. Species at higher risk of extinction were simultaneously at risk from a greater number of threat types. If left unabated, the disproportionate loss of species with certain functional traits and increasing anthropogenic pressures are likely to disrupt ecosystem functions globally. A shift in focus from species‐ to trait‐centric conservation practices will allow for protection of at‐risk functional diversity from regional to global scales.

Status of biodiversity and limno-chemistry of Deepor Beel, a Ramsar site of international importance: Conservation needs and the way forward

Abstract Deepor Beel is a prominent floodplain wetland and a Ramsar site of northeastern (NE) India (Ramsar site no. 1207) covering a 40.14 km2 area and connected to River Brahmaputra. It provides numerous economic (e.g., livelihood and nutritional security) and ecological benefits (e.g. moderation of floods, groundwater recharge, repository of biodiversity) to the locality and its people. Deepor Beel was declared a Wildlife Sanctuary in 1989 and the area covered by the perennial water spread (10.1 km2) was subsequently designated as a Bird Sanctuary by the Government of Assam in 2009. The present paper provides an overview of the rich floral and faunal diversity of the wetland, including aquatic macrophytes (58 species), diatoms (65), zooplankton (171), bryozoans (5), molluscs (15), aquatic insects (55), prawns (3), crabs (2), finfish (68), amphibians (11), reptiles (33), birds (234) and mammals (24), in addition to the beel's limno-chemistry as well its conservation threats and suggested strategies. The wetland supports 18 globally threatened vertebrate species. Eleven bird species occurring in the wetland (Aythya baeri, A. nyroca, Leptoptilos dubius, L. javanicus, Ephippiorhynchus asiaticus, Gyps bengalensis, G. tenuirostris, Haliaeetus leucogaster, Pelecanus philippensis, Sarcogyps calvus, Threskiornis melanocephalus) have been included in the IUCN Red List of Threatened Species. Finfishes reported from the wetland declined from 67 species in 1999 to 64 between 2016 and 2018, which is a matter of concern. According to available information on limno-chemistry, the waters of the wetland have remained within the favorable range of water quality for fish production. Major threats faced by the wetland (e.g. pollution, encroachment, natural processes, etc.) and possible conservation strategies (e.g. control of aquatic pollution, prevention/ control of siltation, enacting and enforcing appropriate legislation, and other strategies) are outlined.

Forests Without Frugivores and Frugivores Without Forests – An Investigation Into the Causes of a Paradox in One of the Last Archipelagos Colonized by Humans

The Mascarenes are sadly famous worldwide for the massive extinction of their native vertebrates since recent human colonization. However, extinction patterns show astonishing disparities between the two main islands and between lineages of forest vertebrates. On Réunion (2,512 km2, 3,070 m) where about a third of native habitats remains, most large-bodied vertebrates, especially frugivores, collapsed by the first half of the 18th century, while several have survived longer and some still exist on Mauritius (1,865 km2, 828 m) where more than 95% of native habitats have been transformed. Considering lineages of forest vertebrates shared by both islands (23 genera, 53 species), we test the hypothesis that differing patterns of lowland suitable habitat destruction is the main cause behind this paradox. Before that, we assess the potential impact of other major drivers of extinctions since first contact with humans. Firstly, Mauritius shows earlier and more numerous introductions of mammal predators known for their devastating impact (except northern islets which have thus become important sanctuaries for several squamates). Secondly, settlers were inveterate hunters on both islands, but while Réunion was overhunted before Mauritius, the burst of human population in the latter in late 18th century has not led to the rapid extinction of all large native vertebrates. These two factors alone therefore cannot explain the observed paradox. Rather, the early destruction of lowland habitats (&amp;lt;400 m) on Réunion is concomitant with most extinctions of forest vertebrate, notably frugivores that rapidly lost most lowland habitats dominated by large fleshy-fruited plants. Moreover, landform-induced fragmentation has likely decreased the ability of adjacent habitats to act as effective refuges. Conversely, Mauritius retained suitable low-fragmented habitats until the late 19th which probably allowed, at least for a time, several native vertebrates to escape from multiple human-induced disturbances. Despite the almost total destruction of native habitats since then on Mauritius, conservation actions have saved several threatened vertebrate species that play a fundamental role in the functioning of native ecosystems. The fact that there are now more favorable habitats on Réunion than on Mauritius argues for the rewilding of Réunion with these extant large vertebrates.

The importance of protected areas and Indigenous lands in securing ecosystem services and biodiversity in the Cerrado

Assessing conservation areas’ effectiveness in safeguarding ecosystem services (ES) and biodiversity is mandatory in a world under intense human pressure. We evaluate the effectiveness of protected areas (PA) and Indigenous lands (IL) to represent five ES and threatened species in the Brazilian Cerrado. We compared the magnitude of ES and biodiversity recorded in each PA and IL (observed values) with that captured by reserves located randomly across the Cerrado (random values). We found that 23.8% of PA and 32.8% of IL were highly effective (i.e. observed value ≥ 75% of the random values) in representing three or more ES. Regarding biodiversity, 42.2% and 40.3% of PA were highly effective in representing threatened plant and vertebrate species, respectively; while 13.1% and 63.9% of IL were highly effective in capturing these species. The entire PA network performed better than expected based on its area for water yield, sediment retention, and threatened species. Likewise, the IL network showed a disproportionately high amount of water yield, carbon storage, and threatened vertebrate species records considering its area. Our results provide evidence that PA and IL are essential for ensuring conservation in the region and call attention to the need to strengthen these reserves politically.

Incorporating coextinction in threat assessments and policy will rapidly improve the accuracy of threatened species lists

Threatened species lists are important policy tools for species conservation, and they are often used to help prioritize recovery actions. Current listing processes, such as the IUCN red list threat assessment, rely on accepted correlates of extinction risk, especially population size. Population size appears to be an effective indicator for threatened vertebrate species, but it is less relevant for invertebrate species, where population sizes are rarely known. Furthermore, many invertebrate species are dependent upon a host species for survival and reproduction, and thus host population sizes would be better indicators of (co-)extinction vulnerability. Improving listing criteria to be relevant to invertebrates is important, because faunal biodiversity consists predominantly of invertebrate species (>98%) and invertebrates likely represent a majority of the current global sixth mass extinction event. Here, we propose including the degree of host specificity of a dependent invertebrate species to threatened host species as an additional correlate of extinction proneness, where there is a continuum from generalist dependents to dependents that are completely specialized to use a single host species per life stage. For highly specialized dependent taxa, we recommend that that threat category assigned to the dependent species should match the threat category assigned to the host species. Incorporating this correlate of extinction risk into threatened species policy, such as the IUCN red list, could rapidly increase the rate of assessments for many invertebrates and justify their inclusion, where previously they may have been considered data deficient.

Global reforestation and biodiversity conservation.

The loss of forest is a leading cause of species extinction, and reforestation is 1 of 2 established interventions for reversing this loss. However, the role of reforestation for biodiversity conservation remains debated, and lacking is an assessment of the potential contribution that reforestation could make to biodiversity conservation globally. We conducted a spatial analysis of overlap between 1,550 forest-obligate threatened species' ranges and land that could be reforested after accounting for socioeconomic and ecological constraints. Reforestation on at least 43% (∼369 million ha) of reforestable area was predicted to potentially benefit threatened vertebrates. This is approximately 15% of the total area where threatened vertebrates occur. The greatest opportunities for conserving threatened vertebrate species are in the tropics, particularly Brazil and Indonesia. Although reforestation is not a substitute for forest conservation, and most of the area containing threatened vertebrates remains forested, our results highlight the need for global conservation strategies to recognize the potentially significant contribution that reforestation could make to biodiversity conservation. If implemented, reforestation of ∼369 million ha would also contribute substantially to climate-change mitigation, offering a way to achieve multiple sustainability commitments at once. Countries must now work to overcome key barriers (e.g., unclear revenue streams, high transaction costs) to investment in reforestation.

Representativeness of threatened terrestrial vertebrates in nature reserves in China

Habitat degradation and loss threaten the survival of many species, especially vertebrates. The establishment of nature reserves is an efficient approach to protect biodiversity. However, fulfilling conservation targets depends largely on the effectiveness of these nature reserves in capturing the targeted species. In this study, we developed a distribution database of threatened terrestrial vertebrates in China, then explored the patterns of the threatened terrestrial vertebrates and evaluated the effectiveness of current nature reserves in terms of their conservation, further identified the conservation priorities for these species in China. We found 452 threatened terrestrial vertebrates widely distributed across China, especially in mountainous and forested regions in southern China. The current nature reserve network is inadequate to capture the threatened vertebrate species. The nature reserves were found to cover 30.47% of the hotspots and 38.65% of the ranges of these species on average, with quite a few conservation hotspot and species gaps. We further propose to either systematically plan and fix current nature reserves or add new protected areas in Yunnan Province and mountainous regions in southeast China.

Accounting for uncertainty in assessing the impact of climate change on biodiversity hotspots in Spain

Our limited understanding of the complexity of nature generates uncertainty in mathematical and cartographical models used to predict the effects of climate change on species’ distributions. We developed predictive models of distributional range shifts of threatened vertebrate species in mainland Spain, and in their accumulation in biodiversity hotspots due to climate change. We considered two relevant sources of climatological uncertainty that affect predictions of future climate: general circulation models and socio–economic scenarios. We also examined the relative importance of climate as a driver of species’ distribution and taxonomic uncertainty as additional biogeographical causes of uncertainty. Uncertainty was detected in all the forecasts derived from models in which climate was a significant explanatory factor, and in the species with taxonomic uncertainty. Uncertainty in forecasts was mainly located in areas not occupied by the species, and increased with time difference from the present. Mapping this uncertainty allowed us to assess the consistency of predictions regarding future changes in the distribution of hotspots of threatened vertebrates in Spain.

Vertebrate biodiversity on indigenous-managed lands in Australia, Brazil, and Canada equals that in protected areas

Declines in global biodiversity due to land conversion and habitat loss are driving a ‘Sixth Mass Extinction’ and many countries fall short of meeting even nominal targets for land protection. We explored how such shortfalls in Australia, Brazil and Canada might be addressed by enhancing partnerships between Indigenous communities and other government agencies that recognize and reward the existing contributions of Indigenous-managed lands to global biodiversity conservation, and their potential contribution to meeting international treaty targets. We found that Indigenous-managed lands were slightly more vertebrate species rich than existing protected areas in all three countries, and in Brazil and Canada, that they supported more threatened vertebrate species than existing protected areas or randomly selected non-protected areas. Our results suggest that overall, Indigenous-managed lands and existing protected areas host similar levels of vertebrate biodiversity in Brazil, Canada, and Australia. Partnerships with Indigenous communities that seek to maintain or enhance Indigenous land tenure practices on Indigenous-managed lands may therefore have some potential to ameliorate national and global shortfalls in land protection for biodiversity conservation using a mix of conventional protected areas and Indigenous-managed lands.

Threatened vertebrate species: associations between conservation actions, funding and population trends

Threatened vertebrate species: associations between conservation actions, funding and population trends

Feral horse impacts on threatened plants and animals in sub‐alpine and montane environments in Victoria, Australia

SummaryFeral herbivores are a major driver of biodiversity loss globally and can alter the structure, composition and functioning of ecosystems. The direct impacts of feral herbivores on plant communities are well studied, but the direct and indirect effect they have on wildlife is not well understood. In Victoria (south‐eastern Australia), a large feral Horse (Equus caballus) population coincides with highly sensitive and nationally endangered Alpine Sphagnum Bogs and Associated Fens communities, and several threatened animal species. We assessed the impact of feral horses on this ecological community and the Alpine Water Skink (Eulamprus kosciuskoi) and the Broad‐toothed Rat (Mastacomys fuscus) at 20 sites with varying levels of horse disturbance. We used scat counts to determine an index of feral horse abundance and quantified impacts associated with their presence in the landscape. Active searches were used for Alpine Water Skink and scat and runway surveys for Broad‐toothed Rat. We also measured the vegetation structure and the abundance of different vegetation types (life forms). Our results suggest that feral horses are associated with vegetation types and characteristics that negatively influence the presence or abundance of Alpine Water Skink and Broad‐toothed Rat. Sites with high horse activity had more low‐growing forbs, and the abundance of Alpine Water Skink was negatively related to this vegetation type. Grasses, sedges, rushes and shrubs were also less dense and lower in height in high horse activity sites, and Broad‐toothed Rat was less likely to be present in areas with these habitat attributes. We recommend that feral horses are controlled to protect these threatened vertebrate species and their Sphagnum bog habitat.

Assessing the spatiotemporal interactions of mesopredators in Sumatra's tropical rainforest.

Co-occurrence between mesopredators can be achieved by differentiation of prey, temporal activity, and spatial habitat use. The study of mesopredator interactions is a growing area of research in tropical forests and shedding new light on inter-guild competition between threatened vertebrate species that were previously little understood. Here, we investigate sympatry between the Sunda clouded leopard (Neofelis diardi) and Asiatic golden cat (Pardofelis temminckii) living in the Sumatran rainforests of Indonesia. We investigate: i) spatial overlap of predator-prey species using a combination of single-species occupancy modelling and Bayesian two-species modelling, while controlling for the possible influence of several confounding landscape variables; and, ii) temporal overlap between mesopredators and their shared prey through calculating their kernel density estimate associations. From four study areas, representing lowland, hill, sub-montane and montane forest, 28,404 camera trap nights were sampled. Clouded leopard and golden cat were respectively detected in 24.3% and 22.6% of the 292 sampling sites (camera stations) and co-occurred in 29.6% of the sites where they were detected. Golden cat occupancy was highest in the study area where clouded leopard occupancy was lowest and conversely lowest in the study area where clouded leopard occupancy was highest. However, our fine-scale (camera trap site) analyses found no evidence of avoidance between these two felid species. While both mesopredators exhibited highest spatial overlap with the larger-bodied prey species, temporal niche separation was also found. Clouded leopard was more nocturnal and, consequently, had higher temporal overlap with the more nocturnal prey species, such as porcupine and mouse deer, whereas the more diurnal golden cat had higher overlap with the strictly diurnal great argus pheasant. The Bayesian two species occupancy modelling approach applied in our study fills several important knowledge gaps of Sumatra’s lesser known mesopredators and provides a replicable methodology for studying interspecific competition for other small-medium sized carnivore species in the tropics.

Bias in protected-area location and its effects on long-term aspirations of biodiversity conventions.

To contribute to the aspirations of recent international biodiversity conventions, protected areas (PAs) must be strategically located and not simply established on economically marginal lands as they have in the past. With refined international commitments under the Convention on Biological Diversity to target protected areas in places of "importance to biodiversity," perhaps they may now be. We analyzed location biases in PAs globally over historic (pre-2004) and recent periods. Specifically, we examined whether the location of protected areas are more closely associated with high concentrations of threatened vertebrate species or with areas of low agricultural opportunity costs. We found that both old and new protected areas did not target places with high concentrations of threatened vertebrate species. Instead, they appeared to be established in locations that minimize conflict with agriculturally suitable lands. This entrenchment of past trends has substantial implications for the contributions these protected areas are making to international commitments to conserve biodiversity. If protected-area growth from 2004 to 2014 had strategically targeted unrepresented threatened vertebrates, >30 times more species (3086 or 2553 potential vs. 85 actual new species represented) would have been protected for the same area or the same cost as the actual expansion. With the land available for conservation declining, nations must urgently focus new protection on places that provide for the conservation outcomes outlined in international treaties.