Targeting Global Protected Area Expansion for Imperiled Biodiversity

Oscar Venter, Richard A. Fuller, Daniel B. Segan, Josie Carwardine, Thomas Brooks, Stuart H. M. Butchart, Moreno Di Marco, Takuya Iwamura, Liana Joseph, Damien O'Grady, Hugh P. Possingham, Carlo Rondinini, Robert J. Smith, Michelle Venter, James E. M. Watson · PLoS Biology · 2014 · conservation-science · 10.1371/journal.pbio.1001891 · record venter2014-targeting-global-protected-area-expansion-biodiversity
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protected areas Aichi target 11 agricultural value imperiled biodiversity biodiversity importance Aichi target 12 distribution of threatened species future establishment of protected areas species conservation gap analysis systematic conservation planning framework World Database on Protected Areas protected area network spatial resolution of 500 m 30 km×30 km pixels category I–VI protected areas protected area layer Critically Endangered Endangered Vulnerable threatened agricultural opportunity cost gross agricultural rents habitat loss agricultural expansion representation target adequacy target geographic range size Marxan simulated annealing algorithm planning unit conservation features conservation costs national protected area target business-as-usual protected area expansion scenario ecologically representative terrestrial ecoregions threatened species efficiency frontier tradeoff curve commission errors overestimation of occurrence simulated commission errors Extent of Suitable Habitat global protected area network agricultural opportunity cost of $4.9 billion threatened vertebrates wide-ranging species narrowly distributed species cheapest land land selected for threatened species low concordance difference in cost nonlinear tradeoff large gains in the number of species potentially protected relatively small increases in cost range maps high-resolution expert-based habitat suitability models cost-efficient manner minimum viable protected areas connectivity additional conservation actions cornerstone of biodiversity conservation societal objectives interdependence of these key commitments residual protected-area placement systematic conservation planning fine-resolution overlay protected-area layer correction opportunity-cost proxy range-scaled adequacy simulated annealing national target constraint efficiency-frontier weighting commission-error simulation spatial mismatch nonlinear tradeoff exploitation protected area coverage conservation targets species adequacy targets commission error rates management costs computational-model
Method, as described: global gap analysis and Marxan systematic conservation planning across national, ecoregional, species-adequacy, and efficiency-frontier scenarios with commission-error sensitivity analyses

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Targeting Global Protected Area Expansion for Imperiled Biodiversity

Abstract

concepts↗concepts↗concepts↗unhoused_relationsunhoused_relationscontent_tags↗Meeting international targets for expanding protected areas could simultaneously contribute to species conservation, but only if the distribution of threatened species informs the future establishment of protected areas.

Introduction

mechanisms↗concepts↗concepts↗concepts↗concepts↗content_tagscontent_tagscontent_tags↗In 2010 the 193 parties to the Convention of Biological Diversity (CBD) adopted a new strategic plan and set of targets to tackle the continuing decline in biodiversity [1] , [2] . A key element of this plan is Aichi target 11, which includes a commitment to expand the global coverage of terrestrial protected areas from the current 13% to 17% by 2020 [1] . This could drive the most rapid expansion of the global protected area network in history [3] , but corresponding biodiversity benefits are far from guaranteed. This is because protected areas are often preferentially established in locations that are remote or have little agricultural value [4] , failing to protect the imperiled biodiversity found on more valuable land.

object_of_studyconcepts↗concepts↗unhoused_relationsRecognizing the failures of past protected area expansion, the current CBD text directs that protected areas should target places of “importance for biodiversity” that are “ecologically representative” [1] . However, these locations can be expensive to protect. For instance, the cost of expanding protected areas to cover all “important bird areas” (IBAs) has been estimated at US$58 billion annually (although these sums are still small compared to government budgets) [5] . Moreover, the majority of terrestrial regions have been identified as important for biodiversity by one or more global prioritization schemes [6] , which provides myriad alternatives for meeting protected area targets in locations that are cheap. Given this, where should new protected areas be located to deliver on the Aichi biodiversity targets? One option could be based on Aichi target 12, which aims to “prevent the extinction of all known threatened species and improve and sustain their conservation status.” In situ conservation of viable populations in natural ecosystems has long been recognized as the fundamental requirement for the maintenance of biodiversity [7] . Hence measuring “biodiversity importance” in terms of protected area coverage of threatened species would help countries to simultaneously meet these two CBD targets.

research_questioncontributionclaim_typemechanisms↗concepts↗concepts↗concepts↗concepts↗claim_relationscontent_tagsUsing new data from the World Database on Protected Areas [3] and distribution maps for 4,118 globally threatened birds [8] , mammals [9] , [10] , and amphibians [10] , [11] , as well as ecoregions [12] , we first perform a gap analysis to determine the representation of these species in the current global protected area network. We then use a systematic conservation planning framework [13] to build scenarios for cost-efficiently expanding the global protected area network to contribute to meeting the protected area and threatened species Aichi targets. Recent works have investigated strategies for achieving Aichi Target 11 by protecting IBAs [5] , [14] or meeting the Global Strategy for Plant Conservation [15] . Our study is the first, to our knowledge, to use an optimization approach to develop scenarios for meeting the Aichi targets in a cost-efficient manner. Incorporating cost efficiency allows the identification of options for meeting Aichi target 11 that contribute optimally to target 12 while minimizing conflict with agricultural production.

Methods

mechanisms↗concepts↗concepts↗content_tagscontent_tagsAll spatial overlays were performed at a spatial resolution of 500 m and then aggregated into 30 km×30 km pixels to identify candidate land for protection. By processing data at the finer resolution, we are able to account for protected areas at the subpixel level, thereby minimizing omission of small-sized protected areas. This resolution of ∼⅓ degree (at the Equator) falls in the midrange between scales of ½ degree [16] and of ⅛ degree [17] typically used in such analyses.

Protected Areas

mechanisms↗concepts↗concepts↗content_tagscontent_tagscontent_tagsTo determine the extent of current protected areas, we extracted data on International Union for Conservation of Nature (IUCN) category I–VI protected areas from the 2012 World Database on Protected Areas [3] , excluding all proposed protected areas and those lacking “national” designation. For terrestrial protected areas with a known areal extent but lacking polygonal representation, we created a circular buffer of the appropriate area around its centroid. To prevent overestimation of the areal coverage of protected areas caused by overlapping designations, we merged buffered points and polygons into a single layer. Our final protected area layer contained 135,062 protected areas covering a total of 17,026,214 km 2 , or 12.9% of the Earth's non-Antarctic land surface ( Figure 1A ).

Key data inputs and output map from the systematic conservation planning framework. (A) Protected areas mapped using polygons and buffered points for nationally designated protected areas [3] . (B) The number of native and extant globally threatened terrestrial and freshwater birds [8] , mammals [10] , and amphibians [10] per grid square. (C) The average annual agricultural opportunity cost of protecting each 30 km grid square in 2012 $US [17] . (D) The distribution of priorities for establishing new protected areas to meet the national-level 17% targets under Aichi target 11 at minimal cost and ignoring ecological representation (red), for covering threatened species (green), and locations selected under both scenarios (yellow). The sizes of the circles in the Venn diagrams are proportional to the area required in each of the three categories.

Distribution of Biodiversity

limitationsconcepts↗concepts↗concepts↗concepts↗unhoused_relationssensescontent_tags↗content_tagscontent_tagscontent_tagsWe used distribution maps for birds [8] , mammals [10] , and amphibians [10] . We focused on these taxa as they are the only major terrestrial taxonomic groups that have been comprehensively assessed for their distribution and extinction risk [10] . We excluded marine species and areas, noting that there are specific coverage targets for protecting the marine realm. For all three taxonomic groups, we focused on those species that are listed by the IUCN Red List as Critically Endangered, Endangered, or Vulnerable, hereafter referred to as “threatened,” resulting in 4,118 species in total (birds = 1,135, mammals = 1,107, amphibians = 1,876; Figure 1B ). We focus only on threatened species as these are by definition the most likely species to go extinct, and therefore are most important for slowing biodiversity loss and contributing to CBD Aichi target 12. We excluded all portions of species ranges where the species was identified as extinct, introduced, or of uncertain origin. In addition to these data, we used data on the distribution of ecoregions as defined by the World Wildlife Fund [12] .

Protected Area Opportunity Cost

mechanisms↗limitationslimitationslimitationsconcepts↗concepts↗concepts↗concepts↗unhoused_relationssensesconcept_edgescontent_tags↗content_tagscontent_tagscontent_tags↗content_tags↗To account for the spatial variation in the cost of protected area expansion, we used a dataset on agricultural opportunity cost [18] , converted to 2012 US$ and with no data values filled using regularized spline interpolation with tension ( Figure 1C ). The dataset provides the estimated gross agricultural rents for terrestrial areas mapped at approximately the 5 km resolution. We use these data as our surrogate for the opportunity costs of establishing new protected areas, as agricultural expansion is the greatest single cause of habitat loss, as well as the one most commonly associated with habitat loss driven by multiple factors [19] , [20] . Agricultural opportunity costs also reflect the reduction in food security and tax revenue that national governments face when implementing protected areas. We applied a fixed cost of US$100 per km 2 to reflect the transaction costs of acquiring new protected areas [21] , although we recognize there is likely to be considerable spatial variation in these costs. We did not attempt to estimate the ongoing management costs of protected areas following establishment, as this metric needs to account for a number of difficult-to-measure social and socioeconomic factors [22] , but a recent analysis estimated that these equate to ∼14% of the agricultural opportunity costs of protection [5] .

Gap Analysis

mechanisms↗concepts↗concepts↗concepts↗unhoused_relationssensessensescontent_tags↗content_tags↗content_tagscontent_tagscontent_tagscontent_tagscontent_tags↗We assessed the occurrence of threatened vertebrates within protected areas using a representation target and an adequacy target. The representation target was achieved if any portion of the species' distribution overlapped with the protected area network. To set adequacy targets we followed the method of Rodrigues et al. [23] to scale the target to the species' overall geographic range size. Complete (i.e., 100%) coverage by protected areas was required for species with a geographic range of <1,000 km 2 . For wide-ranging species (>250,000 km 2 ), the target was reduced to 10% coverage, and where geographic range size was intermediate between these extremes, the target was log-linearly interpolated.

Scenarios for Protected Area Expansion

mechanisms↗concepts↗concepts↗concepts↗concepts↗concepts↗content_tags↗content_tags↗content_tags↗To explore future scenarios for the growth of the global protected area network we used the systematic conservation planning software Marxan [24] . Marxan uses a simulated annealing algorithm to select multiple alternative sets of areas that meet prespecified conservation targets (described in the following section) while trying to minimize overall cost. All spatial data on the distribution of conservation features and conservation costs were summarized into a “planning unit” layer consisting of 30 km×30 km square pixels comprising the world's non-Antarctic terrestrial areas. We intersected this planning unit layer with the protected areas and agricultural opportunity layers and the geographic distribution of each of the 4,118 threatened species and ecoregions at a 500 m resolution. This allowed us to determine the agricultural opportunity cost of the unprotected portion of each planning unit and the protected and unprotected extent of each biodiversity feature within each planning unit.

method↗mechanisms↗concepts↗unhoused_relationscontent_tagscontent_tags↗To explore the costs and benefits of alternate scenarios for achieving 17% protection of terrestrial areas, we developed four separate spatial scenarios using contrasting conservation targets. We accounted for the existing protected area network's contribution to the targets in each scenario, and then added additional protected areas to ensure all targets are met. In each scenario, the aim is to minimize the costs of meeting the conservation targets. However, to avoid the global protected area target being met only through increased protection in low-cost countries, which would reduce the total cost of the target, in all scenarios we maintain the constraint that each country must meet its national protected area target. Moreover, it is at the national level that the target is being interpreted and implemented. For each scenario, we used Marxan to perform 10 runs of 1 billion iterations each, each of which represents an alternate near optimal reserve network for meeting the relevant conservation targets at the lowest overall cost. From these 10 runs, we select and report on the results from the lowest cost solution.

National targets

concepts↗sensescontent_tags↗In the first scenario, we set the conservation target as each country meeting its protected area target at the lowest agricultural opportunity cost. In this scenario, we set all countries' terrestrial protected area target to 17%, except for the 73 countries that have indicated in CBD workshops that they proposed alternative targets [25] , in which case we used these targets. As countries have tended in the past to meet their targets by favoring high, far, and otherwise agriculturally low-value areas [4] , we view this as our business-as-usual protected area expansion scenario. We also determine the conservation benefits of protected area target levels above the current Aichi 17% targets by setting national levels up to 30% of each country.

Ecoregional target

concepts↗concepts↗unhoused_relationscontent_tagscontent_tagsIn this scenario, we maintain the national-level 17% targets from scenario a but add the additional constraint that countries meet their target in a way that ensures that each of the 821 terrestrial ecoregions receive at least 17% protection. We include this scenario as Target 11 calls for areas protected to be “ecologically representative” [1] .

Threatened species target

concepts↗In this scenario, we maintain the national-level 17% targets from scenario a but add the additional constraint that all threatened species must be covered to the level of their adequacy targets [23] .

Threatened species preference

mechanisms↗concepts↗concepts↗sensescontent_tags↗content_tags↗In this scenario, we construct an efficiency frontier between the cost of meeting the 17% target as in a and attaining threatened species conservation targets as in c. The tradeoff curve is established by iteratively increasing the value given to meeting species adequacy targets, from no value to a value equal to that given to the 17% target itself. The 17% target is always met at the national level across the tradeoff frontier.

Commission Errors in Range Maps

mechanisms↗limitationslimitationsconcepts↗concepts↗concepts↗concepts↗unhoused_relationssensesconcept_edgescontent_tags↗content_tagscontent_tagscontent_tags↗The IUCN [10] and Birdlife International and NatureServe [8] range maps used in this study comprise polygons showing distribution of 4,118 globally threatened birds, mammals, and amphibians. These maps may be subject to commission errors [26] – [29] , where the species is mapped as present in locations where it is in fact not present. As they affect range-based species conservation targets and lead to an overestimation of occurrence in existing or prioritized areas, commission errors could influence our study's main conclusions. We performed two analyses to determine the sensitivity of our primary results to commission errors ( Text S1 ). First we created 100 range maps for each of the 4,118 species of birds, mammals, and amphibians that simulated commission error rates [25] by deleting 50% of the range of narrow-ranged species (range<1,000 km 2 ), by deleting 25% of the range of wide-ranging species (range>250,000 km 2 ), and by linearly extrapolating the deletion rate for species of intermediate ranges. Second, we identified the “Extent of Suitable Habitat” (ESH) using high-resolution species distribution models for 1,063 mammals [30] . The ESH maps were used to identify locations in the original maps for mammals that are likely to be commission errors. We then reran our analyses using (a) the maps with simulated commission errors and (b) the ESH maps, to quantify the effects of the simulated and mapped commission errors on our estimated biodiversity value of meeting the 17% protected area target, and the shape of the efficiency frontier between cost and threatened vertebrate coverage.

Results

concepts↗content_tagscontent_tagscontent_tagscontent_tagsWe find that 17% of threatened vertebrates are not found in a single protected area and 85% are not covered to the level of our adequacy targets ( Figure S1A ). A decade ago, 20% of globally threatened terrestrial birds, mammals, and amphibians were not found in a single protected area and 89% were inadequately protected [15] . Our analysis using updated datasets indicates that the global protected area network has made little progress since then toward securing a future for the world's threatened biodiversity.

concepts↗concepts↗content_tagscontent_tagscontent_tagscontent_tagscontent_tagscontent_tagscontent_tagsWe discover that if countries choose to expand their protected areas in a manner that minimizes agricultural opportunity cost, meeting their national-level targets for 17% coverage would entail a once-off transaction cost of US$0.9 billion and an annual agricultural opportunity cost of $4.9 billion ( Table 1 ). As this option aligns with the previous pattern of protected area establishment, we view it as a likely business-as-usual scenario for meeting the terrestrial coverage aspect of Aichi target 11. We find that this would result in only 852 (21%) threatened vertebrates reaching targets for adequate coverage ( Figure S1B ), an increase of only 249 species over existing protection ( Table 1 ) and arguably a failure to meet Aichi target 12. Moreover, even if highly ambitious areal targets were to drive further growth of the global protected area network beyond 2020, the costs of expansion would rise steeply without providing cost-effective coverage for threatened species ( Figure 2 ).

The number of globally threatened vertebrates that reach our adequacy targets (black), and the agricultural opportunity cost of establishing new protected areas (red), as the proportion of global land areas protected increases above 17%.

Costs and benefits of the current protected area network and for future protection scenarios that (a) meet country-level targets for protected area coverage; (b) meet these targets while also achieving 17% protection of each terrestrial ecoregion; (c) meet the targets from scenario a and protect a scaled fraction of the geographic ranges of threatened terrestrial birds, mammals, and amphibians; and (d) achieve the country-level targets for protected area coverage while also achieving five times the level of biodiversity protection relative to scenario a.

concepts↗concepts↗concept_edgescontent_tagscontent_tagscontent_tagscontent_tagsAn alternative is to ensure a representative sample of major vegetation communities is protected, as this would protect a broader range of habitats and could lead to improved conservation outcomes. Target 11 calls for ecologically representative protected area coverage. We find that if countries meet their 17% coverage targets in a way that distributes protection across ecoregions equally, the opportunity cost of establishing the additional protected areas would be 4.5 times higher than the business-as-usual scenario ($24.8 billion annually; Table 1 ), but that coverage of threatened species would increase only marginally ( Figure S1C ). Moreover, the majority of species that reach their adequacy targets are those with a geographic range size ≥250,000 km 2 ( Figure S1C ), as their wide distribution renders them more easily captured when distributing protected areas equitably across ecoregions. The species most likely to be left unprotected are narrowly distributed species, which often are those in greatest need of protection [31] , [32] .

phenomenonmechanisms↗concepts↗concepts↗concepts↗concepts↗unhoused_relationsclaim_relationsconcept_edgesconcept_edgescontent_tagscontent_tagscontent_tagscontent_tagsThese results indicate that protected area expansion targeting either the cheapest land or representation of ecoregions is not an efficient approach for covering threatened species. Alternatively, we find that locating protected areas to ensure they meet targets for adequate coverage of all 4,118 threatened species would cost about $42.5 billion annually ( Table 1 ), which is about 7.5 times more than the cheapest option for meeting the 17% target. This difference in cost is driven by low concordance between areas that are cheap to protect and those that capture the distributions of threatened species ( Figure 1D ). Land selected for threatened species tends to align with tropical forest hotspots ( Figure 1B ), such as the tropical Andes and eastern Madagascar, whereas the cheapest land to protect is remote and often in more arid zones ( Figure 1D ). This lack of overlap helps explain why the existing protected area network, which has favored low-cost areas in each country [4] , represents threatened species rather poorly.

mechanisms↗concepts↗concepts↗concepts↗unhoused_relationsunhoused_relationsclaim_relationscontent_tagscontent_tagscontent_tagscontent_tagsHow can countries reconcile the attraction of low-cost conservation with the benefits of protecting places that contribute to threatened species conservation? By varying the importance placed on meeting targets for adequate coverage of threatened species, we discover a nonlinear tradeoff between the cost of establishing additional protected areas and the proportion of threatened vertebrates covered by these areas ( Figure 3 ). The shape of the curve illustrates that large gains in the number of species potentially protected could be achieved for relatively small increases in cost. For instance, increasing by 5-fold the number of species protected relative to the low-cost, business-us-usual scenario would increase opportunity costs to only $7.4 billion annually (1.5 times as much; Table 1 ).

Efficiency frontier between the cost of establishing additional protected areas to achieve 17% coverage and the number of species covered. The y -axis presents the proportion of each species adequacy target that is met within protected areas, summed across all species, and is not directly comparable to that of the other figures, which only count species whose protected area coverage meets or exceeds their target.

concepts↗concepts↗unhoused_relationscontent_tagscontent_tagsWe find that our primary results are robust to randomly simulated commission errors in the range maps. Although the number of species meeting range-based coverage targets generally decreases once commission errors are simulated ( Text S1 ), this drop averages only 5% across the tradeoff curve ( Figure S2 ). Moreover, both a visual interpretation and a quantitative measure of the shape of the tradeoff curve reveals that the original and commission error updated curves are similarly nonlinear. Moreover, using high-resolution expert-based habitat suitability models for 1,063 threatened mammals, we again find that commission errors are unlikely to alter our primary findings ( Figure S3 ).

Discussion

concepts↗content_tagscontent_tagscontent_tagscontent_tagscontent_tagscontent_tagsA small minority (15%) of threatened vertebrates are adequately covered by existing protected areas. However, the adoption of the Aichi targets marks an historic opportunity for achieving conservation of the world's biodiversity. If countries are to meet the protected area Aichi target, at least 5.8 million km 2 of new protected areas will need to be created by 2020. Although this is a significant opportunity for biodiversity conservation, we have shown that protected area expansion that targets low-cost areas in each country and ignores threatened species is unlikely to protect such species incidentally. This remains the case even if protected areas are further expanded to cover 30% of land areas, or if they are located to cover a representative sample of Earth's terrestrial ecoregions. On the other hand, we find that if protected areas are directed in a cost-efficient manner to protect threatened vertebrates, these species could be protected for an estimated agricultural opportunity cost of about $42.5 billion annually. We also find that there is a nonlinear relationship between cost and species protection, indicating that options exist for increasing threatened species protection above the business-as-usual level at little additional cost.

Our estimate of the cost of reaching adequacy targets for all threatened birds, mammals, and amphibians is lower than the $58 billion annually estimated for protecting the world's IBAs [5] , though each option comprises a similar land area. There are three primary reasons for this. First, the estimated costs of protecting IBAs include management costs, which are estimated at ∼$7 billion annually [5] . Second, IBAs are identified for their contribution to global bird conservation, without consideration of the cost of protecting these areas, whereas we used an optimization approach to identify low-cost options for meeting conservation targets [33] , [34] . Third, IBAs are identified based on the presence of both threatened and nonthreatened species (e.g., congregatory species), while we focused on threatened species alone.

limitationslimitationslimitationslimitationslimitationslimitationslimitationsconcepts↗concepts↗concepts↗unhoused_relationsunhoused_relationsunhoused_relationssensesclaim_relationscontent_tags↗Our analyses are subject to a number of caveats. First, we considered relative cost based on gross agricultural rents, not management costs or the opportunity costs for other land uses [33] , nor the practicalities of establishing reserves among these competing land uses. Second, overlay of coarse scale maps of species distributions onto fine-scale protected area maps generates commission errors [26] , [35] , though these are unlikely to qualitatively change our results. Still, as commission errors mean that species distributions overlap less than these coarse-scale maps suggest, our estimate of the area needed to protect all threatened species is a minimum [30] . Locations identified here should therefore be considered as broad indications of where specific areas for protection might be located, and our estimates of cost and the area requiring protection will be minima. Third, although we recognize that our analyses have limited taxonomic breadth, no other taxonomic groups (e.g., plants) have undergone comprehensive assessment of both extinction risk and distribution at a sufficiently fine scale for a comparable analysis [10] . Yet good indications exist from the literature that protected areas identified for broad taxonomic groups cover the majority of species in other, nontarget groups [36] , [37] . Finally, our species-specific targets for protection do not account for minimum viable protected areas or connectivity and do not guarantee the long-term survival of all species. Moreover, many species are threatened by processes other than habitat loss and therefore require additional conservation actions both inside and outside protected areas [38] .

concepts↗concepts↗concepts↗unhoused_relationsclaim_relationsclaim_relationsFor the global protected area network to fulfill its potential role as the cornerstone of biodiversity conservation [39] , and for governments to meet their commitments on protected areas and species extinctions, the distribution of threatened species must inform future protected area establishment. Preventing the further loss of all threatened species is a lofty goal and will require substantial efforts. But expanding protected areas requires managing tradeoffs among societal objectives [40] , and here we have shown that considerable increases in protected area coverage of species could be achieved at modest additional cost. Exploiting the nonlinearity of this tradeoff will require directly linking the Aichi targets on protected areas and threatened species (as well as other targets, including target 5 on slowing habitat loss), thereby formalizing the interdependence of these key commitments.

Supporting Information

The total extant geographic range size, in logarithmic scale, and the percent of that range in protected areas for 4,118 threatened vertebrates, with the red line detailing the range-based conservation targets used in the analyses. “a” shows the protection afforded by the current protected areas, “b” shows the protection from the current network plus new protected areas necessary to meet national-level 17% targets, and “c” shows the protection from the current network plus new protected areas to meet national-level 17% targets in a way that ensures terrestrial ecoregions are protected to the level of 17%. Numbers in the graphs give the number of threatened species that have their adequacy target fully met in each scenario.

(EPS)

Click here for additional data file.

Efficiency frontier between the cost of establishing additional protected areas to achieve 17% coverage and the number of species potentially covered for the original range maps (black circles) and the randomly reduced species range maps (red stars). The y -axis presents the proportion of each species adequacy target that is met within protected areas, summed across all species. The red stars show the average results from 100 iterations of randomly deleting a portion of each species range; standard deviations for the 100 runs average ±0.82% across the tradeoff frontier and are therefore too small to graph.

(EPS)

Click here for additional data file.

Efficiency frontier between the cost of establishing additional protected areas to achieve 17% coverage and the number of mammal species potentially covered for original range maps (black circles) and the ESH maps (red stars). The y -axis presents the proportion of each species adequacy target that is met within protected areas, summed across all species.

(EPS)

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Analyses of sensitivity to range map commission errors.

(DOCX)

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facettagfieldconf.whose words
object_of_studywhere global protected-area expansion should occur to meet area and threatened-species targets cost-efficientlyobject_of_studythe paper's words
research_questionhow can countries expand terrestrial protected areas while balancing agricultural opportunity cost, ecological representation, and coverage of threatened vertebratesresearch_questionthe paper's words
contributionthe first optimization-based global scenarios jointly targeting Aichi protected-area coverage and threatened-species adequacy at minimum agricultural opportunity costcontributionthe paper's words
claim_typemethodclaim_typethe paper's words
phenomenonlow spatial concordance and a nonlinear cost tradeoff between cheap protected-area expansion and threatened-species coveragephenomenonthe paper's words
methodcomputational-modelmethodthe paper's words
mechanismsPreference for remote or low-value land leaves imperiled biodiversity on valuable land unprotected.mechanisms[0]0.99the tagger's words
mechanismsOptimization locates added protected areas that meet targets while minimizing agricultural conflict.mechanisms[1]0.99the tagger's words
mechanismsProcessing at 500 metres retains subpixel protected areas before aggregation to planning units.mechanisms[2]0.99the tagger's words
mechanismsBuffered nonpolygon records are merged with polygons to avoid double-counting overlapping designations.mechanisms[3]0.99the tagger's words
mechanismsGross agricultural rents represent the foregone production cost of new protected areas.mechanisms[4]0.99the tagger's words
mechanismsCoverage requirements decrease log-linearly as species range size increases.mechanisms[5]0.99the tagger's words
mechanismsMarxan searches alternative planning-unit sets that meet targets at low cost.mechanisms[6]0.99the tagger's words
mechanismsEvery country must meet its own target, preventing global savings through concentration in low-cost countries.mechanisms[7]0.99the tagger's words
mechanismsIncreasing the value assigned to species adequacy traces cost against biodiversity coverage.mechanisms[8]0.99the tagger's words
mechanismsRandom range deletion and suitable-habitat maps test sensitivity of results to false mapped presence.mechanisms[9]0.99the tagger's words
mechanismsLow overlap between cheap land and threatened-species ranges makes comprehensive species targeting more expensive.mechanisms[10]0.99the tagger's words
mechanismsModest added cost can capture a disproportionately large increment in threatened-species coverage.mechanisms[11]0.99the tagger's words
limitationsOnly birds, mammals, and amphibians had sufficiently comprehensive distribution and extinction-risk assessments.limitations[0]0.99the tagger's words
limitationsAgricultural rent is a surrogate rather than a complete measure of protected-area opportunity cost.limitations[1]0.99the tagger's words
limitationsThe fixed transaction cost ignores likely spatial variation.limitations[2]0.99the tagger's words
limitationsOngoing protected-area management costs were not estimated.limitations[3]0.99the tagger's words
limitationsCoarse species-range maps can contain commission errors that overestimate occurrence.limitations[4]0.99the tagger's words
limitationsThe sensitivity simulation deletes fixed fractions of estimated ranges rather than observing true occurrence.limitations[5]0.97the tagger's words
limitationsCost estimates exclude management cost, other land-use opportunity costs, and practical reserve-establishment constraints.limitations[6]0.99the tagger's words
limitationsCoarse species ranges over fine protected-area maps make the required area and cost estimates minima.limitations[7]0.99the tagger's words
limitationsPriority locations are broad indications, not specific reserve boundaries.limitations[8]0.99the tagger's words
limitationsTaxonomic breadth is limited because other groups lack comparable global assessments.limitations[9]0.99the tagger's words
limitationsSpecies targets do not account for minimum viable protected areas or connectivity.limitations[10]0.99the tagger's words
limitationsMeeting habitat-coverage targets does not guarantee long-term survival.limitations[11]0.99the tagger's words
limitationsSpecies threatened by processes other than habitat loss require additional actions inside and outside protected areas.limitations[12]0.99the tagger's words
conceptsprotected areasconcepts[0]0.98the paper's words
conceptsAichi target 11concepts[1]0.98the paper's words
conceptsagricultural valueconcepts[2]0.98the paper's words
conceptsimperiled biodiversityconcepts[3]0.98the paper's words
conceptsbiodiversity importanceconcepts[4]0.98the paper's words
conceptsAichi target 12concepts[5]0.98the paper's words
conceptsdistribution of threatened speciesconcepts[6]0.98the paper's words
conceptsfuture establishment of protected areasconcepts[7]0.98the paper's words
conceptsspecies conservationconcepts[8]0.98the paper's words
conceptsgap analysis
also in: gap analysis
concepts[9]0.98the paper's words
conceptssystematic conservation planning frameworkconcepts[10]0.98the paper's words
conceptsWorld Database on Protected Areasconcepts[11]0.98the paper's words
conceptsprotected area networkconcepts[12]0.98the paper's words
conceptsspatial resolution of 500 mconcepts[13]0.98the paper's words
concepts30 km×30 km pixelsconcepts[14]0.98the paper's words
conceptscategory I–VI protected areasconcepts[15]0.98the paper's words
conceptsprotected area layerconcepts[16]0.98the paper's words
conceptsCritically Endangeredconcepts[17]0.98the paper's words
conceptsEndangered
also in: Endangered
concepts[18]0.98the paper's words
conceptsVulnerable
also in: Vulnerable
concepts[19]0.98the paper's words
conceptsthreatened
also in: threatened
concepts[20]0.98the paper's words
conceptsagricultural opportunity costconcepts[21]0.98the paper's words
conceptsgross agricultural rentsconcepts[22]0.98the paper's words
conceptshabitat loss
also in: habitat loss
concepts[23]0.98the paper's words
conceptsagricultural expansionconcepts[24]0.98the paper's words
conceptsrepresentation targetconcepts[25]0.98the paper's words
conceptsadequacy targetconcepts[26]0.98the paper's words
conceptsgeographic range sizeconcepts[27]0.98the paper's words
conceptsMarxan
also in: Marxan
concepts[28]0.98the paper's words
conceptssimulated annealing algorithmconcepts[29]0.98the paper's words
conceptsplanning unit
also in: planning unit
concepts[30]0.98the paper's words
conceptsconservation featuresconcepts[31]0.98the paper's words
conceptsconservation costsconcepts[32]0.98the paper's words
conceptsnational protected area targetconcepts[33]0.98the paper's words
conceptsbusiness-as-usual protected area expansion scenarioconcepts[34]0.98the paper's words
conceptsecologically representativeconcepts[35]0.98the paper's words
conceptsterrestrial ecoregionsconcepts[36]0.98the paper's words
conceptsthreatened speciesconcepts[37]0.98the paper's words
conceptsefficiency frontierconcepts[38]0.98the paper's words
conceptstradeoff curveconcepts[39]0.98the paper's words
conceptscommission errorsconcepts[40]0.98the paper's words
conceptsoverestimation of occurrenceconcepts[41]0.98the paper's words
conceptssimulated commission errorsconcepts[42]0.98the paper's words
conceptsExtent of Suitable Habitatconcepts[43]0.98the paper's words
conceptsglobal protected area networkconcepts[44]0.98the paper's words
conceptsagricultural opportunity cost of $4.9 billionconcepts[45]0.98the paper's words
conceptsthreatened vertebratesconcepts[46]0.98the paper's words
conceptswide-ranging speciesconcepts[47]0.98the paper's words
conceptsnarrowly distributed speciesconcepts[48]0.98the paper's words
conceptscheapest land
also in: cheapest land
concepts[49]0.98the paper's words
conceptsland selected for threatened speciesconcepts[50]0.98the paper's words
conceptslow concordanceconcepts[51]0.98the paper's words
conceptsdifference in costconcepts[52]0.98the paper's words
conceptsnonlinear tradeoffconcepts[53]0.98the paper's words
conceptslarge gains in the number of species potentially protectedconcepts[54]0.98the paper's words
conceptsrelatively small increases in costconcepts[55]0.98the paper's words
conceptsrange maps
also in: range maps
concepts[56]0.98the paper's words
conceptshigh-resolution expert-based habitat suitability modelsconcepts[57]0.98the paper's words
conceptscost-efficient mannerconcepts[58]0.98the paper's words
conceptsminimum viable protected areasconcepts[59]0.98the paper's words
conceptsconnectivity
also in: connectivity
concepts[60]0.98the paper's words
conceptsadditional conservation actionsconcepts[61]0.98the paper's words
conceptscornerstone of biodiversity conservationconcepts[62]0.98the paper's words
conceptssocietal objectivesconcepts[63]0.98the paper's words
conceptsinterdependence of these key commitmentsconcepts[64]0.98the paper's words
unhoused_relationsdistribution of threatened species → future establishment of protected areas
Threatened-species distributions should inform new protected-area locations.
unhoused_relations[0]0.99the tagger's words
unhoused_relationsprotected areas → species conservation
Area expansion can contribute to species conservation only when species distributions guide siting.
unhoused_relations[1]0.99the tagger's words
unhoused_relationsAichi target 11 → Aichi target 12
Threatened-species coverage can help countries meet both international targets simultaneously.
unhoused_relations[2]0.99the tagger's words
unhoused_relationsCritically Endangered, Endangered, or Vulnerable → threatened
The three IUCN Red List statuses are collectively referred to as threatened.
unhoused_relations[3]0.99the tagger's words
unhoused_relationsagricultural opportunity cost → gross agricultural rents
Gross agricultural rents serve as a surrogate for the opportunity cost of protection.
unhoused_relations[4]0.99the tagger's words
unhoused_relationsgeographic range size → adequacy target
Range size determines the fraction of a species distribution that must be protected.
unhoused_relations[5]0.99the tagger's words
unhoused_relationsnational protected area target → global cost minimization
National constraints prevent all added protection from being concentrated in cheap countries.
unhoused_relations[6]0.99the tagger's words
unhoused_relationsterrestrial ecoregions → ecologically representative
Equal ecoregion coverage implements the policy requirement for ecological representation.
unhoused_relations[7]0.99the tagger's words
unhoused_relationscommission errors → primary conclusions
Commission errors could influence the main conclusions but sensitivity analyses indicate little qualitative change.
unhoused_relations[8]0.99the tagger's words
unhoused_relationscheapest land → threatened species
Cheap protected-area expansion represents threatened species poorly because cheap land and species ranges rarely overlap.
unhoused_relations[9]0.99the tagger's words
unhoused_relationsnonlinear tradeoff → cost-efficient species protection
The frontier exposes options for large species gains at modest additional cost.
unhoused_relations[10]0.99the tagger's words
unhoused_relationslarge gains in the number of species potentially protected → relatively small increases in cost
Large gains in species coverage can be obtained for relatively small cost increases.
unhoused_relations[11]0.99the tagger's words
unhoused_relationscommission errors → primary results
Both simulated deletion and suitable-habitat models leave the main results robust.
unhoused_relations[12]0.99the tagger's words
unhoused_relationspriority locations → specific protected areas
Mapped priorities are broad indications rather than final reserve boundaries.
unhoused_relations[13]0.99the tagger's words
unhoused_relationsspecies-specific targets → long-term survival
Coverage targets do not guarantee long-term species survival.
unhoused_relations[14]0.99the tagger's words
unhoused_relationsprotected areas → additional conservation actions
Non-habitat threats require measures inside and outside protected areas.
unhoused_relations[15]0.99the tagger's words
unhoused_relationsprotected areas target → threatened species target
The commitments should be formally linked to exploit the cost-coverage tradeoff.
unhoused_relations[16]0.99the tagger's words
sensescoverage - representation sense: any protected overlap with a species distributionsenses[0]0.98the tagger's words
sensescoverage - adequacy sense: a range-size-scaled fraction of distribution protectedsenses[1]0.98the tagger's words
sensescost - agricultural-opportunity sense: forgone gross agricultural rentsenses[2]0.98the tagger's words
sensesthreatened - IUCN sense: Critically Endangered, Endangered, or Vulnerablesenses[3]0.98the tagger's words
sensesefficiency - frontier sense: species-target attainment gained per added protection costsenses[4]0.98the tagger's words
sensesbusiness as usual - residual-siting sense: national area targets met on lowest-cost landsenses[5]0.98the tagger's words
sensescommission error - range-map sense: predicted presence where a species is absentsenses[6]0.98the tagger's words
sensespriority location - broad-indication sense: regional guidance rather than a reserve boundarysenses[7]0.98the tagger's words
claim_relationsextends → global protected-area gap analyses
note: The study adds an optimization framework and joint cost/species scenarios.
claim_relations[0]0.99the tagger's words
claim_relationscontrasts-with → low-cost area-only protected-area expansion
note: Species-targeted locations are compared with cheap, often arid and remote locations.
claim_relations[1]0.99the tagger's words
claim_relationssupports → joint implementation of Aichi targets 11 and 12
note: Linking targets exploits the nonlinear cost-species tradeoff.
claim_relations[2]0.99the tagger's words
claim_relationsqualifies → range-map-based priority locations
note: The locations are broad indications and estimated area and cost are minima.
claim_relations[3]0.99the tagger's words
claim_relationsrecommends → threatened-species-informed protected-area establishment
note: The conclusion states that species distributions must inform expansion.
claim_relations[4]0.99the tagger's words
claim_relationssupports → cost-efficient threatened-species targeting
note: The nonlinear frontier permits large biodiversity gains at relatively small added cost.
claim_relations[5]0.99the tagger's words
concept_edgesagricultural expansion —causes→ habitat loss
the relation itself is separately evidenced · strength of the relation: 0.95 · note: Strong flat causal statement. · the relation stated: Agricultural expansion is stated to be the greatest single cause of habitat loss.
concept_edges[0]0.99the tagger's words
concept_edgescommission errors —causes→ overestimation of occurrence
the relation itself is separately evidenced · strength of the relation: 0.95 · note: Strong flat causal statement. · the relation stated: False mapped presence leads to overestimated occurrence in protected and priority areas.
concept_edges[1]0.99the tagger's words
concept_edgeslow concordance —causes→ difference in cost
the relation itself is separately evidenced · strength of the relation: 0.95 · note: Strong flat causal statement. · the relation stated: The cost difference is explicitly driven by spatial discordance.
concept_edges[2]0.99the tagger's words
concept_edgescheapest land —contrasted-with→ land selected for threatened species
the relation itself is separately evidenced · strength of the relation: 0.95 · note: Explicit whereas contrast. · the relation stated: The cheapest remote and arid land is contrasted with tropical forest hotspots selected for threatened species.
concept_edges[3]0.99the tagger's words
concept_edgeswide-ranging species —contrasted-with→ narrowly distributed species
the relation itself is separately evidenced · strength of the relation: 0.95 · note: Explicit cross-group contrast. · the relation stated: Wide-ranging species readily meet ecoregional targets whereas narrow species are left unprotected. Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but the span writes 'species ... with a geographic range size ≥250,000 km 2' and 'their wide distribution', not 'wide-ranging species'.
concept_edges[4]0.99the tagger's words
content_tagsSeventeen percent of threatened vertebrates occur in no protected area and eighty-five percent fall short of adequacy targets.content_tags.findings[0]0.99the tagger's words
content_tagsUpdated data show little global progress relative to the corresponding assessment a decade earlier.content_tags.findings[1]0.99the tagger's words
content_tagsBusiness-as-usual national expansion would cost US$0.9 billion once and US$4.9 billion annually.content_tags.findings[2]0.99the tagger's words
content_tagsThat cheapest expansion would bring only 852 threatened vertebrates to adequate coverage, 249 more than current protection.content_tags.findings[3]0.99the tagger's words
content_tagsExpanding area targets beyond seventeen percent would sharply raise costs without cost-effective threatened-species coverage.content_tags.findings[4]0.98the tagger's words
content_tagsEqual ecoregional representation costs 4.5 times the business-as-usual scenario while only marginally improving threatened-species coverage.content_tags.findings[5]0.99the tagger's words
content_tagsNarrowly distributed species remain the most likely to be unprotected under ecoregional representation.content_tags.findings[6]0.98the tagger's words
content_tagsMeeting adequacy targets for all 4,118 threatened vertebrates would cost about US$42.5 billion annually, 7.5 times the cheapest area-only option.content_tags.findings[7]0.99the tagger's words
content_tagsLow concordance between cheap land and threatened-species distributions drives the cost difference.content_tags.findings[8]0.99the tagger's words
content_tagsThe tradeoff between protected-area cost and threatened-vertebrate coverage is nonlinear.content_tags.findings[9]0.99the tagger's words
content_tagsA fivefold gain in species protected over business as usual would raise annual opportunity cost only to US$7.4 billion.content_tags.findings[10]0.99the tagger's words
content_tagsSimulated commission errors reduce species meeting targets by only five percent on average and do not change the nonlinear frontier.content_tags.findings[11]0.98the tagger's words
content_tagsOnly fifteen percent of threatened vertebrates are adequately covered by the existing network.content_tags.findings[12]0.99the tagger's words
content_tagsAt least 5.8 million square kilometres of new protected areas were required for the Aichi area target.content_tags.findings[13]0.99the tagger's words
content_tagsTargeting low-cost areas, even at thirty-percent land coverage or with ecoregional representation, does not incidentally protect threatened species well.content_tags.findings[14]0.98the tagger's words
content_tagsCost-efficient targeting of threatened vertebrates can substantially improve species coverage at modest additional cost.content_tags.findings[15]0.99the tagger's words
content_tagsspecies listed as Critically Endangered, Endangered, or Vulnerable on the IUCN Red List
also in: threatened
content_tags.definitions[0]0.99the tagger's words
content_tagsa target achieved when any part of a species distribution overlaps the protected-area networkcontent_tags.definitions[1]0.99the tagger's words
content_tagsrange-scaled protected-area coverage, from full coverage for narrow ranges to ten percent for very wide rangescontent_tags.definitions[2]0.99the tagger's words
content_tagsestimated gross agricultural rent used as the cost surrogate for new protectioncontent_tags.definitions[3]0.99the tagger's words
content_tagsa thirty-by-thirty-kilometre square pixel used by the optimization
also in: planning unit
content_tags.definitions[4]0.99the tagger's words
content_tagscountry-level coverage targets met at the lowest agricultural opportunity costcontent_tags.definitions[5]0.99the tagger's words
content_tagsmapped presences at locations where a species is actually absentcontent_tags.definitions[6]0.99the tagger's words
content_tagsthe tradeoff curve between the cost of seventeen-percent coverage and threatened-species target attainmentcontent_tags.definitions[7]0.98the tagger's words
content_tags193 parties — parties adopting the Convention on Biological Diversity strategic plancontent_tags.quantities[0]0.99the tagger's words
content_tags13% to 17% by 2020 — planned increase in global terrestrial protected-area coveragecontent_tags.quantities[1]0.99the tagger's words
content_tags4,118 globally threatened — birds, mammals, and amphibians mapped in the analysiscontent_tags.quantities[2]0.99the tagger's words
content_tags500 m — spatial resolution used for overlayscontent_tags.quantities[3]0.99the tagger's words
content_tags30 km×30 km pixels — aggregation and planning-unit resolutioncontent_tags.quantities[4]0.99the tagger's words
content_tags135,062 protected areas — features in the final protected-area layercontent_tags.quantities[5]0.99the tagger's words
content_tags17,026,214 km² — total area of the final protected-area layercontent_tags.quantities[6]0.99the tagger's words
content_tags12.9% — share of non-Antarctic land in the final layercontent_tags.quantities[7]0.99the tagger's words
content_tags1,135 — threatened bird speciescontent_tags.quantities[8]0.99the tagger's words
content_tags1,107 — threatened mammal speciescontent_tags.quantities[9]0.99the tagger's words
content_tags1,876 — threatened amphibian speciescontent_tags.quantities[10]0.99the tagger's words
content_tagsUS$100 per km² — fixed transaction cost assigned to new protected areascontent_tags.quantities[11]0.99the tagger's words
content_tags∼14% — estimated protected-area management cost relative to agricultural opportunity costcontent_tags.quantities[12]0.99the tagger's words
content_tags<1,000 km² — range threshold receiving complete protected-area coverage targetcontent_tags.quantities[13]0.99the tagger's words
content_tags100% — adequacy coverage target for narrow-ranged speciescontent_tags.quantities[14]0.99the tagger's words
content_tags>250,000 km² — wide-ranging species thresholdcontent_tags.quantities[15]0.99the tagger's words
content_tags10% coverage — adequacy target for wide-ranging speciescontent_tags.quantities[16]0.99the tagger's words
content_tags10 runs of 1 billion iterations each — Marxan search effort per scenariocontent_tags.quantities[17]0.99the tagger's words
content_tags821 terrestrial ecoregions — ecoregions constrained to receive target protectioncontent_tags.quantities[18]0.99the tagger's words
content_tags17% — minimum protection assigned to each terrestrial ecoregioncontent_tags.quantities[19]0.99the tagger's words
content_tags100 range maps — commission-error simulations for each threatened speciescontent_tags.quantities[20]0.99the tagger's words
content_tags1,063 mammals — taxa with high-resolution suitable-habitat sensitivity mapscontent_tags.quantities[21]0.99the tagger's words
content_tags17% — threatened vertebrates absent from all protected areascontent_tags.quantities[22]0.99the tagger's words
content_tags85% — threatened vertebrates below adequacy targetscontent_tags.quantities[23]0.99the tagger's words
content_tagsUS$0.9 billion — one-time transaction cost of business-as-usual expansioncontent_tags.quantities[24]0.99the tagger's words
content_tags$4.9 billion — annual agricultural opportunity cost of business-as-usual expansioncontent_tags.quantities[25]0.99the tagger's words
content_tags852 (21%) — threatened vertebrates adequately covered in business as usualcontent_tags.quantities[26]0.99the tagger's words
content_tags249 species — increment adequately covered over the existing networkcontent_tags.quantities[27]0.99the tagger's words
content_tags4.5 times higher — ecoregional scenario cost multipliercontent_tags.quantities[28]0.99the tagger's words
content_tags$24.8 billion annually — ecoregional scenario opportunity costcontent_tags.quantities[29]0.99the tagger's words
content_tags$42.5 billion annually — annual cost of all-species adequacycontent_tags.quantities[30]0.99the tagger's words
content_tags7.5 times — all-species scenario multiplier over the cheapest optioncontent_tags.quantities[31]0.99the tagger's words
content_tags$7.4 billion annually — cost of a fivefold species-coverage increasecontent_tags.quantities[32]0.99the tagger's words
content_tags1.5 times as much — fivefold-improvement cost relative to business as usualcontent_tags.quantities[33]0.99the tagger's words
content_tags5% — average target-attainment drop under simulated commission errorscontent_tags.quantities[34]0.99the tagger's words
content_tags15% — threatened vertebrates adequately covered by existing areascontent_tags.quantities[35]0.99the tagger's words
content_tags5.8 million km² — new protected area required to meet the Aichi targetcontent_tags.quantities[36]0.99the tagger's words
content_tagsprotected area coveragecontent_tags.variables[0]0.98the tagger's words
content_tagsdistribution of threatened speciescontent_tags.variables[1]0.98the tagger's words
content_tagsagricultural opportunity costcontent_tags.variables[2]0.98the tagger's words
content_tagsgeographic range sizecontent_tags.variables[3]0.98the tagger's words
content_tagsconservation targetscontent_tags.variables[4]0.98the tagger's words
content_tagsconservation costscontent_tags.variables[5]0.98the tagger's words
content_tagsnational protected area targetcontent_tags.variables[6]0.98the tagger's words
content_tagsspecies adequacy targetscontent_tags.variables[8]0.98the tagger's words
content_tagscommission error ratescontent_tags.variables[9]0.98the tagger's words
content_tagsmanagement costscontent_tags.variables[10]0.98the tagger's words
content_tagsconnectivity
also in: connectivity
content_tags.variables[11]0.98the tagger's words

Interpretive choices

Tags that could not be placed in the text:

What was deliberately not tagged, and why, is recorded per paper by the tagger. Where a paper has no such record, this page says so rather than implying that nothing was declined.

© 2026 Shir Sivroni · ORCID 0009-0000-2597-2824