Insect Threats and Conservation Through the Lens of Global Experts

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1 Insect threats and conservation through the lens of global experts 2 3 Short title: Insect threats and conservation 4 5 Marija Miličić1,2*, Snežana Popov3, Vasco Veiga Branco2, Pedro Cardoso2 6 7 1BioSense Institute – Research Institute for Information Technologies in Biosystems, University 8 of Novi Sad, Novi Sad, Serbia 9 2Laboratory for Integrative Biodiversity Research (LIBRe), Finnish Museum of Natural History 10 Luomus, University of Helsinki, Helsinki, Finland 11 3Department of Biology and Ecology, Faculty of Sciences, University of Novi Sad, Novi Sad, 12 Serbia 13 14 Emails: [email protected]; [email protected]; [email protected]; 15 [email protected] 16 17 Corresponding author* Marija Miličić, email: [email protected], 18 telephone: +38164/48-49-13 19 20 Abstract: While many recent studies have focused on global insect population trends, all are 21 limited either in space or taxonomic scope. Since global monitoring programs for insects are not 22 implemented, biased data are therefore the norm. However, expert opinion is both valuable and 23 widely available, and should be fully exploited when hard data are not available. Our aim is to 24 use global expert opinion to provide insights on the root causes of potential insect declines 25 worldwide, as well as on effective conservation strategies that could mitigate insect biodiversity 26 loss. We obtained 753 responses from 413 respondents with a wide variety of expertise. The 27 most relevant threats identified through the survey were agriculture and climate change, followed 28 by pollution, while land management and land protection were recognized as the most significant 29 conservation measures. Nevertheless, there were differences across regions and insect groups, 30 reflecting the variability within the most diverse class of living organisms on our planet. Lack of 31 answers for certain biogeographic regions or taxa also reflects the need for research, particularly 32 in less investigated settings. Our results provide a first step towards understanding global threats 33 and conservation measures for insects.

34 Key words: agriculture, biodiversity decline, climate change, conservation measures, ecosystem 35 services, expert opinion, extinction, land management, pollution, survey.

36 Introduction

37 Insects play a key role in providing numerous irreplaceable services, many of which are critical 38 to human survival and wellbeing. Yet, most insects are non-charismatic at best and perceived as

39 pests at worst. As such, they receive little attention, attracting few resources for monitoring and 40 conservation (Krause & Robinson, 2017). Likewise, recent studies on animal biodiversity 41 confirm an under-representation of insects in the published literature (Titley, Snaddon, & Turner, 42 2017). Nevertheless, declines have not only been observed in relation to insect diversity and 43 abundance (Biesmeijer et al., 2006; Shortall et al., 2009), but also regarding total insect biomass. 44 For example, one pioneer study in Germany showed a dramatic 75% decline in total flying insect 45 biomass, regardless of habitat type (Hallmann et al., 2017). However, all studies quantifying and 46 assessing long-term global trends in insect abundance or diversity to date (e.g. van Klink et al., 47 2020) are biased, both spatially and taxonomically, often leading to misinterpretation (Simmons 48 et al., 2019; Didham et al., 2020; Montgomery et al., 2020).

49 As there is uneven coverage of research across taxa, there is also unequal spatial distribution of 50 studies. The highest density of biodiversity research occurs in temperate climates, markedly in 51 Western Europe (Titley et al., 2017). Bearing in mind that an estimated 85% of insect species are 52 found in the tropics and south temperate regions (Stork, 2018), under-representation of 53 biodiversity and conservation studies in these regions could lead to false notions about the state 54 of insect diversity, threats and their global spatial distribution. Hence, there is an urgent need to 55 know the status of insects across the globe.

56 Global monitoring programs are not implemented, and unbiased data are not available (Cardoso 57 & Leather, 2019). Thus, we must look for alternative strategies to advance our knowledge on 58 insects, namely expert opinion. It frequently plays a fundamental role in conservation, providing 59 substantive information for decision making processes (Cook, Hockings, & Carter, 2010), 60 horizon scanning (Sutherland, Pullin, Dolman, & Knight, 2004) and risk assessments (Patterson, 61 Meek, Strawson, & Liteplo, 2007). Moreover, International Union for the Conservation of 62 Nature (IUCN) species assessments, particularly regarding insects, almost invariably depend on 63 expert scientific judgments as one of the sources of information that is used to measure our 64 progress towards global biodiversity conservation goals.

65 Our research focuses on the latest insights from experts studying a wide variety of insects around 66 the world. The purpose of this work is to identify the root causes of potential insect declines 67 across different biogeographical regions and taxa based on expert opinion. In addition, expert 68 opinion is also used to identify effective conservation strategies that could mitigate insect 69 biodiversity loss at a global level.

70 Material and methods

71 Our query had 16 questions in six main sections, pertaining to demography, expertise, ecosystem 72 (dis)services, threats, conservation measures and supporting information (Supplementary 73 Material). The first set of questions (1–4) aimed to provide information about the demographic 74 structure of the respondents (gender, experience, education and work responsibilities). Questions 75 5–7 were intended to reveal their biogeographic and taxonomic expertise, which was used to

76 sub-set the data. In the case of taxonomic groups, we used order-level. The only exception was 77 for ants, which were analysed as a separate group since we considered them to be a distinct 78 entity. Compared to other Hymenoptera, they provide very different services and suffer from 79 different threats. If the respondents’ expertise covered several biogeographic regions or 80 taxonomic groups, they could either select multiple answers if threats and conservation measures 81 were deemed similar; if answers differed between regions or taxonomic groups, they were 82 encouraged to fill the survey multiple times, one for each region or taxon. Questions 8–9 were 83 related with services and disservices provided by insects. Questions 10–12 focused on the threats 84 to insect populations and on their current trends. Questions 13–14 addressed the current and 85 potential conservation measures focused towards insects, including allocating potential funds for 86 future projects. Finally, questions 15 and 16 were intended to provide references that support the 87 previous answers and leave any additional comments.

88 We created a Google form query with all questions. To ensure geographical coverage, we 89 contacted around 100 entomological societies globally, and asked them to distribute the query 90 among their members. We also contacted approximately 3000 corresponding authors from 91 papers containing the word “insects”, published in international journals in the last 10 years.

92 To identify the possible influence of demographic structure of the respondents, biogeographic 93 regions and taxa on the answers provided, we performed PerMANOVA with 99999 permutations 94 within the R package vegan (Oksanen et al., 2019), using all answers as response variables for 95 the three analyses. In cases where respondents answered for multiple biogeographic regions or 96 insect taxa, the answers were analyzed separately within each of the selected regions or orders. 97 To avoid dominance in answers coming from the same respondent, we applied a weighted 98 average, where the weight of each answer was divided by the number of queries that respondent 99 had filled (for multiple regions or taxa). A weighted average was also used to overcome possible 100 unbalances in the number of answers for different regions or taxa. Answers to questions 101 regarding threats and conservation of insects ranging from 1–5 were rescaled from 0–1 (1-not 102 relevant at all, 2-little relevance, 3-somewhat relevant, 4-very relevant, 5-extremely relevant). 103 The relevance score average was bootstrapped (random selection of elements with repetition) to 104 obtain upper and lower confidence limits. The procedure was conducted for answers from all 105 regions and taxa together for a global analysis, and then separately for all eight biogeographic 106 regions and all taxa for which more than 10 answers were available (the remaining taxa were 107 grouped under “Other”).

108 Results

109 We obtained 439 responses. Several doubtful answers were eliminated, which included selection 110 of only “unknown relevance” options in the entire survey, selection of all regions and all orders, 111 or the indication that answers relate to non-insects (spiders). In total, the final data set contained

112 429 responses from 413 participants. When responses that contained selections of multiple 113 regions or taxa were fragmented, we obtained a total of 753 answers.

114 Demographic structure of the respondents

115 Participants in the survey were predominantly male (298), researchers (359), with more than 10 116 years of experience (291) and with a PhD (318) (Table S1, Supplementary material). The 117 PerMANOVA analysis showed that in most cases, respondent demography did not influence 118 their answers regarding threats and conservation of insects (Table S2, Supplementary material). 119 However, there was within-group variation in answers based on respondents’ education. To 120 account for this, all consecutive analyses were conducted in two ways: 1) analyzing all answers 121 together, and 2) analyzing answers from respondents holding a PhD degree vs. answers from all 122 respondents with other levels of education. Separate analyses are presented in Supplementary 123 material (Figures S1 to S8).

124 Answers per region and taxon

125 Expertise on the Western Palearctic (Figure 1) and on the orders Coleoptera, Lepidoptera and 126 Diptera were dominant (Figure 2). The PerMANOVA analysis showed that there were 127 statistically significant differences in answers based on both biogeographic region and taxon 128 (Table S3, Supplementary material).

129

130 Figure 1. Distribution of answers (with number of respondents) per biogeographical region.

131

132 Figure 2. Distribution of answers (with number of respondents) per insect order.

133 Insect threats

134 At a global level, the most relevant threats for insects identified by the survey were agriculture 135 and climate change, followed by pollution, significantly different from the first two (Figure 3). 136 Natural system modifications, invasive species and residential and commercial development 137 were also identified as important, without statistical difference. The Afrotropical, Neotropical, 138 Western Palearctic and Nearctic followed the global pattern regarding the three top-rated threats. 139 In the Eastern Palearctic, Indomalayan, Australasian and Oceanian regions, residential and 140 commercial development was among the top-rated threats. The most noticeable difference was in 141 Oceania, where co-extinctions were listed among the most significant threats, next to those 142 coinciding with the global pattern (Figure 3).

143

144 Figure3. Global (upper left) and biogeographic section-based significance of most relevant treats 145 for insects. Confidence limits were calculated by bootstrap.

146 In most cases, analyses per insect order revealed that the trends followed the global pattern, with 147 agriculture, climate change and pollution being identified as the most important threats for most 148 orders (Figure 4). For the Ephemeroptera, Plecoptera and Trichoptera (EPT), however, natural 149 system modifications were selected as the most relevant threat. These were also among the most 150 significant responses for Odonata. For ants, in addition to agriculture and natural system 151 modifications, invasive species were among the top-rated threats, not being statistically different 152 from the previous.

153 154 Figure 4.Significance of threatsfor insects by order. Confidence limits were calculated by 155 bootstrap.

156 There was a perceived decreasing trend in almost all parameters, excluding phylogenetic and 157 functional diversities, for which the trends appeared to be stable or are mostly unknown (Figure 158 5). It is also worth mentioning that, except in the cases of species richness and abundance, 159 around one third of the respondents marked “unknown trend” for the different measures.

160

161 Figure 5. Trends of insect populations. Values are represented in percentages.

162 Insect conservation

163 The conservation measures considered most (and equally) relevant for the global preservation of 164 insects were land management and land protection (Figure 6). In most cases, answers per 165 biogeographic region followed the global trend, with slight but non-significant variations 166 between land protection and land management identified as the most important. However, in the 167 case of the Nearctic, Indomalayan and Eastern Palearctic regions, education was listed among 168 the most relevant. In the Nearctic region, law and policy was also recognized among the most 169 relevant conservation measures (Figure 6).

170

171 Figure 6. Global (upper left) and biogeographic section-based significance of conservation 172 measures for the preservation of insects. Confidence limits were calculated by bootstrap.

173 There were some differences in answers regarding the most relevant conservation measures 174 between different insect groups (Figure 7). In Hymenoptera, Hemiptera and Diptera, education 175 was identified among the most relevant conservation measures, next to land protection and land 176 management. In Diptera and Hemiptera, law and policy were also marked as being highly 177 important conservation measures.

178

179 Figure 7.Significance of conservation measures for the preservation of insects by order. 180 Confidence limits were calculated by bootstrap.

181 Regarding the allocation of the conservation investments, respondents would primarily dedicate 182 funds for research of insect biodiversity, followed by monitoring of biodiversity and acquisition 183 of new protected areas. Management of protected areas, education and management of 184 unprotected areas would follow, while the smallest amount of funds would be allocated to ex-situ 185 conservation of insects (Figure 8).

186

187 Figure 8. Allocation of funds for conservation investments.

188 Insects’ services and disservices

189 Respondents of the query considered that the most relevant services provided by insects are 190 within provisioning services, namely monitoring of habitat quality and biocontrol (Figure 9). 191 Among regulating services, pollination was depicted as the most significant, while nutrient 192 cycling through saprophagy and coprophagy were the most relevant supporting services. Serving 193 as models for scientific research was the most significant cultural service provided by insects. 194 Pest damage to agriculture and acting as invasive species were selected as main disservices of 195 insects (Figure 10).

196

197 Figure 9. Services provided by insects.

198

199 Figure10. Disservices provided by insects.

200

201

202 Discussion

203 Agriculture (including livestock farming) and climate change (including extreme weather events) 204 were consistently identified as the main threats for insects, followed by pollution and natural 205 system modifications. Land management and protection were assigned as the most significant 206 measures that could aid in insect preservation globally. However, there was variation in both the 207 threats and conservation measures identified as the most relevant across different biogeographic 208 regions and taxonomic groups. This suggests that each case must be studied individually when 209 addressing threats and designing conservation actions, depending on the target area or taxonomic 210 group of species.

211 Threats

212 When agriculture and insects are put in the same sentence, one would mainly think of insects as 213 pests, causing economic damage. However, different agricultural activities have proven to be a 214 serious threat not only for insects (Showket, Mir, Parry, Yaqob, & Nisar, 2017), but also to 215 biodiversity worldwide. According to the ESA (Earth Space Agency), about 37% of the entire 216 Earth’s land is used for agricultural purposes; roughly 11% of which used for growing crops, and 217 the remainder for pasture. This type of land transformation leads to habitat loss and biotic 218 homogenization (Olden & Rooney, 2006), both of which subsequently drive the loss of species 219 and facilitate the process of ecological drift, i.e. the replacement of specialists by generalists in 220 species assemblages, as shown by Polus, Vandewoestijne, Choutt, & Baguette (2007) with 221 butterflies in Belgium. Likewise, biotic homogenization through the spread of wood plantations 222 has proven to have negative effects on both taxonomic and functional diversity of Odonata 223 studied in the Neotropics (Dalzochio, Périco, Renner, & Sahlén, 2018) and in the Indo-Malayan 224 region (Dolný, Harabiš, Bárta, Lhota, & Drozd, 2012), which is in accordance with our results. 225 Grazing, as a dominant form of agricultural activity (based on the percentages mentioned above), 226 influences insects in several ways. Primarily, it causes alterations to the plant communities in a 227 system (Börschig, Klein, von Wehrden, & Krauss, 2013), thus causing a cascade effect on many 228 insect populations. It also modifies physical properties of soil and induces changes in 229 microclimate. These effects have proven to promote destruction of nesting sites and removal of 230 potential insect foraging and/or sheltering habitats, as shown for grasshoppers (Orthoptera) 231 (Black, Shepherd, & Vaughan, 2011). Studies in the Neotropical region have highlighted the 232 effect of cattle ranching on the homogenization of the landscape (Kappelle & Brown, 2001), 233 which created more severe environmental conditions through opening of drier open areas. 234 Nichols et al. (2007) relate these conditions to the modified assembly structure of dung beetles 235 (Coleoptera), while Ma et al. (2017) associate the decrease in abundances of two Orthopteran 236 and Hemipteran families in Eastern Palearctic with overgrazing. Arid ecosystems, such as those 237 in the Mediterranean zone and in the Afrotropical region are facing the risk of desertification due 238 to overgrazing, driving semi-arid ecosystems towards desert-like landscapes (Louis, 2016).

239 Our results on climate change confirm the global concern about the possible consequences of 240 global warming, not only in the scientific literature, but also in mass media (Legagneux et al., 241 2018). Due to the popularity of the topic, it is not surprising that much research is focused on 242 studying the effects of climate change on living organisms. The most significant effect of climate 243 change on species is the loss of suitable conditions for their survival in a particular ecosystem. 244 We expect that global warming will force species distributions towards “colder” areas at higher 245 elevations and latitudes. Range shifts are documented, for example, in two North American bark 246 beetles (Williams & Liebhold, 2002). Besides range shifts, climate change can cause insects to 247 alter their phenology in order to adapt (Forrest, 2016), either going through evolutionary shifts 248 (Kellermann & van Heerwaarden, 2019) or becoming extinct. However, the biggest concern is 249 that current climate change impacts on biodiversity are several magnitudes lower than those 250 predictable in the future. Although not specifically focused on insects, the work of Kingsford & 251 Watson (2011) and Duffy (2011) address the question of climate change effects in the Oceania 252 region. Duffy (2011) points out the possibility that, due to the all-inclusive consequences of 253 climate change, conservation of biodiversity could be a low priority unless directly related to 254 human needs.

255 Considering the variety of contaminants in the environment that can threaten insects, it is not 256 surprising that pollution was rated as one of the most serious threats among the respondents. 257 However, certain groups of insects might be more prone to specific types of pollution. For 258 example, a significant number of studies have addressed the effect of light pollution on different 259 beetle species. Not only does it influence nocturnal insects such as fireflies (Lampyridae) by 260 affecting their flash signals (Owens, Meyer-Rochow, & Yang, 2018) and intra- and inter-specific 261 interactions (Firebaugh & Haynes, 2019), but it is also related to the reduced occurrence of 262 fireflies in the Neotropics (Hagen, Santos, Schlindwein, & Viviani, 2015). Besides Coleoptera, 263 pollution was depicted as the second most relevant threat for EPT as well. Considering their 264 dependence on water – at least during part of their life cycle – different water effluents possess a 265 severe threat for these communities, as shown in the study of Chi, Hu, Zheng, & Dong (2017), 266 who found the EPT to be particularly sensitive to the effect of arsenic pollution.

267 Natural system modifications were the fourth most relevant threat overall; this was second most 268 relevant for Odonata and the most relevant for the EPT species. These four taxa are driving the 269 overall pattern. As previously mentioned, these insect groups are dependent on water bodies, and 270 are proven to be sensitive to human pressure. Thus, it comes as no surprise that water 271 management and building of dams negatively influences these species. Bredenhand & Samways 272 (2009) found that among all analysed macro-invertebrates in Eerste River, South Africa, the EPT 273 community was the most affected by the existence of a dam, as diversity and abundance of the 274 community dropped to almost zero as a result of the impoundment.

275 In some biogeographic regions or insect groups, certain threats show a contrasting pattern with 276 the global distribution of answers. For example, in the Eastern Palearctic and Indomalayan

277 regions, residential and commercial development was marked as a very significant threat, not 278 significantly lower than the top rated. In the Eastern Palearctic, remarkable economic growth in 279 some regions such as China and Japan, goes hand in hand with infrastructure development, 280 which reflects negatively on biodiversity (Squires, 2014). Investigation of bees in four 281 megacities in Southeast Asia showed that the mean species richness and abundance were 282 significantly higher in the peripheral suburban areas than central business districts (Sing et al., 283 2016), indicating the sensitivity of this insect group towards urbanization, as also reflected in our 284 results. As for the Indomalayan region, Wattanachaiyingcharoen, Nak-eiam, Phanmuangma, 285 Booninkiaew, & Nimlob (2016) highlighted that a low standard of management in the tourist 286 industry in Thailand may cause an accelerated reduction of the firefly fauna in this region. 287 Furthermore, due to recent unplanned developmental activities, some areas in the Indomalayan 288 region have displayed signs of rapid habitat destruction, negatively affecting biodiversity 289 (Bhattacharya, 2019), which might explain why respondents from this region gave increased 290 weight to this threat. Co-extinctions were depicted as a very important threat in Oceania, not 291 significantly lower than climate change and agriculture. This might be due to the insular nature 292 of this region, which supports high levels of localized endemism (Kier et al., 2009). Thus, 293 extinction of host animals or plants for some insect groups would promote the extinction of 294 dependent insects as well. Oceania was also the region where invasive species were given higher 295 weight. Islands are particularly prone to the effects of invasions due to their spatial and temporal 296 isolation (Borges, Rigal, Ros-Prieto, & Cardoso, 2020), with native species often incapable of 297 adapting to new competitor, predator, or parasite species. Invasive species were also depicted as 298 being a relevant threat for ants. The impact of invasive species on indigenous ants in Indonesia 299 was previously documented by Bos, Tylianakis, Steffan-Dewenter, & Tscharntke (2008). A 300 particularly aggressive ant species, the Argentine ant (Linephitema humile) is known to cause 301 major disruptions in native fauna, including outcompeting native ant species (Rowles & 302 O’Dowd, 2007). However, it is worth noting that Oceania was the region with the lowest number 303 of respondents (10). Confidence intervals in this case were proportionally high, so this result 304 should be taken with caution.

305 Due to the extent and severity of threats that insects are facing worldwide (Cardoso et al., 2020; 306 Wagner, 2020), it was somehow expected that respondents would identify a mainly decreasing 307 trend in parameters reflecting the status of insect populations. However, it is noteworthy that in 308 almost one third of the answers, respondents selected unknown trends, which indicates that, even 309 with the recent rise in the number of papers tackling insect declines, the most diverse class of 310 organisms on the planet is still severely understudied, hence a nuanced view of insect declines or 311 increases according to taxon and region is needed (van Klink et al., 2020). Functional and 312 phylogenetic diversity had more neutral answers, which might reflect the fact that unique 313 functions or branches in the tree of life are not necessarily lost at the same pace as other metrics, 314 not necessarily that they are not lost at all.

315

316 Conservation

317 Land management and land protection were recognised as the most relevant conservation 318 measures for the global preservation of insects across regions and taxa. Many regions worldwide 319 have been lagging in the protection of relevant areas for insect conservation (Taylor et al., 2018). 320 Taking a landscape perspective to protection and management of insect biotopes is in fact one of 321 the most effective ways to protect countless taxa, their unique ways of life, evolutionary history 322 and complex networks (Samways et al., 2020). Recent findings show that area size, habitat 323 quality and/or suitability are key points when designating zones that are important for 324 conservation of certain insect species (Janković, Miličić, Ačanski, & Vujić, 2020). Increasing the 325 area and connectivity of protected zones promotes richer insect communities, particularly 326 regarding sensitive orders with specific habitat requirements, such as the EPT (South, DeWalt, & 327 Cao, 2019). Likewise, ant species can be protected most easily by protecting the biotopes in 328 which they occur (Mabelis, 2007). Along with ample land protected, it seems that conservation 329 of insect biodiversity also largely depends on managing sufficient and suitable habitats. For 330 instance, land clearing and removal of over-mature and veteran trees have been identified as 331 threats to some of the orders with many species such as Hymenoptera, Coleoptera, Lepidoptera, 332 and Diptera (Sands, 2018). Therefore, land management practises that avoid harvesting large old 333 growth trees would be of conservation importance for many insect species worldwide (Jörg et al., 334 2016). However, if over-mature trees are present in urban parks, they could be considered as a 335 hazard for public safety and thus get cut by management authorities (Carpaneto, Mazziotta, 336 Coletti, Luiselli, & Audisio, 2010). This is precisely why conservation of insects is not restricted 337 to protected area managers only – tackling this complex challenge requires involvement and 338 collaboration of diverse stakeholder groups, including the general public.

339 One way of gaining an appreciation of insect diversity is to actively involve citizens in scientific 340 projects. Citizen science projects could greatly increase community-based conservation 341 outcomes, providing data with clear conservation relevance (Oberhauser & Prysby, 2008), as 342 well as favouring connection between citizens and scientists. Acquisition of scientific knowledge 343 through environment-based education programs can be especially powerful in increasing 344 biodiversity awareness (Caro, Mulder, & Moore, 2003), which was also recognised by the 345 respondents highlighting education as a conservation measure of a high relevance. This is of 346 particular importance if focused on early childhood experiences, which are one of the drivers of 347 positive or negative human-insect encounters (Nxumalo & Pacini-Ketchabaw, 2017). 348 Interestingly, education is listed as the second most important conservation measure for the 349 Indomalayan and Eastern Palearctic regions, before land protection (albeit not significantly). 350 Insufficient research attention in these regions when it comes to insects leaves a mark on 351 environmental education activities in the region. For example, as noticed by Corlett (2011), 352 information about the Indomalayan fauna is politically and linguistically fragmented among 353 numerous publications. Often it is not easy to obtain information from conservation literature, as 354 it is more focused on vertebrate species while insects are clearly under-represented (Di Marco et

355 al., 2017). Next to land management and land protection, education and education campaigns as 356 a tool for conservation were particularly recognized by those who responded to surveys on the 357 orders Hemiptera and Diptera. Among Hemiptera, species belonging to the families Coccidae, 358 Diaspididae and Pseudococcidae are known as the most serious pests worldwide (Evans & 359 Dooley, 2013).Certain insect groups like pests may be excluded in insect conservation strategies, 360 reflecting both taxonomic and biological ignorance regarding the importance of these insects. 361 Deeming an insect as a “pest” is simply based on social perception and essentially ignores the 362 key ecological roles played by these species (Delibes-Mateos, Smith, Slobodchikoff, & 363 Swenson, 2011). Similarly, the association of many Diptera species with coprophagy (Khofar, 364 Kurahashi, Zainal, Isa, & Heo, 2019), saprophagy, or disease transmission (Sarwar, 2016) makes 365 their role in nutrient recycling, pollination and biological control often overlooked. It seems that 366 ignorance could be listed as one of the leading threats when speaking about insect species. Thus, 367 dealing with large gaps in taxonomical, biological and ecological knowledge of insects should be 368 of high priority on all educational levels.

369 According to Samways (2018), currently insect conservation has a foothold in six interconnected 370 themes: philosophy, research, psychology, practice, validation and policy, the latter of which 371 makes the framework for action. The respondents recognized law and policy as a significant 372 conservation measure in all regions and for all taxa. As an example, protection of insect species 373 at risk, as well as conservation of their natural habitats, was recognized as a strong desire of 374 people in the study of Danks, Smith, Foottit, & Adler (2017). Respondents highlighted the 375 importance of law and policy especially regarding Diptera, Hemiptera and Odonata. Worldwide, 376 dragonflies play a central role in the conservation of freshwater habitats, being used as indicators 377 for environmental health and conservation management (e.g. Chovanec & Waringer, 2005). Yet, 378 a recent study (Kalkman et al., 2018) highlighted a strong mismatch between the distributions of 379 species listed as threatened in the IUCN European Red List and species protected by the EU 380 Habitats Directive. The authors call for re-evaluation of the list of species included in the 381 Directive so it can include Mediterranean threatened species that are neglected under the current 382 policy. The same could be said of other insects and arthropods, with the big, beautiful and 383 common species being privileged in their protection at the European level (Cardoso, Erwin, 384 Borges, & New, 2011) at the expense of the small and “ugly” ones like Diptera and Hemiptera.

385 According to the respondents, the least relevant among all conservation measures was species 386 management. A common challenge in insect species conservation and management is how to 387 manage species of which our knowledge is limited, particularly in relation to life-cycle 388 interventions, which is a pre-requisite for the successful implementation of management actions. 389 Moreover, handling the enormous richness of insect orders makes it difficult to manage species 390 individually. Nevertheless, species management may be a tool to be explored further in insect 391 conservation.

392 Regarding the allocation of conservation investments, the respondents indicated that they would 393 primarily fund research and monitoring of insect biodiversity. Incorporating insects into 394 protected area monitoring activities is possible only with well-documented species inventories 395 (McGeoch et al., 2011), which brings us back to the recognized gaps in taxonomical, biological 396 and ecological knowledge of insects and the need to build unbiased monitoring programs at a 397 global scale (Cardoso & Leather, 2019). Research studies conducted on a variety of spatial and 398 temporal scales are of crucial importance to gather the data needed to inform insect conservation 399 policy and management and, thus properly address the drivers of insect declines. This is the only 400 way to stop insect declines in view of necessarily limited resources for conservation of countless 401 species, described and undescribed.

402 Services and disservices

403 As insects play a key role in various ecosystem services, further research should also be devoted 404 to service quantification. A recent study shows that the most studied ecosystem services provided 405 by insects are pollination, biological control, food provisioning and recycling organic matter 406 (Noriega et al., 2018). Indeed, the respondents selected biocontrol and monitoring of habitat 407 quality as the most relevant services among provisioning services. Among the regulating and 408 supporting services, pollination and nutrient cycling through saprophagy and coprophagy were 409 selected as the most significant. Interestingly, the respondents recognized serving as models for 410 scientific research as the most significant cultural service provided by insects, which is proved 411 by many recent studies worldwide, including evolutionary (Wang et al., 2016), toxicology 412 (Coates et al., 2019), biomonitoring (Miguel, Oliveira-Junior, Ligeiro, & Juen, 2017), robotics 413 (Mintchev, de Rivaz, & Floreano, 2017) and many other studies in different areas. As for 414 disservices, pest damage to agriculture and acting as invasive species were listed as the most 415 significant. The flows of services and disservices, at least in agricultural ecosystems, directly 416 depend on how ecosystems are managed at different landscape scales (Zhang, Ricketts, Kremen, 417 Carney, & Swinton, 2007). Indeed, this should be definitely taken into account when planning 418 policy-relevant entomological research that can influence agricultural practices. The 419 identification of potentially high-risk invasive pest species is the first step in prioritizing species 420 as candidates for further research (Worner & Gevrey, 2006).

421 Conclusion

422 Our study shows that experts consider agriculture, climate change and pollution as the three main 423 drivers for insect declines across different biogeographic regions and orders. Land protection and 424 land management are identified as the most prominent conservation measures that might aid in 425 the preservation of insects and mitigation of negative consequences worldwide. A decreasing 426 trend in parameters reflecting the status of insect populations, as well as the extreme significance 427 of insects, as reflected in the numerous ecosystem services they provide, calls for global 428 monitoring programs and a specialized approach in conservation for each particular case.

429 Acknowledgements

430 We thank Marina Szulc for the help and advice regarding data analysis and visualization. We 431 give attribution to George Starr (Ephemeroptera, link to license 432 https://creativecommons.org/licenses/by-nc-sa/3.0/), Maxime Dahirel (Odonata, 433 https://creativecommons.org/licenses/by/3.0/), Melissa Broussard (Orthoptera, 434 https://creativecommons.org/licenses/by/3.0/), Nico Muñoz (Pleoptera, 435 https://creativecommons.org/licenses/by-nc/3.0/), Didier Descouens (vectorized by T. Michael 436 Keesey) (Trichoptera, https://creativecommons.org/licenses/by-sa/3.0/) for material downloaded 437 from website Phylopic. MM and SP acknowledge financial support of the Ministry of Education, 438 Science and Technological Development of the Republic of Serbia (Grant Nos. 451-03-68/2020- 439 14/ 200358 and 1-0 - 2020-1 / 200125 ) and H2020 Project ANTARES, grant no. 664387. 440 VVB and PC are supported by Kone Foundation. 441 442 Confilct of interest 443 444 The authors declare no conflict of interest. 445 446 References 447 448 1. Bhattacharya, S. (2019). Environmental Crisis in the Eastern Himalayan Landscapes in 449 India. Consilience, 21(1), 66–85. doi: 10.7916/consilience.v0i21.5729 450 2. Biesmeijer, J. C., Roberts, S. P. M., Reemer, M., Ohlemueller, R., Edwards, M., Peeters, 451 ... Kunin, W.E. (2006). Parallel declines in pollinators and insect-pollinated plants in 452 Britain and The Netherlands. Science, 313(5785), 351–354. doi: 453 10.1126/science.1127863 454 3. Black, S. H., Shepherd, M., & Vaughan, M. (2011). Rangeland management for 455 pollinators. Society for Rangeland Management. 9–13. doi: 10.2111/1551-501X-33.3.9 456 4. Borges, P.A.V., Rigal, F., Ros-Prieto, A., & Cardoso, P. (2020) Increase of insular exotic 457 arthropod diversity is a fundamental aspect of the current biodiversity crisis. Insect 458 Conservation and Diversity.doi: 10.1111/icad.12431 459 5. Börschig, C., Klein, A. M., von Wehrden, H., & Krauss, J. (2013). Traits of butterfly 460 communities change from specialist to generalist characteristics with increasing land-use 461 intensity. Basic and Applied Ecology, 14(7), 547–554. doi: 10.1016/j.baae.2013.09.002 462 6. Bos, M. M., Tylianakis, J. M., Steffan-Dewenter, I., & Tscharntke, T. (2008). The 463 invasive Yellow Crazy Ant and the decline of forest ant diversity in Indonesian cacao 464 agroforests. Biological Invasions, 10(8), 1399–1409.doi: 10.1007/s10530-008-9215-4 465 7. Bredenhand, E., & Samways, M. J. (2009). Impact of a dam on benthic 466 macroinvertebrates in a small river in a biodiversity hotspot: Cape Floristic Region, 467 South Africa. Journal of Insect Conservation, 13(3), 297–307. doi: 10.1007/s10841-008- 468 9173-2

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