Ecology

Model substancesWe used the sulfoximine insecticide sulfoxaflor, the methoxy-acrylate fungicide Amistar (azoxystrobin 250 g/l, Suspension Concentrate, see supplementary methods, S1) and the glycine herbicide glyphosate (as active substance, RoundUp ProActive or RoundUp FL, see supplementary methods, S1) as model substances. Our choice was justified by their widespread use, regulatory status and systemic uptake in plants. Because of these characteristics, the likelihood of bees being exposed in the field was considered similarly plausible across model substances. Additionally, we are not aware of published evidence of the acute toxicity of these substances across castes and sexes of B. terrestris and O. bicornis.Sulfoxaflor is a relatively novel insecticide55,56,57, developed to replace or complement the use of older chemical classes against which insect pest populations had developed resistance57. However, because of its risks to bees58, its uses have been recently restricted in the EU to indoor growing conditions. As a nicotinic acetylcholine receptor (nAChR) competitive modulator, sulfoxaflor targets the same neural receptor as the bee-harming neonicotinoid insecticides55,56,57. Despite evidence that sulfoxaflor may target the nAChR in a distinct way compared to recently banned neonicotinoids55,56,57, these substances were shown to be similarly lethal in acute exposure laboratory settings for individuals of Apis mellifera, B. terrestris and O. bicornis38. Additionally, sulfoxaflor was shown to reduce reproduction59,60,61 (but not learning62,63) in bumble bees under field-realistic laboratory settings. When applied pre-flowering in a semi-field study design, sulfoxaflor impacted colony growth, colony size and foraging in bumble bees64 but not honey bees65. Azoxystrobin is a broad-spectrum, systemic fungicide, which has been widely used in agriculture since its first marketing authorisation in 199666. Azoxystrobin shows low acute toxicity to honey bees67. Azoxystrobin residues were found in nectar and pollen from treated crops68,69 and subsequently in the bodies of wild bees70. In a semi-field experimental setting, foraging, but not colony growth, was significantly impaired in B. terrestris exposed to Amistar (azoxystrobin 250 g/L SC)64, while no lethal or sublethal effects could be observed in honey bees65 or in O. bicornis71. However, a recent study showed that, when formulated as Amistar this pesticide induced acute mortality in bumble bees at high doses, which was attributed to the dietary toxicity of the co-formulant C16-18 alcohol ethoxylates50.Glyphosate is a broad-spectrum systemic herbicide and the most widely used pesticide in the world72. Products containing glyphosate may be applied to flowering weeds73 and contaminate their pollen and nectar54, thus driving bee contact and oral exposure. Glyphosate showed low lethal hazards in regulatory-ready laboratory74 and semi-field designs when dosed as pure active substance or as MON 52276 (SL formulation containing 360 g glyphosate/L)75. A recent study found ready-to-use consumer products containing glyphosate to be lethally hazardous to bumble bees73. However, this toxicity was attributed to co-formulants, rather than the active substance itself.We characterised the acute oral and contact toxicity to B. terrestris and O. bicornis of sulfoxaflor, azoxystrobin and glyphosate as either pure active substances or formulation (see supplementary material S2 Table S1). Each test was repeated across castes and sexes of these two species. For bumble bees we used workers, males and gynes (i.e., unmated queens), hereby referred to as queens, whereas for O. bicornis we used males and females. Bumble bee experiments were designed following OECD protocols30,31, while O. bicornis was tested following published76 and ring-tested protocols32, as an OECD protocol for this latter species is not yet available.We used a dose response design whenever the test item was found to drive significant mortality in the tested species. In all other cases, a limit test design using a single, high pesticide dose was used. Details on the methods and results of the limit tests are reported in the supplementary materials (S2 and S4).Pesticide treatmentsAll dose response tests were performed with pure sulfoxaflor, while azoxystrobin was tested as a plant protection product (Amistar 250 g a.s./l, SC, Syngenta, UK) in all oral tests, as its solubility in water was insufficient (6.7 mg a.s./L, see EFSA, 2010) to achieve the desired concentrations. Amistar contains co-formulants with hazard classification (54 C16-18 alcohols, ethoxylated  More

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The interaction between the two factors (substrates and CRF rates) was statistically significant by the F-test (p  More

Deforestation trajectories and economic driversCambodia has undergone significant forest loss in recent decades—with 2.6 million hectares of forest cover loss occurring since 2001, equating to 29.5% of forest cover7 and 1.45 billion tonnes of CO2 emissions8. The deforestation rates have increased by 76% in the last decade (2011–2021) compared to the previous (2001–2010; Fig. 1b)7. We find forest loss has occurred within three distinct Phases demonstrated by changepoint analysis: (1) Phase 1: steady rise from 2000 to 2009 (average = 0.82%/year), (2) Phase 2: peak years from 2010 to 2013 (average = 2.3%/year), (3) Phase 3: moderate phase from 2014 to 2021 (average = 1.6%/year). Whilst the annual rate of deforestation has declined since the Phase 2, Cambodia currently has the highest country-level annual rate of forest loss globally7, illustrating the relentless deforestation spreading across the landscape. Critically, much of this forest loss and degradation is occurring in mature primary forests (Fig. 1b), which hold significant carbon and are home to rich biodiversity and keystone species17,18,19.
This deforestation in Cambodia has been attributed to the widespread development of Economic Land Concessions (ELCs), the expansion of numerous agricultural frontiers and relentless illegal logging20,21,22. These drivers have been abetted by the establishment of an extensive national road network (Fig. 1a)20—developed to promote economic growth and urban–rural connectivity23. The majority (88.4%) of these roads have been funded by foreign governments (the People’s Republic of China: 38.5%, Japan: 37.9%, and Republic of Korea: 12.0%)18—all of whom have established land concessions within Cambodia’s borders24 through the allocation of state land into private land for long-term industrial plantations22,25. The expansion of ELCs across Cambodia (average addition of 105,000 ha/year of ELC land since 1998) has been directly attributed to up to 40% of the country’s deforestation21, with further indirect impacts due to encroachment into rural community lands (indigenous areas, community forests, subsistence agricultural fields). This results in landlessness, poverty, and land conflicts, forcing communities to migrate in search of arable land, further contributing to the growing degradation and destruction of forests22,26,27,28,29.Strategic government interventionProtected areas expanded across Cambodia in 1993 following a royal decree26; the legal details of which were delineated in the 2008 Protected Areas Law, introducing protected categories, wildlife corridors and strict laws prohibiting development9. While over 80 protected areas currently exist covering 35% of Cambodian land10, they are still under substantial threat30. In further efforts to curb deforestation, the Royal Government of Cambodia ordered the suspension of new ELCs and revocation of a subset of existing ELCs in 2012 (Order 01BB)31. This resulted in a reduction of ELCs from a peak of ~ 2.1 million ha in 2012 to ~ 1.6 million ha from 2014 onward (Fig. 1b), with a significant positive correlation between the quantity of land classified as ELCs and the country-level deforestation rate (R = 0.87, p  More

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