BioInvasions Records (2021) Volume 10, Issue 1: 81–90

CORRECTED PROOF

Rapid Communication First incidence of the invasive fall armyworm, Spodoptera frugiperda (J.E. Smith, 1797) attacking maize in Malaysia

Saiful Zaimi Jamil1,*, Mohd Masri Saranum1, Lailatul Jumaiyah Saleh Hudin2 and Wan Khairul Anuar Wan Ali3 1Agrobiodiversity & Environment Research Centre, Malaysian Agricultural Research and Development Institute (MARDI), 43400 Serdang, Selangor, Malaysia 2Plant Biosecurity Division, Department of Agriculture, 50632 Kuala Lumpur, Malaysia 3Industrial Crop Research Centre, Malaysian Agricultural Research and Development Institute (MARDI), 43400 Serdang, Selangor, Malaysia Author e-mails: [email protected] (SZ), [email protected] (MS), [email protected] (LS), [email protected] (WK) *Corresponding author

Citation: Jamil SZ, Saranum MM, Hudin LJS, Wan Ali WKA (2021) First incidence Abstract of the invasive fall armyworm, Spodoptera frugiperda (J.E. Smith, 1797) attacking The impact of invasive species attacking economically important crops is often maize in Malaysia. BioInvasions Records unpredictable, which is why information needs to be provided at the earliest stage 10(1): 81–90, https://doi.org/10.3391/bir.2021. of the attack. This is an incidence report of the fall armyworm, Spodoptera frugiperda 10.1.10 (J.E. Smith, 1797) (Lepidoptera: Noctuidae), a new invasive species attacking maize Received: 4 June 2020 in Malaysia. We found this devastating insect feeding on maize in Changlun, a Accepted: 22 October 2020 district in the state of Kedah in northern Malaysia. Morphological examination on Published: 31 December 2020 the collected specimens and plant damage symptoms due to larval feeding allowed us to confirm the presence of the pest. Hence, we confirm the presence of Handling editor: Luis Reino S. frugiperda in yet another country of South East Asia. Thematic editor: Angeliki Martinou Copyright: © Jamil et al. Key words: invasive species, morphological identification, damage symptoms, South This is an open access article distributed under terms of the Creative Commons Attribution License East Asia, food security (Attribution 4.0 International - CC BY 4.0).

OPEN ACCESS. Introduction Invasive species, in most cases, pose a direct threat to native ecosystems. Indirectly, these invaders also affect humans through crop loss and food insecurity (Paini et al. 2016). The threat is constantly growing, spearheaded by anthropogenic-induced factors such as global trade and transportation – especially in regions where they were previously absent (Tilman et al. 2017). While the issue itself is apparent, predicting the exact cost of loss is challenging due to geographical, culture, socio-economic and legislative complexity. Hence, it is essential to accumulate and disseminate information rapidly to assist in prevention management decisions as a response to new invasive species incursions (Wildemeersch et al. 2019). The fall armyworm, Spodoptera frugiperda (J.E. Smith, 1797) (Lepidoptera: Noctuidae), is a an invasive species that has caused havoc on commercial crops in large parts of the world. Being first detected in the African continent in 2016 (Goergen et al. 2016), the pest has affected at least 28 sub-Saharan African countries by the end of 2017 (Day et al. 2017). Within a year,

Jamil et al. (2021), BioInvasions Records 10(1): 81–90, https://doi.org/10.3391/bir.2021.10.1.10 81 First incidence of Spodoptera frugiperda in Malaysia

S. frugiperda continued its spread across South Asia (Kalleshwaraswamy et al. 2018). It did not take long for the pest to invade South-East Asia. By 2019, S. frugiperda has been reported to attack maize in Vietnam and Indonesia (Hang et al. 2020; Ginting et al. 2020, respectively). In Malaysia, the detection of this pest was first announced in a brief report by the International Plant Protection Convention (IPPC 2019). The pest has caused considerable yield loss to farmers along their invasive pathway. Losses due to S. frugiperda incursions on maize in Africa were estimated between US$ 2,481 million and US$ 6,187 million per annum, translating to a loss ranging from 8.3 to 20.6 million tonnes annually (Shylesha et al. 2018; Day et al. 2017, respectively). Economic losses on maize in South- East region remain unclear. The Food and Agriculture Organization (FAO) of the United Nations has declared S. frugiperda as one of the major invasive pests in the world, emphasizing that serious attention needs to be placed on S. frugiperda as it may threaten global food security due to its polyphagous nature (FAO 2017). This pest possesses various traits to be a successful invader. Being able to complete multiple generations, an adult female moth may lay up to 1,000 eggs throughout its lifetime (CABI 2019). Once emerged, large numbers of S. frugiperda neonate-larvae, persistent like an army, can cause severe damage to maize especially the leaves. The larvae of S. frugiperda potentially feed on more than 350 plant species, including several economically-important crops such as maize, sugarcane and rice (Montezano et al. 2018). Worst damage is typically done by the fourth to sixth instar larval stages where they inflict injury to the maize whorl, stem and cob through extensive feeding. Apart from the larvae, adults are also strong fliers and may fly over 100 km per night (Johnson 1987). All these features contribute directly to the success of S. frugiperda spreading as a global pest. It is vital to collect detailed information on the means of introduction, distribution, host plant feeding patterns and preferences, and the severity of this pest in order to limit its damage on maize in Malaysia. Herein, we report for the first time the presence of S. frugiperda in Malaysia as a prerequisite to develop appropriate management strategies against this recently introduced invasive species.

Materials and methods Study site Following reports by maize growers of devastating attacks by unknown larvae on their crop, MARDI scientists and the Department of Agriculture extension agents organized a joint field visit to a private farm located in Changlun, Kedah (6.339528N; 100.418987E) (Figure 1). Prior to sampling, farm owners were interviewed using a short, structured questionnaire to obtain general information about farm management practices including

Jamil et al. (2021), BioInvasions Records 10(1): 81–90, https://doi.org/10.3391/bir.2021.10.1.10 82 First incidence of Spodoptera frugiperda in Malaysia

Figure 1. Location of the Changlun maize farm in the northern part of Peninsular Malaysia where S. frugiperda was first located (red dot).

Table 1. Mean ± SE of S. frugiperda incidence and severity in Changlun, Malaysia according to pre-determined zones.

Zone* Incidence of infected plants Mean severity Average number of S. frugiperda larvae per plant (plot size) (%) (Scale 1 to 5) Leaf/Stem Cob Total A (10 acres) 100 4.70 ± 0.07 3.40 ± 0.50 2.20 ± 0.51 5.60 ± 0.69 B (15 acres) 100 5.00 ± 0.00 2.70 ± 0.33 3.30 ± 0.40 6.00 ± 0.42 C (20 acres) 100 4.88 ± 0.05 2.80 ± 0.44 2.40 ± 0.40 5.20 ± 0.66 Total 100 4.86 ± 0.03 2.97 ± 0.25 2.63 ± 0.26 – * n = 50 for each zone

planting date, maize variety, source of seeds, pesticides type and frequency, fertilizer type, first experience of damage and type of recent change in infestation. Grain maize was cultivated, but the exact maize variety was not determined. At the time of sampling, most plants were already at the reproductive development stage. Sampling of the S. frugiperda on the farm was undertaken at three pre-determined zones i.e. Zone A, B and C, which differed in plot size at the time of sampling (Table 1). Sampling was done for three consecutive days from the 16th till the 18th of September 2019. The

Jamil et al. (2021), BioInvasions Records 10(1): 81–90, https://doi.org/10.3391/bir.2021.10.1.10 83 First incidence of Spodoptera frugiperda in Malaysia

Table 2. Severity score description based on damage symptoms on maize plant (adapted from Kuate et al. 2019 with slight modifications).

Severity score Description(s) 1 Healthy maize with NO damage. 1–10% leaf damage or presence of damage from S. frugiperda limited to characteristic windows or < 5 mm 2 diameter and or destruction of only the leaf cuticle. S. frugiperda present on flower and shoot of plant. 11–25% leaf damage with presence of chewed areas > 5 mm, funnel leaves still intact. S. frugiperda present on 3 flower, shoot and mature leaves of plant. 26–50% leaf damage with presence chewed areas larger than 1 cm, the funnel slightly damaged or less severe. 4 S. frugiperda present on flower, shoot, mature leaves and stem of plant. > 50% leaf damage, plant stunting and funnel damaged severely. S. frugiperda present on ALL part of plants 5 including flower, shoot, mature leaves, stem and cob.

first day of sampling it rained however rainfall was not recorded. The survey period corresponded to the shift from dry season to wet season in the northern Peninsular Malaysia.

Data collection Insects were collected simultaneously while incidence scouting was conducted. The S. frugiperda incidence scouting was undertaken according to the method described by FAO (FAO 2018). In each zone, scouting was done by inspecting 10 plants per transect. Four transects were established by moving along a W-shape design. Another final transect was established in the middle of each zone. In total, 50 maize plants were sampled per zone in 5 transects, respectively. Distance between two consecutive plants was a function of field size and shape but was representative of the plot area. Spodoptera frugiperda presence was determined using the following indicators: (i) presence of fresh frass in the leaf funnel; (ii) presence of larvae on leaves or in the leaf funnel identifiable with the inverted Y-Shape in the head and the set of four dot forming a square on the upper surface of the last segment of its body; (iii) irregular damage (cuts) on leaves and (iv) presence of egg masses (Day et al. 2017). Field sampling to determine the pest severity was done destructively. Inspected plants (10 plants/zone) were chosen randomly. Plants were cut from the ground level and dissected to record the number of S. frugiperda, egg batches and any other insects present. Damage to the plant was scored according to the severity scoring with slight modifications (Kuate et al. 2019). A rating scale from 1 to 5 was used for scoring of damage severity on whorl-stage plants (Table 2). Collected larvae (third to sixth instar stage) were kept in 90% alcohol and brought to the laboratory for further identification. Another batch of larvae was collected to establish S. frugiperda colonies in the laboratory. All pictures were photographed using the Olympus Tough TG-5 camera.

Results Morphological identification We obtained a total of 223 S. frugiperda larvae collected from the maize plants during the sampling. Morphological characteristics of the S. frugiperda

Jamil et al. (2021), BioInvasions Records 10(1): 81–90, https://doi.org/10.3391/bir.2021.10.1.10 84 First incidence of Spodoptera frugiperda in Malaysia

Figure 2. Morphological characteristics of Figure 3. Morphological characteristics of S. frugiperda larvae (A) the head part with an S. frugiperda (A) pupae (B) male adult (C) female inverted clear Y-shaped marking on the adult. Photographs by Saiful ZJ. anterior (B) trapezoidal-patterned black spots on the first to seventh abdominal segment (C) squarish-pattern black spots on eighth segment. Photographs by Saiful ZJ.

are presented in Figure 2 and Figure 3. Mature larvae were marked with clear inverted “Y” on their dark-coloured head (Figure 2A). They also consistently exhibit distinct black spots (pinacula) on the body which were arranged in a trapezoidal pattern for the first seven abdominal segments and a rather square pattern on the 8th abdominal segment (Figure 2B, C). These two unique larval characteristics distinguishes this species from other

Jamil et al. (2021), BioInvasions Records 10(1): 81–90, https://doi.org/10.3391/bir.2021.10.1.10 85 First incidence of Spodoptera frugiperda in Malaysia

Figure 4. Damage symptoms by the S. frugiperda on maize (A) consistent damage on almost every plant in the farm (B) holes and windowpanes on leaf (C) damage on the stalk and tassel (D) damage on the whorl (E) frass around the feeding site (F) Asian corn borer, Ostrinia furnacalis feeding on maize. Photographs by Saiful ZJ.

species. The pupa is brightly reddish-brown in colour (Figure 3A). It measures between 13 to 18 mm in length and about 4.3 mm in width. Pupation normally takes place in the soil where the 6th instar larva drops off or travel down from the plant. However, we did discover a few pupae specimens being suspended between the whorl during our sampling. Emerging adults have a wingspan between 30 to 40 mm. The forewings of females (Figure 3C) are dull, greyish brown and less distinctly marked in comparison to forewings of the males (Figure 3B). Male moths on the other hand possess a forewing which is generally brown, with distinct white spots at the tip and centre of the wing (Figure 3B). For both sexes, the hind wing is silvery and iridescent with a thin dark border.

Plant damage symptoms Damage symptoms by the S. frugiperda were apparent on almost every plant at the farm (Figure 4A). Most plant leaves are tattered with small holes and “windowpanes” (Figure 4B). Symptoms of damage due to larvae attack were also obvious on most part of the plant including the stalk, whorl, tassel, ear and cob (Figure 4A–E). Feeding activities were also confirmed by evidence of yellowish to brown frass around the feeding site (Figure 4E). We attribute the damage of the leaves and stalk mainly to the S. frugiperda. However, we did observe the larvae of the common Asian

Jamil et al. (2021), BioInvasions Records 10(1): 81–90, https://doi.org/10.3391/bir.2021.10.1.10 86 First incidence of Spodoptera frugiperda in Malaysia

corn borer, Ostrinia furnacalis (Guenée, 1854) (Lepidoptera: Noctuidae) co-infesting the cobs of maize on several occasions (approximately 13 cases) (Figure 4F).

Spodoptera frugiperda incidence and severity Spodoptera frugiperda was recorded in all zones at the Changlun Farm during the survey (Table 1). While S. frugiperda incidence was recorded in all zones, there was a slight variation in the severity score among the zones. Severity was generally high with score ranging from 4.70 ± 0.07 in Zone A (least severe) to 5.00 ± 0.00 in Zone B (most severe). The differences are probably contributed to different planting dates between the zones in which Zone B was planted much earlier than Zone A and C (in fact Zone B was ready for harvest at the time of sampling). Damage was consistent on all plant parts. Average larva count on the leaf/stem ranged from 2.70 ± 0.33 to 3.40 ± 0.50. In contrast, average larva count on the cob ranged from 2.20 ± 0.51 to 3.30 ± 0.40. Altogether, the highest number of larvae were recorded in Zone B (6.00 ± 0.42) followed by Zone A (5.60 ± 0.69). Zone C (5.20 ± 0.66) recorded the lowest number of larvae infesting the maize.

Discussion Spodoptera frugiperda is a well-known polyphagous invasive insect known for its destructive feeding. The pest is a nocturnal insect preferring to feed, mate and fly at night (Schöfl et al. 2009; Rojas et al. 2004). The females lay their eggs on the upper side of the leaf. The egg mass is normally covered with a layer of whitish-grey hairy scales as an additional layer of protection from egg parasitism or other environmental influences (Carneiro and Fernandes 2012). Heavy rain on the first day of sampling may have washed off the eggs from the leaves onto the ground as we only managed to sample one cluster of S. frugiperda egg mass during the sampling. Unfortunately, no larvae emerged from the collected cluster in our laboratory. It may be possible that the eggs were not fertile or were parasitized by an egg parasitoid. Further studies would need to be done to elucidate plausible reasons why the S. frugiperda larvae did not emerge from the collected egg cluster. The potential discovery of natural enemies such as parasitoid and predators would greatly help the management of this serious pest sustainably. In addition, further study on the impact of environmental factors (especially precipitation) towards S. frugiperda population as the influence of such factors remain understudied especially in this region where precipitation is considerably high all year round. Damage by the S. frugiperda larvae was substantially serious in the farm with injury visible on all plant parts. Under normal circumstances, a single plant may be infested by two to three larvae. We found at least three large larvae on a plant indicating the significance of the S. frugiperda attack on the farm. The nature of damage includes moist sawdust-like frass which

Jamil et al. (2021), BioInvasions Records 10(1): 81–90, https://doi.org/10.3391/bir.2021.10.1.10 87 First incidence of Spodoptera frugiperda in Malaysia

was observed on the funnel and upper leaves. Larvae were found feeding on leaf gregariously till the leaf becomes dry and subsequently move to other leaf for feeding. Early symptoms of S. frugiperda infestation resemble other stem borers damage with the likes of holes or “windowpane” on the leaves emerging from the whorl similar to Deole and Paul (2018). Likewise, the matured larvae digested large amounts of leaf tissue resulting in a ragged appearance to the leaves similarly to grasshopper damage. These larger larvae were then found to feed into the corn ear and seek shelter from natural enemies and other environmental factors. The colour of larger S. frugiperda larvae varied substantially from yellowish to greenish to brownish while maintaining the unique characteristics of the species such described previously in the morphological identification section. It was recently established that the S. frugiperda occurs in two races: The R strain which prefers to feed on rice and the C strain which favours maize, cotton and sorghum (Gouin et al. 2017; Nagoshi et al. 2007). Both strains are morphologically identical and can only be distinguished by molecular techniques. Hence, there is an urgent need to apply molecular techniques. Elucidating which strain is currently invading a rice-producer country such as Malaysia would greatly help researchers and stakeholders to develop appropriate management strategies. In summary, S. frugiperda is undoubtedly a threat to the food security and agricultural production in Malaysia. Currently, the impact has been limited to maize but taking into consideration the polyphagous nature of the S. frugiperda, there is a high risk of the pest invading other economically important crops such as rice as well. We anticipate that both maize small holders and large scale farmers will be affected by this pest. Interestingly, the Changlun farmer applied various pesticides according to recommendations by agricultural extension officers, but control was not effective as damage incidence were still noticeable on every plant. The efficacy of chemical control may have been limited by behaviour of the S. frugiperda larvae which prefers to remain inside the maize whorl and cob, making it difficult to reach the target with insecticidal sprays. The current practice where farmers continuously rely on a pesticides with the same active ingredient is not sustainable and may lead to the S. frugiperda developing resistance towards these pesticides. Hence, urgent research, including studies focusing on the management of this pest, assessment of the impact on maize yield, early warning systems and biological control based Integrated Pest Management (IPM) is therefore imperative.

Acknowledgements

Both MARDI Agrobiodiversity and Environment Research Centre and Industrial Crop Research Centre, are gratefully acknowledged for technical and funding contributions. The Department of Agriculture are acknowledged for assistance in the field. Mohd Fuad Mohd Nor and Shahidan Shohaimi is thanked for sampling and laboratory works. Jayprakash Pertabrai Keshavla is thanked for confirming the initial identification of the specimens which was based on description available in literature and internet mining. Fadilah Bakri and one other reviewer is thanked for their constructive comments during the review process.

Jamil et al. (2021), BioInvasions Records 10(1): 81–90, https://doi.org/10.3391/bir.2021.10.1.10 88 First incidence of Spodoptera frugiperda in Malaysia

Funding Declaration

This project was supported by the MARDI Special Research Grant (Grain Corn Project) K-GB167-1001. Funding was specifically awarded to authors SZ, MS and WK who were involved largely with the research and collectively decided to publish in this journal.

References

CABI (2019) CABI Invasive Species Compendium - Spodoptera frugiperda. https://www.cabi. org/isc/datasheet/29810#tobiologyAndEcology (assessed 29 May 2020) Carneiro TR, Fernandes OA (2012) Interspecific interaction between Telenomus remus (Hymenoptera: Platygastridae) and Trichogramma pretiosum (Hymenoptera: Trichogrammatidae) on Spodoptera frugiperda (Lepidoptera: Noctuidae) eggs. Anais da Academia Brasileira de Ciências 84: 1127–1135, https://doi.org/10.1590/S0001-37652012000400027 Day R, Abrahams P, Bateman M, Beale T, Clottey V, Cock M, Colmenarez Y, Corniani N, Early R, Godwin J, Gomez J (2017) Fall armyworm: impacts and implications for Africa. Outlooks on Pest Management 28: 196–201, https://doi.org/10.1564/v28_oct_02 Deole S, Paul N (2018) First report of fall army worm, Spodoptera frugiperda (J.E. Smith), their nature of damage and biology on maize crop at Raipur, Chhattisgarh. Journal of Entomology and Zoology Studies 6(6): 219–221 FAO (2017) FAO advisory note on fall armyworm (S. frugiperda) in Africa, 2017. http://www. fao.org/3/a-i7470e.pdf (assessed 30 May 2020) FAO (2018) FAO Guidance Note 2: Fall armyworm scouting. http://www.fao.org/3/I8321EN/ i8321en.pdf (assessed 31 May 2020) Ginting S, Agustin Z, Risky HW, Sipriyadi S (2020) New invasive pest, Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae) attacking corns in Bengkulu, Indonesia. Serangga 25(1): 105–117 Goergen G, Kumar PL, Sankung SB, Togola A, Tamò M (2016) First report of outbreaks of the fall armyworm Spodoptera frugiperda (J.E. Smith) (Lepidoptera, Noctuidae), a new alien invasive pest in West and Central Africa. PLoS ONE 11: 1–9, https://doi.org/10.1371/journal. pone.0165632 Gouin A, Bretaudeau A, Nam K, Gimenez S, Aury JM, Duvic B, Hilliou F, Durand N, Montagné N, Darboux I, Kuwar S (2017) Two genomes of highly polyphagous lepidopteran pests (Spodoptera frugiperda, Noctuidae) with different host-plant ranges. Scientific Reports 7: 11816, https://doi.org/10.1038/s41598-017-10461-4 Hang DT, Van Liem NG, Lam PV, Wyckhuys KA (2020) First record of fall armyworm Spodoptera frugiperda (J.E. Smith (Lepidoptera: Noctuidae) on maize in Viet Nam. Zootaxa 4772: 396–400, https://doi.org/10.11646/zootaxa.4772.2.11 IPPC (2019) Fall armyworm report. Report No. MYS-02/4. International Plant Protection Convention. https://www.ippc.int/en/countries/malaysia/pestreports/2019/11/fall-armyworm-spodoptera- frugiperda-control/ (assessed 31 May 2020) Johnson SJ (1987) Migration and the life history strategy of the fall armyworm, Spodoptera frugiperda in the Western Hemisphere. International Journal of Tropical Insect Science 8: 543–549, https://doi.org/10.1017/S1742758400022591 Kalleshwaraswamy CM, Asokan R, Swamy HM, Maruthi MS, Pavithra HB, Hegde K, Navi S, Prabhu ST, Goergen G (2018) First report of the fall armyworm, Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae), an alien invasive pest on maize in India. Pest Management in Horticultural Ecosystems 24(1): 23–29 Kuate AF, Hanna R, Fotio AR, Abang AF, Nanga SN, Ngatat S, Tindo M, Masso C, Ndemah R, Suh C, Fiaboe KK (2019) Spodoptera frugiperda Smith (Lepidoptera: Noctuidae) in Cameroon: Case study on its distribution, damage, pesticide use, genetic differentiation and host plants. PLoS ONE 14: e0217653, https://doi.org/10.1371/journal.pone.0215749 Montezano DG, Specht A, Sosa-Gómez DR, Roque-Specht VF, Sousa-Silva JC, Paula-Moraes SV, Peterson JA, Hunt TE (2018) Host plants of Spodoptera frugiperda (Lepidoptera: Noctuidae) in the Americas. African Entomology 26: 286–300, https://doi.org/10.4001/ 003.026.0286 Nagoshi RN, Silvie P, Meagher RL, Lopez J, Machado V (2007) Identification and comparison of fall armyworm (Lepidoptera: Noctuidae) host strains in Brazil, Texas, and Florida. Annals of the Entomological Society of America 100: 394–402, https://doi.org/10.1603/0013- 8746(2007)100[394:IACOFA]2.0.CO;2 Paini DR, Sheppard AW, Cook DC, De Barro PJ, Worner SP, Thomas MB (2016) Global threat to agriculture from invasive species. Proceedings of the National Academy of Sciences 113: 7575–7579, https://doi.org/10.1073/pnas.1602205113 Rojas JC, Virgen A, Malo EA (2004) Seasonal and nocturnal flight activity of Spodoptera frugiperda males (Lepidoptera: Noctuidae) monitored by pheromone traps in the coast of Chiapas, Mexico. Florida Entomologist 87: 496–503, https://doi.org/10.1653/0015-4040(2004) 087[0496:SANFAO]2.0.CO;2

Jamil et al. (2021), BioInvasions Records 10(1): 81–90, https://doi.org/10.3391/bir.2021.10.1.10 89 First incidence of Spodoptera frugiperda in Malaysia

Schöfl G, Heckel DG, Groot AT (2009) Time‐shifted reproductive behaviours among fall armyworm (Noctuidae: Spodoptera frugiperda) host strains: evidence for differing modes of inheritance. Journal of Evolutionary Biology 22: 1447–1459, https://doi.org/10.1111/j.1420- 9101.2009.01759.x Shylesha AN, Jalali SK, Gupta AN, Varshney RI, Venkatesan T, Shetty PR, Ojha RA, Ganiger PC, Navik OM, Subaharan K, Bakthavatsalam N (2018) Studies on new invasive pest Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae) and its natural enemies. Journal of Biological Control 32: 145–51, https://doi.org/10.18311/jbc/2018/21707 Tilman D, Clark M, Williams DR, Kimmel K, Polasky S, Packer C (2017) Future threats to biodiversity and pathways to their prevention. Nature 546: 73–81, https://doi.org/10.1038/ nature22900 Wildemeersch M, Franklin O, Seidl R, Rogelj J, Moorthy I, Thurner S (2019) Modelling the multi-scaled nature of pest outbreaks. Ecological Modelling 409: 108745, https://doi.org/10. 1016/j.ecolmodel.2019.108745

Jamil et al. (2021), BioInvasions Records 10(1): 81–90, https://doi.org/10.3391/bir.2021.10.1.10 90