The spotted lanternfly, Lycorma delicatula (SLF, Hemiptera: Fulgoridae), is considered a significant ecological and economic threat to agricultural, environmental and amenity plant hosts (Dara et al., 2015). Recently, SLF has expanded its international distribution from its native range of China, Vietnam and India with invasive populations establishing in South Korea (Bourgoin, 2014; Han et al., 2008), Japan (Kim et al., 2013) and, most recently, the north-eastern part of the United States (US) (Barringer et al., 2015; Dara et al., 2015).
SLF feeds on plants using piercing-sucking mouthparts, which can result in wilting or the death of branches under high pest population levels (Barringer et al., 2015; Han et al., 2008). Large amounts of honeydew are excreted and deposited on host trees and the understories during feeding, promoting the growth of sooty mould that hinders plant photosynthesis and contaminates agricultural and forest crops (Barringer et al., 2015; Han et al., 2008). Significant damage has been recorded in vineyards in Korea (Han et al., 2008; Lee et al., 2009; Park et al., 2009), while in the US, a vineyard at the core of the original infestation, reported a 90% yield loss with a corresponding loss in fruit quality (Urban, 2020). Orchard trees have not, as yet, observed any significant damage (Urban, 2020). In countries like Australia, where SLF is not known to occur, it is expected to threaten nursery, fruit, landscape and hardwood industries. Impacts can also be expected on businesses and residents as it is a nuisance pest, congregating in large numbers on wood piles, garden furniture, fence posts and other inanimate objects (Liu, 2019).
Several studies modelling the environmental niche of SLF have estimated significant establishment potential in many regions where SLF is currently not known to occur (Namgung et al., 2020; Wakie et al., 2020). A south Korean study estimated that maximum temperatures in the warmest month of the year were an important predictor with some evidence that the occurrence followed areas of high human impact (Namgung et al., 2020). A study on the global distribution potential of SLF identified highly suitable areas for SLF in Asia, Oceania, South America, North America, Africa, and Europe (Wakie et al., 2020). The authors found that the most important environmental variables for predicting the potential distribution of L. delicatula were mean temperature of the driest quarter, elevation, degree-days with a lower developmental threshold value of 11°C, iso-thermality, and precipitation in the coldest quarter. Further, contrary to other published predictions (e.g., Jung et al., 2017), Wakie et al. (2020) predicted that tropical habitats were not suitable for L. delicatula establishment. While researchers attempt to further refine predictions to incorporate important life-stage events, such as egg hatch following winter diapause (Smyers et al., 2021), modelling the timing of development throughout the entire life cycle of SLF across its global range remains unstudied.
In the northern hemisphere, SLF is generally considered univoltine – undergoing one generation per year – however there is some uncertainty surrounding whether it may become multivoltine if it is able to establish in warmer climates (D.-H. Lee et al., 2019). SLF is a hemimetabolous insect, passing through an egg stage, four immature instars (nymphs), and an adult stage. In the northern hemisphere, SLF eggs are typically laid during the autumn (Dara, Barringer, and Arthurs 2015; Lee et al. 2011; Park et al. 2009) in masses of 30–50 eggs in an egg case (ootheca) (Dara et al., 2015; J. G. Kim et al., 2011; J.-D. Park et al., 2009), with a large variation in the number of the masses found per tree. The eggs are approximately 2.6 mm in length and 1.4 mm in width, greyish and ovoid in shape with a distinctive ‘stem-like’ structure at the anterior end that runs back across the dorsal surface of the egg (Park et al., 2009). Embryonic development represents a crucial life stage as a significant duration is spent overwinter suspended in diapause under cold conditions not tolerable in other life stages (Dara et al., 2015; Park et al., 2009). First instars are black with white spots and emerge as early as the following mid-spring (Dara, Barringer, and Arthurs 2015; Kim et al. 2011; Lee et al. 2014; Park et al. 2009). The fourth instar is predominantly red, retaining the white spots of earlier stages but acquires a black longitudinal pattern. The average size of each instar is 3.6–4.4 mm, 5.1–6.4mm, 6.9–9.4 mm, and 10.9–14.8 mm for instars 1 to 4, respectively (Dara et al., 2015; J.-D. Park et al., 2009). Adult female L. delicatula are 24–27 mm in length; males are smaller at 21–22 mm (Dara et al., 2015). Adult males emerge first, followed by females, and may live up to four months (Liu 2019).
Recently, significant temperature response data has accumulated from measurements of SLF development under different conditions. The reported lower temperature threshold for egg development varies between studies, including 8.1°C (Choi et al., 2012), 11.1°C (M. Park, 2015) and 13.2°C (Smyers et al., 2021). The reason(s) underpinning this variability between the studies remain unclear, and could be due to regional/population differences, physiological pre-conditioning and/or inconsistencies in the experimental methods. Egg duration is typically several months from autumn through to spring, while the first, second, third, and fourth instar durations are 18.8, 20.9, 20.8, and 22.2 d, respectively at approximately 20°C (Park et al., 2009). The lower developmental threshold was estimated at 13.00 ± 0.42°C, 12.43 ± 2.09°C, 8.48 ± 2.99°C, and 6.29 ± 2.12°C for instars 1 to 4, respectively (Kreitman et al., 2021).
Here we build upon this recently acquired knowledge of the temperature response of SLF development to map the expected life-history and survival of SLF in both occupied and uninvaded ranges. Specifically, we address the following research questions: 1) Can compiled temperature response data on the development and survival of SLF life stages predict the seasonal observations of SLF occurrences in native and invaded ranges? 2) What is the predicted life-history and associated survival of different life-stages in the uninvaded ranges of Australia and Europe?