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Received: 16 May 2020 | Revised: 31 August 2020 | Accepted: 4 September 2020 DOI: 10.1002/ece3.6820 ORIGINAL RESEARCH Climate change has different predicted effects on the range shifts of two hybridizing ambush bug (Phymata, Family Reduviidae, Order Hemiptera) species Vicki Mengyuan Zhang1,2 | David Punzalan1,3 | Locke Rowe1 1Department of Ecology and Evolutionary Biology, University of Toronto, Toronto, ON, Abstract Canada Aim: A universal attribute of species is that their distributions are limited by numer- 2 Department of Biology, University of ous factors that may be difficult to quantify. Furthermore, climate change-induced Toronto, Mississauga, ON, Canada 3Department of Biology, University of range shifts have been reported in many taxa, and understanding the implications Victoria, Victoria, BC, Canada of these shifts remains a priority and a challenge. Here, we use Maxent to predict Correspondence current suitable habitat and to project future distributions of two closely related, Vicki Mengyuan Zhang, Department of parapatrically distributed Phymata species in light of anthropogenic climate change. Ecology and Evolutionary Biology, University of Toronto, 25 Willcocks St. Toronto, ON Location: North America. M5S 3B2, Canada. Taxon: Phymata americana Melin 1930 and Phymata pennsylvanica Handlirsch 1897, Email: [email protected] Family: Reduviidae, Order: Hemiptera. Funding information Methods: We used the maximum entropy modeling software Maxent to identify en- Theodore Roosevelt Collection Study Grant; Canadian Network for Research and vironmental variables maintaining the distribution of two Phymata species, Phymata Innovation in Machining Technology, Natural americana and Phymata pennsylvanica. Species occurrence data were collected from Sciences and Engineering Research Council of Canada museum databases, and environmental data were collected from WorldClim. Once we gathered distribution maps for both species, we created binary suitability maps of current distributions. To predict future distributions in 2050 and 2070, the same environmental variables were used, this time under four different representative concentration pathways: RCP2.6, RCP4.5, RCP6.0, and RCP8.5; as well, binary suit- ability maps of future distributions were also created. To visualize potential future hy- bridization, the degree of overlap between the two Phymata species was calculated. Results: The strongest predictor to P. americana ranges was the mean temperature of the warmest quarter, while precipitation of the driest month and mean temperature of the warmest quarter were strong predictors of P. pennsylvanica ranges. Future ranges for P. americana are predicted to increase northwestward at higher CO2 con- centrations. Suitable ranges for P. pennsylvanica are predicted to decrease with slight fluctuations around range edges. There is an increase in overlapping ranges of the two species in all future predictions. Main conclusions: These evidences for different environmental requirements for P. americana and P. pennsylvanica account for their distinct ranges. Because these This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. © 2020 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd 12036 | www.ecolevol.org Ecology and Evolution. 2020;10:12036–12048. ZHANG ET AL. | 12037 species are ecologically similar and can hybridize, climate change has potentially im- portant eco-evolutionary ramifications. Overall, our results are consistent with ef- fects of climate change that are highly variable across species, geographic regions, and over time. KEYWORDS abiotic, bioclimatic variables, climate change, Maxent, range shifts, species distribution modeling 1 | INTRODUCTION (a) A universal attribute of all species is that their geographic distribu- tions are limited. The abiotic and biotic factors that jointly deter- mine this distribution are expected to be numerous, posing a serious empirical challenge to their identification and quantification. One approach is to employ species distribution models (SDMs), which at- tempt to explain species presence data with a large set of predictor variables (Elith & Leathwick, 2009). Although the approach is gen- erally limited to the use of environmental variables (or their proxies) as predictors of suitable habitat, SDM have provided new insights into species requirements that are akin to the “fundamental niche” (Hutchinson, 1957) by incorporating constraints set by biotic inter- actions (Leach et al., 2016). Geographic distributions are dynamic and dependent upon (b) changing environmental conditions. Species also vary in their sensitivity to shifting environmental conditions and will respond differently to the same changes (Hickling et al., 2006; Malcolm et al., 2002), including the possibility of failure to track new condi- tions altogether (Loarie et al., 2009). While there is growing evidence of climate change-induced range shifts in many taxa, predicting its ecological and evolutionary implications remains a central challenge (Parmesan, 2006). For example, climatic variation is undoubtedly linked to natural changes in community composition over geological timescales, and there is growing evidence of rapid changes in climate being linked to the invasion and expansion of alien species (Bellard et al., 2013; Guo et al., 2012). Changing climatic conditions has also led to more frequent contact between historically separate species, and this can result in hybridization (Vallejo-Marín & Hiscock, 2016) and, in some cases, species collapse (Njiru et al., 2010). For these FIGURE 1 Photographs of the two organisms in the present study: (a) P. americana waiting for prey on a Black-eyed Susan reasons, the responses of hybridizing species to environmental (Rudbeckia hirta); (b) P. pennsylvanica consuming a bald-faced hornet change have been touted as a particularly important “window” on (Dolichovespula maculate). Images by David Punzalan climate change (Taylor et al., 2015). In the present paper, we evaluate potential range shifts in a para- patric pair of insect species. Phymata americana Melin 1930 and P. Swanson, 2013); consistent with this, current molecular phyloge- pennsylvanica Handlirsch 1897 (Figure 1) are two of the most com- netic data fail to distinguish between the two (Masonick et al., 2017; mon North American species in the genus (Family: Reduviidae, Order: Masonick & Weirauch, 2020), despite substantial morphological Hemiptera), with the former more northerly in distribution, extend- divergence (Punzalan & Rowe, 2017). Both species are general- ing west across the American Midwest and Canadian prairies, and ist predators occurring in temperate habitats, where they utilize the latter mostly concentrated in the northeastern United States. a wide range of plant species as hunting sites (Balduf, 1939, 1941; Hybridization in wild populations has been suspected or inferred Yong, 2005), suggesting considerable niche overlap. In at least one in overlapping regions of their ranges (Punzalan & Rowe, 2017; of the species, climatic variables (e.g., environmental temperature) 12038 | ZHANG ET AL. play a key role in thermoregulation, which is linked to mating activity Museum (https://www.zoolo gy.ubc.ca/entom ology/) and the Plant (Punzalan et al., 2008a). Thermoregulatory abilities have been linked Bug Inventory maintained by the AMNH (http://resea rch.amnh.org/ to melanic traits, and within- and between-species latitudinal varia- pbi/). We also gathered data from two citizen science websites, iN- tion in these traits (Punzalan & Rowe, 2015) indirectly supports the aturalist.org and BugGuide.net. Data collected from iNaturalist.org importance of climatic variables in the ecology of Phymata. Although were included if the species identity was verified by Paul Masonick there is evidence that their ecological requirements are consequen- (UC Riverside), an authority on Phymata systematics and curator of tial to their life histories, there is a shortage of knowledge regarding the iNaturalist project “Uncovering the ambush bugs” (https://www. their ecology. Thus, there is value in understanding their habitat re- inatu ralist.org/proje cts/uncov ering-the-ambush-bugs). Data col- quirements and predicting their current and future ranges. lected from BugGuide.net require secondary identification verifica- We used biogeographic climate data and SDM to characterize tion before publishing on the Phymata webpage and were assumed the current and recent historical range of P. americana and P. penn- to be accurate. sylvanica and forecast future distributions under several scenarios of For accessions lacking latitude and longitude data, we supple- anthropogenic climate change. We hypothesized that different sets mented these data manually using Google Earth Pro version 7.3.1 of candidate abiotic factors limit the respective ranges of the two (Google Earth, 2018). Given that bug sightings occurred at a specific species, resulting in their current distributions. Given anthropogenic point, but its accuracy was not reflected on Google Earth, coordi- climate warming, we also hypothesized that their future overlapping nates were rounded to include degree and hour only (i.e., minutes ranges would increase. and seconds were not used). This is because locality data were only accurate down to the city level and, in some cases, were accurate down to the location of the

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