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Predicting Habitat Suitability of Coptotermes Gestroi (Isoptera: Rhinotermitidae) with Species Distribution Models

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Predicting Habitat Suitability of Coptotermes Gestroi (Isoptera: Rhinotermitidae) with Species Distribution Models HOUSEHOLD AND STRUCTURAL INSECTS Predicting Habitat Suitability of Coptotermes gestroi (Isoptera: Rhinotermitidae) With Species Distribution Models 1,2 3 1 HOU-FENG LI, IKUKO FUJISAKI, AND NAN-YAO SU J. Econ. Entomol. 106(1): 311Ð321 (2013); DOI: http://dx.doi.org/10.1603/EC12309 ABSTRACT Coptotermes gestroi (Wasmann) is an important structural pest reported from Asia, PaciÞc islands, North America, Caribbean islands, South America, and Indian Ocean islands. This study summarized previous records of C. gestroi and its synonyms, presenting 184 infested counties from 24 countries. Based on the geo-references occurrence locations and global raster data of climate, geography, and human population, C. gestroi were found most commonly in warm, high precipitation, low altitude, and human populated areas. By using species distribution models, we predicted its current infested area (model 1), habitat suitability (model 2), and probability of introduction (model 3) on a global scale. The results showed its recorded locations and the predicted distribution of the present day are similar, but the suitable habitat is larger than its current distribution. The patterns of the introduction frequency (model 3) and habitat suitability (model 2) are inconsistent. Temperate cities with high introduction risk are located in Europe, United Sates, northeastern China, and Japan where habitat suitability is low and hence successful colonization is unlikely. In tropics and subtropics, habitat suitability of C. gestroi is high. We speculate that continuous urbanization and increasing human population will increase its introduction frequency and cause further extension in fast developing tropical and subtropical countries. KEY WORDS invasive species, species distribution model, urban pest, subterranean termite Urban environments are characterized by manmade (King and Spink 1969, Messenger et al. 2005). Only a construction and designed landscape but are also few termite species reach cosmopolitan distribution shaped by natural climate and geography. Some ar- (Gay 1967, Evans 2010). Their peridomestic habitat is thropods adapted to urban habitats could reproduce in affected by outdoor climatic and geographic factors large numbers, resulting in esthetic damage, economic that probably limit their colonization and further ex- loss, and human health threats (Robinson 2005). Do- tension in introduced area. mestic pests, such as Germen cockroach (Appel 1995), In tropical and subtropical areas, the genus, Cop- Pharaoh ant (Wheeler 1910), and stored food beetles totermes, is the most signiÞcant group, including Ϸ70 (Hill 2002) complete their life cycles indoors with described species (Vargo and Husseneder 2009, Con- limited food, water, and harborage, and have been stantino 2011) and one third of which considered pests spread around the world through commerce. Subter- (Edwards and Mill 1986, Su and Scheffrahn 2000). Six ranean termites (Rhinotermitidae) are important ur- Coptotermes species have been recorded as invasive ban pests. The most damaging subterranean species pests, and Coptotermes gestroi (Wasmann) and Cop- occur in three genera, Coptotermes, Heterotermes, and, totermes formosanus Shiraki, have the largest distribu- Reticulitermes (Edwards and Mill 1986), with a com- tion (Evans 2010), and are the most often occurring bined global economic impact of $40 billion annually in tropical and temperate areas, respectively (Su (Rust and Su 2012). In urban environments, subter- 2003). The distribution of C. formosanus had been well ranean termites live in the peridomestic area and usu- documented (Tamashiro and Su 1987). However, no ally nest in soil. Their subterranean gallery system and global distribution of C. gestroi has been available aboveground mud tubes could extend over a hundred because of the long-standing confusion of its taxon- meters connecting underground nests and food omy. C. gestroi was Þrst described from Myanmar, and sources such as tree stumps, dead branches of living then several synonyms were created from other Asian tree, gardening wood, and wooded constructions countries, such as C. heimi (Wasmann 1902) and C. parvulus Holmgren 1913a from India, C. havilandi Hol- 1 Department of Entomology and Nematology, Fort Lauderdale mgren 1911a from Malaysia, C. vastator Light 1929 Research and Education Center, University of Florida, 3205 College from the Philippines, C. pacificus Light 1932 from In- Ave., Ft. Lauderdale, FL 33314. donesia, and C. obliquus Xia and He 1986 and C. yax- 2 Department of Wildlife Ecology and Conservation, Fort Lauder- ianensis Li 1986 from China. In the 20th century, C. dale Research and Education Center, University of Florida, 3205 College Ave., Ft. Lauderdale, FL 33314. gestroi was introduced to other geographic areas in- 3 Corresponding author, e-mail: houfeng@uß.edu. cluding PaciÞc islands, North America, Caribbean is- 0022-0493/13/0311Ð0321$04.00/0 ᭧ 2013 Entomological Society of America 312 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 106, no. 1 lands, South American, and Indian Ocean islands (Gay Materials and Methods 1967). The synonyms of C. gestroi were adopted in- Locations Sampled. Published literature containing dividually at different geographic areas for many de- collection records of C. gestroi and its synonyms were cades. In light of recent taxonomic studies (Roonwal reviewed. The Þrst documented year in each country, and Chhotani 1962; Gay 1967; Kirton and Brown 2003; recorded species name, identiÞed material, and ref- Yeap et al. 2007, 2010; Li et al. 2011), the C. parvulus, erences were tabulated (Table 1). To establish a reg- C. pacificus, C. havilandi, C. vastator, C. heimi, C. ular resolution database for worldwide distribution obliquus, and C. yaxianensis were proven to be junior simulation, only county records or smaller adminis- synonyms of C. gestroi. Currently, C. gestroi is consid- trative regions were used for current study. If multiple ered the most destructive structural pest in South- locations were documented in a county, we randomly east Asia (Kirton and Azmi 2005, Lee et al. 2007), selected one location. If no speciÞc location was men- India (Roonwal and Chhotani 1989, referred as C. tioned in a county, physical middle point was chosen. heimi), and Brazil (Constantino 2002, referred as C. In total, 184 county records of 24 countries were ob- havilandi). C. gestroi continually extended its infes- tained from 66 publications (Table 1). Coordinates of tation in newly invaded areas, including Taiwan (Li the 184 observation points were estimated by using et al. 2009) and Florida (Su et al. 1997, Scheffrahn Google Earth software. In addition, background ref- and Su 2005). erence points were selected at 0.5 degree interval, In addition to the taxonomic confusion, obtaining a which approximately correspond to a county size, on global distribution map through Þeld survey is always terrestrial area between 40 degrees north and south time-consuming and Þnancially challenging. Previous latitude where most termite species were found (Egg- termite surveys were geographically scattered and de- leton 2000). In total, 30,385 background reference pendant on work of few scientists. Sampling bias points were obtained. caused by different investigators and different original Environmental Factors. To describe ecological research purposes remain uncorrected because of niche of C. gestroi and to predict its current distribu- sparse information on previous sampling methods. We tion area, habitat suitability, and possibility of intro- have no idea if many of the areas that show an absence duction in worldwide scale, publicly available global of C. gestroi truly lack this species or just have not been raster data of environmental factors were used. properly surveyed yet. To overcome these limitations, Monthly minimum and maximum temperatures, pre- the current study reviews published literature with cipitation and altitude, which were representative of several hundreds of collection records of C. gestroi 1950Ð2000 (Hijimas et al. 2005), were obtained from (Table 1) to obtain its recorded occurrence locations WorldClim (2010). From monthly minimum and (Fig. 1A), and by using species distribution models to maximum temperatures and precipitation, we de- predict its current areas of infestation (Fig. 1B). Spe- rived annual minimum and maximum temperatures cies distribution models or ecological niche models and precipitation. Moderate Resolution Imaging are frequently used to predict the potential range of Spectroradiometer (MODIS)/terra land cover type exotic species (Peterson and Vieglais 2001, Welk et al. 2007 data with the International Geosphere-Biosphere 2002) to assess invasion potential. A common ap- Programme (IGBP) vegetation classiÞcation scheme, proach is to parameterize models based on the native which differentiates 16 land cover classes (Friedl et al. 2002), were obtained from NASA the Warehouse In- range of species and use the deÞned environmental ventory Search Tool (2010). Gridded Population of correlates to predict the adventive range (Welk et al. the World (GPW) version three (Center for Interna- 2002, Richardson and Thuiller 2007). Various methods tional Earth Science Information Network [CIESIN] have been developed that allow predictions using ex- 2005), which depicts the human population density in isting species occurrence data to understand the 2010, was obtained from Web site of the Center for mechanism of speciesÕ arrange shifts in the face of International Earth Science Information Network
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