Platymeris Biguttatus (Linnaeus, 1767) with Description of Nymphs Dominik Chłond, Agnieszka Bugaj-Nawrocka* and Łukasz Junkiert
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Chłond et al. Zoological Studies (2015) 54:9 DOI 10.1186/s40555-014-0092-5 RESEARCH Open Access Current and potential geographical distribution of Platymeris biguttatus (Linnaeus, 1767) with description of nymphs Dominik Chłond, Agnieszka Bugaj-Nawrocka* and Łukasz Junkiert Abstract Background: The description of Platymeris biguttatus (Linnaeus 1767) nymphal instars as well as the prediction of the potentially suitable ecological niche was the main goal of this study. Our research was based on 258 specimens of P. biguttatus species of museum collections. A set of 23 environmental predictor variables covering Africa was used at ecological niche modeling - a method performed using the Maxent software to prepare potential distribution maps for this species. Results: The results suggested the most suitable areas seen as potentially suitable ecological niche for P. biguttatus in Africa. A jackknife test showed that temperature seasonality and percentage of tree cover were among the most important environmental variables affecting the distribution of the species. The analysis of climate preferences shows that most of the potentially suitable niches for this species were located in the area of tropical savanna climate, with a small participation of tree vegetation. Conclusions: P. biguttatus was only known to be widely distributed in the tropical part of continental Africa. Thanks to the ecological niche modeling methods and the museum data on the occurrence of the species, we introduced new information about potentially suitable ecological niches and the possible range of distribution. Keywords: Africa; Comparison of niches; Ecological niche modeling; Habitat suitability; Species distribution Background of a large number of individuals of this species deposited Platymeris biguttatus (Linnaeus 1767) is one of thirteen in European museum collections as well as numerous live species of the genus Platymeris Laporte 1833 described specimens breed in captivity in our laboratory, examin- so far. It is a medium-sized genus belonging to the sub- ation and description of morphology of each nymphal in- family Reduviinae, and all known species of these assas- stars of this species was possible, and it is provided in the sin bugs are large-sized insects, distributed exclusively in present paper. Africa (Maldonado Capriles 1990). P. biguttatus (L.) is Ecological niche modeling was introduced into this widely distributed in the tropical part of continental study to learn more about some ecological aspects of the Africa, and it can be easily recognized amongst the other mentioned species. This method is widely used in the representatives of this genus by the white spots on the faunal and floral studies, and it is very useful for bio- hemelytra, yellow bands on the legs and black connexiva. geography, e.g., to identify potential migration routes of Redescription of the species as well as notes on its biol- species (mostly invasive ones), to plan the establishment ogy in captivity were published by Li et al. (2010). Al- of protected areas, or even to predict the effects of climate though the article contains pictures and presents the change (Beck 2012; Berzitis et al. 2014; Ochoa-Ochoa biology of nymphs of this species, it does not provide a et al. 2009; Wieczorek and Bugaj-Nawrocka 2014). description of each nymphal instars. Due to the availability As many factors, both biotic and abiotic, affect the dis- tribution of species or plant communities, it is important to choose the right method and environment variables. * Correspondence: [email protected] Department of Zoology, Faculty of Biology and Environmental Protection, Our study was aimed at determining which niches might University of Silesia, Bankowa 9, 40-007 Katowice, Poland be potentially suitable for P. biguttatus, interpreting the © 2015 Chłond et al.; licensee Springer. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly credited. Chłond et al. Zoological Studies (2015) 54:9 Page 2 of 11 influence of predictors on the distribution and comparison using a camera lucida. Measurements are given in milli- with the results obtained for Platymeris rhadamanthus meters. Measurements were performed on 30 individuals (Chłond and Bugaj-Nawrocka 2014). The maximum en- of each nymphal stage - a total of 150 individuals were tropy model (Maxent) is based on a machine learning measured. Photos were taken with a Sony SLT-A37 cam- method and has many advantages (e.g., it avoids commis- era with additional 13-mm and 31-mm extension tubes. sion errors, incorporates interactions between variables, utilizes continuous and categorical data) that make it a Material well-suited method for species distribution modeling See Additional file 1 for a detailed list of all examined (Phillips et al. 2006). It is also worth to note that Maxent individuals of P. biguttatus. This list, as well as the list in seems to work better than other software and has a better Additional file 2, provides the information about occur- performance in predicting the distribution of species, es- rence localities. pecially when a limited number of sample localities is available (Elith et al. 2006; Hernandez et al. 2006; Papeş Occurrence data and Gaubert 2007). A total of 50 occurrence localities were assembled for the species P. biguttatus. It has been localized in Senegal, Methods Gambia, Guinea, Mali, Ivory Coast, Togo, Benin, Niger, Taxonomy Nigeria, Chad, Sudan, Ethiopia, Somalia, the Democratic Numerous live specimens breed in captivity in our labora- Republic of Congo, Uganda, Kenya, Tanzania, Zambia, tory were available at various stages of nymphs. External Zimbabwe, and Mozambique (Figure 1). All occurrence structures of nymph specimens were examined using a data was based on the detailed review of 258 specimens stereoscopic microscope Olympus SZH10 (Olympus studied in museum collections (see abbreviations for de- Corporation, Shinjuku-ku, Japan). All drawings were made positories in Additional file 1). Records with unspecified Figure 1 Dorsal habitus of the first three instar nymphs of P. biguttatus. (A) first instar nymph; (B) second instar nymph; (C) third instar nymph. Chłond et al. Zoological Studies (2015) 54:9 Page 3 of 11 or unknown localities were not used. All localities were indicate that the model accuracy (AUC values) depends on georeferenced using Google Earth 7.1.2.2041 (Google Inc. the number of records, the used algorithm, and the type of 2013, Mountain View, CA, USA) (geographical projection, data used (Raes and ter Steege 2007; Wisz et al. 2008; decimal degrees, datum: WGS84). Aguirre-Gutiérrez et al. 2013), so it should not be consid- ered to be the most important determinant of the accuracy Environmental predictors of the model. As potential predictors of habitat distribution for P. bigut- Raw environmental data was extracted from all raster tatus, environmental variables were used. The study layers at species occurrence records using SAGA GIS was based on bioclimatic variables which were down- 2.1.0 (SAGA Development Team 2013). A Spearman rank loaded from WorldClim 1.4 dataset (Hijmans et al. 2005; correlation was performed in the R software (version http://www.worldclim.org/bioclim). The Global Land 3.0.2) (R Development Core Team 2011) using Rattle One-km Base Elevation Project (GLOBE) (GLOBE Task package (version 3.0.2 r169) (Williams 2011) to minimize Team and others 1999; http://www.ngdc.noaa.gov/mgg/ the number of variables by discarding these which were topo/globe.html) provided a digital elevation model highly correlated (r > 0.75). The variables that did not have (DEM). Also a layer of the tree cover continuous fields any significant contribution to the model were removed, was used. It shows vegetation as continuous fields of land and six uncorrelated environmental factors were selected cover, where every pixel has a percentage value for tree (Table 1). cover (DeFries et al. 2000; http://glcf.umd.edu/data/ As species occurrence points came from the museum treecover). All variables were cut to the boundaries of data and were not collected randomly, a bias file was Africa and a spatial resolution of 30 arc-seconds provided during Maxent modeling. The bias grid file was (approximately 1 km) was chosen. generated in SAGA GIS, weighted by a Gaussian kernel To determine the possible climatic preferences, results with a standard deviation (SD) of 200 km (see Elith et al. for potential ecological niche were compared to the 2010). To avoid extreme values, the resulting grid was Köppen-Geiger climate classification system updated by scaled to have a maximum of 20 and a minimum of 1. Peel et al. (2007). Moreover, to identify niche overlap from Maxent model predications for P. biguttatus and P. rhadamanthus Ecological niche modeling (Chłond and Bugaj-Nawrocka 2014), SAGA GIS 2.1.0 Maxent software (version 3.3.3 k), based on maximum en- software was used. The ASCII median output files of tropy algorithm, was used to develop a species distribution Maxent for both species were reclassified and then added model. The following settings were chosen: maximum together to obtain a raster that contained the suitable number of background points = 10,000, convergence niche classes for both species. threshold = 0.00001, regularization multiplier = 1, maximum number of iterations = 5,000. These settings allowed for ad- Results equate time for model convergence (Stohlgren et al. 2011; Taxonomy Elmore et al. 2013; Solhjouy-Fard and Sarafrazi 2013). Description of nymphs A 50-fold cross-validation was run using the full occur- rence dataset randomly split into a 25% test and 75% First instar (Figure 1A) Color: Head and thorax black training dataset (Phillips et al. 2006; Phillips and Dudík with dark brown ventral parts and red ecdysial lines. An- 2008). The logistic output of Maxent was used, which tennae with black scapus and pedicellus, both segments gives the results for the prediction values ranging from 0 covered by black setae.