Zootaxa, Description of the Male of the Spider Dubiaranea Difficilis
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TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Zootaxa 2405: 55–62 (2010) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2010 · Magnolia Press ISSN 1175-5334 (online edition) Description of the male of the spider Dubiaranea difficilis (Araneae: Linyphiidae), with new records and modeling of its potential geographic distribution GONZALO D. RUBIO1,3, EVERTON N. L. RODRIGUES2 & LUIS E. ACOSTA1,3 1CONICET. Diversidad Animal I. Facultad de Ciencias Exactas, Físicas y Naturales. Universidad Nacional de Córdoba. Av. Vélez Sarsfield 299, X5000JJC Córdoba, Argentina. E-mails: [email protected]; [email protected] 2Programa de Pós-Graduação em Biologia Animal, Departamento de Zoologia, Instituto de Biociências, Universidade Federal do Rio Grande do Sul. Av. Bento Gonçalves, 9500, Bloco IV, Prédio 43435, 91501-970 Porto Alegre, RS, Brazil. E-mail: [email protected] Abstract The male of Dubiaranea difficilis (Mello-Leitão 1944) is described and illustrated for the first time. New geographic records of this species are provided, and its potential distribution is modeled using MAXENT. The actual and potential distribution of Dubiaranea difficilis proved to be much larger than the hitherto known records. This species inhabits at least in three ecoregional sectors in Argentina (northwestern area, central Sierras, and the Pampas), and is also likely that inhabits in Brazil and Bolivia. Possible causes of its wide and environmentally heterogeneous distribution are discussed. Key words: Argentina; Dubiaraneinae; Neotropical region; spider taxonomy Introduction The spider genus Dubiaranea Mello-Leitão 1943, currently including about 100 species, is by far the largest linyphiid genus in South America, although the number of species is suspected to be much higher, possibly doubling that amount (Millidge 1991). Of the known species 36 occur in Peru, 18 in Colombia, 17 in Ecuador, 15 in Chile, seven in Bolivia, six in Venezuela, five in Brazil, and only three have been recorded from Argentina: D. difficilis (Mello-Leitão 1944), D. remota Millidge 1991 and D. tristis (Mello-Leitão 1941). In addition, two further species are known from such distant places like Borneo and Juan Fernández islands, with one species each (Platnick 2009). The latest revision of Dubiaranea was carried out by Millidge (1991), and the 80 new species treated there represent 80% of the species known today. However, only 27 species are known from both males and females, and 48 are just based on females. Moreover, several of the species were described on a single specimen, so that intraspecific variability is completely unknown. This is the case of D. difficilis, which was described by Mello-Leitão (1944) based on a female and originally placed in the tetragnathid genus Paranesticus Mello- Leitão (1944). Millidge (1991) transferred difficilis to its current generic placement and illustrated the female genitalia for the first time. Nevertheless, the species was hitherto recorded only from two localities in Argentina (Millidge 1991; Grismado 2007), so that our knowledge on its distribution is clearly incomplete. As a result of an ecological study in northwestern Argentina, together with miscellaneous samples obtained in the central region of the country, additional specimens of both sexes were collected, providing evidence of a much larger range than previously known. Mating couples observed in the field (Fig. 1) provided definitive support to the conspecificity of males and females in our samples. Geographic distribution can help with species identification in speciose genera, and Dubiaranea is no exception. Based on the present knowledge, most species of this genus appear to be quite limited in their ranges, and a species collected at 10º S, for example, is deemed unlikely to be present at 5º N or 30º S, Accepted by G. Homiga: 22 Feb. 2010; published: 22 Mar. 2010 55 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. although it might be found at nearer latitudes (Millidge 1991). Such an intuitive approach, however, is normally based on incomplete distribution data and contains some implied circularity. As a sounder alternative, the predictive range modeling based on actual occurrence records of species is a useful tool to complement incomplete recording and to contrast the actual versus the potential distribution (Maddock & Du Plessis 1999; Raxworthy et al. 2003; Acosta 2008). This approach models the distribution of a species as a function of a set of environmental variables (Maes et al. 2005). The available information on the geographic distribution of most spider species is no more than a few points on a map. For this reason the modeling of potential distribution offers a first step to infer the basic dimensions of the distribution range of a species (Acosta 2008). Recent advances in collection of climatic data, GIS technology, and design of diverse modeling algorithms (e.g. GARP, Stockwell & Peters 1999; BIOCLIM, Fischer et al. 2001; MAXENT, Phillips et al. 2009) make this type of study very feasible and accurate (Elith et al. 2006; Hijmans & Graham 2006; Acosta 2008; Echarri et al. 2009). In this paper, the male of D. difficilis is described for the first time and its somatic and palpal morphology is illustrated. New records complete the deficient knowledge of the geographic distribution of this species and, together with the previous records, they are used to model the potential distribution of D. difficilis. Methods Sampling. Specimens were collected in different sites in northwestern Argentina, using a Garden-Vacuum method to suck spiders from the vegetation (for details on the method, see Bolger et al. 2000; Bell et al. 2002). Additional samples were obtained by manual collecting in Córdoba and San Luis Provinces, Argentina. Taxonomic description. The taxonomic descriptions follow Millidge (1991), with updated terminology as used by Hormiga (2000). Males of Dubiaranea are diagnosed by the form of the suprategulum and the lamella characteristica (Millidge 1991, referred there as “suprategular apophysis” and “embolic division”, respectively). These two palpal components were observed and illustrated after clearing in clove oil and examined from more than one angle. The trichobothrium position on metatarsus I was described following Denis (1949). The tibial spine formula is based on Roberts (1987). Illustrations were made on photographs obtained with a Sony® DSC-W290 digital camera attached to a Leica® MS5 stereomicroscope. Photographs in nature were taken with a Nikon® D80 digital camera using an 18-135mm lens. All measurements were taken with a micrometric ocular and are in millimeters. The specimens examined are deposited in the following institutions (abbreviations and curators in parentheses): Museo de La Plata, La Plata (MLP, L. Pereira); and Colección Aracnológica de la Cátedra de Diversidad Animal I, Facultad de Ciencias Exactas, Físicas y Naturales, Universidad Nacional de Córdoba, Córdoba (CDA, L.E. Acosta). Modeling. The records dataset consists of 15 locality points and was arranged to be used within a geographic information system (DIVA-GIS 5.4, Hijmans et al. 2005a). Only one occurrence record per grid cell was retained as valid in the analysis, resulting in 14 valid (non duplicate) points. The potential distribution of D. difficilis was modeled using 19 bioclimatic parameters extracted from the WorldClim database (Hijmans et al. 2005b), at a resolution of 30 arc-seconds (1 km²). The total area modeled was between the coordinates 9º28'01''S to 43º17'60''S, and 43º47'60''W to 74º21'00''W. The model was built with MAXENT software, version 3.3.0 (Phillips et al. 2009). This method estimates the likelihood of a species being present by finding the distribution of maximum entropy (closest to uniform to a certain number of iterations), subject to the constraint that the expected value of each environmental variable under this estimated distribution matches its empirical average (Phillips et al. 2006). MAXENT is a widely used method for modeling species potential distributions and has been shown to perform well in comparison with alternative approaches (Elith et al. 2006; Hijmans & Graham 2006; Echarri et al. 2009; Tognelli et al. 2009). The model was run applying the “equal training sensitivity plus specificity” threshold rule (Liu et al. 2005). Other relevant settings were used in their default values: the convergence threshold (10-5); maximum background points (10,000); maximum iterations (1,500); replicated run type (subsample), and output format (logistic). MAXENT produces a continuous 56 · Zootaxa 2405 © 2010 Magnolia Press RUBIO ET AL. TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. prediction of specific presence that ranges from 0 to 1 (Phillips et al. 2006). Resulting predictions were visualized importing the ASCII files into DIVA-GIS 5.4 (Hijmans et al. 2005a). The accuracy of the predictive range generated was evaluated by calculating the AUC in a ROC (receiver operating characteristic) plot. To calculate the accuracy 70% of the original points were randomly re-sampled as training data to perform 20 iterations of the model (Acosta 2008). Results Taxonomy Family Linyphiidae Blackwall 1859 Dubiaranea Mello-Leitão 1943 Dubiaranea difficilis (Mello-Leitão 1944) (Figs. 1–6) Paranesticus difficilis Mello-Leitão 1944: 333, figs. 19–20. Dubiaranea difficilis: Millidge 1991: 52, fig. 173. Type material. Holotype female (MLP 15980), ARGENTINA: Buenos Aires: General Guido (36º38'22''S, 57º47'24''W), February 1941, M. Birabén coll. (examined). FIGURE 1. Dubiaranea difficilis (Mello-Leitão 1944), male and female mating in nature; spiders are located in the sheet-web of the female. Left photo from La Falda, Córdoba (November 2008); right photo from Huerta Grande, Córdoba (September 2009). New records. ARGENTINA: Córdoba: La Falda (31º05'21''S, 64º27'42''W), 24 October 2008 (G.