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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 Dubiaranea difficilis (Araneae: ), with new records and modeling of its potential geographic distribution

GONZALO D. RUBIO1,3, EVERTON N. L. RODRIGUES2 & LUIS E. ACOSTA1,3 1CONICET. Diversidad 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

Introduction

The spider 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 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. Rubio), 1 male (MLP 17986); Huerta Grande, site 1 (31º04'38''S, 64º29'28''W), 02 May 2009 (G. Rubio), 1 female (CDA 000.812); same loc., site 2 (31º04'52''S, 64º29'12''W), 09 May 2009 (same coll.), 3 females (CDA 000.813), 06 September 2009 (same coll.), 1 male (CDA 000.816). Jujuy: Potrero de Yala (24º07'13''S, 65º27'24''W), 29 November 2008 (M. Guerra & G. Rubio), 2 females (CDA 000.814); Calilegua, Ríos Aguas Negras–San Lorenzo intersection (23º45'47''S, 64º51'01''W), 19 October 2009 (M. Pocco & G. Rubio), 1 female (CDA 000.837); same loc., near Monolito (23º40'27''S, 64º54'01''W), same date (same coll.), 1 female (CDA 000.838). Salta: Posta de Yatasto (25º35'44''S, 64º57'09''W), 26-28 April 2006 (G. Rubio, J. Corronca, B. Cava, V. Olivo & A. González-Reyes), 1 female (MLP 18017); Arroyo Las Tipas (25º24'37''S, 64º56'58''W), 01 August 2006 (same coll.), 1 female (MLP 18018); Quebrada San Lorenzo: site 1 (24º43'16''S, 65º30'94''W), 01 August 2006 (same coll.), 1 male (MLP 18019); La Caldera: site 1 (24º30'19''S,

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65º19'61''W), 01 August 2006 (same coll.), 1 female (MLP 18020). San Luis: Merlo, Rincón del Este (32°20'51"S, 64°57'56"W), 10 January 2010 (M. Pocco & G. Rubio), 2 females (CDA 000.817) and 2 male (CDA 000.818). Tucumán: Sierra de San Javier (26º48'04''S, 65º20'56''W), 24 November 2008 (G. Rubio) 1 female (CDA 000.815); Parque Provincial El Cochuna (27º19'23''S, 65º55'40''W), 12 October 2009 (M. Pocco & G. Rubio) 3 females and 1 male (CDA 000.839).

FIGURE 2–6. Dubiaranea difficilis (Mello-Leitão 1944). 2. Female from Huerta Grande, Córdoba, 09 May 2009 (CDA 000.813), ventral view of epigynum. 3–6: Male from La Falda, Córdoba, 24 October 2008 (MLP 17986). 3. Left palpus, ectal view. 4. Left palpus, mesal view. 5. Habitus, dorsal view. 6. Habitus, lateral view. (C = abdominal chevrons; E = embolus; LC = lamella characteristica; CLC = caudal apophysis of LC; G = thoracic groove; MLC = mesal apophysis of LC; MSA = marginal suprategular apophysis; P = paracymbium; SPT = suprategulum). Scale bars: (2–4) 0.25 mm; (5, 6) 1 mm.

Diagnosis: The male of D. difficilis (Figs. 3, 4) resembles that of D. vetusta Millidge 1991 (Millidge 1991: figs. 52, 53) and of D. fruticola Millidge 1991 (Millidge 1991: figs. 80, 81) from Ecuador and Peru, respectively. It differs from both species in that the suprategulum of D. difficilis carries a small acute marginal suprategular apophysis (Fig. 3), and the lamella characteristica presents a well developed mesal apophysis (Fig. 4). The caudal apophysis of the lamella characteristica is slightly longer and slender than in D. fruticola, and the paracymbium tip is acute, not rounded as in D. vetusta. The female of D. difficilis is diagnosed by the scape of epigynum, which is broaden anteriorly and widened posteriorly, with a wide socket and wide atria (Fig. 2; Millidge 1991: fig. 173). Description. Male from La Falda, Córdoba (MLP 17986): Total length 4.16; carapace length 1.80; carapace width 1.27; clypeus height 0.32; sternum length 1.00; sternum width 0.84; abdomen length 2.30; abdomen width 0.77. Carapace low and elongated, pale yellowish-brown, with three blackish longitudinal bands starting at posterior eyes toward the posterior margin (Figs. 5, 6). Cephalic region elevated, slightly separated from thoracic region by a V-shaped furrow immediately anterior to the groove (Figs. 5, 6). Groove

58 · 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. located in carapace posterior half. Eyes on shallow black tubercles (Fig. 6); both rows of eyes recurved; the first row more than the second row; posterior median eyes almost twice larger than the lateral ones; anterior median eyes smaller than the rest; anterior lateral eyes and posterior lateral eyes touching (Fig. 5). Sternum orange, truncated posteriorly. Chelicerae orange-brown, with about 20 stridulatory ridges; promargin with 3 teeth (the middle tooth much larger), and retromargin with 6 subequal teeth. Abdomen with parallel sides; ventrally dark grey; dorsally pale, with 3 blackish chevrons in the posterior half (Figs. 5, 6), and with some lateral silver spots. Abdomen black distally. Legs yellowish (darker in the distal part of the tibiae), long and feeble, with some spines (Fig. 6). Coxa III less developed. Position of Tm I 0.16. Legs I-III with one trichobothrium on each metatarsus, absent on metatarsus IV. Tibial spine formula: 2-2-2-2. Palpus with embolic membrane well developed; paracymbium relatively simple, robust distally; tegulum not well developed with sinuous edge, embolus long and filiform (Figs. 3, 4). Female (Holotype, MLP 15980): See Millidge (1991: 52). Sexual dimorphism. Males and females differ only slightly in their somatic morphology. Females are slightly larger than males, mainly due to their larger and more globular abdomen. The cephalic/ocular region is somewhat higher in males than in females. Variation. Males (n=4): Total length 3.80–4.62; carapace length 1.49–1.87; carapace width 1.10–1.28; sternum length 0.82–1.02; sternum width 0.62–0.85; abdomen length 2.20–2.56; abdomen width 0.77–0.95. Female (n=14): Total length 4.00–5.31; carapace length 1.54–1.90; carapace width 1.05–1.45; abdomen length 2.46–3.48; abdomen width 1.10–2.06. In both males and females, the dorsal coloration of abdomen may have pink-silvery spots. Geographic distribution. Dubiaranea difficilis was only known from General Guido, which is the type locality (Mello-Leitão 1944; Millidge 1991), and Reserva Natural Otamendi (Grismado 2007), both in Buenos Aires province, Argentina (Fig. 7). The 13 new records here provided revealed that D. difficilis has a much more extensive distribution indeed. This species has been recorded in three different sectors in Argentina: (1) the Pampean sector, comprising both the type locality and the record for Reserva Natural Otamendi, corresponding to the “Pampa Inundable” (flooding Pampas) and “Deltas e Islas del Paraná” eco- regions respectively (Brown et al. 2006) (southeastern area in figure 7); (2) the central Sierras sector, with three records in the Sierras Chicas and one in the Sierras de Comechingones of Córdoba and San Luis Provinces respectively (Fig. 7), bearing sierra Chaco vegetation; and (3) the northwestern sector, with most records corresponding to mountain forests and rainforests (Yungas eco-region), from Tucumán, Salta and Jujuy (Fig. 7). In the distribution model, the presence is set over the 0.213 logistic threshold, and the algorithm converged after 220 iterations. The potential range of D. difficilis under the model parameters is displayed in figure 7. The map shows that this species has a distribution potentially larger than the actual occurrences of Argentina, and suggests different probabilities from each sector. The model assigns the highest suitability in the northwestern sector, matching the Yungas area. Another sector with good probabilities corresponds to the Sierras de Córdoba. The Pampean sector shows low suitability in spite of having records. According to the model, the species might reach Tarija in Bolivia, and eastern São Paulo and southern Minas Gerais in Brazil (Fig. 7). In general, the predicted range looks quite consistent with the expected distribution, considering the known records and the ecoregions involved. However, there is one meaningful region that is unlikely to be inhabited by the species, but is predicted by the model as present: the xeric valleys west of the Aconquija chain in Salta, Tucumán and Catamarca (Fig. 7). Both sides of this mountain are markedly contrasting, the east-faced slopes bearing dense yungas vegetation, so these results should be taken with caution. It should be noted that the model assigns a similar low probability to extensive regions in the Pampas, among them the two sites in Buenos Aires Province with actual records. The AUC evaluation values from 20 stochastic iterations ranged from 0.9777 to 0.9953 (mean = 0.9849); thus, individual and average AUC values resulted a high accuracy and good performance model. These values are comparable with performances obtained by Echarri et al. (2009), and Boubli & de Lima (2009).

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FIGURE 7. Locality records (yellow circles) and potential predictive range (blue colored areaa) of the spider Dubiaranea difficilis (Mello-Leitão 1944)(Linyphiidae) in Argentina (Northwest, central Sierras and Pampean sectors), Bolivia and Brazil. The arrow indicates the type locality. The darkest color indicates highest probability assigned by the MAXENT model (four intervals: 0.98–0.85; 0.85–0.60; 0.60–0.43 and 0.43–0.21). (BA = Buenos Aires Province; Co = Córdoba Province; Ju = Jujuy Province; Sa = Salta Province; SL = San Luis Province; Tu = Tucumán Province).

Discussion.

Both the actual records of D. difficilis and the modeled area show a remarkable distribution feature: the species seems to spread over a large and heterogeneous area, which makes difficult to get a simple definition on its environmental requirements. On one side, the presence of this spider in the Yungas area and in the central sierras may be explained by geographic proximity and topographical similarity. While the Yungas are humid and dense rainforests (Brown et al. 2006), the original sierra-forest vegetation has been since long devastated in the central sierras (Luti et al. 1979), although some forested patches with humid habitats still remain. This might be in accordance with our observations on the preference of D. difficilis for slightly humid and shady conditions (G.D.R. pers. obs.). In contrast, the two records in Buenos Aires Province —especially the type locality, in the middle of a treeless Pampa— seem not as easy to account for. Even the modeling yielded curious results there, with one point actually outside the suitable area, the other in a very low probability sector. This leads to speculate that these points might represent peripheral populations of the species. In any case, it should be kept in mind that the model is based on climatic features alone (i.e., on a subset of the fundamental niche; Guisan & Zimmermann 2000; Austin 2002) and neither biological (e.g., vegetation type or coverage) nor topographical variables have been computed. At this point one can wonder whether D. difficilis, with records spreading over approximately 2000 km and embracing such different regions, represents a case of wide-range species (like similar cases mentioned in the literature, for example in Ott & Brescovit 2003), or some kind of dispersal

60 · 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. ability or accidental introduction, or yet a combination of those phenomena, are involved. In particular, the aerial dispersal of linyphiid spiders by ‘ballooning’ on silk threads is well documented (Blackwall 1827; Duffey 1998; Thomas et al. 2003). Members of this family are known to be able to travel considerable distances, as linyphiids are among the first colonists of newly available habitats, including reclaimed land (Meijer 1977). The geographic area that embraces the distribution modeling of D. difficilis is consistent with the species records. Although this seems to be an obvious aspect because the model uses the information of the same records, the species was first known in some localities that fall within the area assigned of low probability of occurrence. It is likely that particular habitat conditions in these sites allowed the species inhabiting these places (pampas in Buenos Aires Province as mentioned before) and that such conditions are not fully detected by the climatic scale used by the model. With data at hand it is not easy to corroborate the accurate range limits of the species neither the forces driving its dispersion. However, if everything above is true the pampas are likely to represent the southernmost distribution limit of D. difficilis, and knowing the environment and habitats where D. difficilis was found, we consider that the model is relevant to the species. So we think that it is necessary to address samplings in the South East of Brazil where the species is modeled, and/or to look for specimens evidence that could be deposited in collections of this country.

Acknowledgments

We thank Cristian Grismado (MACN-Ar, Argentina) for his kind help in providing information of localities; to Gustavo Hormiga (George Washington University, USA) and an anonymous reviewer for the constructive comments; to José Corronca (IEBI-Universidad Nacional de Salta, Argentina) for enabling us to study samples from Salta Province obtained during the project directed by M.C. Coscarón (P.I.P. Nº 5261, Argentinean Consejo Nacional de Investigaciones Científicas y Técnicas - CONICET). This work was supported by a research scholarship given to G.D.R. by CONICET and by CNPq (Brazil) to E.N.L.R. Additional funding was given to L.E.A. by CONICET (P.I.P. Nº 6319) and SECyT-UNC (Secretaría de Ciencia y Técnica, Universidad Nacional de Córdoba, 2008-2009). L.E.A. is a researcher of CONICET. This contribution is a part of the Ph.D. Thesis of G.D.R. (in course, at Universidad Nacional de Córdoba under advice of L.E.A.).

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