Comments on the genus Diplura C. L. Koch, 1850, with description of two new species (Araneae, Mygalomorphae, Dipluridae)

Abstract Two new species of Diplura C. L. Koch 1850 are described from Brazil: Diplura mapinguari sp. n., from the state of Rondônia in southeastern Amazonia, northern Brazil, and Diplura rodrigoi sp. n., known from southeastern and central west regions of Brazil. Diplura rodrigoi sp. n. is morphologically similar to D. lineata (Lucas, 1857), D. sanguinea (F. O. Pickard-Cambridge, 1896), and D. mapinguari sp. n. Comments on diagnostic characters of Diplura are included. The synonymy of D. maculata (Mello-Leitão, 1927) with D. catharinensis (Mello-Leitão, 1923) is corroborated. A classification of color pattern of the dorsum of the abdomen is given.


Introduction
Diplura C. L. Koch, 1850 is a Neotropical mygalomorph genus currently including 17 species distributed from Panama to Argentina World Spider Catalog 2018). Most species were described from southeastern and southern Brazil, with additional records from Panamá and numerous South American countries: Argentina, Bolivia, Colombia, Ecuador, Paraguay, and Venezuela . Recently, we have examined many specimens of several species from Peru. The genus is currently recognized by the following combination of characters : simple lyra, formed by a single series of strong and thickened setae, with tip spatulate and not curved (Maréchal and Marty 1998, Drolshagen and Bäckstam 2011, Pedroso and Baptista 2014, legs with thin and restricted scopula (Drolshagen andBäckstam 2011, Pedroso andBaptista 2014), tarsi with only a few cracks (Drolshagen andBäckstam 2011, Pedroso andBaptista 2014) and males with short and thickened palp tibia (Pedroso and Baptista 2014).
Herein we describe two new species of Diplura, one from the state of Rondônia, southeastern Amazonia, northern Brazil, and the other from southeastern and central west Brazil, both based on male and female specimens.

Materials and methods
The description of color pattern is based on specimens preserved in 75% ethanol. Information and photos of living animals were included, when available. Observations, photographs and measurements were made with a Leica DFC295 camera attached to a Leica M205C stereoscopic microscope. All photos were edited in the Photoshop CS5 software and plates were prepared with CorelDraw X7 software. Measurements are given in millimeters, unless otherwise noted. Scale bars represent 1 mm, unless otherwise noted. Body length was measured from the anterior margin of the chelicerae to the posterior border of the abdomen, without spinnerets. Carapace length was measured from anterior margin of the clypeus to the posterior border. Each article of the pedipalp and legs was measured in retrolateral view, from the basal condylus to the distal one. The receptaculum seminis (spermathecae) was examined through dissection of the genital region of the females, cleaned, and immersed in clove oil for clearing. Geographical coordinates were obtained from Geonames (2018). The distribution map was created using ESRI ARCGIS 10 software.
Abbreviations. Institutions (and curators): The specific name is taken from the folklore of Amazonian Indian tribes. "Mapinguari" is a magical creature, a huge long-haired animal, with long arms and large claws.
Diagnosis. Diplura mapinguari sp. n. shares the same oblique continuous light stripes on the dorsum of the abdomen with other species (D. lineata, D. sanguinea, and D. rodrigoi sp. n.), but differs by having the stripes unequally spaced. The three median stripes of D. mapinguari sp. n are wider, almost touching each other near the middle and merging with the ventral light background (Fig. 1). The male of D. mapinguari sp. n. has the longest embolus in Diplura, 2.5× longer than the bulb (Fig.  5). Females also have an elongated receptaculum seminis, with a thin stalk, and only three lobes: the internal lobe is lateral and larger; the other two are a pair at the apex (Fig. 10).
Female (paratype, IBSP 12336). Measurements: Body length 28.1; carapace length 11.3, carapace 9.2; abdomen length 12.3, abdomen width 8.7. Leg formula 4123, total length: I 37.3, II 34.7, III 35.2, IV 44.8. Females resemble males, except by the following characteristics. Carapace: Clypeus length around 1/2 the diameter of AME, with anterior margin bearing four thick setae, elongated, and turned forward. Eye tubercle with three elongated setae at anterior margin and five setae between the posterior eyes. Chelicerae with 12 promarginal teeth. Maxilae with 26 cuspules. Lyra with 13 elongated setae, slightly curved medially. Genitalia: Receptaculum seminis ( Fig. 10) paired, elongated, separated by a little less than its height, with thin and elongated stalk, bearing one large and elongated internal lobe and two distal lobes that are not as large as the basal one.
Color pattern. Carapace reddish brown, with thoracic sulci a bit darker. Eye region black. In live juveniles, the carapace presents an almost black color and the colors of all body parts are more vivid (Fig. 1). Chelicerae, labium, sternum and coxae reddish brown. Legs mostly reddish brown, with darker femurs. Abdomen black, dorsum with five light transverse stripes, of different sizes and shapes (Fig. 1). First and last stripes shorter and thinner (disappearing in some specimens). Three median stripes wider and shorter, widest at their middle region, where they almost touch one another. Compared to the last median stripe, the two first median stripes are wider, longer and are connected to the venter. The carapace is covered with abundant light brown setae.
Distribution. Known only from Porto Velho and Guajará-Mirim, Rondônia state, Amazon area, northern Brazil (Fig. 29). Etymology. This species is named after the biologist, Rodrigo de Cerqueira da Costa, who first directed Denis Pedroso and Alessandro Giupponi in their studies of zoology.
Diagnosis. Diplura rodrigoi sp. n. differs from other species displaying several oblique continuous light stripes on the dorsum of the abdomen (D. lineata, D. sanguinea, and Diplura mapinguari sp. n.) by its very characteristic stripe pattern. Its elongated oblique stripes are very thin near the median line of the dorsum but continuously widening and approaching each other towards the venter, where they merge in the light background (Fig. 25). The three median stripes are longer and not as wide and confluent at their middle portion as in Diplura mapinguari sp. n. In contrast to D. lineata and D. sanguinea, the light stripes are longer and clearly delimited, with well-defined borders. Males of D. rodrigoi sp. n. are the only Diplurinae with a prolateroventral swelling at the distal portion of metatarsus I, which is glabrous and lighter than the surrounding areas (Fig. 13). As in D. sanguinea, males of Diplura rodrigoi sp. n. bear a relatively wide sperm duct from the base of the embolus towards the apex (Fig. 17), in contrast to Diplura mapinguari sp. n. and D. lineata, where the sperm duct is strongly constricted and becomes almost filiform towards the apex. Diplura rodrigoi sp. n. males differ from D. sanguinea by having the megaseta of tibial spur of leg I a bit shorter than the basal portion of the spur (Fig. 12) and 10-11 setae in the lyra (Fig.  19), in contrast with megaseta longer than the spur and seven setae at the lyra in the latter species.
Female (Paratype, UFRJ 0102): Measurements: Body length 26.4; carapace length 9.6, carapace width 8.2; abdomen length 12.3; abdomen width 7.6. Leg formula 4123, length: I 30.4, II 24.8, III 23.7, IV 30.6. Females resemble males, except by the following characteristics. Carapace: Clypeus length similar to AME diameter, with anterior margin bearing five thick setae, elongated and turned forward. Eye tubercle with three thick and long setae and ten smaller setae, of variable size, at anterior margin. Area between posterior eyes with single long and thick setae. AME separated from each other by 80% of their diameter. Chelicera with 12-13 promarginal teeth. Plectrum with five thick and elongated setae. Maxillae with 15-17 cuspules. Lyra (Fig.  19) formed by 11-12 subequal setae, except for the first one, which is shorter and thinner. Setae slightly spatulated, with a thin and pointed apex. Genitalia: Receptaculum seminis (Fig. 20) paired, separated by a distance similar to its own height. Stalk relatively wide, its diameter a little smaller at the distal third, near the five or six apical large lobes, with all of them being similar in size.
Color pattern. Carapace reddish, with thoracic sulci a little darker. Eye region darkened. Chelicerae reddish brown. Labium, sternum and coxae orange brown. Sigillae slightly darker than sternum. Legs light brown. Dorsum of abdomen dark brown with five elongated pale brown transverse stripes, with their width diminishing posteriorly. Stripes continuous, with well-defined borders, thin and far from each other near the midline of the dorsum, but growing wide and closer toward the venter, where they blend into the pale background. When alive, they bear a vivid red carapace and dark brown legs and abdomen.
Natural history. The female paratype from Mendes (UFRJ 0102) was collected in a short tunnel, with a small silk lined opening, on the slope of an old road through second-growth Atlantic Forest. Other specimens have been collected under rocks and logs on forest floor. Most specimens were caught by pitfall traps in forested areas.
Records Distribution: Widespread and common in a wide area of southeastern and midwestern Brazil, from north Espírito Santo state and central-south Minas Gerais state, reaching west to the Distrito Federal and south, down to southeastern São Paulo state (Fig. 29). It is sympatric with D. lineata (Lucas, 1957), a similar species, in the municipalities of Casimiro de Abreu and Rio de Janeiro, both in Rio de Janeiro state, Brazil (see Pedroso et al. 2016).

Notes on the diagnosis and composition of Diplura
The diagnosis of Diplura in relation to other Diplurinae needs some comments and amendments. The presence of a lyra in the inner face of the palp maxilla is a key character that allows for separation between lyrate (Diplura C. L. Koch, 1850, Harmonicon  , 1879). The single line of stiff setae found in lyra of Diplura and Harmonicon is quite different from the plate-like lyra, with several series of setae, in Trechona (Pedroso et al., 2008). In Harmonicon, the lyra is formed by four to seven setae in adult specimens, with a flattened and curved tip (Drolshagen andBäckstam 2011, Pedroso andBaptista 2014). In Diplura, the number of setae in the lyra is quite variable. The described species of Diplura have stiff setae in the lyra ranging from four to seven in Diplura taunayi (Mello-Leitão, 1923) to 15 in Diplura studiosa (Mello-Leitão, 1920). However, we found specimens of Diplura from southern Brazil with up to 18 setae. On the other hand, specimens from southeastern and northeastern Brazil may have only one to three stiff setae in the lyra. One specimen we examined had only one seta in one maxilla, but the other maxilla was devoid of stiff setae. Those findings show that the number of setae in the lyra may increase or decrease in different species, and even the complete absence of the lyra may happen in some specimens. Therefore, as pointed out by Drolshagen and Bäckstam (2011) and Pedroso and Baptista (2014), the number of stiff setae in the lyra is not diagnostic for the genera.
The shape of the stiff setae of the lyra is much more variable than previously pointed out in the literature. In many described species of Harmonicon, including the typespecies Harmonicon rufescens F. P.-Cambridge, 1896, the setae in the lyra are strongly hook-shaped, with a strong distal curvature and an acute and not widened tip (F.  fig. 6), similar to some undescribed species we have examined from Brazil. The lyra of one of those undescribed species has a very wide and clearly spatulated tip. On the other hand, there are species of Diplura with curved stiff setae (ex. D. mapinguari sp. n., Fig. 9) and the tip not clearly spatulated (see D. lineata, Pedroso et al. 2016: fig. 22-24, and D. rodrigoi sp. n., Fig. 19). Therefore, we consider that the variation found on the shape of the stiff setae of the lyra in both Diplura and Harmonicon are too wide and clearly overlapping.
Besides the lyra, there are also other diagnostic characters in the maxilla. On its ventral face, there is a transversal suture just below the lyra. In Diplura, the area just below the basal portion of the maxillar suture is either glabrous or has just some small thin setae (Figs. 9, 19). On the other hand, there is a field of spiniform setae in that area in Harmonicon (Pedroso and Baptista 2014: fig. 6) and Trechona (Guadanucci et al. 2016: fig. 1g). Additionally, there is also a dense fringe of long and thin setae between the lyra and the basal portion of the suture in Trechona (Guadanucci et al. 2016: fig. 1g).
Specimens of Diplura have relatively short and thin scopula, sometimes almost absent in the metatarsus and tarsus of the legs. Scopulae in the other lyrate Diplurinae are formed by long numerous setae and cover most of the distal articles of the legs. In Diplura, the tarsi of the legs are not pseudosegmented and not very flexible, lacking large and numerous cracks; they have only a few ventral, thin cracks, that allow just a limited bending of the article (Drolshagen andBäckstam 2011, Pedroso andBaptista 2014).
An easy way to separate males of Diplura from other Diplurinae is by the short and incrassate pedipalp tibia (Fig. 21), which contrasts strongly with the elongated and thin tibia of other Diplurinae (Fig. 22). The ratio length/width of the pedipalp tibia reaches 3.6× in Diplura and is at least 5× in the other Diplurinae. This character was already pointed out by Pedroso and Baptista (2014) and Pedroso et al. (2016).
The receptaculum seminis of most Diplura species is formed by a short and relatively wide stalk topped by a cluster of many globular to ovoid lobes (fundus), of variable sizes (Fig. 20, Pedroso et al. 2016: fig. 13, 25). In D. mapinguari sp. n., the stalk is elongated and thinner and the lobes are separated in two groups (Fig. 10). In Harmonicon (Drolshagen and Bäckstam 2011: fig. 5) and Trechona Baptista 2004: fig. 1, Pedroso et al. 2008: fig. 4), there are usually two branches: the first one simple and at the inner side of the stalk, and the second one distal and topped by one large fundus, sometimes with lateral flaps. However, there are no clear multiple lobes as in Diplura.
Currently, 17 species of Diplura are considered as valid, distributed from Panama to Argentina World Spider, Catalog 2018). However, Diplura maculata (Mello-Leitão, 1937) has been already synonymized with Diplura catharinensis (Mello-Leitão, 1927) by Bücherl (1962: 261). Although this synonymy had been completely overlooked in the literature and catalogs, we agree with Bücherl after examining the types and additional material from the type-localities or nearby areas in Santa Catarina state, southern Brazil. As a result, Diplura now includes 18 species, taking into consideration the synonymy referred to above and the two new Brazilian species herein described.
Most Diplurinae species have a pattern of light stripes or dots over the dark dorsum of the abdomen. The exceptions are Harmonicon, and some species of Diplura, whose abdomen are uniformly dark, without a contrasting light pattern. The light pattern of most Diplurinae may be present only at the sides of the dorsum, but usually extend either to the middle area of the dorsum or to the venter, even merging with the background color in that area. The species of Diplura may be grouped below in six color pattern types following the different markings on the dorsum of abdomen, numbered from I to VI (Figs 23-28). These pattern types are an easy way to group species and also allows a clear distinction between these groups or isolated species.
Type IV (Fig. 26): Pattern formed by elongated transverse light blotches, sometimes more or less restricted to the sides of the dorsum: D. garbei (Mello-Leitão, 1923), D. petrunkevitchi (Caporiacco, 1955), D. sanguinea (F. O. Pickard-Cambridge, 1896). Type V (Fig. 27): Pattern formed by irregular large light spots, with most of them connected to one another, forming an irregular longitudinal stripe on the sides of the dorsum: D. lineata (Lucas, 1857). Type VI (Fig. 28): Pattern formed by three irregular very wide light spots, widest at their middle region, where they almost touch one another: D. mapinguari sp. n.
The only described species of Diplura not listed above is D. garleppi (Simon, 1892), from San Mateo, Bolivia. As its holotype has not been found in the Museum national d´Histoire naturelle, Paris (E. Leguin, pers. comm.), the short description does not give enough detail on the color pattern and we have not examined any specimen ascribable to D. garleppi, we were not able to include it in any of the proposed color pattern types.