Three new species of the spider genus Nopsma (Araneae, Caponiidae, Nopinae) from Colombia

Three new Colombian species of the spider genus Nopsma Sánchez-Ruiz, Brescovit & Bonaldo, 2020 are described and illustrated: Nopsma leticia sp. nov. (male) from Amazonas department, Nopsma macagual sp. nov. (male) from Caquetá department and Nopsma paya sp. nov. (male and female) from Putumayo department. The collection data of the holotype of Nopsma florencia Sánchez-Ruiz, Brescovit & Bonaldo are corrected. Additionally, an updated identification key for all species of the genus and a distribution map for the Colombian species are included.


Introduction
The two-eyed spider genus Nopsma was recently proposed by Sánchez-Ruiz et al. (2020) for four species occurring in Colombia, Ecuador, Peru and Nicaragua. The Ecuadorian species Nopsma juchuy (Dupérré, 2014) was transferred from Nyetnops Platnick & Lise, 2007 and was elected as the type species. Members of this genus can be distinguished from other two-eyed Nopinae (except Nyetnops), by lacking a crista and an arolium on anterior metatarsi, and from Nyetnops by the presence of a gladius. Additionally, Nopsma species are characterized by having an elongated, prolaterally protruded embolus, unique among nopines, and by the shape of endites, without projected outer sides on anterior margin (Sánchez-Ruiz et al. 2020).
Members of Nopsma are poorly known in Colombia, being represented by only one species, N. florencia Sánchez-Ruiz, Brescovit & Bonaldo, 2020. The species was described from male specimens revised and photo-graphed prior to the publication by the second author of this paper (L.M.), who unfortunately confused the type´s data label with those of other Colombian Nopsma specimens from Macagual in Caquetá department, while the actual locality of N. florencia is in Chocó department. The collection data of this Colombian species is corrected below. On the bright side, this mislabeling event led to the discovery of a new Nopsma species, since the specimens from Macagual belong to an undescribed species. This finding also triggered the opportunity to review caponiids from three Colombian collections, leading to the discovery of two additional undescribed Nopsma species. These three species are herein described, along with the correction of the type locality of N. florencia. Photos of the habitus, male palpal morphology and female genital organs, line drawings of the male copulatory bulbs and schematic drawings of the female internal genitalia are provided. Additionally, a distribution map for all Colombian species and an updated identification key for all species of the genus are presented.
Morphological observations and illustrations were made using a Leica MC125 stereomicroscope with a camera lucida. Multifocal images were taken with the Leica MC-190 HD and Leica MC-170 HD digital cameras attached to Leica S8AP0 and Leica MC125 stereomicroscopes respectively with extended focal range. All multifocal images were assembled using Helicon Focus Pro ver. 5.3.14. The measurements are in millimeters (mm) and were made using an ocular micrometer. Descriptions and measurements follow Sánchez-Ruiz and Brescovit (2018). Coloration patterns were described based on specimens preserved in 70-80% ethanol. The internal female genitalia was dissected with fine forceps and their soft tissues were digested for 24 hours with Ultrazyme enzymatic eye lens cleaner, diluted with distilled water at the proportion of 1 tablet/5 ml. After cleaning, samples were immersed in clove oil for visualization of internal structures. The terminology for copulatory structures follows Sánchez-Ruiz et al. (2015). All digital photos were edited using Adobe Photoshop CS ver. 12.0 and the distribution map was prepared in QGIS (QGIS 2021). Plates were edited with Corel Draw X7 ver. 17.1. Geographic coordinates were extracted from original labels. Locality elevations refer to meters above sea level.
The following abbreviations are used in the text and figures: ap = anterior plate, as = anterior tracheal spiracles, dmr = distal margin of receptaculum, e = embolus, ess = external sclerotization around spiracles, go = genital opening (gonopore), mk = membranous keel on embolus, re = receptaculun, pmr = proximal margin of receptaculum, pp = posterior plate, ps = posterior tracheal spiracles, t = tegulum.  Remark. The type locality of this species is here corrected. The data labels of the holotype and paratype reported in the original description are actually those belonging to Nopsma macagual sp. nov.
Diagnosis. Males of Nopsma florencia resemble those of Nopsma leticia sp. nov. by the similarly shaped membranous keel on embolus tip (Fig. 6A, B, E, F), but can be distinguished by the conspicuous small oval tegulum, with only one-fifth the cymbium length (Fig. 1C, D),  Distribution. Known only from the type locality in Chocó, Colombia (Fig. 8).
Preservation status. Preserved in 70% ethanol. Male holotype in good condition, left palp dissected in a separate microvial. Diagnosis. Males of Nopsma leticia sp. nov. resemble those of Nopsma florencia by having a similarly shaped membranous keel on embolus tip (Fig. 6A, B, E, F), but can be distinguished by having a larger tegulum, one-third of the cymbium length (Fig. 2C, D), and by the thicker embolus and membranous keel (Fig. 6E, F Fig. 2A). Chelicerae, palps, sternum, endites, labium and legs orangish brown (Fig. 2B). Abdomen dorsally gray with tenuous light stripes ( Fig. 2A), ventrally yellowish (Fig. 2B). Anal tubercle and spinnerets pale yellowish. Palp with rounded tegulum and a posteriorly directed embolus (Fig. 2C, D), with a long membranous keel, reaching more than onethird of the embolus tip (Fig. 6F). Female: unknown. Etymology. The specific name is a noun in apposition taken from the type locality.
Etymology. The specific name is a noun in apposition taken from the type locality.  Distribution. Known only from the type locality (Fig. 8).
Etymology. The specific name is a noun in apposition taken from the type locality.
Distribution. Known only from the type locality (Fig. 8).
Preservation status. Preserved in 80% ethanol. Male holotype with only a half of the abdomen, left palp dissected in a separate microvial. Female paratype IAvH-I 3796 in good condition, genitalia dissected in a separate microvial.

Discussion
The three new Colombian species of Nopsma described here increase to seven the number of known species of this genus. These new species are probably endemic, being currently known from only one or two localities  within the country. Until now, only the males of Nopsma are well known since the few known female specimens belong to the type species, N. juchuy (Dupérré), which was internally studied by (Sánchez-Ruiz et al. 2020). In this paper we present the internal genitalia of Nopsma paya sp. nov., the second species represented by females within the genus. The internal genitalia of the Nopinae females are weakly sclerotized (Sánchez-Ruiz and Brescovit 2018) and sometimes it is necessary to dissect several females in order to resolve the internal morphology. Generally, internal female genitalia of Nopinae genera tend to be very similar among its species and only differences in the morphology of the receptaculum were observed in previous studies on these genera (see Platnick 1995;Sánchez-Ruiz et al. 2010, 2020Sánchez-Ruiz andBrescovit 2017, 2018). The genus Nopsma seems to be no exception, since the female genitalia of N. juchuy and N. paya are very similar internally, with only few differences in the distal margin of the receptaculum (Fig. 7A, B). The straight distal margin of the receptaculum in N. juchuy (Fig. 7A, Sánchez-Ruiz et al. 2020: fig. 16G-J) appears to be diagnostic, while N. paya sp. nov. shows an invagination on distal margin of receptaculum with both sides slightly sloping (Fig. 7B). Furthermore, comparing the external genital area of both species we found also some discrete differences that are diagnostic for each of these species (see diagnosis of N. paya sp. nov.).
Members of Nopsma lack crista and arolium but retain the gladius on the anterior legs. Sánchez-Ruiz et al. (2020) suggested that this genus could be phylogenetically related to Nyetnops Platnick & Lise and Cubanops Sánchez-Ruiz, Platnick & Dupérré, since these three genera have several common characteristics in the shape and stain patterns of the cephalothorax, sharing a   distinct sub-circular, broad carapace shape, with narrow pars cephalica on dorsal view. Besides, the pars thoracica is elevated near the middle and slops abruptly posteriorly on lateral view. The only known species of Nopsma that diverges from those patterns is Nopsma armandoi Sánchez-Ruiz, Brescovit & Bonaldo (see Sánchez-Ruiz et al. 2020: fig. 17A). This species, however, exhibits the Nopsma diagnostic characteristics mentioned by Sánchez-Ruiz et al. (2020). Thus, although lacking an arolium on the pretarsi, N. armandoi has the cephalothorax similar to that presented by genera of the clade with arolium (Nops MacLeay, Medionops Sánchez-Ruiz & Brescovit and Nopsides Chamberlin). The absence of a membranous keel on the embolus tip is another important characteristic that set apart N. armandoi from other members of Nopsma. All presently known Nopsma species have this membranous keel, however N. armandoi has three, very thin, long projections at the embolus tip (see Sánchez-Ruiz et al. 2020: fig. 17B, C, F). These projections at the embolus tip again relate this species to those representatives from the arolium clade. Therefore, this species must be better studied, preferably with additional specimens, including the females which are currently unknown. An updated phylogeny of Nopinae that includes Nopsma, must address the hypothesis that this species would not belong to the genus, but instead to a hitherto undescribed lineage of Nopinae in which the loss of arolium has occurred independently.