Strange new spiders: on Roddenberryus , a new and unusual caponiid genus (Araneae, Caponiidae)

. The new nopine genus Roddenberryus gen. nov. is erected to include in total ﬁ ve species: three new species, R. kirk gen. et sp. nov. (male and female) from Costa Rica, R. spock gen. et sp. nov. (female) from Campeche, Mexico and R. mccoy gen. et sp. nov. (male) from Baja California Sur, Mexico, as well as two species previously misplaced in Caponina Simon, i.e., R. sargi (Pickard-Cambridge, 1899) gen. et comb. nov. from Guatemala and R. pelegrina (Bryant, 1940) gen. et comb. nov. from Cuba. A male specimen reported as C. sargi Pickard-Cambridge from Costa Rica by E. Kritscher (1957) is assigned to Roddenberryus kirk together with one female collected at the same locality. The new genus is characterized by the distally projected endites and a triangular, projected labium, a conformation unique among nopines; also by a triangular, very short, scaly gladius, serrula with interspersed multiple tooth rows, and a tarsal organ with strongly projected margins. Roddenberryus shares with Tarsonops Chamberlin the cracked tarsi and metatarsi, with multiple adesmatic joints intertwined on the cuticle and an unusual internal respiratory system with both posterior tracheae fused in a single trunk.

of setae with elongated sockets in Tisentnops Platnick, 1994, or the presence of a projected distally clypeal horn in Nasutonops Brescovit & Sánchez-Ruiz, 2016 (Brescovit & Sánchez-Ruiz 2016). Nonnopines show a great variety in the number of eyes, ranging from eight, as in Calponia Platnick, 1993 andin Caponia Simon, 1887, to the eyeless troglobitic genus Carajas Brescovit & Sánchez-Ruiz, 2016(Brescovit & Sánchez-Ruiz 2016. Despite this wide morphological variety, some structures appear to be conservative across caponiid taxa. For example, ultrastructures such as the bases of the trichobothria, the tarsal organs, the serrula, the sensilla, the cheliceral membranous lobes and some types of bristles are morphologically very similar in both nopinae and non-nopine genera. Also, the structures of the internal respiratory system are homogeneous in almost all known genera of caponiids. Among the leg membranous modifications of Nopinae, the crista and arolium show the greatest morphological variety among genera, species and even between sexes, as in species of Aamunops Galán-Sánchez & Álvarez-Padilla, 2022, where the crista is sexually dimorphic (Galán-Sánchez & Álvarez-Padilla 2022). However, the morphology of the gladius is almost constant in all nopines, except for representatives of Tarsonops Chamberlin, 1924 (see Sánchez-Ruiz & Brescovit 2015: fig. 50).
Studying the internal respiratory system of a wide range of caponiids, specimens of three undescribed nopine species from Mexico and Costa Rica were found to have the two posterior tracheae fused in a unique large tracheal trunk. These specimens also bear modifications in some structures that generally do not change in caponiids, such as the tarsal organ, the gladius and the serrula. Additionally, the morphology of their endites and labium are unique among the genera of Nopinae. This paper proposes the new nopine genus Roddenberryus to accommodate these three new Central American species as well as two species previously misplaced in Caponina Simon, 1892 (C. sargi Pickard-Cambridge, 1899 and C. pelegrina Bryant, 1940). In addition, male and female specimens from Costa Rica ascribed to C. sargi by Kritscher (1957) are assigned to Roddenberryus kirk gen. et sp. nov. Some aspects of the morphological diversity of the new genus and its closest relatives are discussed.

Repositories
The specimens examined were supplied by the following collections (curators in parentheses):

Morphological examinations
Observations were made using a Leica MZ12 stereo microscope. Coloration patterns are described based on specimens preserved in 80% ethanol. Internal female genitalia were dissected, and 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. Descriptions and terminology for copulatory structures mostly follow  and Sánchez-Ruiz & Brescovit (2018), but some terms for the female internal genitalia are newly introduced. Measurements are in millimeters (mm) and were made using a microscope micrometer eyepiece. Digital SEM micrographs were taken using a Quanta 250 scanning electron microscope with attached SLR digital camera at Instituto Butantan (São Paulo, Brazil) and a Tescan Mira3 scanning electron microscope at Museu Paraense Emilio Goeldi (Belém, Brazil), following standard procedures for sample preparation. Multifocal photos were taken with a Leica MC170 HD digital camera attached to a Leica M205 C stereo microscope, using Leica Application Suite ver. 4.10 software at Museu Paraense Emilio Goeldi.

Geographical distribution
Distribution maps were generated with ArcView ® ver. 9.0. Locality coordinates are in parentheses when obtained from specimen labels, and square brackets when inferred from Google Maps ® or converted to the DMS format (degrees, minutes and seconds). All figures were edited using Adobe Photoshop CC 2015 ver. 5.1.

Diagnosis
Members of Roddenberryus gen. nov. can be distinguished from all non-nopine genera by the presence of tarsal adesmatic joints (Figs 1H,5G,9H,12H) and from all other Nopinae by a conformation of the unique characters of the endites and labium among these genera, in which the endites have an accentuated finger-shaped forward projection and the labium is triangular and projected (Figs 1E, 2C,  3C, 4C, 8F, 9E, 12E, 14C). Additionally, members of Roddenberryus gen. nov. can be distinguished by the triangular, very short, scaly gladius (Figs 5G, H, 9H, 12H), strongly projected tarsal organ margins ( Fig. 6E-F) and the serrula composed by interspersed multiple rows of teeth (Fig. 4F).

Etymology
The generic name, masculine, is a patronymic honoring Eugene Wesley Roddenberry Sr, the creator of Star Trek, a science fiction media franchise that inspired generations of kids to pursue scientific careers.  (Bryant, 1940) gen. et comb. nov.

Description
Caponiids with only two eyes (Fig. 1D). Carapace orange, elongate oval, widest at rear of coxae II, only gradually narrowed anterior of eyes, without conspicuous pattern (  External female genitalia with a sclerotized anterior plate ( Fig. 11A, C, E); posterior plate narrower, sclerotized; external sclerotization around spiracles (ess) pointy anteriorly on anterior spiracles and tear-shaped on posterior spiracles (Fig. 11A, C, E). Internal female genitalia consisting of a transverse, sclerotized, anteriorly directed posterior receptacle (pr) or interpulmonary fold, a presumed uterus externus (ue) and a membranous anteromedian receptacle formed by a membranous duct at base (arb) protruding anteriorly from bursa, leading to large, oval, membranous sac-like structure (sac) (  2. Embolus short, not reaching half the length of tegulum; elongated tibia, width less than half the length (Fig. 8G-H)  5. Apical sector of tibia with dense tuft of black setae (Fig. 14A) .........R. pelegrina (Bryant, 1940) - . Additionally, males can be recognized by the long embolus, with a length greater than half the length of the tegulum, and a thick tibia, width of which is almost equal to its length ( Fig. 2D-F).

Variation
The Costa Rican males and females from San Jose (Figs 2A-F, 3A-F) were identified by Kritscher (1957) as Caponina sargi, but they actually belong to Roddenberryus kirk gen. et sp. nov.; these specimens have dorsal and ventral patterns of several dark gray stains on the abdomen (Figs 2A-B, 3A-B), which does not appear in any of the R. kirk specimens examined from Guanacaste, Costa Rica (Fig. 1A).

Female
Unknown.

Distribution
Known only from the type locality in Baja California Sur, Mexico (Fig. 15).  (Fig. 9D), but can be distinguished by its larger size and by having several adesmatic joints occupying almost the entire tarsi and almost a half of the metatarsi (Fig. 9H). Additionally, members of R. spock gen. et sp. nov. lack a dense tuft of black setae on the apical part of tibia.

Etymology
The specific name refers to the fictional character S'Chn T'Gai Spock, a Vulcan/Human hybrid from the Star Trek universe, who served aboard the starship USS Enterprise as the chief science officer, played in the original series by American actor Leonard Nimoy.

Distribution
Known only from Campeche and Quintana Roo in Mexico (Fig. 15).

Diagnosis
Rodeenberryus sargi resembles R. spock gen. et sp. nov., R. mccoy gen. et sp. nov. and R. pelegrina by having invaginations in the margins of the clypeus (Fig. 12D), but can be distinguished by its smaller size and by having a few adesmatic joints occupying only the last third of the tarsi and the last third of the metatarsi (Fig. 12H); females can be distinguished by the epigastric furrow having a median invagination and rounded laterals (Figs 11E, 12F).

Diagnosis
Roddenberryus pelegrina resembles R. spock gen. et sp. nov., R. mccoy gen. et sp. nov. and R. sargi gen. nov. by having invaginations in the margins of the clypeus (Fig. 14A), but can be distinguished by having a dense tuft of black setae on the apical part of the tibia (Fig. 14A).

Type material
Holotype CUBA • ♀; Santiago de las Vegas; N. Banks leg.; MCZ 22577. (The holotype was examined and photographed in 2002 by the first author; further attempts to examine this specimen were unsuccessful.)

Description Male
Unknown.

Distribution
Known only from the type locality in Cuba (Fig. 15).

Discussion
The descriptions of new species and genera in Caponiidae have been based primarily in male specimens, because most of the diagnostic characteristics came from the male palpal morphology. However, female specimens from the new genus Roddenberryus have unusual characteristics for a caponiid, and even species can be differentiated only with females. At first glance, members of this genus could be recognized  (Bryant, 1940)  as belonging to a large group of nopine genera, characterized by a narrowly ovate carapace and a flattened pars thoracica (Aamunops, Nops MacLeay, 1839, Orthonops Chamberlin, 1924, Medionops Sánchez-Ruiz & Brescovit, 2017and Nopsides Chamberlin, 1924see Sánchez-Ruiz & Brescovit 2018: fig. 77 and Galán-Sánchez & Álvarez-Padilla 2022: fig. 81). However, the presence of adesmatic metatarsal joints in Roddenberryus gen. nov. complicates this assessment, since this character is also shared by members of Tarsonops and Cubanops Sánchez-Ruiz, Platnick & Dupérré, 2010, which have a broad, almost subcircular carapace and an elevated pars thoracica. These three genera also share the presence of an anteromedian receptaculum in the female internal genitalia. Thus, members of Roddenberryus, sharing characteristics with representatives of both groups of nopines, may be of great importance to unveiling the relationships among nopine genera.
Despite the narrowly ovate carapace, Roddenberryus shares with Tarsonops the tarsi and metatarsi cracked with several adesmatic joints intertwined on the cuticle, as well as the unusual respiratory system with a single tracheal trunk on posterior spiracles. These are the only two nopines with this combination of characters. The other nopines with metatarsal adesmatic joints are Cubanops and Aamunops, but representatives of those genera have only a few closely spaced ademastic joints on the lower half of metatarsi IV, which make this segment appear to be divided (Sánchez-Ruiz et al. 2010: fig. 131). On the other hand, the respiratory system of nopines usually has two tracheal trunks protruding from the posterior spiracles (Sánchez-Ruiz & Brescovit 2017: fig. 4J-L) rather than just a single tracheal trunk, as occurs in Tarsonops and Roddenberryus. Therefore, at least two primary lineages (Tarsonops and Roddenberryus) in the assemblage of Nopinae are defined by basic differences in the metatarsal cuticle continuity on all legs and the internal respiratory system. Considering this, we could assume that members of Roddenberryus would be much closer to the group that includes Tarsonops (broad,  (Bryant, 1940) comb. nov. (star). subcircular carapace and elevated pars thoracica) than to the group with a narrowly ovate carapace and flattened pars thoracica.
Another unusual characteristic among nopines is the presence of an anteromedian receptacle in the female internal genitalia. The only nopines possessing an anteromedian receptacle are Roddenberryus, Tarsonops and Aamunops; they generally resemble those shown by the non-nopine genera Tisentnops, Caponia, Diploglena Purcell, 1904 andCarajas (Brescovit &Sánchez-Ruiz 2016;Haddad 2015;Dippenaar-Schoeman et al. 2020). This structure is composed of a membranous duct (base) leading to an oval membranous sac-like structure. All other nopines do not have an anteromedian receptacle, and the internal genitalia are limited to the presence of a transverse, sclerotized posterior receptacle and a presumed uterus externus (Sánchez-Ruiz & Brescovit 2017, 2018. Roddenberryus is readily recognized by four characteristics exclusive among nopines. Representatives of this new genus have a triangular, very short, scaly gladius, which sets them apart from most nopines with a short-sword gladius (Sánchez-Ruiz & Brescovit 2018: fig. 21l). This structure is morphologically constant within the nopines, and only in Tarsonops is this structure shown as a large, wide, semicircular shape (see Sánchez-Ruiz & Brescovit 2015: fig. 50). Additionally, the new genus presents strongly projected tarsal organ margins and a strong distal serrula, consisting of interspersed multiple tooth rows. In all other nopines, the tarsal organ is exposed, roundish, with the marginal ring only slightly pronounced, and the serrula consists of a single tooth row (Sánchez-Ruiz & Brescovit 2018: figs 21c, 22e). The conformation of the endites and labium are also unique for species of Roddenberryus; the endites have an accentuated finger-shaped forward projection and the labium is triangular and projected. Indeed, endites and labium shape in Nopinae are quite remarkable, since each genus apparently has a unique combination.