Taxonomic revision and cladistic analysis of ghost spiders of the genus Tafana Simon, 1903 (Araneae: Dionycha, Anyphaenidae), with the descriptions of twelve new species

Abstract. The diversity of the genus Tafana Simon, 1903 is poorly known in the Neotropical regions. In this work we provide a taxonomic review of the genus as well as a phylogenetic analysis. The ingroup of the analysis is composed of sixteen species of Tafana and the outgroup is composed of fi ve representatives of Anyphaenidae. The sister-group recovered for Tafana is the clade Aysha + Xiruana, being supported by the embolic process on the male bulb. Two species groups within Tafana are herein proposed, the silhavyi group and the riveti group, based on two exclusive synapomorphies in the male bulb. We redescribe Tafana quelchi and present a description of the previously unknown female of Tafana silhavyi, both from Venezuela. In addition, we describe the fi rst adult specimens of Tafana straminea. Twelve new species, along with several previously described species, are described, illustrated and mapped: T. riveti, T. straminea, T. quelchi, T. kunturmarqa sp. nov., T. humahuaca sp. nov., T. pastaza sp. nov., T. nevada sp. nov., T. huatanay sp. nov. and T. ruizi sp. nov. from the riveti species group; T. maracay sp. nov., T. arawak sp. nov., T. chimire sp. nov. and T. pitieri sp. nov. from the silhavyi species group; T. oliviae sp. nov. from Argentina and T. orinoco sp. nov. from Venezuela, neither of which belongs to any species group. We also discuss the genital morphology of the species groups based on the results of the phylogenetic analysis. Furthermore, distribution maps for all species, including new records for T. riveti, T. straminea and T. quelchi, are presented.

In the sensitivity analysis using different values of the concavity constant (k) for implied weighting of characters, different k values result in different numbers of MPTs and steps. These results are provided in Table 1.
The analysis under implied weighting resulted in four similar topologies (Fig. 2). The main difference between the trees involves the relationships and monophyly of Tafana orinoco sp. nov. and clade G (Figs 2,5). We decided to choose the tree obtained with k = 7, because it shares the highest number of nodes with the other trees and was considered the most stable (Figs 2,5). The analysis with k = 7 resulted in one MPT with total fi t 27.319 (CI = 0.46; RI = 0.60). Support values and sensitivity of this tree to concavity changes are presented in Figs 3 and 4. The synapomorphies of Tafana and its internal clades are listed in Table 2.  The silhavyi group presents one exclusive synapomorphy Clade D): the laminar shape of the apophysis at the base of the embolus (character 25 [1]), and three homoplastic synapomorphies: tegulum, ventral position, sperm duct an inverted S-shape (character 15 [2]); tegulum, ventral position, secondary horizontal loop of the sperm duct present (character 16 [1]); and long length of the embolus (character 27 [0]). This clade presents RB < 50% (Fig. 3). The riveti group presents one exclusive synapomorphy Clade G): ventral tegular process, positioned near median apophysis (character 19 [1]), and one homoplastic synapomorphy, primary spermathecae contiguous (character 38 [1]), convergent in other clades. This clade presents RB > 50% (Fig. 3).     Simon, 1903 Tafana Simon, 1903:124 (type species by original designation: T. riveti Simon, 1903: 124).

The riveti group Diagnosis
Males of this group resemble those of the silhavyi group by the apophysis at the base of the embolus, but differ by presenting an apophysis at the base of the embolus that is short and conical ( Fig. 12B-C); the sperm ducts are 'U-shaped' in the male palp when it is not expanded (Figs 9B, 10C). The females can be recognized by the short atrium, presenting a septum or not, and copulatory ducts narrow (Figs 10F-G, 11F, 15F-G).

Diagnosis
Males of Tafana riveti resemble those of T. straminea, T. kunturmarqa sp. nov., T. humahuaca sp. nov. and T. nevada sp. nov., by the apophysis at the base of the embolus short, but differ by retrolateral tibial apophysis short and with large base in retrolateral view and visible subtegulum on palp when not expanded ( Fig. 10C-E). Females differ from those of other species by the short atrium, triangular hood and lateral lobe curved in the anterior region and sclerotized (Fig. 10F).

Remark
The holotype is an immature in last instar. In comparison with other adult specimens from Colombia, we noticed that the reddish brown color of the carapace is unique for this species. The spots on the abdomen and spination of the legs are similar. Therefore, we consider here that these specimens represent the adults of the species Tafana straminea.
PALP. Retrolateral tibial apophysis in vertical position long and conical; cymbium with prolateral projection; ventral tegular process short near of the base of the median apophysis; apophysis at the base of the embolus short; sperm duct in vertical position with secondary looping without reaching the apex of the tegulum; median apophysis sharp and curved at the apex (Figs 11C-E, 12A-C). Female (IBSP 258181) COLOR. As in males of this species (Fig. 11B).
EPIGYNUM. Hood circular and sclerotized; epigynal plate with posterior region covered by cuticle; atrium with septum; lateral lobes extremely curved in the anterior region. Internally, copulatory ducts large in the anterior region; secondary spermathecae rounded in middle of the copulatory ducts, oval primary spermathecae; fertilization ducts long (Figs 11F-G, 32G).

Diagnosis
Males of Tafana kunturmarqa sp. nov. resemble those of T. straminea and T. ruizi sp. nov. by having the retrolateral tibial apophysis conical, but differs by having the embolus short and the apophysis at the base of the embolus larger ( Fig. 14C-E). Females resemble those of T. straminea and T. ruizi sp. nov. by having the epigynal plate with atrial septum, but differ by having the secondary spermathecae above the primary spermathecae, rounded, and fertilization ducts smaller than primary spermathecae (Figs 14F-G, 32I).

Etymology
The specifi c name, a noun in apposition, is a tribute to the Pre-Colombian language 'Quechua', originated in the central Andes. The meaning of the expression 'Kuntur marqa' in this language is 'Condor's nest'.
PALP. Retrolateral tibial apophysis conical with apex lightly curved; cymbium with retrolateral projection; short embolus; ventral tegular process near the median apophysis; sperm duct in vertical position with secondary loop reaching the apex of the tegulum; median apophysis long and curved at the apex ( Fig. 14C-E).

Etymology
The specifi c name, a noun in apposition, is a tribute to 'Quebrada de Humahuaca', a deep canyon located in Jujuy Province, the type locality.
PALP. Retrolateral tibial apophysis conical with apex curved; cymbium with retrolateral projection more rounded; embolus short; ventral tegular process near the median apophysis and sclerotized; sperm duct in vertical position with secondary looping reaching the apex of the tegulum; median apophysis slender and curved apically ( Fig. 15C EPIGYNUM. Hood semicircular and sclerotized; short atrium; lateral lobes over copulatory ducts. Internally, copulatory ducts narrow; primary spermathecae large and oval; secondary spermathecae above the primary spermathecae and rounded; fertilization ducts long ( Fig. 15F-G).

Diagnosis
Males of Tafana pastaza sp. nov. differ from those of the other species of this genus by having the apophysis at the base of the embolus bifi d, primary apophysis 'S-shaped' and secondary apophysis large, retrolateral projection of the cymbium keel-shaped ( Fig. 16C-E). Females differ from the others by presenting internal copulatory ducts long and sinuous, with a globose extension proximally of the copulatory ducts (Figs 16F-G, 32K).

Etymology
The specifi c name is a noun in apposition taken from the type locality, Pastaza Valley. Paratype ECUADOR • 1 ♀; same collection data as for holotype; AMNH.

Remark
The male specimen is in very bad shape, having lost the chelicerae and almost all the legs, and with the abdomen partially deteriorated (Fig. 16A).
PALP. Retrolateral tibial apophysis conical and curved, apical protuberance on the tibia in retrolateral view; cymbium with retrolateral projection keel-shaped; embolus long; subtegulum prolateral projected; ventral tegular process not present; sperm duct secondary loop reaching apex of the tegulum; median apophysis thick and curved at the apex ( Fig. 16C-E).
EPIGYNUM. Hood circular; atrium small; epigynal plate without posterior region covered with sclerotized cuticle; lateral lobes chalice-shaped. Internally, copulatory ducts long and narrow; primary spermathecae oval; secondary spermatechae in the middle of the copulatory ducts; fertilization ducts long, two times the length of primary spermathecae (Figs 16F-G, 32K).

Etymology
The specifi c name is a noun in apposition taken from type locality, Sierra Nevada de Santa Marta.

Etymology
The specifi c name, a noun in apposition, is a tribute to the 'Sacred Valley of the Incas' called Huatanay.

Diagnosis
Males of Tafana ruizi sp. nov. resemble those of T. straminea and T. kunturmarqa sp. nov. by having the base of the embolus large and inclined, but differ by having the ventral tegular process large and apex of tegulum not rounded (Fig. 21C). Females resemble those of T. straminea and T. kunturmarqa sp. nov. by having the atrial septum, but differ by having the wide atrium, showing the copulation openings and hood long and circular (Figs 21F-G, 33B).

Etymology
The specifi c name is a patronym in honor of the Cuban arachnologist Alexandre Sánchez-Ruiz for his contributions to the taxonomy of spiders.
Female (paratype, IBSP 258183) COLOR. Coloration as in males of this species, except for the reddish carapace and chelicerae reddish brown and robust (Fig. 21B). Abdomen gray with spots in dorsal view (Fig. 21B).
EPIGYNUM. Hood circular; atrial septum present; epigynal plate covered by cuticle in the posterior region; lateral lobes extremely curved in anterior region. Internally, copulatory ducts sinuous and laminar; primary spermathecae large and oval; secondary spermathecae short and rounded in the anterior region on the copulatory ducts; fertilization ducts long over the primary spermathecae (Figs 21F-G, 33B).

Diagnosis
Males of this group can be recognized by having the apophysis at the base of the embolus curved and laminar (Figs 9C-D, 22C, 23B, 26A-B); the sperm duct forming an inverted 'S-shape' in the male palp not expanded, with secondary looping in the middle of the tegulum (Figs 9A, 22C, 25C). The females can be recognized by having the atrium wide and copulatory ducts large in the anterior region (Figs 22G, 26D, 33D-F).

Diagnosis
Males of Tafana silhavyi resemble those of T. chimirae sp. nov., T. maracay sp. nov., T. arawak sp. nov. and T. pittieri sp. nov. by having the laminar apophysis at the base of the embolus, but differ by presenting a secondary looping with wide curvature (Figs 9A, 22C). Females resemble those of T. maracay sp. nov. and T. arawak sp. nov. by having the wide atrium, but differ by having the copulatory ducts extremely large in the anterior region (Figs 22F-G, 33C). sternum and endites yellow. Chelicerae reddish. Legs I-IV yellow, darker at tarsi. Abdomen gray with dark spots (Fig. 22A).
PALP. Retrolateral tibial apophysis short and curved; cymbium with retrolateral projection; projection at the base of the embolus laminar and coiled; embolus long, with large base and narrow apex; median apophysis long and curved apically; ventral tegular process hyaline and short; sperm duct an inverted 'S-shape' (Figs 22C-E, 23A-C).

Diagnosis
Males of Tafana chimire sp. nov. resemble other males of the silhavyi group by the laminar base of the apophysis of the embolus, but differ from them by the base of the retrolateral tibial apophysis with a hook shape apically and the presence of a large basal prominent protuberance on the tegulum (Fig. 24C-E).

Etymology
The specifi c name is a noun in apposition taken from the name of the type locality, Chimire.
PALP. Retrolateral tibial apophysis short; cymbium with basal retrolateral projection small; apophysis at the base of the embolus laminar and coiled; embolus long with large base and narrow apex; median apophysis long and curved apically; ventral tegular process hyaline apically in the bulb; sperm duct with secondary looping horizontal position (Fig. 24C-E).

Female
Unknown.

Diagnosis
Males of T. maracay sp. nov. resemble those of the silhavyi group by having the apophysis at the base of the embolus laminar, but differ from the other species by having the retrolateral tibial apophysis short, curved apically and presenting a basal projection rounded (Figs 25C-E, 26C). Females resemble those of T. silhavyi and T. arawak sp. nov. by having the epigynal plate with a wide atrium, but differ by having the copulatory ducts slightly narrowing in the anterior region (Figs 25F-G, 33D).

Etymology
The specifi c name is a noun in apposition taken from the type locality Maracay, currently the capital of the State of Aragua in Venezuela. Other material VENEZUELA -Aragua • 1 ♂, 2 ♀♀; same collection data as for holotype; MNHN 11314.
PALP. Retrolateral tibial apophysis short, extremely curved at the apex; cymbium with basal retrolateral projection; apophysis at the base of the embolus laminar and coiled; embolus long with large base and narrow apex; median apophysis long and curved apically; ventral tegular process short and hyaline; subtegulum hardly visible; sperm duct with primary looping large (Figs 25C-E, 26A-C).

Etymology
The specifi c name is derived from the Brazilian Portuguese word 'Aruaques' (in Spanish 'Arahuaco') for the name of an indigenous group from the Americas, named 'Arawak', meaning 'fl our eater'. The name should be treated as a noun in apposition.
PALP. Retrolateral tibial apophysis bifi d with anterior branch truncated and posterior branch thin; cymbium with retrolateral projection; apophysis at the base of the embolus laminar and coiled; embolus long with large base and narrow apex; median apophysis long and curved apically; ventral tegular process long and hyaline; basal protuberance of the tegulum short; sperm duct with secondary looping in oblique position in the tegulum (Fig. 27C-E). Female (paratype, MNHN 11320) COLOR. Exemplar probably discolored. Coloration as in males of this species (Fig. 27B).
EPIGYNUM. Hood triangular; wide atrium; epigynal plate covered with hyaline cuticle in the posterior region; lateral lobes sinuous. Internally, copulatory ducts extremely large in the anterior region; primary spermathecae oval and separated; secondary spermathecae small, lateral of the primary spermathecae, and fertilization ducts longer than the primary spermathecae (Figs 27F-G, 33E).

Diagnosis
Males of Tafana pittieri sp. nov. resemble those of the silhavyi group by having the secondary looping in the middle in the tegulum, but differ from them by having the large laminar, ring-shaped apophysis at the base of the embolus and retrolateral tibial apophysis short and truncated ( Fig. 28C-E). Females differ from the other species by having the lateral lobes wing-shaped in the median region, covering part of the atrium, and secondary spermathecae with slender shape (Fig. 28F-G, 33F).

Etymology
The specifi c name is a patronym in honor of Henri Pittier, a botanist, geographer and ethnologist who devoted himself to the study of the fl ora and fauna in the National Park Henri Pittier, where the specimen was collected. In the year of 1953 the park (until then called Rancho Grande) was renamed in honor of H. Pittier. Paratype VENEZUELA • 1 ♀; same collection data as for holotype; AMNH.
PALP. Cymbium with large retrolateral projection, embolus long with large base and narrow apex; median apophysis long and curved apically; ventral tegular process short and hyaline; tegulum without basal protuberance; sperm duct with secondary looping in horizontal position; median apophysis laminar, long and curved at the apex (Fig. 28C-E).
EPIGYNUM. Triangular hood with sclerotized border; atrium short; epigynal plate not covered by cuticle, covered in the posterior region; lateral lobes sinuous with wing form in the median region. Internally, copulatory ducts short and curved; primary spermathecae oval and separated; secondary spermathecae small in the posterior region of the copulatory ducts; fertilization ducts longer than primary spermathecae (Figs 28F-G, 33F). ABDOMEN. Length 3.2, epigastric furrow 0.6 from tracheal spiracle, spiracle 1.1 from base of spinnerets.

Diagnosis
Males of Tafana oliviae sp. nov. differ from those of the other species in the silhavyi group by having the apophysis at the base of the embolus fi nger-shaped, retrolateral tibial apophysis bifi d with anterior branch short and fl attened, posterior branch triangular and median tibial apophysis bifi d in dorsal position on the tibia (Fig. 29C-E). Females differ by having the epigynal plate with lateral lobes extremely projected to the sides and by having the small primary spermathecae (Figs 29F-G, 33G).

Etymology
The specifi c name is a matronymic in memory of Olivia de Moura Oliveira, the fi rst author's mother.

Remark
The male and female were matched by their close localities in Misiones, Argentina. Despite the different colorations of the male and female, the morphology of the genitalia suggests that both exemplars belong to Tafana oliviae sp. nov.
PALP. Retrolateral tibial apophysis bifi d; cymbium without retrolateral projection; embolus short and bifurcate at the apex; median apophysis long and curved apically; sperm duct with secondary looping reaching the apex of the tegulum (Fig. 29C-E).
EPIGYNUM. Hood semicircular and thick; lateral lobes curved in anterior region; wide atrium; epigynal plate covered by cuticle in the posterior region. Internally, copulatory ducts long and narrow; secondary spermathecae in the anterior region of the copulatory ducts; primary spermathecae small and fertilization ducts longer than primary spermathecae (Figs 29F-G, 33G).

Etymology
The specifi c name, a noun in apposition, is a tribute to 'River Orinoco', one of the most important rivers of the South American continent.
PALP. Retrolateral tibial apophysis short and curved; cymbium without retrolateral projection; apophysis at the base of the embolus bifi d with secondary apophysis furrowed; embolus short; median apophysis long and curved apically; sperm duct with secondary looping not reaching the apex of the tegulum; subtegulum strongly projected; ventral tegular process inconspicuous (Figs 30C-E, 31A-C).
EPIGYNUM. Hood narrow and rounded, connected to the lateral lobes; wide atrium; epigynal plate not covered by hyaline cuticle in the posterior region; sinuous and narrow lateral lobes. Internally, copulatory

Discussion
The genus Tafana is predominantly distributed along the northern and western edges of South America, with some species associated with high elevation regions of the Andes mountain range (Figs 34-35). Tafana oliviae sp. nov., the sister species of all Tafana, occurs in the lowland regions of Misiones, Argentina. The next species to diverge from the main clade, Tafana orinoco sp. nov., occurs in both lowland and high elevation areas like Cerro de La Neblina at 1690 m a.s.l. in Venezuela. One of the major species groups of Tafana, the silhavyi group, includes species that occur in lowland regions of far northern Venezuela. The other, the riveti group, includes species endemic to the Andean region (T. riveti, T. straminea, T. huatanay sp. nov., T. nevada sp. nov.), with those species generally occurring at elevations from 800 to 3600 m, as well as species from the Amazonian region (T. quelchi and T. pastaza sp. nov.). Tafana humahuaca sp. nov., also in this group, occurs in the Argentinian region of Jujuy, at altitudes from 1259 to 4000 m. To fully understand the distributional patterns of the genus, inclusion of more species and a biogeographical analysis are needed.
The monophyly of the genus Tafana is strongly supported, although the internal branches have low support values for several monophyletic groups (Figs 3-4). Our results and topology are corroborated by other works based on morphological data (Ramírez 1995b;Izquierdo & Ramírez 2008;Werenkraut & Ramírez 2009;Soto & Ramírez 2012;Cabra-García & Brescovit 2016). In addition, our results recovered the close relationship of Aysha and Xiruana, based on the features of the embolic process in the male bulb copulatory. The close relationship of these species is also found in other works (Brescovit 1997;Ramírez , 2007Labarque et al. 2015).
Apophysis at the base of the embolus After revising the genus-level taxonomy, we found that all known species are diagnosed by the presence of an apophysis at the base of the embolus (Figs 18A-C, 23A-C, 31A-C). This character is shared by all species of Tafana, but it is not restricted to this genus. Some species of Anyphaeninae, such as Xiruana tetraseta (see Oliveira & Brescovit 2015a: 209, fi g. 5a-b), which were not included in this analysis, also have this structure. In the phylogeny of the genus presented here, T. orinoco sp. nov. and T. pastaza sp. nov. present a bilobed apophysis at the base of the embolus, and in T. orinoco sp. nov. the secondary apophysis ( Fig. 31B-C) has a furrow, which we suggest may function as a conductor. In the silhavyi group, species only have one apophysis, with a laminar shape, while in the riveti group, species also have one apophysis, generally with a conical shape (except in T. pastaza sp. nov.).

Cymbium, retrolateral projection
Another character that is not limited to Tafana is the retrolateral projection on the cymbium (Figs 10C, 12A, 22C, 26A, C). The absence of this character is plesiomorphic for Tafana and the structure is absent in T. oliviae sp. nov., T. orinoco sp. nov. and T. quelchi, although it is present in all other species in the genus. Some other genera of the Anyphaeninae also present some retrolateral projection on the cymbium, as in the genera Katissa Brescovit, 1997 andOsoriella Mello-Leitão, 1922. For these genera, the retrolateral projection is restricted to the basal region of the cymbium (Brescovit 1997: 180, fi g. 270). Apparently, this retrolateral projection is associated with a repose of the retrolateral tibial apophysis in the cymbium and would likely not be homologous to the paracymbium of others families, but deeper analysis would be necessary to test this (Platnick & Gertsch 1976;Coddington 1990;Griswold et al. 1998).

Ventral tegular process
The genus was originally diagnosed by Brescovit (1997: 88) based on sexual characters such as a distal ventral tegular process. The ventral tegular process (Figs 6J, 10C, 12B, 22C, 26A-B) is present in most Anyphaeninae, and changes position on the copulatory bulb. In Xiruana gracilipes and Anyphaena accentuate, it is located on the apical portion of the copulatory bulb. The silhavyi group also shares this apical position. Yet, the position of the ventral tegular process near the median apophysis (Figs 9B, 10C, 12B-C, 18B-C) is present in the riveti group (Clade G) and could be homologous to the one found, for example, in the genus Umuara (see Oliveira & Brescovit 2015b: 438, fi g. 2a-c).

Hood
The hood of the epigynum (Figs 7D, 10F, 22F, 32E-F, 33D-E) supports Anyphaena accentuata and Patrera fulvastra as the fi srt lineages to diverge in Anyphaeninae. Although Anyphaena and Patrera are not part of the Aysha group, as they do not have an embolic process on the male copulatory bulb, the anterior region of the epigyne, however, is provided with a hood. This was mentioned by Brescovit (1997: 119), who stated that the hood in the female genitalia is not restricted to the Aysha group, but is also present in the genera Anyphaena and Patrera. This character is plesiomorphic in Tafana and may be homologous to the anterior pouch of the median fi eld present in many Amaurobioidinae such as Monapia Simon, 1897, Gayenna Nicolet, 1849and Oxysoma Nicolet, 1849(Ramírez 1995a1999: 423, fi gs 4-5;2003: 21;Ramírez et al. 2004: 6, fi gs 6-7).

Lateral lobes, shape
The shape of the lateral lobes is diffi cult to standardize in the Anyphaeninae, because they have a great variation in female genitalia (Brescovit 1997). However, all species of Tafana are united by having a sinuous shape of the lateral lobes (Figs 10F, 11F, 15F, 27F), while in Anyphaena accentuata, Patrera fulvastra and Aysha prospera the shape is straight (Fig. 7D), and in Xiruana gracilipes the lateral lobes are curved (Fig. 7E).

Data used in the phylogenetic analysis
Similarity obtained through SPR-distances for (EW) trees and with different concavity for (IW). Values in bold below indicate the average between the topologies.