The first Paratropididae (Araneae, Mygalomorphae) from Colombia: new genus, species and records

Abstract The family of mygalomorph spiders Paratropididae Simon, 1889 is here reported for the first time for Colombia, where it is represented by three genera (Anisaspis, Paratropis, Stormtropisgen. n.) and eight species. One genus, Stormtropis, and six species constitute new taxa that are here diagnosed, described and illustrated. The geographical distribution of Paratropispapilligera FO Pickard-Cambridge, 1896 and Paratropiselicioi Dupérré, 2015 are also redescribed and expanded on the basis of new material examined. The diagnosis of the subfamily Paratropidinae, Paratropis Simon, 1889 and Anisaspis Simon, 1892 are emended including the variations of the new species. Likewise, a geographic distribution map for the entire family and a taxonomic key for the males of Paratropidinae are included. Other biogeographic, morphological, and taxonomic aspects are discussed.


Introduction
Paratropididae Simon, 1889, known as bald legged spiders, is one of the most enigmatic groups of Mygalomorphae due to its cryptic habits, singular biology, and controversial phylogenetic position. Paratropidids constitute a small family of spiders, currently comprising four genera and eleven species, distributed in Mexico, Central America, Lesser Antilles, and northern South America except Colombia (Raven 1985, WSC 2018. Their presence in Venezuela, Mexico, and Ecuador has been recently reported by Bertani (2013), Valdez-Mondragón et al. (2014) and Dupérré (2015), respectively. Paratropidids are cursorial, medium to small-sized mygalomorphs, which hide themselves in the surface layers of the soil (West in Raven 1999); found in leaf litter and rotten logs, under rocks, moss, and burrows in ravines (personal observations). The main paratropidid morphological characteristic is the presence of a scaly cuticle adapted to adhere to soil particles (Raven 1985, Jocqué andDippenaar-Schoeman 2006).
Formerly, Paratropididae was considered the sister-group of Theraphosidae into Theraphosoidea, and this group was related with Barychelidae in Theraphosoidina (Raven 1985, Goloboff 1993. However, results of some molecular and total evidence studies suggest a distant phylogenetic relationship of Paratropididae with the Theraphosoidina clade, not closely related with Theraphosidae or Barychelidae Hedin 2006, Bond et al. 2012). Conversely, in recent studies, Paratropididae was recovered as sister group of some ctenizids (Wheeler et al. 2017), or some nemesiids (Garrison et al. 2016); although, both studies pointing out this position as questionable. Likewise, in Fernandez et al. (2018) Paratropididae was resolved as a sister group of Macrothele (Macrothelidae), but this study focused in Araneomorphae, and mygalomorph sampling was very limited. Hamilton et al. (2016) recovered Paratropididae as the sister group to all other Aviculariodea taxa, with the exception of Bymainiella sp. (Hexathelidae); this grouping was followed by Hedin et al. (2018).
Paratropidids are characterized by the soil encrusted or scaly cuticle, weakly or ascopulate tarsi I and II and absence of scopulae elsewhere, maxillary lobes elongated, and the presence of labial cuspules usually arranged in an anterior rectangular group (Raven 1985). Pérez-Miles et al. (2017) found that the putative scopula of paratropidids was in fact a pseudoscopula, constituted of chemosensory not-adhesive setae. The family comprises two subfamilies, Paratropidinae Simon, 1889 andGlabropelmatinae Raven, 1985. Paratropidinae is characterized by a long single tooth on the superior tarsal claws, the steeply elevated eye tubercle, very long anterior maxillary lobes, and absence of a tibial apophysis and claw tufts, unlike Glabropelmatinae that has thin claw tufts (without scopula), a bifurcated tibial apophysis, a typical elevated eye tubercle, and shorter anterior maxillary lobes (Raven 1985).
Paratropidinae includes three genera, Paratropis Simon, 1889 with six species, distributed in Mexico, Venezuela, Brazil and Peru; Anisaspis Simon, 1892 with one species from Saint Vincent island; and Anisaspoides FO Pickard-Cambridge, 1896 with one species from Brazil. Anisaspis and Anisaspoides are only known by females. Glabropelmatinae includes only one genus Melloina Brignoli, 1985, with three species, distributed in Venezuela andPanama (WSC 2018).
Two characteristics have been important in Paratropidinae taxonomy, the number of spinnerets, four (Paratropis) or two (Anisaspis, Anisaspoides), and the presence of a third tarsal claw, only on leg I (most Paratropis), only on legs I and II (Anisaspoides), or absent (Anisaspis) (Raven 1985, Pickard-Cambridge 1896, Valdez-Mondragón et al. 2014. The presence of Paratropididae in Colombia was informally indicated by de Mello-Leitão (1941) and in unpublished studies Sánchez-C. 1995, Perafán et al. 2013); however, this is the first formal report for the country. A review of myga-lomorph spiders from Colombia including field and collection data produced some important results for Paratropididae, which we present here: a new genus, Stormtropis gen. n., with four new species; a new species of Anisaspis, which represent the first male description for the genus and the first record for South America; one new species of Paratropis; and a new record of P. papilligera FO Pickard-Cambridge, 1896 andP. elicioi Duperré, 2015. We discuss the geographic distribution and diversity of paratropidids as well as some biological and morphological characteristics.

Materials and methods
The general description format follows Raven (1999) and Bertani (2013), with modifications. Specimens were examined using an Advanced Optical stereomicroscope. Photographs were taken with an Olympus LC30 camera adapted to a stereomicroscope. All measurements are in millimeters and were taken with an ocular micrometer. Legs and palp measurements were taken along a dorsal longitudinal line of the left side. Total body length excludes chelicerae and spinnerets. Spination: we consider as spines the thick, sclerotized setae, with acute and non-translucent apex; similar setae but with translucent apex were not counted. Male palpal bulbs (usually left) were removed from cymbium for study and illustrations. Records without geographic coordinates were determined using GoogleMaps and the gazetteers GeoLocator and GeoNames. The distribution map was produced using the Geographic Information System QGIS 'Girona' (version 3.0, QGIS team, www.qgis. org), with raster files from NaturalEarth and DivaGis. WSC means World Spider Catalog. Tibial apophysis with numerous spines on the proximal row (12) ( Figure 7D, E); presence of a sclerotized dark mark on proximal dorsal tibia, with a slight excavation ( Figure 7A)  Tibial apophysis with less spines on the proximal row (6) ( Figure 5D, E); without sclerotized dark mark on tibia ( Figure 5A)  Type species. Anisaspis tuberculata Simon, 1892, deposited in NHM, examined. Diagnosis. Anisaspis differs from other paratropidid genera by the presence of only two spinnerets (PLS) ( Figure 1D) and all legs with only two tarsal claws (STC), males without tibial apophysis and palpal bulb with sinuous embolus ( Figure 1E, F). PLS relatively long ( Figure 1A, B). Females remain unknown.
Remarks. The specimen described as holotype female of A. tuberculata is actually a juvenile specimen. Therefore, females of this genus remain unknown.
Distribution. Lesser Antilles, Saint Vincent island and Colombia, on the eastern flank of the Eastern Cordillera, Meta Department, Llanos foothills ( Figure 10); from sea level to 570 m altitude.
Diagnosis. Anisaspis camarita sp. n. differs from A. tuberculata by the absence of a spine on dorsal tarsi distally, longer spinnerets (PLS) with apical segment digitiform (domed in A. tuberculata) ( Figure 1A, B) and AME on a super-tubercle (another higher tubercle on the ocular tubercle, Figure 1D).
Legs: cuticle with soil particles encrusted. Leg and palpal segments measurements provided in Table 1. Leg I clearly thicker than the others. Bristles, plumose and thornlike setae and spines evident. Trichobothria: filiform, on central 2/3 of tarsi, palp 5, leg I 8, II 7, III 6, IV 8; on distal 1/4 of metatarsi, legs I-III 4, IV 5; and on proximal 1/3 of tibiae, palp two rows of 3 each, legs I-III two rows of 4 each, IV two rows of 4 and 1 respectively. Scopula absent. Pseudoscopula weak and divided by conical longer setae, only present on distal tarsi I and II; tarsi III and IV with few sparse pseudoscopula setae. Claw tufts absent. Tarsal claws: ITC absent on all legs but a very small tooth present on right leg I in the same position of ITC; STC with one medial tooth on all legs. Tibial apophysis absent. Spination: principally thorn-like setae on all segments. Spines: palp and legs I-II 0; leg III, fe 0, pa 0, ti 0, me 1pd, 3v, ta 1p; leg IV, fe 0, pa 0, ti 0, me 2v, ta 1p. Palp: cymbium with two unequal lobes separated by a sclerotized groove; tibia with shallow distoventral groove. Palpal bulb pyriform elongated; embolus curved, long, tapering to the apex, apex wide; a triangular translucent tooth on the subapical region, close to the apex ( Figure 1E, F).
Female. Unknown. Distribution. Only known from its type locality, in the foothills of the Eastern Cordillera of Colombian Andes (at 570 m altitude), Meta Department, Bavaria forest ( Figure 10).
Etymology. The specific epithet camarita is a noun in apposition which means friend, in the colloquial way of the Llanos Region of Colombia, where this species is distributed.  Distribution. Mexico, Lesser Antilles, and northern South America (Brazil, Colombia, Ecuador, Peru and Venezuela). In Colombia it is widely distributed in the three mountain ranges that make up the Andes, the inter-Andean valleys and lowlands of the Amazon, Llanos, and Caribbean regions (Perafán 2017) ( Figure 10).  Emended diagnosis. Males of P. elicioi differ from those of other Paratropis species by the morphology of the palpal bulb with very long and straight embolus ( Figure 2E, F). Females can be distinguished by the morphology of spermathecal receptacles with long neck, longitudinal dorsal fold and numerous apical lobes ( Figure 2H).

Paratropis elicioi Dupérré, 2015
Remarks. The examination of new material from Colombia and Ecuador allowed us to infer that both the diagnosis and the descriptions of the two sexes of P. elicioi were inaccurate (Dupérré 2015). Below we present an emended description.
Palp: cymbium with two unequal lobes separated by a sclerotized groove; tibia with distoventral groove. Palpal bulb pyriform very elongated; embolus as long as tibia and half patella, straight then tapering to the apex, apex stout but flattened ( Figure 2E, F).
Abdomen: with four longitudinal dorsal rows of seven small tubercles, each emitting from its apex a plumose, bacilliform seta. Book lung apertures projected, oval, sclerotized. Two spermathecal receptacles with a long neck, with a longitudinal dorsal fold, ended in a multilobed fundus ( Figure 2H). Spinnerets: PMS length 0.60; PLS length 3.0, apical segment digitiform. Basal segment of PLS divided in two unequal cuticle plates.
Distribution. South of Colombia and north of Ecuador, on the Western Andean montane forest, between 500-1700 m altitudes. In Colombia it's distributed on Nariño Department (Barbacoas, Reserva Natural Río Ñambí) and Ecuador distributed on Cotopaxi Province (Otonga Biological Reserve) and Pichincha Province (Santo Domingo; Nanegalito; Las Tolas) ( Figure 10).
Abdomen: with four longitudinal dorsal rows of seven small tubercles, each emitting from them a plumose, bacilliform seta. Book lung apertures projected, oval, sclerotized ( Figure 3B). Two spermathecal receptacles with a long neck, with the base widened and apex narrow, ended in a fundus with several projected lobes ( Figure 3D). Spinnerets: PMS length 0.6; PLS length 2.7, apical segment digitiform. Basal segment of PLS divided in two unequal cuticle plates ( Figure 3B).

Male. Unknown.
Distribution. Only known from its type locality, in the western Cordillera of Colombian Andes, Valle del Cauca Department, km 16 road Cali-Buenaventura, at 1800 m altitude (Figure 10). Natural history. The females of Paratropis florezi sp. n. live in shallow burrows that they dig in the substrate of the ravines of the road.
Etymology. The species epithet is a noun in genitive, in honor of Dr Eduardo Flórez Daza (ICN), in recognition of his friendship, teachings, and vast contributions to Colombian arachnology. Paratropis papilligera FO Pickard-Cambridge, 1896 Figure 4 Type material. Holotype male and paratype female from Santarem, Pará, Brazil, deposited in NHM, only male examined. Additional material examined. Colombia, Amazonas, Leticia, km 11 road to Tarapacá, 100 m, 25-iv-2002, it was collected manually in the day on leaf litter, col. G Amat and Estudiantes Introducción Sistemática Animal -Universidad Nacional de Colombia, 1 male (ICN-Ar 2315).
Emended diagnosis. Males of P. papilligera differ from those of other Paratropis species by the morphology of the palpal bulb with very thin and long embolus, distally curved ( Figure 4F, G), and by the tibia I with a basal retrolateral conic process with spiniform setae (Figure 4D, E).
Distribution. Amazonas of Brazil and Colombia. Brazil, Santarem (Pará); Colombia, Amazonas (Leticia) (Figure 10). Description. Carapace round, almost glabrous, light to dark brown. Caput arched. Fovea shallow, transverse, straight to slightly procurved. Eye group subquadrate, wider than long, tubercle well defined, elevated. Clypeus absent. Chelicerae without rastellum, cheliceral furrow narrow with teeth on both margins: promargin 7-13, retromargin 6-13, fangs long. Labium subquadrate with 20-70 cuspules restricted to anterior edge. Maxillae longer than wide with the anterior prolateral lobe very elongated, conical; few cuspules (24-77) throughout the prolateral diagonal half of the maxillae. Labio-sternal groove narrow in the middle and wider laterally. Sternum heart shaped, slightly wider than long, sigillae oval, submarginal. Legs, thin and long, pair I slightly stouter than II-IV; clubbed setae present. Few filiform tricobothria on tarsus, metatarsus, and tibia in males. Long paired claws (STC) with one medial long tooth ventrally; third unpaired claw (ITS) absent on all legs of males; ITS present on leg I of females. Claw tufts absent, tarsal scopula absent, pseudoscopula setae generally present on the distal third of anterior tarsi. Males with spinose apophysis (similar to Aviculariinae) on prolateral distal tibiae I (except S. muisca). Abdomen oval, glabrous, with clubbed setae present on dorsum. Four spinnerets; PLS well developed, PMS small (half of the basal segment of PLS). All body encrusted by soil particles. Males without spines, and females with few spines on all legs. Males with cymbium with two unequal lobes separated by a sclerotized groove; palpal tibia with shallow distoventral groove; and palpal bulb pyriform elongated, with embolus slightly curved tapering to the apex, and a subapical triangular tooth. Females with spermathecal receptacles with a tubular neck and globose fundus.
Distribution. Stormtropis gen. n. is distributed in the central and eastern Cordilleras of Colombia, on the montane forests of the Magdalena Valley and Cauca Valley, between 1400-3400 m altitudes, in the Departments of Antioquia (Santa Elena), Boyacá (Sotaquirá and Santuario de Fauna y Flora Iguaque), Caldas (Pensilvania) and Cundinamarca (Topaipí) (Figure 10).
Etymology. The name Stormtropis is a Latin declension (neuter) of the noun Stormtrooper from the fictional universe of the Star Wars films. The stormtroopers are the soldiers of the main ground force of the Galactic Empire. These soldiers are very similar to each other, with some capacity for camouflage but with unskillful movements, like this group of spiders.  Diagnosis. Stormtropis colima sp. n. differs from the other species of the genus by the presence of a tibial apophysis with shorter base and not so much separated from the tibia as in the other species ( Figure 5D, E). Additionally, S. colima sp. n. differs from S. parvum sp. n. by the presence of a continuous row and more numerous cheliceral teeth on promargin (13-11) (7 in S. parvum sp. n.) and from S. paisa sp. n. by the absence of a sclerotized dark mark on proximal dorsal tibia, without a slight excavation.
Palp: cymbium with two unequal lobes separated by a sclerotized groove; tibia with shallow distoventral groove. Palpal bulb pyriform elongated; embolus curved, very long, tapering to the apex, apex wide; a triangular translucent tooth on subapical region, close to apex ( Figure 5F, G).
Female: unknown. Distribution. Only known from its type locality, Topaipí in the Río Negro Province (Cundimarca), in the Eastern Cordillera of Colombian Andes, at 1300 m altitude ( Figure 10).
Etymology. The species epithet colima is a noun in apposition which means warrior in the extinct Muisca language. The Colimas were an indigenous tribe that inhabited in the central highlands of Colombia, where the species occurs.
Legs: cuticle normal. Leg and palpal segments measurements provided in Table 6. Leg I clearly thicker than the others. Bristles and thorn-like setae present. Trichobothria: filiform, on central 2/3 of tarsi, palp 4, leg I 7, II 5, III 4, IV 4; on distal 1/4 of metatarsi, leg I 4, II 3, III 3, IV 3; on proximal 1/3 of tibiae, palp and legs I-III two rows of 2 each, IV a row of 3p and 1d. Scopulae absent. Pseudoscopula weak and divided by conical longer setae, only present on distal tarsi I, II and III; tarsi IV with few sparse pseudoscopula setae. Claw tufts absent. Tarsi distally incrassate, mainly on anterior legs. Tarsal claws: ITC absent on all legs, but a very small tooth present on legs I; STC with one curved tooth on all legs. Spination: principally thorn-like setae on all segments. Spines absent.
Palp: cymbium with two unequal lobes separated by a sclerotized groove; tibia with shallow distoventral groove. Palpal bulb pyriform; embolus slightly curved tapering to the apex, apex stout but flattened, a subapical triangular tooth on embolus ( Figure 6D, E).
Etymology. The species epithet muisca is a noun in apposition which refers to the indigenous tribe who inhabit in the same region where this species occur. Diagnosis. Stormtropis paisa sp. n. differs from the other species of the genus by the presence of a sclerotized dark mark on proximal dorsal tibia, with a slight excavation ( Figure 7A). Males additionally differ from the other species by the presence of a tibial apophysis bearing more numerous spines on the proximal row (12) ( Figure 7D, E); in the other species with tibial apophysis the number of such spines is 5-6; and by the palpal bulb with the embolus sinuous, distally twisted and more widened on the apex (Figure 7F, G). Females can be distinguished by the morphology of the spermathecal receptacles with a tubular neck and a wide globose fundus (mushroom shaped) ( Figure 8D).
Legs: cuticle with soil particles encrusted. Leg and palpal segments measurements provided in Table 7. Legs I slightly stouter than II-IV. Clubbed plumose and thorn-like setae. Trichobothria: filiform, on central 2/3 of tarsi, palp and all legs 5; on distal 1/4 of metatarsi, leg I 4, II-IV 3; on proximal 1/3 of tibiae, palp and legs I-III 5 in two rows (2/3 each), IV 4. Scopula absent, pseudoscopula slightly denser on tarsi I and II, sparse pseudoscopula setae on tarsi III and IV. Claw tufts absent. Tarsal claws: ITC absent on all legs; STC long, with one tooth on all legs. Tibial apophysis on leg I present: only one flattened branch on distal prolateral side, with spines on two parallel rows, 15 dis-  tal and 12 proximal ( Figure 7D, E). Thorn-like setae on tibiae, metatarsi, and tarsi of all legs, more dense on legs III and IV. Spines absent. Palp: cymbium with two unequal lobes separated by a sclerotized groove; tibia with a distoventral groove. Palpal bulb pyriform elongated; embolus sinuous and distally twisted, long, tapering to the apex; a triangular translucent tooth on subapical region ( Figure 7F, G).
Distribution. Only known from its type locality, Central Cordillera of the Colombian Andes, Antioquia Department, Medellin (Santa Elena), at 2400 m altitude ( Figure 10).
Etymology. The species epithet paisa is a noun in apposition which means the vernacular name given to the people of Medellín, where the species occurs.
Diagnosis. Stormtropis parvum sp. n. differs from the other species of the genus by the less numerous and discontinuous row of cheliceral teeth on promargin (2-2-3) ( Figure 9B), and by the presence of a tibial apophysis, with a larger base, more separated from the tibia than on the other species ( Figure 9D, E). Additionally, differs from S. paisa by the absence of a sclerotized dark mark on proximal dorsal tibia, without excavation, and lower number of spines in the spur (eleven on the distal row and six on proximal).
Although Colombian paratropidid species richness is the highest found in a country (WSC 2018), several undescribed species are expected to be found in the future, considering the enormous and unexplored ecological diversity of mygalomorph spiders in this region (Perafán 2017, Perafán andValencia-Cuéllar 2018), coupled with the evidence of numerous specimens not yet studied.
Stormtropis gen. n. includes some species in which males have a tibial apophysis, a feature not found among paratropidids (except Melloina) until now. It is probably a recent acquisition of this group of species, which should be more closely related with Paratropis, but this hypothesis must be tested with a rigorous phylogenetic analysis. Another remarkable morphological characteristic of this genus is the sexual dimorphism in tarsal claws: ITC on leg I is only present in females. The presence of ITC on leg I was originally used to diagnose Paratropis from other paratropidids, but Valdéz-Mondragón et al. (2014) found that females of Paratropis tuxtlensis have ITC on legs I and II but males lack them. Likewise, Dupérré (2015) described Paratropis elicioi, lacking ITC in both sexes, and considered this character as ambiguous. We consider that there is a great inter-and intraspecific variation in this character, so this independent character should be carefully taken into account in the taxonomy of the family. Another sexual difference found in the Paratropidinae is related to the pseudoscopula, present in males and absent in females, as was reported for several Mygalomorphae (Pérez-Miles et al. 2017).
An important characteristic of Paratropididae, unique among Mygalomorphae, is the ability to adhere soil particles to their scaly cuticle. Although the natural history of the group is poorly known, paratropidids were considered cursorial spiders which hide themselves in the surface layers of the soil (West in Raven 1999). This condition should be compatible with the presence of soil encrusted on the cuticle, facilitating the camouflage of individuals on the ground. However, and unexpectedly in our field study, we found several females of P. florezi sp. n. and other undetermined species inhabiting tubular burrows on ravine walls or soil. Considering that burrowing paratropidids maintain the encrusted cuticle, burrowing habits could be a secondary adaptation in order to exploit different habitats.