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Zootaxa 3814 (1): 121 132 www.mapress.com/zootaxa/ Copyright 2014 Magnolia Press Article http://dx.doi.org/10.11646/zootaxa.3814.1.7 http://zoobank.org/urn:lsid:zoobank.org:pub:78219981-7610-4584-b9be-2d226a87fa55 A new Colombian species of Cryptocellus (Arachnida, Ricinulei), with notes on the taxonomy of the genus ISSN 1175-5326 (print edition) ZOOTAXA ISSN 1175-5334 (online edition) RICARDO BOTERO-TRUJILLO 1,2 1 División Aracnología, Museo Argentino de Ciencias Naturales Bernardino Rivadavia, Avenida Ángel Gallardo 470, CP: 1405DJR, C.A.B.A., Buenos Aires, Argentina. E-mail: rbt@macn.gov.ar 2 Associate Researcher, Laboratorio de Entomología, Unidad de Ecología y Sistemática UNESIS, Departamento de Biología, Pontificia Universidad Javeriana, Bogotá, Colombia Abstract Cryptocellus sofiae sp. nov. is described based on males and females obtained from the easternmost part of Colombia, in Vichada Department. The new species is placed in the adisi species-group, based on the morphology of the male copulatory apparatus and the presence of polygonal (navicular or calyx-like) setae. With this addition, the group now comprises four species. A key for the identification of the members of this group is provided. Some taxonomic remarks about the adisi group and the genus Cryptocellus Westwood, 1874 are made. Key words: adisi group, Cryptocellus sofiae sp. nov., Vichada Resumen Se describe a Cryptocellus sofiae sp. nov. en base a machos y hembras provenientes de la región más oriental de Colombia, en el Departamento de Vichada. La nueva especie es asignada al grupo de especies adisi, en consideración a la morfología del aparato copulador masculino y la presencia de setas poligonales (naviculares o en forma de cáliz). Con esta adición, el grupo ahora reúne cuatro especies. Se proporcionan una clave para la identificación de los miembros del grupo. Se presentan algunas consideraciones respecto al grupo adisi y el género Cryptocellus Westwood, 1874. Introduction Ricinulei is a rare arachnid order comprising both extant and fossil species. There are 74 recognized extant species, placed in three genera. Ricinoides Ewing, 1929, currently consisting of 11 species, is known from 14 countries of western and central Africa. Pseudocellus Platnick, 1980 contains 25 species and occurs from southern U.S.A. (Texas) south to Panama and the Caribbean Islands, with most species having been described from Mexico. Cryptocellus Westwood, 1874 is so far known from 38 named species (not including the new species described here) and exhibits a distribution that overlaps with that of Pseudocellus in southern Central America, occurring from Honduras southward through tropical South America to Brazil (Harvey 2003; Naskrecki 2008; Penney et al. 2009; Tourinho & Saturnino 2010; Tourinho et al. 2010, in press; Murienne et al. 2012; Pinto-da-Rocha & Andrade 2012; Valdez-Mondragon & Francke 2013). Since Platnick s (1980) phylogenetic hypotheses for extant ricinuleids, the above-mentioned three genera have been recognized, and their monophyly was recently supported by molecular data (Murienne et al. 2012). During the 1980s, Cryptocellus received special attention in a series of publications by N.I. Platnick and coworkers (Platnick & Shadab 1976, 1977, 1981a, 1981b; Platnick & Paz 1979; Platnick 1980), wherein the knowledge of the genus was significantly increased, including the establishment of the foedus, centralis and magnus species-groups. More recently, Tourinho & Saturnino (2010) reviewed the species-group divisions of Cryptocellus and proposed the peckorum and adisi species-groups. Accepted by M. Judson: 12 May 2014; published: 6 Jun. 2014 121

Several authors in recent decades have paid attention to the integumental structures found in the cuticle of ricinuleids (Pittard & Mitchell 1972; Tuxen 1974; Legg 1976; Platnick & Shadab 1976, 1977; Adis et al. 1999; Talarico et al. 2005, 2006, 2008b; Salvatierra et al. 2013). These studies have concentrated on the legs and pedipalps, while other body regions remain understudied. No uniform classification of the integumental structures of ricinuleids has yet been proposed. Consequently, the cuticular structures have not been sufficiently used, in spite of their potential for taxonomy and phylogeny (Talarico et al. 2006; Salvatierra et al. 2013). Seven species of Cryptocellus have been described from Colombia: C. magnus Ewing, 1929, C. glenoides Cooke & Shadab, 1973, C. peckorum Platnick & Shadab, 1977, C. narino Platnick & Paz, 1979, C. florezi Platnick & García, 2008, C. platnicki Botero-Trujillo & Pérez, 2008, and C. luisedieri Botero-Trujillo & Pérez, 2009. An expedition to the easternmost extreme of Colombia, in Vichada Department (Fig. 1), yielded four ricinuleid specimens of a new species of Cryptocellus, herein described. With this addition, the number of known species of Cryptocellus is raised to 39. This number may still be long way from representing the true diversity of the group, however, due to the paucity of specimens gathered by field expeditions and the incomplete study of cave systems, where these animals are now known to be fairly common (Cokendolpher & Enríquez 2004; Talarico et al. 2005; Platnick & García 2008; Valdez-Mondragon & Francke 2011, 2013). Material and methods The specimens used in the species description belong to the arachnological collection of the Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá, Colombia (ICN). General terminology follows Platnick & Shadab (1977), except for male leg III, which follows Talarico et al. (2008a). Measurements are provided for a single specimen of each sex. Specimens were examined and photographed with LEICA M205 and LEICA M165 C stereomicroscopes. Measurements were obtained following the methodology outlined by Cooke & Shadab (1973), using an ocular micrometer fitted to a calibrated Advanced Optical JSZ-6S Zoom stereomicroscope. The distribution map was produced using SimpleMappr (Shorthouse 2010). For Scanning Electron Microscope (SEM) preparations, male (ICN-Ari-011) and female (ICN-Ari-012) specimens were dissected, dehydrated in 100% ethanol, fixed to aluminum stubs, and gold-palladium coated in a VG Scientific SC 7620 mini sputter-coater. SEM micrographs were taken under high vacuum with a Philips FEI XL30 TMP at the MACN. Taxonomy Family Ricinoididae Ewing, 1929 Genus Cryptocellus Westwood, 1874 Cryptocellus Westwood, 1874: 201. Type species. Cryptocellus foedus Westwood, 1874, by monotypy. Adisi species-group of Cryptocellus This group of species was created by Tourinho & Saturnino (2010) to include all the species having the tarsal process (=fixed accessory piece) of male leg III thin, curved and with a rounded contour. With the addition of C. sofiae sp. nov., the adisi group encompasses species in which the tarsal process of the male copulatory apparatus is boat- or canoe-shaped (C. florezi, C. lisbethae González-Sponga, 1997 and C. adisi Platnick, 1988), or else spoonlike, meaning that it has a moderate distal expansion (C. sofiae sp. nov.). All species currently assigned to this group are less than 5 mm in length, have polygonal (navicular or calyx-like) setae, and have the basal segment of the pygidium unnotched. At this time, however, these features are not included in the diagnosis for the group, because some of them (e.g. polygonal setae) also occur in species currently assigned to other groups. 122 Zootaxa 3814 (1) 2014 Magnolia Press BOTERO-TRUJILLO

Identification key to the species of the adisi group Ricinuleids are speciose, with new locality records often yielding a new species (Selden 1992). Moreover, the species of Cryptocellus are generally conserved morphologically, making their identification challenging (Salvatierra et al. 2013). It is therefore recommended that the original descriptions be consulted to confirm whether a specimen belongs to any of the species included in the key given below. The distribution of the species should also be taken into consideration, since most Cryptocellus have very narrow geographic ranges (Platnick 2002). The key avoids using morphological features that have been studied with SEM in only some species of this group (e.g. pits and other particularities of the cuticle sculpture and small details of the male copulatory apparatus). 1. Median plates of tergites XI and XII approximately as long as wide; male pedipalp femur noticeably bulkier than that of the female and armed with an enlarged basal tubercle directed ventrally on the retrolateral face. C. lisbethae (Bolívar, Venezuela) - Median plates of tergites XI and XII noticeably wider than long; pedipalp femur similar in male and female, not bearing an enlarged tubercle in either sex (Brazil, Colombia)............................................................ 2 2. Female spermathecae well separated from each other; males with a long and thin metatarsal process (about four times longer than wide) and a massive accessory piece of the copulatory apparatus...................... C. adisi (Amazonas, Brazil) - Female spermathecae very close together (C. florezi) or touching (C. sofiae sp. nov.); males with small and robust metatarsal process (about twice longer than wide) and a filiform accessory piece of the copulatory apparatus (Colombia)............ 3 3. Femur of leg II about five times longer than wide; tarsal process of male copulatory apparatus boat-shaped........................................................................................... C. florezi (Caquetá, Colombia) - Femur of leg II about 3.5 times longer than wide; tarsal process of male copulatory apparatus spoon-shaped, with moderate ventral expansion in distal half............................................ C. sofiae sp. nov. (Vichada, Colombia) Cryptocellus sofiae sp. nov. Figures 1 22 Type material. Holotype male (adult): COLOMBIA: Vichada Department, Cumaribo, El Tuparro Natural National Park, Administrative Center, 0.5º21 15.1 N 67º51 43.7 W, 80 m elev., 21 March 2012, in litter sample from forest associated with rocky outcrop, D. A. Luna & C. Romero (ICN-Ari-009). Paratypes: COLOMBIA: 2 adult females (ICN-Ari-010 and ICN-Ari-012) and 1 adult male (ICN-Ari-011), same data as holotype. Specimens ICN-Ari-009 and 010 preserved in 70% ethanol; specimens ICN-Ari-011 and 012 dehydrated and fixed to aluminum stubs for SEM. Distribution. Known only from the type locality (Fig. 1). Etymology. The species is named after my lovely 7 year old niece, Sofia Carlosama Botero, who always kneels on the chair to take a curious glance at the cute bugs (as she says) I work with. It is a celebration of her precious company. Diagnosis. A member of the adisi species-group, as here defined, based on the morphology of male leg III, which exhibits a relatively thin and curved tarsal process of the copulatory apparatus (Fig. 15). It can be distinguished from other species in the group in having the median plates of tergites XI and XII wider than long (Figs. 2, 4, 7), a moderate ventral expansion on distal half of male tarsal process (=spoon-like shaped) (Figs. 14 16), and the female spermathecae touching together (Figs. 19 20). Description (male and female). The description is based on the four specimens available. Coloration (in preserved specimens). Body and appendages with cuticle uniformly chestnut to light reddish, except for medial region of sternite XI in male, which is dark-red; carapace without pale or yellowish areas in ocular region; opisthosomal membranes yellow. Polygonal (navicular or calyx-like) setae whitish (Figs. 2 7, 8 10). Setation. Body entirely covered with setae. Most conspicuous are the large whitish polygonal (navicular or calyx-like) setae, that vary in shape (Figs. 2 5, 21 22), on external surface of cucullus (except for distal fifth), carapace (including ocular areas and median longitudinal furrow), opisthosoma (except for medial region of sternites X and XI), pygidium, dorsal surfaces of pedipalp trochanter II and femur, leg coxae (only distally), leg trochanters I and II, femora, patellae, tibiae and metatarsi. These setae are large and not very numerous (e.g. about 60 70 setae on median plate of tergite XII; Fig. 7). Distal fifth of external surface of cucullus, medial region of sternites X and XI, tritosternum, pedipalp coxae, pedipalp trochanter I, prolateral, retrolateral and ventral surfaces of pedipalp trochanter II and femur, pedipalp tibia, most of leg coxae and leg tarsomeres lack these large setae, being covered instead with smaller and thinner setae (Figs. 3, 5 6, 8 9). SEM observations revealed a range of A NEW COLOMBIAN CRYPTOCELLUS Zootaxa 3814 (1) 2014 Magnolia Press 123

setal types on the cucullus, carapace and opisthosoma (Figs. 21 22), similar to that present in C. adisi (Adis et al. 1999). Carapace. Lateral margins not parallel (carapace narrowing anteriorly); anterior margin slightly concave, posterior margin convex; carapace approximately as long as wide, widest at level of leg III (Figs. 2, 4); ocelli completely absent, but corresponding ocular regions noticeably elevated, with an evident depression behind them; abundant cuticular pits of variable size, randomly distributed (most containing tubercles); very few tubercles present outside the pits; a shallow, median, longitudinal furrow is present (Fig. 6), beginning at level of coxae I and ending at level of coxae III. Cucullus. Wider than long, with lateral margins not parallel (cucullus narrowing posteriorly) (Fig. 6); with abundant cuticular pits of variable size, randomly distributed (most of them containing tubercles); very few tubercles present outside the pits (Fig. 22). Chelicera. Description based on left chelicera of paratype male (Fig. 11). Movable finger longer than fixed finger; movable finger armed with ten separate small teeth, sub-equal in size except for third basalmost tooth and the two distalmost teeth, which are twice the size of the others; fixed finger with markedly enlarged distal tooth, followed by two separate smaller teeth (slightly larger than the largest of the movable finger) and a small basal tooth, latter similar in size to most teeth of movable finger. Sternal region. Without cuticular pits; coxae I not meeting tritosternum; coxae II IV meeting entirely, subrectangular, progressively decreasing in length; coxae IV pear-shaped, with posterior margin slightly curved (female) or almost straight (male); suture line of coxae II about three fifths length of that of coxae III (coxae III being larger) (Figs. 8 9); coxae I with minute tubercles along posterior margin; coxae II III with tubercles along the distal two thirds of anterior and posterior margins; coxae IV without tubercles. Opisthosoma. Longer than wide, widest at level of junction between tergites XI and XII (Figs. 2 5); with abundant cuticular pits of variable size, randomly distributed on dorsal surface (most containing tubercles); very few tubercles present outside pits (Fig. 21); median plates of tergites XI XIII with paired, shallow, lateral depressions beside median elevation; median plates of tergites XI XIII progressively decreasing in width, lateral margins not parallel on XI XII (narrowing posteriorly); median plate of tergites XI XII wider than long, median plate of tergite XIII approximately as wide as long (Figs. 2, 4). Sternites without cuticular pits and devoid of tubercles; male with median region of sternite XI slightly protuberant and more sclerotized (unlike that of female, which is not elevated) (Figs. 3, 5). Basal segment of pygidium unnotched on either dorsal or ventral borders (Fig. 10). Pedipalps. Without cuticular pits; with some minute tubercles on ventral surface of trochanters I and II; femur curved and widened in basal half; approximately the same shape in male and female (no distinct sexual dimorphism); femur without any trace of basal tubercle in either sex; tibia longer than femur, slightly widened in basal third (Figs. 8 9); both movable and fixed claws armed with minute teeth, more evident on fixed claw; movable claw noticeably larger than fixed claw. Legs. Legs I IV similar in shape in male and female (Figs. 2 5); without cuticular pits and with few tubercles; completely devoid of any spine-like projection or similar structures; leg II not widened; leg III metatarsus not inflated (noticeably longer than wide), trochanter IV unmodified; tarsal claws strongly curved. Male copulatory apparatus. Tarsal process connected basally to a bone-shaped (phalanx-like) base. Tarsal process bulkier than accessory piece, spoon-like, with a moderate ventral expansion in distal half forming a curved cavity along the prolateral surface where the accessory piece rests; apically with two distinct lobes (L1 and L2) forming a single canal. Accessory piece movably jointed with basal prolateral aspect of tarsal process; progressively becoming thinner and bifurcated into two prominent, sharp-pointed lobes (L and L ) of similar length (snake-tongue-shaped); the accessory piece is rolled up, leaving visible the sperm transfer groove as a foldlike gap running longitudinally from the base towards the tip of L. The bifurcation of the piece into L and L starts at about one third of the piece from the base (its beginning is indicated by an arrow in Figs. 15 17). Tarsal process and accessory piece strongly curved upwards (Figs. 14 18). Spermathecae. Touching together, slightly wider than long, constricted medially to give a rounded protruding section (Figs. 12 13, 19 20). Measurements (in mm). Male holotype (ICN-Ari-009) and female paratype (ICN-Ari-010, in parentheses): body length, excluding pygidium 3.06 (3.50); cucullus 0.50 (0.58) long, greatest width 0.66 (0.74); carapace 1.03 (1.17) long, 1.06 (1.15) wide at level of leg III (where widest); opisthosoma 1.56 (1.75) long (excluding pygidium), 124 Zootaxa 3814 (1) 2014 Magnolia Press BOTERO-TRUJILLO

1.39 (1.57) wide at level of junction between tergites XI and XII (where widest); median plate of tergite XI 0.44 (0.49) long, 0.83 (0.97) wide (where widest); median plate of tergite XII 0.39 (0.43) long, 0.67 (0.74) wide; median plate of tergite XIII 0.53 (0.60) long, 0.56 (0.62) wide; suture line of coxae II 0.08 (0.08) long, of coxae III 0.14 (0.13) long; pedipalp femur 0.49 (0.52) long, greatest diameter 0.22 (0.24); pedipalp tibia 0.71 (0.87) long, greatest diameter 0.10 (0.12); femur I 0.47 (0.49) long, 0.21 (0.23) in diameter; femur II 0.69 (0.78) long, 0.21 (0.23) in diameter. Generic placement of the new species. Cryptocellus sofiae sp. nov., is herein assigned to Cryptocellus based on the presence of morphological features of this genus as defined by Platnick (1980): body short and wide (median plate of tergite XII wider than long); and ocular areas small in size and directed laterally at the edge of the carapace near the second legs. Some aspects of its morphology do not completely match the features described for Cryptocellus, however, and are briefly discussed below. The coloration pattern of the cuticle of C. sofiae sp. nov. is chestnut to light reddish (Figs. 2 7, 8 10), which is a condition deemed diagnostic for Pseudocellus species, as opposed to Cryptocellus whose members are supposed to display a dark reddish-black coloration (Platnick 1980). The light coloration of the new species is not considered to link it to Pseudocellus, however, based on the above-referred features shared with other Cryptocellus, and because adults of other species in the later genus also display a color that is not particularly dark but rather chestnut-reddish [e.g. Cryptocellus iaci Tourinho, Lo Man-Hung & Bonaldo, 2010 (Tourinho et al. 2010: figs. 7 12; Salvatierra et al. 2013: fig. 2c)]. FIGURE 1. Known distribution records of the genus Cryptocellus in Colombia. A NEW COLOMBIAN CRYPTOCELLUS Zootaxa 3814 (1) 2014 Magnolia Press 125

FIGURES 2 7. Cryptocellus sofiae sp. nov. 2, holotype male (ICN-Ari-009), dorsal; 3, holotype male, ventral; 4, paratype female (ICN-Ari-010), dorsal; 5, paratype female, ventral (some setae accidentally removed from the opisthosoma); 6, paratype female (ICN-Ari-012), frontal aspect (median longitudinal furrow of carapace indicated by arrow); 7, holotype male, polygonal setae on tergites XI and XII. 126 Zootaxa 3814 (1) 2014 Magnolia Press BOTERO-TRUJILLO

FIGURES 8 13. Cryptocellus sofiae sp. nov. 8, holotype male (ICN-Ari-009), sternal region, pedipalps and sternites X XI; 9, paratype female (ICN-Ari-012), sternal region, pedipalps and sternites X XI; 10, holotype male, pygidium, dorsal posterior aspect; 11, paratype male (ICN-Ari-011), dorsal aspect of left chelicera; 12, paratype female (ICN-Ari-012), genital lip and spermathecae, anterior aspect; 13, paratype female, genital lip and spermathecae, posterior aspect. A NEW COLOMBIAN CRYPTOCELLUS Zootaxa 3814 (1) 2014 Magnolia Press 127

FIGURES 14 20. Cryptocellus sofiae sp. nov., SEM micrographs. 14 18, paratype male (ICN-Ari-011), right leg III; 14, prolateral aspect; 15, copulatory apparatus, prolateral aspect; 16, prolaterodorsal aspect; 17, apex of tarsal process and accessory piece; 18, level of junction between accessory piece and tarsal process; 19 20, paratype female (ICN-Ari-012), spermathecae; 19, posterior aspect; 20, inclined posterior aspect. Abbreviations: ac, accessory piece (sperm transfer groove indicated by arrowheads, beginning of bifurcation indicated by arrow); Lc, lamina cyathiformis; L1 2, lobes of tarsal process; L, L, lobes of accessory piece; mc, medial constriction of spermathecae; mp, metatarsal process; MT, metatarsus; pe, prolateral expansion of tarsal process; tp, tarsal process. 128 Zootaxa 3814 (1) 2014 Magnolia Press BOTERO-TRUJILLO

FIGURES 21 22. Cryptocellus sofiae sp. nov., SEM micrographs, different setal types. 21, paratype female (ICN-Ari-012), part of tergite XIII; 22, paratype male (ICN-Ari-011), part of cucullus. Although the bifurcate accessory piece of the copulatory apparatus in the male C. sofiae sp. nov. (Figs. 14 17) resembles that of some Pseudocellus species, this condition is now also known to occur in some other members of Cryptocellus, as discussed at the end of this paper. Based on these considerations, and taking into account the fact that no species of Pseudocellus is so far known to inhabit South America, the new species is here considered to be a member of Cryptocellus as currently defined. Affinities of the new species and remarks on the adisi group The five species-groups currently recognized within Cryptocellus are primarily defined by characters of the male copulatory apparatus (Tourinho & Saturnino 2010). The shape of the copulatory apparatus of C. sofiae sp. nov., among other features, prevents it from being assigned to the foedus, centralis or magnus groups, as defined by Tourinho & Saturnino (2010). The peckorum and adisi groups include the species with male features that mostly resemble those of the new species. Cryptocellus sofiae sp. nov. is similar to both of these groups in having a relatively thin and curved tarsal process (= fixed piece) of male leg III (Fig. 14). It also resembles the peckorum group in having a curved accessory piece, but differs in the shape of the metatarsus (=basitarsus) of leg III, which is not inflated in the new species (noticeably longer than wide; Fig. 14), as opposed to being moderately inflated in the peckorum species-group. Given that the adisi group is currently supported by the presence of a tarsal process that is thin, curved and with a rounded contour, C. sofiae sp. nov. is herein allocated to this group. This decision is also supported by the fact that the three species currently included in this group, namely C. florezi, C. adisi and C. lisbethae, share with the new species the presence of large polygonal (navicular or calyx-like) setae covering the body, as well as having the basal segment of the pygidium unnotched. Although neither of these features has so far been used to support the adisi group, polygonal setae are only present outside this group in three species: C. narino, C. albosquamatus Cooke, 1967 and C. canga Pinto-da-Rocha & Andrade, 2012. The first of these species is presently placed in the magnus group (Tourinho & Saturnino 2010). With respect to it, Pinto-da-Rocha & Andrade (2012: 477) noted that some species of Magnus-group also shares [sic] with Adisi-group a fixed accessory piece curved and with rounded contour, such as C. narino. Interestingly, C. albosquamatus was considered to be related to C. adisi by Platnick (1988), but it was not placed in any species-group by Tourinho & Saturnino (2010), due to the lack of males. As for C. canga, its describers commented that this species shares morphological features with members of both the adisi and the magnus groups, without assigning it to either of them (Pinto-da-Rocha & Andrade 2012). It is herein hypothesized that the presence of polygonal (navicular or calyx-like) setae has relevance for the classification of Cryptocellus species, as implied by Pinto-da-Rocha & Andrade (2012). For instance, the presence of these setae may provide additional support for the adisi group, suggesting that C. narino may actually be more A NEW COLOMBIAN CRYPTOCELLUS Zootaxa 3814 (1) 2014 Magnolia Press 129

closely related to the species in this group. It is also plausible that C. albosquamatus may prove to be a member of the same group when the male is discovered. Nonetheless, it is appropriate not to draw any definite conclusions in this respect for the moment, since the set of morphological features useful for the classification of Cryptocellus species is no doubt still far from complete. To conclude, the results of the present study support the need to further revise the species-group definitions (particularly those of the adisi and magnus groups) within this genus, as previously indicated by Pinto-da-Rocha & Andrade (2012). The copulatory apparatus of the new species and some considerations on Cryptocellus An interesting morphological feature of C. sofiae sp. nov. is the bifurcate accessory piece of the male copulatory apparatus (Figs. 14 17). The bifurcate condition has repeatedly been depicted for Pseudocellus species. For instance, Pittard & Mitchell (1972: figs. 106, 107) and Cooke (1971: figs. 10, 11) provided illustrations of the bifurcated piece of P. pelaezi (Coronado-Gutiérrez, 1970) and, more recently, Valdez-Mondragón & Francke (2011: figs. 5, 14, 21, 28) showed the same condition in four Mexican species of this genus. A bifurcate accessory piece has also been depicted previously for three Cryptocellus species. Platnick & Paz (1979: figs. 10 11) provided an illustration for C. narino that suggested that this piece was bifurcate, as subsequently confirmed by Talarico et al. (2008a). Afterwards, Platnick (1988: figs. 3 4) illustrated the copulatory apparatus of C. adisi in which the bifid condition at the tip of the accessory piece was evident. More recently, a bifurcate accessory piece was reported for C. luisedieri by Botero-Trujillo & Pérez (2009: fig. 9). It is important to note that, although the accessory piece has not been reported to be bifurcate in two species of the adisi group (C. florezi and C. lisbethae), there is a doubt about its form in those species since they have not yet been studied with SEM. Because the bifurcate condition of the accessory piece can be difficult to notice (it was found to occur in C. luisedieri only after dissection of the piece), it now comes unclear whether this structure does indeed end in a simple, non-bifurcate apex in some other species of Cryptocellus as supposed in the past, since it might have gone unnoticed. Interestingly, however, SEM micrographs of C. canga showed that the accessory piece is not bifurcate in that species (Pinto-da-Rocha & Andrade 2012: figs. 7 9), while the same is true for the accessory piece of P. pearsei (Chamberlin & Ivie, 1938) (Talarico et al. 2008a). Thus, a bifurcate accessory piece is not unique to members of Pseudocellus, just as a single-tipped piece is not unique to Cryptocellus. Traditionally, the monophyly of Pseudocellus and Ricinoides are relatively well-defended hypotheses, whereas that of Cryptocellus has been regarded as questionable (Platnick 1980). The intergeneric relationships of the extant ricinuleids are highly controversial. For instance, Platnick (1980) hypothesized that Cryptocellus is more closely related to Ricinoides than to Pseudocellus, while Selden (1992) hypothesized Ricinoides to be the sister group of the two New World genera. In a recent contribution, Murienne et al. (2012) obtained the first molecular phylogeny of Ricinulei, wherein the monophyly of each of the three extant genera was recovered and their relationships proposed to be (Cryptocellus (Pseudocellus + Ricinoides)). All the possible different phylogenetic hypotheses have thus been proposed by different authors, while features contradicting one or the other continue to be uncovered. For instance, the findings concerning sperm morphology in the studies of Talarico et al. (2008a) and Talarico & Michalik (2010) conflict with the phylogenetic proposals of Platnick (1980) and Murienne et al. (2012), tending instead to be more consistent to that of Selden (1992). The various studies that have been carried out on different aspects of ricinuleid morphology (e.g. cuticular characters, morphology of the copulatory apparatus, sperm morphology), along with the aforementioned molecular phylogeny, have used very few out of the 75 currently known species. Most importantly, species of the five speciesgroups of Cryptocellus have not yet been studied for these aspects, or combined in a phylogenetic analysis. Taking this into account, it is possible that the species so far studied do not adequately cover the morphological and molecular diversity existing within Ricinoididae. The search for phylogenetically useful features of the genitalic structures was encouraged by Platnick (1980), but these have not yet been sufficiently studied to allow a distinction between the two New World genera. Although Murienne et al. s (2012) phylogeny confirmed monophyly of the three extant genera of ricinuleids, our knowledge about the male copulatory apparatus (i.e. bifurcate vs. single tipped accessory piece) in Cryptocellus and Pseudocellus species conflicts in some degree with the existing hypotheses, specifically in relation to how the New World genera are defined. The same is true for the coloration pattern (chestnut to light reddish vs. dark reddish black). 130 Zootaxa 3814 (1) 2014 Magnolia Press BOTERO-TRUJILLO

A representative selection of species from the different species-groups of Cryptocellus for phylogenetic analyses that also include members of the other genera, will be necessary to unravel the relationships existing between those groups with the most varied morphologies. This will not be an easy task, though, due the rarity of ricinuleids and the absence of suitable fresh material of many species for DNA isolation. Once additional data becomes available, it should be possible to address the question of whether additional genera might need to be recognized (i.e. for species of Cryptocellus formerly gathered in species-groups). Acknowledgments I am particularly grateful to Ana Lúcia Tourinho (Museum of Comparative Zoology, Harvard University), Ricardo Pinto da Rocha (Universidade de São Paulo, Brazil), Giovanni Talarico (Ernst Moritz Arndt Universität Greifswald, Germany) and Andrés Alejandro Ojanguren Affilastro (MACN) for reading earlier versions of the manuscript and providing valuable insights. I would like to acknowledge Eduardo Flórez D., David A. Luna Sarmiento and Catalina Romero Ortiz (Universidad Nacional de Colombia, Bogotá) for making the specimens herein used available for study. I thank Andrés Sánchez (Pontificia Universidad Javeriana, Bogotá) for his help in producing and editing some photographs. Luis N. Piacentini and Hernán A. Iuri (MACN) provided assistance in obtaining SEM micrographs. References Adis, J., Messner, B. & Platnick, N. (1999) Morphological structures and vertical distribution in the soil indicate facultative plastron respiration in Cryptocellus adisi (Arachnida, Ricinulei) from Central Amazonia. Studies on Neotropical Fauna and Environment, 34, 1 9. http://dx.doi.org/10.1076/snfe.34.1.1.8915 Botero-Trujillo, R. & Pérez, G.A. (2008) A new species of Cryptocellus (Arachnida, Ricinulei) from northwestern Colombia. Journal of Arachnology, 36, 468 471. http://dx.doi.org/10.1636/ch07-95sc.1 Botero-Trujillo, R. & Pérez, G.A. (2009) A new species of Cryptocellus (Arachnida, Ricinulei) from the Kofán Territory in southwestern Colombia. Zootaxa, 2050, 56 64. Chamberlin, R.V. & Ivie, W. (1938) Arachnida of the orders Pedipalpida, Scorpionida and Ricinulida. Carnegie Institute Washington, Publications, 491, 101 107. Cokendolpher, J.C. & Enríquez, T. (2004) A new species and records of Pseudocellus (Arachnida: Ricinulei: Ricinoididae) from caves in Yucatán, Mexico and Belize. Texas Memorial Museum, Speleological Monographs, 6, 95 99. Cooke, J.A.L. & Shadab, M.U. (1973) New and little known ricinuleids of the genus Cryptocellus (Arachnida, Ricinulei). American Museum Novitates, 2530, 1 25. Cooke, J.W. (1967) Observations on the biology of Ricinulei (Arachnida) with descriptions of two new species of Cryptocellus. Journal of Zoology, 151, 31 42. http://dx.doi.org/10.1111/j.1469-7998.1967.tb02864.x Cooke, J.W. (1971) Mating behavior and the functional morphology of the male copulatory apparatus in Cryptocellus pelaezi (Arachnida, Ricinulei). Unpublished M.Sc. thesis, Graduate Faculty of Texas Tech University 50 pp. Coronado-Gutiérrez, L. (1970) Estudio de un Cryptocellus de cavernas de Mexico (Arachn., Ricin.). Ciencia, México, 27, 47 62. Ewing, H.E. (1929) A synopsis of the American arachnids of the primitive order Ricinulei. Annals of the Entomological Society of America, 22 (4), 583 600. González-Sponga, M.A. (1997) Arácnidos de Venezuela. Cryptocellus lisbethae nueva especie de Ricinulei del Estado Bolívar (Arachnida: Ricinulidae). Memoria de la Sociedad de Ciencias Naturales La Salle, 57 (148), 49 54. Harvey, M.S. (2003) Catalogue of the Smaller Arachnid Orders of the World. CSIRO Publishing, Collinwood, Victoria, Australia, 385 pp. Legg, G. (1976) The external morphology of a new species of ricinuleid (Arachnida) from Sierra Leone. Zoological Journal of the Linnean Society, 59, 1 58. http://dx.doi.org/10.1111/j.1096-3642.1976.tb01007.x Murienne, J., Benavides, L.R., Prendini, L., Hormiga, G. & Giribet, G. (2012) Forest refugia in Western and Central Africa as Museums of Mesozoic biodiversity. Biology Letters, 9, 20120932, 1 7. http://dx.doi.org/10.1098/rsbl.2012.0932 Naskrecki, P. (2008) A new ricinuleid of the genus Ricinoides Ewing (Arachnida, Ricinulei) from Ghana. Zootaxa, 1698, 57 64. Penney, D., Marusik, Y., Wheater, C.P. & Langan, A.M. (2009) First Gambian Ricinulei (Arachnida: Ricinoididae): northernmost African record for the order. Zootaxa, 2021, 66 68. A NEW COLOMBIAN CRYPTOCELLUS Zootaxa 3814 (1) 2014 Magnolia Press 131

Pinto-da-Rocha, R. & Andrade, R. (2012) A new species of Cryptocellus (Arachnida: Ricinulei) from Eastern Amazonia. Zoologia, 29 (5), 474 478. http://dx.doi.org/10.1590/s1984-46702012000500012 Pittard, K. & Mitchell, R.W. (1972) Comparative morphology of the life stages of Cryptocellus pelaezi (Arachnida, Ricinulei). Graduate Studies Texas Tech University, 1, 1 77. Platnick, N.I. (1980) On the phylogeny of Ricinulei. In: Gruber, J. (Ed), Verhandlungen des 8. Internationalen Arachnologen- Kongress, Wien. H. Egermann, Wien, pp. 349 353. Platnick, N.I. (1988) A new Cryptocellus (Arachnida: Ricinulei) from Brazil. Journal of the New York Entomological Society, 96(3), 363 366. Platnick, N.I. (2002) Ricinulei. In: Adis, J. (Ed), Amazonian Arachnida and Myriapoda. PENSOFT Publishers, Sofia-Moscow, pp. 381 386. Platnick, N.I. & García, L.F. (2008) Taxonomic notes on Colombian Cryptocellus (Arachnida, Ricinulei). Journal of Arachnology, 36, 145 149. http://dx.doi.org/10.1636/h07-27.1 Platnick, N.I. & Paz, N. (1979) On the Cryptocellus magnus group (Arachnida, Ricinulei). American Museum Novitates, 2677, 1 9. Platnick, N.I. & Shadab, M.U. (1976) On Colombian Cryptocellus (Arachnida, Ricinulei). American Museum Novitates, 2605, 1 18. Platnick, N.I. & Shadab, M.U. (1977) On Amazonian Cryptocellus (Arachnida, Ricinulei). American Museum Novitates, 2633, 1 17. Platnick, N.I. & Shadab, M.U. (1981a) On Central American Cryptocellus (Arachnida, Ricinulei). American Museum Novitates, 2711, 1 13. Platnick, N.I. & Shadab, M.U. (1981b) On the Cryptocellus centralis group (Arachnida, Ricinulei). Bulletin American Museum of Natural History, 170, 18 22. Salvatierra, L., Tourinho, A.L. & Giribet, G. (2013) Description of the male, larva and nymphal stages of Cryptocellus iaci (Arachnida, Ricinulei), with an overview of tarsal sensilla and other integumental structures. Zootaxa, 3709 (2), 149 161. http://dx.doi.org/10.11646/zootaxa.3709.2.3 Selden, P.A. (1992) Revision of the fossil ricinuleids. Transactions of the Royal Society of Edinburgh: Earth Sciences, 83, 595 634. http://dx.doi.org/10.1017/s0263593300003333 Shorthouse D.P. (2010) SimpleMappr, a web-enabled tool to produce publication-quality point maps. Available from: http:// www.simplemappr.net (accessed 10 October 2013). Talarico, G., García, L.F. & Michalik, P. (2008a) The male genital system of the New World Ricinulei (Arachnida): ultrastructure of spermatozoa and spermiogenesis with special emphasis on its phylogenetic implications. Arthropod Structure & Development, 37, 396 409. http://dx.doi.org/10.1016/j.asd.2008.01.006 Talarico, G. & Michalik, P. (2010) Spermatozoa of an Old World Ricinulei (Ricinoides karschii, Ricinoididae) with notes about the relationships of Ricinulei within the Arachnida. Tissue and Cell, 42, 383 390. http://dx.doi.org/10.1016/j.tice.2010.10.003 Talarico, G., Palacios-Vargas, J.G. & Alberti, G. (2008b) The pedipalp of Pseudocellus pearsei (Ricinulei, Arachnida) - ultrastructure of a multifunctional organ. Arthropod Structure & Development, 37, 511 521. http://dx.doi.org/10.1016/j.asd.2008.02.001 Talarico, G., Palacios-Vargas, J.G., Fuentes Silva, M. & Alberti, G. (2005) First ultrastructural observations on the tarsal pore organ of Pseudocellus pearsei and P. boneti (Arachnida, Ricinulei). Journal of Arachnology, 33, 604 612. http://dx.doi.org/10.1636/04-110.1 Talarico, G., Palacios-Vargas, J.G., Fuentes Silva, M. & Alberti, G. (2006) Ultrastructure of tarsal sensilla and other integument structures of two Pseudocellus species (Ricinulei, Arachnida). Journal of Morphology, 267, 441 463. http://dx.doi.org/10.1002/jmor.10415 Tourinho, A.L., Lo-Man-Hung, N.F. & Bonaldo, A.B. (2010) A new species of Ricinulei of the genus Cryptocellus Westwood (Arachnida) from northern Brazil. Zootaxa, 2684, 63 68. Tourinho, A.L., Lo-Man-Hung, N.F. & Salvatierra, L. (in press) A new Amazonian species of Cryptocellus (Arachnida, Ricinulei), with descriptions of its integumental structures and all free-living life stages. Zootaxa. Tourinho, A.L. & Saturnino, R. (2010) On the Cryptocellus peckorum and Cryptocellus adisi groups, and description of a new species of Cryptocellus from Brazil (Arachnida: Ricinulei). Journal of Arachnology, 38, 425 432. http://dx.doi.org/10.1636/ha09-108.1 Tuxen, S.L. (1974) The African genus Ricinoides (Arachnida, Ricinulei). Journal of Arachnology, 1, 85 106. Valdez-Mondragón, A. & Francke, O.F. (2011) Four new species of the genus Pseudocellus (Arachnida: Ricinulei: Ricinoididae) from Mexico. Journal of Arachnology, 39, 365 377. http://dx.doi.org/10.1636/ha11-02.1 Valdez-Mondragón, A. & Francke, O.F. (2013) Two new species of ricinuleids of the genus Pseudocellus (Arachnida: Ricinulei: Ricinoididae) from southern Mexico. Zootaxa, 3635 (5), 545 556. http://dx.doi.org/10.11646/zootaxa.3635.5.4 Westwood, J.O. (1874) Thesaurus Entomologicus Oxoniensis. Clarendon Press, Oxford, UK, 205 pp. 132 Zootaxa 3814 (1) 2014 Magnolia Press BOTERO-TRUJILLO