Unveiling the diversity of Phalangodus Gervais, 1842 (Opiliones: Cranaidae): descriptions of four new species from Colombia

The Cranaidae genus Phalangodus Gervais, 1842 is revisited and its species are diagnosed. An identifi cation key for the males of Phalangodus is provided. Four Colombian species are described as new: Phalangodus briareos sp. nov. from a cave in Zapatoca, Santander department, P. cottus sp. nov. from Villavicencio, Meta department, P. gyes sp. nov. from Ibagué, Tolima department and P. kuryi sp. nov. from Sierra Nevada de Santa Marta, Magdalena department. SEM images of the penis of Phalangodus, including the type species of the genus, P. anacosmetus Gervais, 1842, are provided for the fi rst time. Sexual dimorphism and the morphology of stylar caps and ovipositor in Cranaidae are discussed. Additionally, we revised the geographical distribution of P. anacosmetus, including a map with the geographical records of the genus.


Introduction
Cranaidae is the third most numerous family among Gonyleptoidea with 166 species (Kury 2013;Hara et al. 2014;Villarreal et al. 2015). The taxonomy and systematics of this group are complicated and the generic boundaries are poorly defi ned, as partially explained by Villarreal et al. (2015). The three main issues that lead to this situation are: 1) antique diagnoses of the genera that still follow the Roewerian system of classifi cation, i.e., use of variable characters to establish artifi cial groups, as shown by several authors (e.g., Piza 1942;Kury 1990;Villarreal et al. 2015); 2) underestimation of the specifi c richness of many cranaid groups (the fi rst author is working on an ongoing revision of the family) and 3) lack of revision, only a few genera have been revised according to modern standards (i.e., Pinto-da-Rocha & Kury 2003;Pinto-da-Rocha & Bonaldo 2011;Kury 2012a;Hara et al. 2014;Villarreal et al. 2015).
In the present article, four new species from Colombia are described and diagnosed, based on material of Phalangodus studied from four arachnological collections. A key to identify males of the genus is provided. Sexual dimorphism and morphology of the stylar caps in Cranaidae is discussed.

Material and methods
The description standard and morphological terminology follow Villarreal et al. (2015) and integumentary ornamentation follows Da Silva & Gnaspini (2009). Ovipositor terminology follows Townsend et al. (2015) with modifi cations here proposed. Outline shape of dorsal scutum follows Kury et al. (2007) extended in Kury & Medrano (2016). Pedipalpal tibia and tarsus setation formula follows Mello-Leitão (1939), as explained and used in Pinto-da-Rocha (1997) and Villarreal-Manzanilla & Pinto-da-Rocha (2006). Tarsal formula (numbers of tarsomeres from tarsus I-IV): counts are given from left to right side of each specimen; numbers in parentheses denote quantity of tarsomeres in distitarsus I-II.
The nomenclature of the macrosetae of penis ventral plate follows  with the following modifi cation: in Phalangodus there is an increase of the number of macrosetae that makes it diffi cult to distinguish between some MS groups (e.g., MS D from MS C or MS B from MS A). We are not able to identify homologies between these MS groups, but we can observe, at least, one proximal and one distal cluster. By using relative position and comparison with other related genera, these groups are here called MS A/B and MS C/D respectively, without ruling out the presence or absence of any groups, i.e., postulating either (1)  Color descriptions use the standard names of the 267 Color Centroids of the NBS/IBCC Color System, using the Descriptive Color Names Dictionary (http://people.csail.mit.edu/jaffer/Color/Dictionaries#nbsiscc). For the complete synonymic list on each genus and species, see Kury's catalogue (2003). Only references from 2003 on are mentioned here.
Scanning Electron Microscopy of genitalia was carried out with a JEOL JSM-6390LV at the Center for Scanning Electron Microscopy at the Museu Nacional/UFRJ. The pictures of specimens were made using a digital camera Sony Cybershot DSC-v1 and assembled and edited the images of limited focal planes using Combine ZP Suite (by Alan Hadley) and Photoshop CC softwares. The drawings were made using a camera lucida attached to a stereomicroscope and processed with the software Inkscape v.0.91. Measurements were made using ImageJ v.1.50i Suite (by Wayne Rasband). The map with distribution of the species was made using Quantum Gis 2.2.0 software (QGIS Development Team 2014). Coloured areas mean WWF Terrestrial Eco-regions of the World (Olson et al. 2001, here abbreviated as WWF).

Remarks
The material collected in Cúcuta (that corresponds to the syntypes of Allocranaus giganteus) presents the following differences compared with those from Cundinamarca: (1) ocularium not surpassing the scutal (2) narrower ocularium (ratio wide/length = 1.79 in Cundinamarca specimens vs 1.46 in Cúcuta specimens); (3) pedipalp of the subadult male from Cúcuta sub cylindrical in lateral view ( Fig. 2b) vs dorsally curved from Cundinamarca males. (4) Females from Cundinamarca have a higher ocularium and wider dorsal scutum than Cúcuta´s females.
The lack of adult males from Cúcuta in collections makes it is impossible for a comparison of male sexual secondary characters or genital structures. On the other hand, there is a considerable gap between the localities of both populations, which indicates an uncommon disjunctive distribution, in neotropical harvestmen (Fig. 18).
Nonetheless, these data suggests the possibility of two species, more detailed studies including males are necessary to taxonomic decisions about identity of Allocranaus giganteus.

Distribution
Known from Magdalena valley montane forests in southwestern Cundinamarca and Cúcuta, Norte de Santander (Fig. 18) (see remarks above and discussion for details).

Etymology
Briareos (from Greek Βριάρεως), the vigorous, also called as Aegaeon, one of the three Hecatonchires.  Table 1. DORSUM. Dorsal scutum type alpha (Figs 4a, 5a). Abdominal scutum widest at scutal groove II; lateral borders of dorsal scutum mostly smooth with few minute granules only on the middle region. Carapace mostly smooth, with a group of granules on the anterolateral region. Ocularium high, without median depression, with a paramedian pair of acuminate tubercles (Figs 4a-c, 5a). Integumentary dome of ozopore raised and conspicuous. Abdominal scutum well delimited, divided into four well-marked scutal areas; scutal area I divided into left and right halves by invasion of the scutal area II, with few granules (Fig. 5a); scutal area III with a pair of paramedian acuminate, short, subparallel acuminate tubercles (Figs 4a, c, 5a). Posterior border of scutum straight and with few granules on a row. Free tergites I-III with a row of granules. Stigmatic area smooth. Stigmata large, oval and oblique. Coxa I with a row of large tubercles of different size; coxa II longer than coxae I and III, with a median row of low tubercles; coxa III with the posterior border sigmoid, a median row of small tubercles; coxa IV strongly backward, with group of low tubercles close to the stigmatic suture. Free sternites each with a row of small granules.
CHELICERA (Figs 4a-b, 5a). Chelicera swollen, bulla with tubercles in the proximal region, one ectal row and one group of dorsomesal tubercles. Fixed fi nger with a large subproximal tooth and 3 subdistal small teeth. Movable fi nger with a small subproximal tooth, one large median and one tridentate subdistal lamella. Mesal side of the base of fi xed fi nger and near the base of the movable fi nger with setiferous tubercles.
LEGS. Coxa I with two dorsal tubercles, the retrolateral one reaching the dorsal tubercle of the coxa II which has one prolateral and two retrolateral tubercles; III unarmed; IV with the prolateral face slightly granulate and with one dorsodistal domed large tubercle. Trochanter I-III unarmed; trochanter I-II with a large ventral, two ventrodistal tubercles and some prolateral granules; trochanter III with a prolateral row of granules and a slight retrolateral expansion; trochanter IV proximally swollen ( Fig. 4a-b) with a ventrodistal retrolateral tubercle and many granules irregularly distributed. Femora I-III straight and with longitudinal rows of granules; femur II with a large proximal tubercle on the ventrolateral side ( Fig. 5e-f); femur IV sub-straight, granulate, with a distal pair of large spiniform tubercles, one pointing retrolaterally and the other, prolaterally (Fig. 5c). Tibia I-IV straight and densely granulate, unarmed (Fig. 5d). Claws III-IV smooth. Ratio Fe IV/scutum = 1.44. Tarsal counts in Table 2. GENITALIA. Ventral plate (VP) subrectangular, with mid-constriction, the basal lobes curved and laterodistal border slightly projected, anterior border concave, ventrally with a well-marked longitudinal median elevation (Fig. 6a). Proximal ventral surface of VP with two inverted lung-shaped fi elds of small, needle-like microsetae ( Fig. 6a-b). MS A/B forming a proximal group of seven pairs of large, acuminate and cylindrical setae, pointing towards to the base of the penis trunk; MS C composed by 11-12 pairs of  . 6b-c). Two pairs of very small MS E, placed on latero-distal fl ange (Fig. 6b). Glans sac columnar elongate, with proximal folds and circular small protuberances on the base (Fig. 6b- Female (MNRJ 2163) Similar to male, differing by: ocularium slightly narrower; carapace larger; coda wider; tubercles of area III higher. Chelicerae with movable fi nger thinner. Pedipalpal femur lower and thinner in lateral view, with ventroproximal smaller tubercles, pedipalpal claw not swollen. Stigmatic area shorter, without the groups of granules anteriorly to stigmata. Genital operculum wider. Trochanters III-IV narrower; femur III without ventroproximal large tubercle; femur IV slightly curved proximally, thinner, without retrolateral distal spine and with a reduced prolateral tubercle; tibia IV with retrolateral granules smaller.
Ovipositor ( Fig. 7a-d): Dorsal lobes (dl) and ventral lobes (vl) rounded, with four and three pairs of large, acuminated, single-tipped setae respectively. Dl with three pairs of dorsal setae (ds) distally located and one pair basally located (Fig. 7c). Lateral region of the ovipositor with one dorso-lateral group of seven short setae (dls), two of them rounded (Fig. 7b, d).

Distribution
Known only from the type locality .

Etymology
Cottus (from the Greek Κόττος), the striker, one of the three Hecatonchires.  (2016) DORSUM. Dorsal scutum type alpha (Figs 8a, 9a). Abdominal scutum widest at scutal groove III level; lateral borders of dorsal scutum smooth. Carapace mostly smooth, with a group of small tubercles on the anterolateral region. Ocularium high, without median depression, with a paramedian pair of large, acuminate tubercles (Figs 8a-b, 9a). Integumentary dome of ozopore raised and conspicuous. Abdominal scutum well delimited, divided into four well-marked scutal areas: scutal area I divided into left and right halves by invasion of the scutal area II, with a pair of small granules on each side; scutal area III with a pair of paramedian acuminate high subparallel tubercles (Figs 8a, 9a). Posterior border of scutum straight, with only a pair of paramedian tubercles (Figs 8a, 9a), unarmed. Free tergites I-III with granules sparsely distributed, tergite II with a paramedian pair of large granules.
VENTER. Stigmatic area smooth. Stigmata large, oval and oblique. Coxa I with a row of large tubercles of different size; coxa II longer than coxae I and III, with a median row of large tubercles; coxa III with a median row of small tubercles and with the posterior border sigmoid; IV strongly backward, with some dark tubercles sparsely distributed and a group close to the stigmatic area. Free sternites with a row of small dark tubercles.
CHELICERA (Figs 8a, 9a). Chelicera swollen, bulla with curved tubercles on its proximal border and ectal face, and a group of mesodorsal tubercles. Hand with a longitudinal row of downward curved acuminate tubercles (distal largest). Fixed fi nger with a proximal conical tooth and a subdistal laminar tooth. Movable fi nger with a proximal rounded tooth, a median conical tooth and four low subdistal teeth. With many setiferous tubercles on the base of fi xed fi nger .   (2016) PEDIPALPUS. Trochanter with two-three tubercles on a dorsal protuberance, ventrally with two tubercles, the median largest and curved. Femur (Fig. 9b) slightly compressed, dorsally curved and ventrally straight in lateral view, with a dorsal row of four-fi ve forward curved tubercles, ventrally with a pair of geminated large tubercles. Patella short (ratio FePp/PaPp = 1.75), cylindrical and curved with small dorsal granules. Tibia dorsally with scarce and small granules, ventrally smooth with two longitudinal medial row of small granules; tibial setation: ectal (IiIi) (3 > 1 > 4 > 2) and accessory distal tubercle (the last ones St share a common base), mesal (IiIi) (3 > 1 > 4 > 2). Tarsus ectally (IiIi) (3 > 1 > 2 = 4) and mesally (IIi) (1 = 2 > 3), the mesal and ectal faces with a row of proximal tubercles at the base of the St forming a crenulate lamella. Claw not swollen.
LEGS. Coxa I with one dorsal tubercle; II with a dorsal tubercle; III unarmed; IV with few prolateral tubercles and one dorsodistal, prolateral, curved, acuminated tubercle. Trochanters I-IV unarmed; ventrally, I with three tubercles (one medial and two distal), and one proximal retrolateral tubercle; II with retrolateral group of tubercles and one retrolateral ventral, large, acuminate tubercle; III enlarged in the middle, dorsally unarmed and ventrally with a median large tubercle and four retrolateral tubercles; IV proximally enlarged (Fig. 8a), dorsally smooth, with few ventral, small, dark granules and two retrolateral, distal, large tubercles. Femora I-III straight, with longitudinal rows of granules; femur II with two short rows of large ventral tubercles (Fig. 9d, e); femur III with a retrolateral, ventral proximal row of large tubercles; femur IV slightly curved, densely granulate, with a ventral subdistal pair of large spiniform tubercles pointing opposite directions, the retrolateral one oblique (Fig. 9h) and the prolateral one hook-like shaped (Fig. 9f). With one retrolateral proximal tubercle (Fig. 9f). Patella IV ventrally with prolateral row of tubercles (Fig. 9f). Tibiae III-IV straight and densely granulate; tibia III with a retroventral row of tubercles; tibia IV with a retrolateral row of large, spiniform tubercles, and two distal ventral rows of large tubercles. (Fig. 9f, g). Metatarsus IV with pale ring markings. Claws III-IV smooth. Ratio Fe IV/scutum = 1.18. Tarsal counts in Table 4.
GENITALIA (MNRJ 17942). Ventral plate (VP) subrectangular, with mid-constriction, the proximal lobes curved and laterodistal border sligthly projected, anterior border with smooth concavity (Fig. 10b-c). Ventral surface of VP with two inverted? lung-shaped fi elds of minute and needle-like microsetae (Fig. 10c). MS A/Bforming a proximal group of two pairs of large, acuminate and cylindrical setae, pointing towards to the base of the penis trunk; MS C/D composed by 11 pairs of setae with similar shape to MS A/B. (Fig. 10a-b, e). Two pairs of very small MS E located on latero-distal fl ange (Fig.  10a). Glans sac columnar elongate, with proximal folds. Stylus straight, without processes ( Fig. 10a- Female (MNRJ 17942) Similar to male, differing by: ocularium slightly narrower; carapace slightly larger; coda wider; chelicerae with movable fi nger thinner; pedipalpal femur, patella and tibia thinner, with ventroproximal tubercles smaller; spines of ocularium and tubercles of area III higher; stigmatic area shorter, without the groups of granules anteriorly to stigmata; genital operculum wider; trochanters III-IV narrower, trochanter IV without retrolateral tubercles; femur IV thinner, without proximal tubercle; tibia IV without retrolateral row of granules and the ventral rows with smaller tubercles; tergite II with a pair of large paramedian tubercles.

VILLARREAL O. & GARCÍA M.A.F., Four new species of Phalangodus from Colombia
OVIPOSITOR. dl and vl rounded with fi ve and two pairs of large, acuminated, single-tipped setae respectively. Dl with fi ve pairs of ds, one of them basally located and the lateral region of the ovipositor with two pairs of dorso-lateral group of short setae.

Distribution
Known only from the type locality (Fig. 18).

Etymology
Gyes (from the Greek Γύης), the big-limbed, one of the three Hecatonchires.

Description
Male (MNRJ 8655) Measurements of body and appendage in Table 5. DORSUM. Dorsal scutum type alpha (Figs 8c, 11a). Abdominal scutum wider at scutal groove II; lateral borders of dorsal scutum smooth. Carapace mostly smooth, with a row of granules on the anterolateral region. Ocularium high, without median depression, with a paramedian pair of short and acuminate tubercles ( Fig. 8c-d). Integumentary dome of ozopore raised and conspicuous. Abdominal scutum well delimited, divided into four well-marked scutal areas: scutal area I divided into left and right halves by invasion of the scutal area II, with a pair of small dark granules (Figs 8c, 11a); scutal area II with a transverse row of six dark granules; scutal area III with a pair of paramedian conical, dark and short tubercles (Fig. 8d). Posterior border of scutum straight, with a row of seven low dark tubercles (Fig. 8c). Free tergites I-III each with a row of few dark granules.
VENTER. Stigmatic area smooth. Stigmata large, oval and slightly oblique. Coxa I densely covered with irregular rows of tubercles; coxa II longer than coxae I and III, with two rows of small tubercles; coxa III with a median row of tubercles and few distal small tubercles sparsely distributed and with the posterior border sigmoid; coxa IV strongly backward, with some granules sparsely distributed and one group of tubercles on a prominence close to the stigmatic area. Free sternites each with a row of small granules. CHELICERA (Fig. 11a). Chelicera swollen, bulla with an ectal row of four-fi ve tubercles curved anteriorly and a group of dorsodistal mesal tubercles. Hand with small frontal tubercles. Fixed fi nger with four conical teeth decreasing in size subdistally. Movable fi nger with a proximal wide lamella, a median truncate tooth and two fused subdistal teeth.
LEGS. Coxa I with one dorsal and one retrolateral tubercles; II with one prolateral and one retrolateral tubercles; III with retrolateral tubercle partially fused with coxa IV; IV with few lateral tubercles and one prodorsal triangular small tubercle. Trochanter I-IV dorsally unarmed; trochanter I ventrally with three tubercles (one medial and two distal), and one proximal retrolateral tubercle; trochanters II-III retrolaterally swollen and with a group of tubercles; trochanter II with a retrolateral ventral large tubercle; trochanter IV proximally enlarged with retrolateral tubercles. Femora I-IV substraight (Fig. 11e) with longitudinal rows of small granules or tubercles, femur II very elongated. Patellae I-IV unarmed. Tibia I-III unarmed, II very elongated, IV with proximal groups of retrolateral and large prolateral tubercles  . Ventral plate subrectangular, with mid-constriction, the proximal lobes curved and laterodistal border sligthly projected, anterior border concave. Ventral surface of VP with two inverted lung-shaped fi elds of small, needle-like microsetae separated by a wellmarked longitudinal median elevation (Fig. 12b-c). MS forming two lateral groups: one proximal group of four pairs of large, acuminate and cylindrical setae, pointing towards to the base of the penis trunk (MS A/B); and one distal (MS C/D) group composed by 11 pairs of setae with similar shape to the distal group (Fig. 12a, c). Two pairs of very small MS E located on latero-distal fl ange (Fig. 12b-c). Glans sac columnar elongate, with proximal folders. Stylus straight, without processes (Fig. 12a). Stylar caps ring-shaped without lateral or ventral projections (Fig. 12d). Female (MNRJ 17907) Similar to male; differing by: carapace slightly larger; coda wider; chelicerae smaller; pedipalpal femur, patella and tibia thinner, with ventroproximal tubercles smaller; stigmatic area shorter, without the groups of granules anteriorly to stigmata; genital operculum wider; trochanters III-IV narrower; femur IV slightly thinner; tibia IV without proximal group of granules; tergite I-III with a pair of large paramedian tubercles. OVIPOSITOR (Fig. 7e-f). dl and vl rounded with four and two pairs of large, acuminated, single-tipped setae respectively. Dl with four pairs of ds one of them basally located; and the lateral region of the ovipositor with one group of four very long and thin dorso-lateral setae, reaching the base of ds (Fig. 7f).

Etymology
The species is named after the Brazilian arachnologist Adriano B. Kury, advisor and friend, who immensely contributes to the knowledge of Opiliones worldwide.  Table 7. DORSUM. Dorsal scutum type alpha (Fig. 13a). Abdominal scutum widest at scutal groove II; lateral borders of dorsal scutum smooth. Carapace mostly smooth, with a group of granules on the anterolateral region. Ocularium unarmed, very low, without median depression. Integumentary dome of ozopore raised and conspicuous. Abdominal scutum well delimited, divided into four marked areas: scutal area I divided into left and right halves by invasion of the scutal area II; scutal areas I-II each with two pair of granules; scutal area III with a paramedian pair of low tubercles (Fig. 13a-b). Posterior border of scutum straight and unarmed, with a row of four minute tubercles on a row (Figs 13a, 14a). Free tergites I-III with few granules.
VENTER. Stigmatic area with two pairs of paramedian granules. Stigmata large, oval and slightly oblique. Coxa I densely covered with irregular rows of tubercles; coxa II longer than coxae I and III, with three rows of small tubercles, the median ones more conspicuous; coxa III with a curved median row of granules and distal minute tubercles sparsely distributed,with the posterior border sigmoid; coxa IV   CHELICERA (Fig. 13a). Basichelicerite very swollen, bulla with sparsely distributed dorsal tubercles. Hand slightly enlarged, its frontal region covered by tubercles of different sizes. Fixed fi nger with a proximal minute tooth, three median teeth decreasing in size distally and one subdistal laminar tooth. Movable fi nger with a proximal wide and low tooth, a small gap and three teeth (the proximal one rounded and the two distal forming a lamella). PEDIPALPUS (Fig. 14c). Trochanter with two tubercles on a dorsal protuberance, ventrally with two fused median tubercles and two ventrolateral tubercles. Femur strongly infl ated, dorsally curved and ventrally straight in lateral view, with a dorsoproximal row of forward curved tubercles, ventrally with a proximal group of three small tubercles in a common base and two ventral rows of tubercles, ectal largest. Patella short (ratio FePp/PaPp = 2.0), subsquare in dorsal view, smooth. Tibia infl ated, dorsally smooth; tibial  LEGS. Coxa I with one prolateral and one retrolateral tubercles; II with two prolateral, two dorsal and three retrolateral tubercles; III with one prolateral and one retrolateral tubercles; IV with few lateral tubercles and a prolateral group of conical granules. Trochanters I-IV fi nely granulate; trochanter I with one large mesoventral tubercle and few retrolateral tubercles; trochanter II ditto, with a group Fig. 15 (Fig. 14d). Patellae I-IV unarmed; the III-IV slightly swollen. Tibia I-IV unarmed, straight (Fig. 14d). Metatarsus IV without clear ringed marks on the subdistal portion. Claws III-IV smooth. Ratio Fe IV/scutum = 1.14. Tarsal counts in Table 8.
GENITALIA. Ventral plate sub trapezoidal with subdistal constriction and distal parabolic concavity., Ventral surface of VP with two elongated fi elds of small and needle-like microsetae with a longitudinal medial well marked elevation ( Fig. 15a-b). MS forming a single group distally positioned (A/D), composed by 12-13 pairs of acuminate and cylindrical setae (Fig. 15a-c, e). Two pairs of very small MS E close together in the distal region of the fl ange (Fig. 15b). Glans sac columnar elongate, with numerous proximal folders (Fig. 15a, e) Stylus curved, without processes (Fig. 15c). Stylar caps ringshaped without lateral or ventral projections (Fig. 15d).
COLORATION (in alcohol). Carapace with two paramedian areas of homogenous color paprika 38, three longitudinal stripes and posterior area reticulated squash yellow 83. Abdominal scutal areas reticulated between squash yellow 83 and pastel yellow 84. Paramedian tubercles of scutal area III, free tergites, anal operculum and coxae I--IV venetian red 41. Chelicerae and pedipalps reticulated dark wine 44 on light amber 71.
Female (ICN AO 1436.1) Similar to males, differing by: carapace shorter and lower, with ocularium and spines higher in lateral view (Fig. 14b, e); coda wider and longer; chelicerae smaller; pedipalpal femur, patella and tibia thinner; femur IV curved and slightly thinner. Sexual dimorphism evident in alpha males with pedipalpal femora much thicker than female (Fig. 14c, f-g).
Ovipositor. dl and vl rounded with three and two large, acuminated, single-tippedsetae respectively. Dl with three pairs of ds without the basal located pair. and the lateral region of the ovipositor with a pair of short dorso-lateral setae.

Diagnosis
Lateral border of DS and scutal areas I and IV densely granulate (see, e.g., Hara et al. 2014: fi gs 2-3); ocularium high, rounded; femur IV straight; Scutal area III with a pair of paramedian acuminate high subparallel tubercles (shared with P. cottus sp. nov.); pedipalpal claw of the males swollen (shared with P. anacosmetus and P. briareos sp. nov.); MS of the VP without obvious gap (slightly similar to P. kuryi sp. nov.).

Sexual dimorphism in Phalangodus
Sexual dimorphism is frequent in Gonyleptoidea, and the differences between the sexes are evident in numerous structures, mainly in males. The main characters that present striking differences are: chelicera hand swollen or armed with spines, tarsomeres infl ated, presence of modifi ed setae on the metatarsus, tibiae and femora of legs III and IV armed and swollen, stigmatic area elongated or armed posteriorly with processes or tubercles; carapace expanded posteriorly, scutal areas elevated (in females) or armed with bizarre tubercles (in males), anal operculum with differentiated tubercles and coloration pattern with additional spots on the females. More detailed information is summarized in table 9.
Within the Cranaidae, interesting cases of sexual and intra-sexual dimorphism can be found, with alpha and beta males recognized (see, e.g., Orrico & Kury 2009;Villarreal et al. 2015). The pedipalps of cranaids particularly, undergoes extreme modifi cation in some genera, acquiring an astonishing elongation in many Stygnicranainae, where some males have pedipalps longer than females or larger than females and other males (beta males), e.g., Tryferos Roewer, 1931and Stygnicranaus (Orrico & Kury 2009, 2009).
As explained by Villarreal et al. (2015), an intermediary degree of this elongation can be observed in some species of Ventrifurca Roewer, 1913, where only sexual dimorphism is known. Differences in size and stockiness of the femur between both sexes occur at least in Panalus Goodnight &Goodnight, 1947 andSpirunius Roewer, 1932 where males are larger and a dorsal crest is observed in males of Chiriboga Roewer, 1959 andCarsevennia Roewer, 1913 (unpublished data).
In Phalangodus both kinds of dimorphism occur, sexual and intrasexual. Sexually dimorphic pedipalps are present in all known species of the genus (e.g., Figs 9b-c, 11b-c, 14c, f-g) and dimorphic in "alpha" and "beta" males pedipalps are present at least in P. kuryi sp. nov. (Fig. 14c, g). Differences in length and height of the carapace between males and females are also present in some species of the genus. P. anacosmetus and some other species have males with ocularium-carapace ratio (h1/h3) higher than females (Fig. 1b, d). The male carapace of P. kuryi sp. nov.) is extended posteriorly, invading the scutal area I (Figs 13a, 14a). This expansion of the carapace however, is not exclusive to Phalangodus, being also present homoplasically in Chiriboga as noticed by Orrico & Kury (2009) char. 7(0),  char. 3(1) and Pinto-da-Rocha & Bonaldo (2011) char. 10(1). Three species of Phalangodus (P. anacosmetus, P. briareos sp. nov. and P. palpiconus) have sexual dimorphic claws in the pedipalps, being strongly swollen and dark at the base on the males. Some dimorphic characters like pedipalpal tarsal claw and extremely swollen femora on alpha males, seem to be restricted to a subgroup of Phalangodus, while other cases of probably homoplasic dimorphism appear in other genera in the family, like sexual differences in coloration (Stygnicranaus), presences of ventral apophysis on the coxae IV (Phareicranaus, Stygnicranaus), elongation of pedipalps (Stygnicranainae) and size of the carapace (Chiriboga). Interestingly, many kind of sexual dimorphism are found in Stygnicranainae species.

About the ovipositor morphology
Genital morphology of the females is poorly known in Gonyleptoidea. It has been neglected and rarely used as source of taxonomic characters. Although some authors illustrated the ovipositor of some species (e.g., Goodnight & Goodnight 1977;Šilhavý 1979;Kury 1990), only a few researchers have studied and used these characters (e.g., Martens et al. 1981;Bennett & Townsend 2013;Walker & Townsend 2014;Townsend et al. 2015;Brooks et al. 2015 (2016) and Brooks et al. (2015) have proposed and used names for the external structures associated to the ovipositor (lobes, setae, pores, etc.).
Nonetheless, we found issues regarding the structures proposed so far. Martens et al. (1981) referred to the presence of four external lobes with groups of peripheral sensory setae on the tip of the ovipositor of Laniatores. The fi rst ones to assign names for these lobes were Townsend et al. (2015), who used "anterior lobes" and "posterior lobes" (fi g. 1B) and "anterior row" and "posterior row" to name the setae associated to them (fi g. 1A). In the present article, after a topological analysis we propose alternative names, on the basis of topology of these structures on the ovipositor and its relative position to the body.
Following the widely used topology for the male genitalia structures, such as the ventral process, dorsal process and the ventral plate (Martens 1986;Kury 1994;Townsend et al. 2010;, the "anterior lobes" would be located on the dorsal face and the "posterior lobes" on the ventral face of the ovipositor. Therefore, we propose that the dorsal lobes (dl) and the ventral lobes (vl) be called "anterior lobes" and "posterior lobes" and consequently, dorsal rows (dr) and ventral rows (vr) instead of "anterior" and "posterior rows" (Fig. 7a-h). The peripheral setae (ds and vs) of all studied species are similar to those of P. calcariferus, with striated surface and straight and undivided tip.
The only cranaids for which SEM pictures of the ovipositor have been published is Phareicranaus calcariferus (Simon, 1879) (Bennett & Townsend 2013: fi g. 2A-F). Those images show at least one pair of accessory setae not assignable to any of the dorsal or ventral rows, positioned dorsolaterally and located slightly proximal on the ovipositor. That pair is named here as dorso-lateral setae (dls) (Fig.  7a-f). From six species of cranaids studied in the present article, the Dls group was found in all new species of Phalangodus here described, but was absent in Homocranaus sp. and Phareicranaus hermosa Pinto-da-Rocha & Kury, 2003 (Fig. 7g-h). Additionally, in all the studied species of Phalangodus, but not in P. kuryi sp. nov., an increase of the quantity of Dls occurs, (Fig. 7b, d, f) and could be a condition associated to some species of the genus.

Stylar caps in Cranaidae
Although the genital morphology of many species and genera of Cranaidae is still unknown, some characters from the cranaid penis have been proposed and increasingly used (e.g., stylar caps, shape of distal portion of the stylus, proximal folds of the glans, shape and number of MS C; Pinto-da-Rocha & Bonaldo 2011; Kury 2012a; Hara et al. 2014;Villarreal et al. 2015). Orrico & Kury (2009) referred to the presence of a laminar structure on the tips of the stylus in Stygnicranaus for the fi rst time, naming it "capellum" and the "lateral paired beaks of the stylus" associated to the capellum (characters 34 and 35). Pinto-da-Rocha & Kury (2003) and Pinto-da-Rocha & Bonaldo (2011) called this structure "stylar subdistal pointed apophysis" (character 16 and 19, respectively). Kury (2012a) described these structures in detail, renaming it as "Stylar caps". He also indicated their presence in many cranaids, including Phalangodus anacosmetus. In the particular case of P. anacosmetus, Kury (2012a) classifi ed its Stylar cap as "elongate-erythrocyte cap". Hara et al. (2014) and  interpreted the stylar caps as absent in Phalangodus, due to its unique shape. However, as the SEM images of Phalangodus show, these are present as an elongate-erythrocyte cap, corroborating Kury's previous description (Kury 2012a (2016) The shape and size of the Stylar caps are highly variable among the Cranaidae genera and at least three main types can be recognized: 1) elongate-erythrocyte SC; this is found in Phalangodus (Figs 15c-d, 16a), two species of Stygnicranaus (Stygnicranaus alessandroi Orrico &Kury, 2009 andS. poncedeleoni Orrico &Kury, 2009) and Tryferos elegans Roewer, 1931, as referred by Kury (2012a). This author proposed a similarity between the SC of these species and those found in Phareicranaus gracilis (Pinto-da-Rocha & Kury, 2003), P. singularis (H. Soares, 1970) and Zannicranaus morlacus Kury, 2012. Nonetheless, we consider the SC of these three species more similar to the kind described for other Amazonian Phareicranaus (e.g., P. divisor Pinto-da-Rocha & Bonaldo, 2011;P. onorei Pinto-da-Rocha & Kury, 2003;P. rohei Colmenares & Tourinho, 2014) and the Andean species Zannicranaus monoclonius Kury, 2012. 2) small, laminar and sub-square stylar cap; this is present in some species of Phareicranaus (Fig. 16d-e).
Additionally, a particular structure associated to the tip of stylus is known in Prostygnus Roewer, 1913 (Fig. 17a), which is more similar to those found in Cosmetidae (Fig. 17b), as mentioned in Kury (2012a). Possibly this is not homologous to the stylar caps of other cranaids.

Remarks about geographical records of P. anacosmetus
According to Kury (1996Kury ( , 2003, P. anacosmetus is recorded from three Colombian localities, two of them associated to P. anacosmetus (Cundinamarca and Boyacá) and the third linked to the type locality of Allocranaus giganteus (North of Santander), a species currently under synonymy with P. anacosmetus (see remarks on the species complementary description).