Taxonomic revision of the Andean genus Eulibitia Roewer, 1912 (Arachnida, Opiliones, Cosmetidae), with the description of fi ve new species

Abstract. The genus Eulibitia was described by Roewer including three species from Colombia and Ecuador: Eulibitia maculata Roewer, 1912, E. annulipes Roewer, 1912 and E. sexpunctata Roewer, 1914. Herein, the genus is revisited and only the type species, Eulibitia maculata Roewer, 1912 is maintained from the previous confi guration. Eulibitia sexpunctata is transferred to Ambatoiella MelloLeitão, 1943 and E. annulipes is considered as Cosmetinae incertae sedis. Acromares lateralis Goodnight & Goodnight, 1943 and Paramessa castanea (Sørensen, 1932) are here considered junior subjective synonyms of Messatana scalaris (Sørensen, 1932) and Eulibitia maculata, respectively. Brachylibitia Mello-Leitão, 1941, currently a junior synonym of Platymessa, is regarded as a synonym of Eulibitia. Platymessa Mello-Leitão, 1941, with two species, and the monotypic genera Messatana Strand, 1942 and Paramessa Mello-Leitão, 1933, are synonymized with Eulibitia Roewer, 1912, yielding the following new combinations: Eulibitia ectroxantha (for Brachylibitia ectroxantha Mello-Leitão, 1941), E. h-inscriptum (for Platymessa h-inscripta Mello-Leitão, 1941) and E. scalaris (for Libitia (Messa) scalaris Sørensen, 1932). Eulibitia is re-diagnosed and fi ve new species are described: E. castor sp. nov., E. pollux sp. nov., E. clytemnestra sp. nov., E. leda sp. nov. and E. helena sp. nov. A key to the nine Eulibitia species is presented; the morphology of Eulibitia is compared with that of Libitia, Libitiella and Ambatoiella.


Material and methods
Although the taxonomy of Cosmetidae is still unstable, an effort has been made here to provide a diagnosis of Eulibitia comparing it with genera that are morphologically similar (see Discussion). Therefore, a choice of the genera for comparison was made among cosmetids with the following characteristics: (1) small body, not exceeding 5 mm in length; (2) monomorphic chelicerae; (3) fl at body in lateral view, that is, without large tubercles or spines in the scutal areas; and (4) short legs, femur length not exceeding twice the body length. The types of Sørensen were photographed in the Sackler Biodiversity Imaging Laboratory of BMNH through their image capture system.
Descriptions follow  for external morphology; DaSilva & Gnaspini (2009) for the dorsal armature;  for the outline of the dorsal scutum; Kury & Villarreal (2015) and  for the chaetotaxy of macrosetae on the ventral plate of the penis; and Kury (2016b) for terminology of the microsetae. The order of numbers of tarsomeres in tarsus I to IV is from left to right side (fi gures in parentheses denote number of tarsomeres only in the distitarsi I-II). The caruncle (Ferreira & Kury 2010), also called wattle (Kury & Barros 2014), is the distal serrate crest of the stylus of Cosmetidae and closely related families that resembles a cockscomb. All measurements are in millimeters unless otherwise noted. The color patterns were based only on specimens preserved in 70% ethanol, using the standard names of the 267 color centroids of the NBS/IBCC Color System (Jaffer 2001-) as explained in Kury & Orrico (2006). Information on the material examined enclosed in square brackets [ ] was obtained during this study and is not part of the original label data. The geographical coordinates were obtained directly from the labels (unless in square brackets).

European Journal of
The specimens were submerged in 70% alcohol on blue sand to improve contrast and avoid movement. Dry specimens were necessary to illustrate tubercles of small size. The illustrations were made using a stereo microscope with a camera lucida and digitalized using the software Inkscape v. 0.48. Photographs were taken with a Sony Cybershot DSC-V1 camera attached to the stereo microscope. The software package COMBINEZ, version COMBINEZP (Hadley 2015), was used to create composite images with extended depth of fi eld. The resulting images were edited with Photoshop CS5. The plates were prepared in Inkscape for line drawings and Corel Draw X7 for photographs.
The treatment of the male genitalia follows Acosta et al. (2007). Drawings were made in Inkscape based on the series of photographs taken using a stereo microscope equipped with a digital camera and composed with COMBINEZ. Scanning electron microscopy was carried out with a JEOL JSM-6390LV at the Center for Scanning Electron Microscopy of MNRJ, with an accelerating voltage of 10 kV after sputter-coating with gold-palladium.

Diagnosis
Paramedian tubercles of areas I, III and posterior margin of scutum acuminate (Fig. 5E), tubercles of posterior margin higher than wide and slanted backwards (Fig. 5A, D-E). Ladder mask only present in the scutal groove between areas I and II (Fig. 6A). Coda short and mid-bulge symmetrical (Fig. 6A), clavi inguines composed of two tubercles. DS shape is very similar to that in E. pollux sp. nov., but differs because it is slightly more slender and the paramedian tubercles of the posterior margin are separated, with independent basis.   Roewer, 1912 (ICN-AO 875), dorsal and lateral view (from left to right) showing parts and interpretation of macrosetae, labeled with letters A-E.
DORSUM (Figs 5,6A). Dorsal scutum beta-shaped, with symmetrical mid-bulge. Areas I, III and posterior margin of scutum with two paramedian acuminate tubercles. Lateral margins with irregular row of minute granules at mid-bulge. Posterior margin of scutum with row of small tubercles. Tergites with row of small tubercles and anal operculum fi nely granular. VENTER ( Fig. 5C-D). Free sternites fi nely granular; coxae II-IV fi nely and uniformly granular; coxa I with longitudinal row of tubercles and smooth area corresponding to lace area of pedipalp.  (Fig. 7F). Basichelicerite uniformly occupied with tubercles of different sizes, some basal and one mesodistal larger; movable fi nger with row of eleven tubercles, giving a serrated appearance, fi xed fi nger with seven tubercles decreasing in size from basal to distal part of fi nger. PEDIPALPS ( Fig. 7D-E). Trochanter with strong ventral process. Femur with pronounced dorsal keel formed by fusion of six tubercles, with ventral row of nine setiferous tubercles and mesodistal process. Patella with mesal keel formed by six small tubercles.  Fig. 7A-C, G). Coxa IV granulated, with well-marked clavi inguines, distal apophysis of coxa IV slightly tuberculate and directed laterally. Trochanter IV with small retro-distal apophysis. Femur IV straight, with two longitudinal ventral rows of small tubercles along entire length, tubercles increasingly larger distally in prolateral row. Patella IV substraight, with small setiferous tubercles. Tarsal counts: 6(3)/14 (3)

Distribution
Only known from the type locality, in the WWF ecoregion Cordillera Oriental montane forest (NT0118), tropical and subtropical moist broadleaf forest biome, in Boyacá Department (Fig. 34).

Diagnosis
Paramedian tubercles of posterior margin of scutum rounded and contiguous, other areas fi nely granulated, without tubercles (Fig. 9A). Ladder mask only present in area II, entering only a little into scutal grooves (Fig. 8A). Differs from other species of the genus because of the smooth lateral margins (Fig. 9A). Coda very short and mid-bulge asymmetrical, coxa IV slightly granulated, without clavi inguines (Figs 8A, 9A). Femur IV straight, without conspicuous ornamentation (Figs 8D, F, 10A-C).

Etymology
Noun in apposition from Clytemnestra, daughter of Leda and sister of Helen of Troy, from Greek mythology. Two of the new species of Eulibitia have been named after Helen and Clytemnestra to complete the four siblings originating from the union of Zeus and Leda. of scutum with row of small tubercles. Tergites with row of small tubercles and anal operculum fi nely granular. VENTER (Fig. 8C, E). Free sternites fi nely granular; coxae II-IV fi nely and uniformly granular; coxa I with longitudinal row of tubercles and smooth area corresponding to locking area of Pp Tr. CHELICERAE (Fig. 10F). Basichelicerite with minute tubercles on ecto-basal side and two on antero-mesal corner; movable fi nger with row of tubercles, giving a serrated appearance, and large basal tubercle; fi xed fi nger with fi ve tubercles decreasing in size from basal to distal part of fi nger. PEDIPALPS ( Fig. 10D-E). Trochanter with strong ventral process. Femur with pronounced dorsal row of seven tubercles, with ventral row of ten setiferous tubercles and mesodistal process. Patella with low mesal keel. LEGS (Figs 8F, 10A-C, G). Coxa IV fi nely granulated, without clavi inguines, distal apophysis of coxa IV slightly tuberculate and directed laterally. Trochanter IV with small retro-distal apophysis. Femur IV straight, with two longitudinal ventral rows of small tubercles along entire length. Patella IV substraight, with small tubercles. Tarsal counts: 6(3)/12-11(3)/7-?/8. COLOR (Fig. 8). Body and appendages color background 68 (Strong Orange Yellow), ladder mask 104 (Pale Greenish Yellow). Trochanter and tarsomeres 98 (Brilliant Greenish Yellow). GENITALIA ( Fig. 9B-C). Ventral plate subrectangular, narrower basally and distal border hardly concave; dorsal apophysis of glans long and rounded, wattle long. Shapes and organization of macrosetae as follows: MS C1-C2 large, curved and fl at; MS D1-D2 large and straight, D2 smaller than D1 and located closer to MS A1; MS A1-A2 large, cylindrical, straight, located in basal middle of ventral plate (asymmetry in number of MS A already seen in E. maculata); MS B and MS E1-E2 ventral, very small and immersed in microsetae. Pair of MS B at base of ventral plate.

Female
Unknown.

Diagnosis
Paramedian tubercles of posterior margin of scutum acuminate. Differs from other species of the genus in the presence of paramedian tubercles in area III but without any armature in area I, and the ladder mask with rungs only, no rails in scutal grooves I, II and III (see Mello-Leitão 1941 andKury 2016).

Remarks
This species was discussed and tagged as species inquirenda in  and the available information is suffi cient to include it within the new diagnosis of Eulibitia. (Mello-Leitão, 1941)

Diagnosis
E. h-inscriptum has a pair of paramedian tubercles in areas I, III and on the posterior margin, but differs from its congenerics because its ladder mask is not confi ned to the mesotergum, reaching the lateral margins of the scutum.

Remarks
This species was recently redescribed by  and their morphological features match the new diagnosis of Eulibitia.

Etymology
Noun in apposition for Helen of Troy, daughter of Leda and cause of the Trojan War in Greek mythology.
DORSUM (Figs 11,12A). Dorsal scutum beta-shaped, with slightly asymmetrical mid-bulge. Areas I, III and posterior margin with two paramedian rounded tubercles. Lateral margins of scutum irregularly granulate. Posterior margin of scutum with row of small tubercles. Tergites with row of small tubercles and anal operculum fi nely granular.  Fig. 11C-E). Free sternites fi nely granular; coxae II-IV fi nely and uniformly granular; coxa I with longitudinal row of tubercles and smooth area corresponding to lace area of pedipalp. CHELICERAE (Fig. 13A). Basichelicerite with small tubercles on dorsal surface and some larger ones on ecto-basal side, movable fi nger with row of ten small tubercles, giving a serrated appearance; fi xed fi nger with fi ve tubercles, decreasing in size from basal to distal part of fi nger. PEDIPALPS ( Fig. 13B-C). Trochanter with strong ventral process. Femur with pronounced dorsal row of seven tubercles, with ventral row of fi fteen setiferous tubercles and mesodistal process. Patella with low mesal keel ending in setiferous tubercle. LEGS ( Fig. 12B-E). Coxa IV fi nely granulated, with clavi inguines, distal apophysis of coxa IV slightly tuberculate and directed laterally. Trochanter IV with small retro-distal apophysis. Femur IV straight, with two longitudinal ventral rows of small tubercles along entire length, dorsal row of tubercles increasing in size, with large apical spine. Patella IV substraight, with small tubercles. Tarsal counts: 6(3)/14-?(3)/7/8. COLOR (Fig. 11). Body color background 49 (Brilliant Orange), ladder mask 104 (Pale Greenish Yellow). Appendages 67 (Brilliant Orange Yellow). GENITALIA (Fig. 14). Ventral plate subrectangular, narrower basally and distal border substraight, dorsoapical membranous invaginations basal to MSC; dorsal apophysis of glans long and rounded, wattle long. Shapes and organization of macrosetae as follows: MS C1-C2 large, curved and fl at; MS D1-D2 large and straight, D2 smaller than D1 and located closer to MS A1; MS A1-A2 large, cylindrical, straight, located on basal middle of ventral plate; MS B and MS E1-E2 ventral, very small and immersed in microsetae. Pair of MS B at base of ventral plate.

Variation
There is a limited sample of blots (n = 7), the parallel bars (rails) may be more or less wide and the posterior stripe is sometimes interrupted by the paramedian tubercles (Figs 11A, 12A). The dorso-distal tubercle of Fe IV may seem bifurcate by the growth of the tubercles of its own base. Tarsal counts: 5-6; 8-12; 5-6; 6-7. Variation of measurements is given in Table 1.

Distribution
This species is very similar to E. scalaris in the shape of the ladder mask, but differs by having a short coda with the constriction poorly marked (Fig. 15A), the basichelicerite bears larger tubercles (Figs 15A, 17B) and the femur of leg IV is more tuberculated in the ventro-distal region ( Fig. 17G-H, J). Apophyses of coxa IV acuminate and poorly tuberculated (Figs 15, 16A).

Etymology
Noun in apposition of Leda, mother of Clytemnestra, Castor, Pollux and Helen of Troy, from Greek mythology. This species has been named Leda to round up the family formed from the union of Zeus and Leda. Paratypes COLOMBIA: 5 ♂♂, 5 ♀♀, same collection data as for holotype (ICN-AO 1165.1); 1 ♂, 1 ♀, same collection data as for holotype (MNRJ 9264).
DORSUM (Figs 15, 16A). Dorsal scutum beta-shaped, with asymmetrical mid-bulge. Areas I, III and posterior margin with two paramedian tubercles, tubercles of posterior margin bifi d. Lateral margins with irregular row of minute granules at mid-bulge. Posterior margin of scutum with row of small tubercles. Tergites with row of small tubercles and anal operculum fi nely granular.   (2017) 26 CHELICERAE (Fig. 17A-B). Basichelicerite with row of large tubercles from posterior border to ectal side; movable fi nger with row of ten tubercles, fi xed fi nger with two small basal tubercles and fi ve large tubercles decreasing in size from basal to distal part of fi nger. PEDIPALPS (Fig. 17C-F). Trochanter with strong ventral process. Femur with pronounced dorsal keel formed by row of tubercles, with ventral row of twelve setiferous tubercles and mesodistal process. Patella with mesal keel without tubercles but with distal process. Shallow slit along tibia mesal surface, separating dorsal and ventral sides.

Female
With larger clavi inguines and longer coda than males. Males with thicker basitarsus (but not notably swollen) (Fig. 16B).

Diagnosis
This species differs from the other species of Eulibitia by the absence of paramedian tubercles on the posterior margin of the scutum (Fig. 19D-E). The ladder mask can occupy the groove between areas III and IV (Fig. 20A, C-H). This species is similar to E. helena sp. nov. and E. clytemnestra sp. nov. by the shape of DS, with mid-bulge asymmetrical, but differs from the former by the absence of clavi inguines (Figs 19A,20A) and by not having larger tubercles on distal femur IV (Fig. 21B-C), and from the latter by the presence of a row of tubercles on the lateral margins of DS (Fig. 20A).

Variation
Pattern of yellow spots as in Fig. 20C-H. MS A of VP of penis may be variable and asymmetrical in number from two to three, as shown in Figs 3 and 22, corresponding to two specimens from same lot. Tarsal counts: 5-6; 9-10; 6; 6-7. Variation of measurements is given in Table 3.

Distribution
Eulibitia maculata occurs in the WWF ecoregions: (1) Magdalena Valley montane forests (NT0136), tropical and subtropical moist broadleaf forest biome, and (2) Northern Andean páramo (NT1006), Roewer, 1912, ♂ (ICN-AO 875, different specimen than used in Fig. 2A-B montane grassland and shrubland biome; in Tolima, Cundinamarca and Boyacá Departments (Fig. 32). The localities given by Roewer for Eulibitia maculata are rather imprecise: Tolima (Roewer 1912) and "Neu Granada" ; probably referring to Viceroyalty of New Granada, territory corresponding to modern Colombia, Ecuador, Panama, and Venezuela under the jurisdiction of the Spanish Empire until 1819). Tolima Department has an area of 23.562 km 2 and four WWF ecoregions can be distinguished in it; here, a point in NT0136 of the eastern region of the department was chosen to coincide with the other points of occurrence of the species.

Remarks
Although Roewer (1912) described various characteristics of the specimen ZMH 458, he labeled the lectotype SMF RI/471 as "Type", which does not entirely match the original description and certainly is a different species. Therefore, the use of ZMH 458 ("ex-typo", probably belonging to Ambatoiella) has masked the identity of the real Eulibitia (see for example Pinto-Da-Rocha & Hara 2011, who mentioned characteristics of ZMH 458 in their table 1, although they cited SMF RI/471 as the "holotype" in their text).
The paralectotype ZMH 458 (used for original description) differs from the lectotype (designated here) by: (1) abdominal scutum reaching its maximum width in scutal area II (vs area III in the lectotype); (2) scutal area IV and the posterior margin with two small tubercles; (3) coda long, of same length as the mid-bulge; (4) ectal border of protoglyphs bifurcated, and (5) presence of clavi inguines. The use of those features by Roewer (1928) and Mello-Leitão (1932, 1933 to diagnose the genus, masked the identity of Eulibitia and made the detection of synonymies problematic. The species Paramessa castanea was considered a different species from Eulibitia maculata, possibly for two reasons: (1) the blot pattern of DS, which is absent in the holotype of P. castanea, and the full ladder mask in the lectotype of E. maculata (see Figs 20C-H for variation); or (2) the designation of different species in the type series, since Roewer designated ZMH specimens of a species more similar to Ambatoiella as paratypes, as explained above.
Eulibitia pollux sp. nov. urn:lsid:zoobank.org:act:522C7A02-61AC-45D7-8F84-C42E01F1EC46 Figs 23-26, 31B, 34; Table 4 Diagnosis This species is similar to E. castor sp. nov. in having the paramedian tubercles on the posterior margin of the scutum acuminate, but differs from it because the other areas are unarmed (Figs 23A-B, D-E, 24A). Ladder mask often as paired points in scutal grooves (Fig. 24C); in its full form it covers the scutal groove between areas I and II with a spot with a shape of a mustache (Fig. 24F). Coda short, with symmetrical mid-bulge (Figs 23A, 24A); coxa IV slightly granulated, without clavi inguines (Fig. 24A). Femur IV curved in dorsal view, without conspicuous ornamentation ( Fig. 25A-C). This is the only species of Eulibitia in which macrosetae D2 are minute, of the same size as MS E and MS B (Fig. 26B).

Etymology
Noun in apposition from Pollux, son of Leda and Zeus, and twin of Castor. From Greek mythology.   (Figs 23, 24A). Dorsal scutum beta-shaped, with symmetrical mid-bulge. Posterior margin of scutum with two contiguous acuminate paramedian tubercles. Lateral margins of scutum granulated. Posterior margin of scutum with a row of small tubercles. Tergites with row of small tubercles and anal operculum fi nely granular. VENTER (Fig. 23B, E). Free sternites fi nely granular; coxae II-IV fi nely and uniformly granular; coxa I with longitudinal row of tubercles and smooth area matching interlocking Tr of pedipalpus. CHELICERAE (Fig. 25F). Basichelicerite with row of seven small tubercles on basal side and some disperse tubercles on dorsal side; movable fi nger with row of eleven tubercles, giving a serrated appearance; fi xed fi nger with fi ve tubercles, decreasing in size from basal to distal part of fi nger. PEDIPALPS (Fig. 25D, E). Trochanter with strong ventral process. Femur with pronounced dorsal row of seven separated tubercles, with ventral row of seven setiferous tubercles and mesodistal process. Patella with low mesal keel.

Diagnosis
This species is similar to E. h-inscriptum by having paramedian tubercles in areas I, III and on the posterior margin of the scutum, but differs from it because its ladder mask does not reach the lateral margins of the scutum (Fig. 28C-H), and from the other species of Eulibitia by the size of the coda (Fig. 27A), ⅓ of the mid-bulge length (vs ¼ of the mid-bulge length in other species, except E. h-inscriptum). It is similar to E. castor sp. nov. by having two paramedian tubercles in areas I, III and on the posterior margin, and for the symmetrical mid-bulge, but it differs from it by the shape of the tubercles on the posterior margin, being rounded instead of acuminate (Fig. 28A).

Etymology
Scalaris: refers to the shape of the ladder of white blots on DS. Lateralis: refers to the rails of the ladder mask on DS (visible in the holotype of Acromares lateralis).

Material examined
Syntypes COLOMBIA: 2 ♂♂, 2 ♀♀, Cundinamarca Department, Bogotá (BMNH, examined by photograph).  mid-bulge. Posterior margin of scutum with row of small tubercles. Tergites with row of small tubercles and anal operculum fi nely granular. VENTER (Fig. 27D-E). Free sternites fi nely granular; coxae II-IV fi nely and uniformly granular; coxa I with longitudinal row of tubercles and smooth area corresponding to lace area of pedipalp.  Fig. 29F-G). Basichelicerite uniformly covered with tubercles of different sizes, some basal and one mesodistal larger; movable fi nger with row of ten tubercles; fi xed fi nger with four tubercles, decreasing in size from basal to distal part of fi nger. PEDIPALPS (Fig. 29D-E). Trochanter with strong ventral apophysis. Femur with pronounced dorsal keel, with ventral row of eight setiferous tubercles and mesodistal process. Patella with mesal keel formed by fi ve small tubercles and large one distally. Shallow slit along tibia mesal surface, separating dorsal and ventral sides.

Female
With larger clavi inguines and longer coda than males ( Fig. 28F-G). Males with thicker basitarsus (but not notably swollen) (Fig. 28B).   Goodnight & Goodnight (1943) described Acromares lateralis and placed it in that genus because they thought that the outstanding tubercles of the dorsal scutum were localized in area IV. However, examination of a photograph of the type material allows us to conclude that the tubercles are on the posterior margin of the scutum instead of on area IV and to recognize this species as a synonym of E. scalaris comb. nov.

Remarks
This species is morphologically similar to Ambatoiella vigilans Mello-Leitão, 1943 by: (1) coxa IV only apically visible in dorsal view, and (2) coda large, about one third the length of DS. Likewise, the type localities of the two species are only 50 k m apart from each other and are in the same mountain range. There is a strong possibility that they are male and female of the same species; however, we decided to maintain them as separate species until further analyses have been carried out.

Remarks
The size of the body, shape of the outline of DS and the strong and uniform granularity of the scutum do not correspond to the genus Eulibitia. It was impossible to fi t this species in any known genus; the most similar genus is Ambatoiella based on the alpha-shaped scutum, but an analysis of the type material or topotypes is needed to allow an accurate decision.

Morphology
Eulibitia seems to be a good example of the conservative external morphology of Cosmetidae mentioned above in the Introduction; the variations in shape of outline of DS and the morphology of pedipalpus, legs and chelicera are extremely subtle, and the delimitation of species may be diffi cult. Nonetheless, some characteristics have proved useful to differentiate species of Eulibitia, such as: (1) the armature of the scutal areas, commonly useless for diagnosing genera; (2) the armature of femur IV; and (3) the occurrence of clavi inguines.
The morphology of the reproductive organs in Gonyleptoidea has been shown to be very valuable in phylogenetic analyses (i.e., Kury & Villarreal 2015;Cruz-López & Francke 2015), and has been well explored in detail for some species (i.e., Bennett & Townsend 2013;Walker & Townsend 2014;Townsend et al. 2015). In Cosmetidae, however, the genital morphology is somewhat uniform, if compared with Gonyleptidae for example. In the species of Eulibitia only the size of MS D2 varies. It is probable that the genital morphology in Cosmetidae is more useful in defi ning genera or suprageneric groups (e.g., the enormous caruncle of the stylus in Libitia, the small MS A in Ambatoiella and the long VP in Discosomaticinae such as Gryne).  Roewer, 1912 andE. h-inscriptum (Mello-Leitão, 1941) comb. nov. Colored patches on the background represent WWF ecoregions.

Relationships of Eulibitia
The Andean genera Ambatoiella, Libitia and Libitiella seem to be related to Eulibitia in having: (1) legs short and femur IV sometimes slightly curved; (2) body almost unarmed, with tubercles or spines short or minute in the scutal areas; (3) absence of marked sexual dimorphism, such as hypertelic chelicera in males; (4) basitarsomeres of leg I of males clearly thicker than distitarsomeres (except in Ambatoiella, where they are monomorphic, equally sturdy), but not enormously enlarged as in other more distantly related genera (e.g., Cumbalia Roewer, 1963, Fig 2); (5) macrosetae D1 closer to C2 than to D2; and (6) presence of a soft latero-distal area on VP (at macrosetae D1) which may be sunken, forming a noticeable notch (Figs 4A, E, 14A, 18A, 22A, 26A, 30B). Although no phylogenetic analyses have been carried out in Cosmetidae, these genera seem to form a natural group. Cumbalia shows other differences to the core of the Eulibitia-like Andean genera, such as: (1) the basalmost basitarsomere is much longer than the second one (as opposed to being subequal, regardless of whether counting two or three articles) and (2) Mt III and IV of male tapering (as opposed to ordinarily cylindrical rods).
On the other hand, the genus Eulibitia also seems to be related to the Amazonian genus Taito, which exhibits common characteristics like beta-type DS, clavi inguines sometimes present, slight sexual dimorphism and a transverse yellow stripe on the posterior margin of the scutum. However, this genus differs from Eulibitia by: (1) presence of equuleus in DS; (2) posterior margin of scutum never with paramedian tubercles; (3) anal operculum of males sometimes with robust tubercles; (4) legs long and slender; and (5) dorsal apophysis of coxa IV large and directed backwards; in addition to this are the features mentioned in Kury & Barros (2014) as possible synapomorphies, such as the calli (latero-distal projections of the truncus), which are not present in species of Eulibitia (Fig. 26).

Remarks on distribution
The genus Eulibitia is confi ned to the northern Andean montane forests of Colombia, with a marked preference for the Magdalena Valley montane forest (NT0136) and Northern Andean páramo (NT1006) Fig. 34. Northern Andes in Colombia, showing localities for Eulibitia ectroxantha (Mello-Leitão, 1941) comb. nov., E. pollux sp. nov., E. castor sp. nov. and E. helena sp. nov. Colored patches on the background represent WWF ecoregions. ecoregions, in two biomes: tropical and subtropical moist broadleaf forests, and montane grasslands and shrubland.
Species of Eulibitia are widely sympatric, which is why they are presented here on three maps (Figs 32-34) to avoid a muddled overlap. Eulibitia scalaris is the most abundant species in collections and the most widespread, occurring in three different biomes corresponding to three ecoregions: tropical and subtropical moist broadleaf forests (Magdalena Valley montane forests -NT0136), tropical and subtropical dry broadleaf forests (Magdalena Valley dry forests -NT0221) and montane grasslands and shrubland (Northern Andean páramo -NT1006).
There is a single point of occurrence of E. pollux sp. nov. in the Cauca Valley montane forest ecoregion (NT0109), in Palmira (Valle del Cauca), a place isolated from the other records of the species and even the genus. The marked preference of species of Eulibitia for the Northern Andean páramo (NT1006) ecoregion, contrasting with the lowland city Palmira (1040 m a.s.l.) and the Magdalena River and the Central Cordillera as physical barriers, suggest that this record should be considered doubtful.