A revisionary approach of Colombian Ananteris (Scorpiones, Buthidae): two new species, a new synonymy, and notes on the value of trichobothria and hemispermatophore for the taxonomy of the group Author Botero-Trujillo, Ricardo Laboratorio de Entomología, Unidad de Ecología y Sistemática-UNESIS, Departamento de Biología, Pontificia Universidad Javeriana, Bogotá, Colombia. E-mail: pachyurus @ yahoo. com Author Flórez, Eduardo Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá, Colombia. E-mail: aeflorezd @ unal. edu. co text Zootaxa 2011 2011-06-02 2904 1 1 44 https://biotaxa.org/Zootaxa/article/view/zootaxa.2904.1.1 journal article 10.11646/zootaxa.2904.1.1 1175-5326 5284774 9594B61B-DF8E-460B-BFA1-374E5F472C4A Ananteris arcadioi Botero-Trujillo, 2008 Figures 1–5 , 32 , 40 , 48 , 60 ; Tables 1 , 2 Ananteris arcadioi Botero-Trujillo, 2008a: 287 , 288–292, 293, 295, 297, figs. 1–11, tables 1–2; Botero-Trujillo 2009: 93, fig. 1; Botero-Trujillo & Noriega, 2011: 38 , 40, 44, figs. 7–10. Type material. Holotype male: COLOMBIA : Meta Department : Puerto Gaitán , Altamira , Club Los Llaneros , 04°19’N 72°05’W , 140 m asl , into forest, direct collection, at night, 19 October 2006 , I. Gélvez (MPUJ-SCO-356) (examined). Further material examined. COLOMBIA : Meta Department : 3 adult males , 1 adult female, 1 subadult female, Puerto López , Remolinos , Cafam Llanos , 04°16’28’’N 72°32’19’’W , 183 m asl , UV light, 03–05 September 2008 , J. A. Ochoa & R . Botero-Trujillo (MPUJ-SCO-383) . 2 adult males, 2 adult females, Puerto López , Remolinos , Cafam Llanos , 04°16’28’’N 72°32’19’’W , 183 m asl , UV light, 03–05 September 2008 , J. A. Ochoa & R . Botero-Trujillo (ICN-As-773) . 6 adult males, 2 adult females, between Villavicencio and Restrepo , Farm Con Esto Tengo , 04°11’51’’N 73°35’37’’W , 407 m asl , UV light, 07 September 2008 , R . Botero-Trujillo & J. A. Ochoa (MPUJ-SCO-384) . 7 adult males, 1 adult female, 1 juvenile female, between Villavicencio and Restrepo , Farm Con Esto Tengo , 04°11’51’’N 73°35’37’’W , 407 m asl , UV light, 07 September 2008 , R . Botero-Trujillo & J. A. Ochoa (ICN-As-774) . 1 adult male, Serranía de La Macarena , 05 March 1992 (ICN-As-641) . 1 adult male, 2 adult females, San Martín , Rey Zamuro Natural Reserve , 03°31’57.46’’N 73°30’52.35’’W , 317 m asl , UV light, March 2010 , J. A. Noriega (MPUJ-SCO-398) . Revised diagnosis. Carapace with well-developed anteromedian projection ( Fig. 32 ); metasomal carinal formula 10:10:10:6:5, with ventrosubmedian and median lateral carinae absent on segment IV; median lateral carinae on metasomal segment III only weakly developed, evident in females but vestigial in males, especially posteriorly; V 1 and V 2 trichobothria unaligned axially, with V 2 located on an external position in relation to V 1 ( Fig. 40 ); fixed finger trichobothria in the order eb : esb : est / db : et : dt , with est located beside db or nearly so ( Fig. 48 ); dorsal surface of chelicerae with incomplete reticular pattern ( Fig. 32 ); pectines with 17–21 teeth on males (mode = 18), 16–18 on females (mode = 18); sternite VII with paramedian longitudinal carinae vestigial and incomplete; pedipalp hand base color yellow most usually with conspicuous brown areas ventrally, internally and externally (brown may be absent on the external surface), but infrequently completely yellow; coxosternal region entirely yellow and spotless. Hemispermatophore: capsular region with median, external and internal lobes; flagellum with long and strongly coiled pars reflexa ( Fig. 60 ). Male. Holotype described by Botero-Trujillo (2008a) ( Figs. 2–3 ). Hemispermatophore: Description based on specimens from MPUJ-SCO-383 and ICN-As-774: Flagelliform, thin and poorly sclerotized. Foot narrow and flat. Pedal flexure inconspicuous but movable. Body very long, wider on basal third. Capsular region with median, external and internal lobes, subequal and strong. Flagellum long, with short pars recta and long, coiled pars reflexa ( Fig. 60 ). Carapace length to hemispermatophore body length ratio = 1:1.08 to 1:1.04. There was no variation between the four hemispermatophores examined. Female. Description based on adult female from MPUJ-SCO-383. Measurements: see Table 2 . Coloration: general coloration yellowish with variegated pigmentation over almost the entire body and appendages ( Figs. 4–5 ). Carapace predominantly brown with some yellow spots and stripes; anterior and posterior margins brown; arising posteriorly to each lateral ocular tubercle there is a thin and almost straight yellow stripe that ends behind the median ocular tubercle; median ocular tubercle black, surrounded on its base by thin yellow lines on anterior, anterolateral and posterolateral margins. Chelicerae with coxa predominantly yellow with abundant pale-brown mottling; hand with incomplete reticular pattern but abundant reticulations on dorsal surface ( Fig. 32 ); fixed finger yellow with reddish teeth, movable finger with a basal brown area and reddish teeth. Coxosternal region, genital operculum, pectinal basal piece, pectines and sternite III entirely yellow and spotless; sternites IV– VI predominantly yellow, with some minute and inconspicuous brownish spots paramedially; sternite VII with large darkbrown lateral areas; sternite V with a small flattened posterior median hyaline area. Tergites predominantly brown; twο ¹οngitudina¹ ye¹¹οw StripeS crοSSing tergiteS I— VI are οn¹y evident οn the pοSteriοr ha¹f οf each Segment beSide the midline; each side of tergites I– VI with two transverse yellow lines converging near the longitudinal lines, arrow-like; tergite VII with variegated pigmentation; lateral margins of tergites I–VII yellow, with an anterior small brown area. Metasoma predominantly yellowish; dorsal intercarinal spaces of segments I–IV with a median brown design wider anteriorly, triangle-like on segments I–III but arrow-like on segment IV; segments I– V with variegated pigmentation on all surfaces; segment V reddish with brownish areas. Telson vesicle yellowish-red with brown areas ventrally and laterally; subaculear tubercle yellowish. Pedipalp femur and patella predominantly brown dorsally and externally, with conspicuous yellow areas; primarily yellow ventrally and externally with abundant brown areas; both segments with trichobothrial pits yellow. Pedipalp coxa and trochanter yellow with large brown regions dorsally; hand with base color yellow and conspicuous brown areas ventrally, internally and externally; fingers entirely brown. Legs with variegated pigmentation. Carapace: densely covered with rounded granules especially on the brown areas; lateral margins converging anteriorly; anterior margin with well-developed median projection ( Fig. 32 ); ocular carinae moderately strong and granulose, others inconspicuous; with a median depression anterior to the ocular tubercle; postocular furrow evident; median ocular tubercle low, located on the posterior half of the anterior third of carapace; lateral ocular tubercles each with three evident ocelli. Chelicerae: with abundant fine white setae on the internal and ventral surfaces; cheliceral dentition characteristic of the family Buthidae ( Vachon 1963 ) . Movable finger dorsally with two small basal teeth, one median pronounced, one subdistal slightly shorter than the median, and one distal tooth. Ventrally with three strong teeth, one basal, one median, and one distal tooth larger than its dorsal counterpart. Fixed finger dorsally with one basal and one median tooth forming a bicuspid, one subdistal, and one distal tooth. Ventrally with only one subdistal tooth slightly basal to the dorsal subdistal. Coxosternal region: sternum subtriangular, with deep median depression and two anterolateral furrows; all the components of this region smooth, with few setae; coxapophyses I–II densely setose anteriorly. Genital operculum and pectines: genital operculum completely divided longitudinally; pectinal basal piece as long as wide, pentagonal and deeply notched anteriorly; pectines small, not reaching posterior end of coxae IV; count of pieces on the pectines: marginal lamellae 3:3, middle lamellae 8:10, teeth 17:18, fulcra absent. Sternites: with abundant setae; III– VI smooth, except for some granulations on the lateral margins of V – VI ; VII with vestigial and incomplete paramedian longitudinal carinae and some granules on the brown regions, lateral carinae completely absent; V with a small flattened posterior median smooth area; spiracles oval elongated, approximately twice longer than wide. TABLE 2. Meristic data for Ananteris species. Measurements in millimeters. 1 Sum of prosoma, mesosoma and metasoma. 2 Sum of tergites I–VII. 3 Sum of metasomal segments I–V and telson. 4 Sum of femur, patella and chela. 5 Measured from the commissure of the junction with the movable finger to the finger tip. 6 Measured along the marginal lamella. See for other measurements: Lourenço (1991) and Teruel & Roncallo (2008) for A. columbiana , Lourenço (1994) for A. ehrlichi , Botero-Trujillo (2007) for A. myriamae , Botero-Trujillo (2008a) for A. arcadioi and A. dorae .

A. arcadioi	A. dorae	A. ehrlichi	A. myriamae	A. ochoai sp. nov.	A. solimariae sp. nov.
♀ (MPUJ- SCO-383)	♂ (ICN- As-816)	♂ (IAvH-E 100814)	♂ (MPUJ- SCO-382)	♂holo/♀allo	♂holo/♀allo
Total body L:1	29,47	15,06	31,84	21,20	29,05/41,80	25,53/34,32
Carapace:	L:	3,53	1,83	3,58	2,51	3,21/4,70	3,03/4,00
Anterior W:	2,42	1,17	2,16	1,60	2,26/3,40	1,87/2,79
Posterior W:	3,63	1,75	3,42	2,44	3,05/5,00	2,79/4,16
Mesosoma:	Total L:2	8,21	3,79	8,05	5,20	6,79/11,15	6,11/9,58
Metasoma:	Total L:3	17,74	9,45	20,21	13,49	19,05/25,95	16,39/20,74
Metasomal	L:	1,84	0,85	1,89	1,27	1,84/2,65	1,53/2,05
segment I:	W:	2,16	1,09	2,00	1,38	1,79/2,70	1,68/2,53
H:	1,89	0,95	1,79	1,24	1,55/2,40	1,58/2,16
Metasomal	L:	2,00	0,97	2,26	1,45	2,11/3,10	1,79/2,32
segment II:	W:	2,05	1,03	1,84	1,27	1,68/2,50	1,63/2,26
H:	1,84	0,99	1,74	1,20	1,53/2,30	1,53/2,11
Metasomal	L:	2,16	1,07	2,53	1,60	2,42/3,40	2,00/2,58
segment III:	W:	2,05	1,03	1,84	1,27	1,68/2,50	1,63/2,26
H:	1,79	0,99	1,74	1,16	1,58/2,35	1,53/2,05
Metasomal	L:	2,95	1,48	3,63	2,18	3,47/4,70	2,79/3,42
segment IV:	W:	2,00	1,07	1,84	1,27	1,68/2,60	1,61/2,26
H:	1,84	1,03	1,79	1,20	1,71/2,45	1,58/2,26
Metasomal	L:	4,63	2,64	5,42	3,53	5,11/6,70	4,39/5,37
segment V:	W:	2,05	1,07	1,79	1,35	1,66/2,55	1,68/2,32
H:	1,95	1,01	1,79	1,27	1,63/2,40	1,66/2,21
Telson:	L:	4,16	2,44	4,47	3,45	4,11/5,40	3,89/5,00
Vesicle W:	1,21	0,76	0,89	0,73	0,84/1,50	0,84/1,47
Vesicle H:	1,11	0,68	0,89	0,69	0,97/1,55	0,84/1,26
Pedipalp:	Total L:4	11,68	6,33	12,11	9,13	12,16/17,58	10,21/13,92
Pedipalp femur:	L:	3,16	1,69	3,42	2,51	3,42/4,90	2,74/3,74
W:	0,89	4,27	0,84	0,58	0,76/1,15	0,74/1,05
H:	0,68	3,30	0,63	0,44	0,58/0,90	0,58/0,79
Pedipalp patella:	L:	3,84	2,12	4,00	2,95	4,00/5,78	3,37/4,58
W:	1,21	5,83	1,00	0,73	0,89/1,50	1,00/1,47
H:	1,05	5,44	0,79	0,58	0,74/1,30	0,84/1,21
Pedipalp chela:	L:	4,68	2,53	4,68	3,67	4,74/6,90	4,11/5,60
W:	0,82	0,39	0,68	0,55	0,63/1,10	0,63/1,00
H:	0,84	0,39	0,74	0,55	0,68/1,20	0,68/1,05
Fixed finger L:5	3,16	1,78	3,21	2,51	3,37/4,75	2,79/3,74
Movable finger L:	3,58	1,94	3,55	2,76	3,74/5,30	3,11/4,21
Palm L:	1,11	0,58	1,16	0,91	1,05/1,60	1,00/1,53
Pectines:	L:6	2,58	1,46	3,11	2,22	2,89/3,60	2,47/2,74

FIGURE 1. Known distribution of Colombian Ananteris . () Ananteris arcadioi Botero-Trujillo, 2008 . () Ananteris columbiana Lourenço, 1991 . () Ananteris dorae Botero-Trujillo, 2008 . () A. dorae and Ananteris ochoai sp. nov. () Ananteris ehrlichi Lourenço, 1994 . () Ananteris faguasi Botero-Trujillo, 2009 . () Ananteris gorgonae Lourenço & Flórez, 1989 . () Ananteris leilae Lourenço, 1999 . () Ananteris meridana González-Sponga, 2006 (also type locality in Venezuela). () Ananteris myriamae Botero-Trujillo, 2007 . () Ananteris solimariae sp. nov. () Ananteris tolimana Teruel & García, 2007 . () Ananteris volschenki Botero-Trujillo, 2009 . FIGURES 2–5. Ananteris arcadioi Botero-Trujillo, 2008 . 2–3. Holotype male, dorsal and ventral aspects (MPUJ-SCO-356). 4–5. Adult female, dorsal and ventral aspects (MPUJ-SCO-383). Scale bars = 5 mm. Tergites: with similar granulation to that of carapace; axial carina only evident on the posterior half of tergites II– VI ; obsolete dorsolateral carinae present on tergites II– VI ; tergite VII tetracarinate, with incomplete paired dorsolateral and lateral carinae, and low granulose median elevation on the position of the axial carina. Metasoma: with abundant setae; segments I–III with ten carinae (paired ventrosubmedian, ventrolateral, median lateral, dorsolateral and dorsosubmedian carinae); segment IV with six (ventrosubmedian and median lateral carinae absent); segment V with five (ventromedian, paired ventrolateral and dorsolateral carinae); ventrolateral and median lateral carinae converge distally in segments I–II; all carinae serrulate; intercarinal spaces with abundant granulation. Telson: vesicle elongated, smooth dorsally, with three ventral longitudinal carinae made up of conspicuous granulation; subaculear tubercle strong and spine-like; aculeus long and curved. Pedipalps: femur with five vestigial longitudinal carinae (dorsoexternal, dorsointernal, ventroexternal, ventrointernal and internal median carinae); patella without distinct carinae but with few small granules on the position of the dorsointernal and ventrointernal carinae; chela acarinate; fixed finger with six almost linear rows of granules, being the basal the longest; movable finger with seven rows (including the short apical row). Trichobothriotaxy type A, femur with β configuration ( Vachon 1974 , 1975 ). Legs: tibia, basitarsus and telotarsus with numerous ventral setae; tibial spur present on legs III–IV; prolateral pedal spur single on legs I–II, bifid on legs III–IV; retrolateral pedal spur present on all legs. Variability. Fixed finger trichobothria (n = 62): 37 (59.7%) fingers with trichobothria in the order eb : esb : est / db : et : dt ; 17 (27.4%) in the order eb : esb : est : db : et : dt , with est very slightly basal to db ; seven (11.3%) in the order eb : esb : db : est : et : dt , with db very slightly basal to est ; one (1.6%) lacking est (anomalous condition). Cheliceral reticulation (n = 62): incomplete reticular pattern present on all chelicerae studied, but slightly less reticulated in the holotype (3.2%), probably due to normal interpopulational variability. Pectinal teeth count : 16 to 18 on females (n = 20; mode = 18); 17 to 21 on males (n = 42; mode = 18). Coloration of the pedipalp hand (n = 62): 50 (80.6%) hands with brown areas ventrally, internally and externally; ten (16.1%) completely yellow; two (3.2%) with brown areas ventrally and internally. Total body length (including telson) : male 21–26 mm , female 27–33 mm . Anomalies. See ‘Fixed finger trichobothria’ in the ‘variability’ section above. Distribution. So far known from five localities in Meta Department ( Fig. 1 ). Probably widely distributed within this department’s limits and a typical element of open savanna-like formations. New locality records: COLOMBIA : Meta Department : i) Puerto López; ii) Villavicencio–Restrepo; iii) Serranía de La Macarena; iv) San Martín. Ecological notes. All specimens of A. arcadioi from ‘Cafam Llanos’ and ‘Con Esto Tengo’ were hand-captured at night, after detection with UV light when actively moving. In the former locality, most specimens were found in leaf litter or sandy walls of small streams inside gallery forests, whereas few were collected in borders of forests surrounded by extensive grasslands. In the latter locality, some specimens were found on the walls of a stream or the leaf litter close to it, but most were found on the border of a road in an area strongly influenced by human activities. Even though A. arcadioi was easily encountered in both places, it was noticeably less abundant than Tityus bastosi Lourenço, 1984 in ‘Cafam Llanos’ and Tityus nematochirus Mello-Leitão, 1940 in ‘Con Esto Tengo’, species with which it was found in sympatry. However, Ananteris specimens were very scarce in the places where Tityus Koch, 1836 was abundant.