Tityus bastosi Lourenço, 1984

Moreno-González, Jairo A., O, Ranulfo González & D, Eduardo Flórez, 2019, Taxonomic revision of the Colombian Tityus (Archaeotityus) (Scorpiones, Buthidae) species: a morphological and morphometric approach, with a description of a new species, Zootaxa 4660 (1), pp. 1-94 : 8-29

Tityus bastosi Lourenço, 1984

Figures 4 , 5A, B , 6A, D , 7A, B , 8A, B , 9A, B , 10A, B , 11 A–C, 12, 13, 14A–D, 15A, 16A, B, 17A, 18A, 19A, 20A, 29, 46A, 47A, 48, 49; Tables 1–3, 6–9.

Tityus bastosi Lourenço 1984: 358–359 , figures 17, 18; tables I–II.

References after Fet & Lowe (2000: 235). Tityus bastosi: Lourenço 2000: 456 , 458, figure 13; Flórez 2001a: 56, 57; Flórez 2001b: 28; Lourenço 2002a: 132; Lourenço 2002b: 115, 116, 117, 126, figure 257, 258, 259, 260, 261, 262, 263; Lourenço 2005: 224, 225, 226, figure 7; Lourenço 2006: 60, figure 17; Lourenço et al. 2006: 118, 119; Botero-Trujillo & Fagua 2006: 130; Botero-Trujillo 2007a: 67; Botero-Trujillo 2007b: 32; Teruel & García 2008: 9, 12, Botero-Trujillo & Flórez 2011: 9, 26; Botero-Trujillo & Noriega 2011: 50; Brito & Borges 2015: 2, 7, 14, figure 4; Lourenço & Ythier 2013: figure 18; Ojanguren-Affilastro et al. 2017: 40, 41, 42, 43, figure 1.

Type material. Tityus bastosi : Holotype: ECUADOR: Morona Santiago province: one adult male, Los Tayos, 30.vii.1976 ( NHMUK LTE-976 ) (not examined) ; Paratypes: ECUADOR: Morona Santiago province: adult female, Los Tayos, 27.vii.1976 ( NHMUK LTE-664 ) (not examined) . BRAZIL: Amazonas state: one adult female, São Paulo de Olivença, de Mathau ( MNHNH-RS-8499 ) (not examined) ; one juvenile male, Tefé, de Mathau ( MNHNH-RS- 8499 ) (not examined) . PERU: Iquitos province: one adult female and one juvenile male, de Mathau ( MNHNH-RS- 8499 ) (not examined) . Pebas province: one adult female, de Mathau ( MNHNH-RS-8499 ) (not examined) . Tarapoto province: one adult male, de Mathau ( MNHNH-RS-8499 ) (not examined) .

Examined material. COLOMBIA: Boyacá department: one adult female, Santa María, Vereda Caño Negro , Finca San Joaquín , 4°51’27.58’’ N 73°16’00.77’’ W, 1100 masl, in forest leaf litter with 11 juveniles on its back, 5.x.2003, D. M. Valencia ( ICN-As-521 ) ; one adult female, Santa María, 4°51’28.71’’ N 73°16’00.92’’ W, 860 masl, night manual capture with UV light, 15.v.2009, E. Flórez & D. Luna ( ICN-As-792 ) . Caqueta department: one adult female, one adult male and one juvenile female, Solano, Río Mesay , 0°14’24’’ N 72°56’2’’ W, 350 masl, collected in a floodplain forest inside a dead tree, 20.i.2000, F. Quevedo ( ICN-As-362 ) . Casanare department: one adult male, Yopal , 5°20’45.23’’ N 72°24’19.60’’ W, 370 masl, found inside a house, 5.iv.2009, C. Rincón & H. Rojas ( ICN- As-805 ) . Meta department: one adult male, Puerto López, Hacienda Moneque , 4°5’51.51’’ N 72°57’19.24’’ W, 181 masl, pitfall, 2.xi.2008, A. Neira ( ICN-As-741 ) ; one adult male, Puerto López, Inspección Remolinos , Hacienda CAFAM Llanos , 4°5’51.51’’ N 72°57’19.24’’ W, 270 masl, Riparian forest near Mata-Mata pool , pitfall tramp, iii.2004, A. Sánchez ( ICN-As-603 ) ; two adult females, San Martín, Finca Caduceo , near Camoa river, 3°41’39.46’’ N 73°41’37.16’’ W, 420 masl, 19.v.2006, C. Sarmiento ( ICN-As-740 ) ; two adult females, two adult males and one juvenile male, San Martín, Finca Caduceo , near the river Camoa , between palms, 3°41’39.46’’ N 73°41’37.16’’ W, 420 masl, 19.v.2006, R. Sarmiento ( ICN-As-738 ) ; five adult females and one adult male, Villavicencio, Vereda El Carmen , 4°9’38.73’’ N 73°41’5.66’’ W, 1263 masl, night manual capture with UV ligth, 7-10.viii.2016, J. A. Moreno & N. Herreño ( MZUSP 71740 ) .

Diagnosis. This species can be readily recognized from other Tityus (Archaeotityus) species by the following combination of character states: Total length: 33.12–40.33 (males) and 35.42–41.17 mm (females). General body coloration ( Figure 4 ) dark yellow background densely covered with dark reddish-brown variegated spots and metasomal segment V completely black ( Figure 6A, D ). Metasomal segments II–IV ( Figures 15A , 16A, B ) each has a distoterminal medium-sized acute granule on the DL carinae. Metasomal segment V with the VSM2 macrosetae slightly distal or at the same level of the VL2 macrosetae ( Figures 17A , 18A ). Subaculear tubercle large-sized pyramidal, flat, and crest-like with a blunt apex in lateral view ( Figures 19A , 20A ). Dorsal margin of the subaculear tubercle with a pair of ventrodistal small rounded granules with apices pointing to the basal portion of the aculeus. Vesicle and subaculear tubercle with VM carina composed of fine granules and feebly crenulate in lateral view ( Figures 19A , 20A ). Telson with a well-marked (female) ( Figure 19A ) or moderately marked (male) carinae ( Figure 20A ). Metasomal intercarinal areas densely covered with fine granules ( Figures 15A , 16A, B ). Pectinal tooth counts and movable finger oblique rows of denticles: 13–16/ 12–14 (females) and 15–17/ 13–15 (males) ( Table 2, 3). Metasomal segments II–V ( Figures 15A , 16A, B ), female pedipalp ( Figure 11 A–C), and male pedipalp femur and patella ( Figure 10A, B ) short and of a medium-width. Chela ( Figure 12 A–C) and metasomal segment V ( Figure 16A ) incrassate in large males.

Affinities with other Colombian species. Tityus bastosi is most similar to T. tayrona , sharing with it the presence of a subaculear tubercle that is large-sized pyramidal, flat, and crest-like with a blunt apex in lateral view ( Figures 19A , 20A ) and metasomal intercarinal areas densely covered with fine granules ( Figures 15A , 16A, B ). Tityus bastosi can be readily distinguished by the presence of a general body coloration that is dark with a yellow background densely covered with dark reddish-brown spots ( Figure 4 ); metasomal segment V completely black ( Figure 6A, D ) with the VSM2 macrosetae slightly distal or at the same level of the VL2 macrosetae ( Figures 17A , 18A ) and metasomal segments II–IV each with a distoterminal medium-sized acute granule on the DL carinae ( Figures 15A , 16A, B ).

Conversely, in T. tayrona the general body coloration is dark yellow in the background and moderately covered with dark brown spots ( Figure 41 ); the metasomal segment V has a dark yellow background moderately covered with brown variegated spots, a VL pair of dark brown spots, ventral yellow spots and a dorsomedian area without spots ( Figure 33B, D ). In addition, T. tayrona has the VSM2 macrosetae proximal to the level of the VL2 macrosetae on the metasomal segment V ( Figures 17E , 18E ) and the metasomal segments II–IV do not have conspicuous distoterminal granules on the DL carinae ( Figures 39B , 40C, D ).

Remarks. Lourenço (1997) recorded T. bastosi from Socaina, Casanare department, but there is no locality with this name. Despite this, the presence of this species was confirmed in Casanare department (see examined material). Also, Flórez (2001a) recorded T. bastosi from Bochalema and Toledo, Norte de Santander department, and Chalará, Santander department; however, a detailed examination of these materials revealed these populations possibly belong to another species.

Redescription. Based on an adult female (ICN-As-740, #1) and an adult male (ICN-As-741). For female live habitus see Figure 46A .

Total length. Female: 38.01 mm; male: 40.34 mm.

Coloration. General body coloration (in ethanol 70%) ( Figure 4 ) with a dark yellow background densely covered with dark reddish-brown variegated spots. Carapace ( Figure 5A, B ) densely covered with dark reddish-brown variegated spots, except for the anterior margin that is dark yellow; lateral and median eyes surrounded by black spots; median ocular tubercle surrounded by a lateral black stripe on each side; posterior area to the median ocular tubercle with an inverted ‘’Y’’-shaped spot. Chelicerae ( Figure 7A, B ) coxa and hand with a light yellow background; hand with dark brown reticulated spots on the anterior half and small areas without spots on the internal surface, median surface, and the adjacent area to the movable finger articulation; movable and fixed fingers with dark brown spots on their posterior halves; teeth dark reddish-brown. Pedipalps ( Figure 4 ) densely covered with dark reddish-brown variegated spots and yellow spots; trochanter, femur, and patella ventrally yellow; trichobothrial pits yellow. Legs with all the segments having a light yellow background moderately covered with dark brown variegated spots on their prolateral surfaces. Coxosternal region ( Figure 4B, E, F ) coxae I–IV, sternum, genital operculum, pectines (middle lamellae, fulcra, and teeth), and basal pectinal piece light yellow; basal pectinal piece with an anterior white area (female); coxapophyses I–II and pectines (marginal lamellae) with a light yellow background moderately covered with dark brown variegated spots. Mesosoma ( Figure 4 ) tergites I–VII and sternites III–VII each with a dark yellow background densely covered with dark reddish-brown variegated spots; tergites I–VII with three posterior dark yellow spots (two submedian and one median); spiracles yellow. Metasoma ( Figure 6A, D ) with a dark yellow background densely covered with dark reddish-brown variegated spots and with ventral yellow spots; each segment progressively darker towards segment V; segment V completely black ( Figure 6A, D ). DL intercarinal areas of segments I–IV each with an anterior median arrow-shaped brown spot and a posterior median brown spot; ML, VSM, and VM intercarinal areas of segments I–IV each with a pair of distal brown spots. Telson ( Figure 4 ) almost completely black with a lateral light reddish-brown stripe; subaculear tubercle and aculeus dark reddish-brown.

Morphology. Carapace ( Figure 8A, B ) densely covered with fine granules and a few coarse granules; anterior margin with a shallow median notch; carinae (anterior median, lateral, and posterior median) and furrows (anterior, lateral, median ocular, posterior median, posterior marginal, and posterior lateral) well-marked; median ocular tubercle well-marked and located on the anterior half of the carapace; median eyes separated by about 0.9 ocular diameters. Lateral eye pattern type 4A: three pairs of major ocelli present (PLMa, MLMa and ALMa ) and one pair of minor ocelli (ADMi).

Chelicerae ( Figure 7A, B ) dentition characteristic of the family Buthidae ( Vachon 1963) ; hand and fingers densely covered with setae on the internal and ventral surfaces.

Pedipalps. Chela short and of a medium-width (female) (Cl/Cw= 4.72) or incrassate (male) (Cl/Cw= 3.48). Trichobothriotaxic pattern Type A with alfa configuration (hand: Eb3:Eb2:Eb1:Esb:Est:Et; fixed finger: eb:esb:est: et:db:dt:it) (e.g., Figure 3 ). Femur ( Figures 9A , 10A ) with five well-marked and complete carinae: VI, DI, and DE crenulate; VE and IM serratocrenulate; intercarinal areas densely covered with fine granules. Patella ( Figures 9B , 10B ) with seven carinae: VI, VE, DI, DE, and EM complete and crenulate; DM incomplete and crenulate; IM complete and serratocrenulate with a spur near the segment base; intercarinal areas densely covered with fine granules. Chela (Tibia) ( Figures 11 A–C, 12A–F) with eight carinae: VI, VE, D, DS, DMA, and ES complete and crenulate; DI complete and serratocrenulate (female) ( Figure 11 A–C) or feebly crenulate (male) ( Figures 12 A–F); SA incomplete and crenulate, only present on the anterior half of the hand. Pedipalp movable and fixed fingers with an obsolete basal lobe (male) ( Figure 12A ) or without a basal lobe (female) ( Figure 11A ); dorsal surface of the movable finger with 13–14 (female) and 13–13 (male) oblique rows of denticles.

Coxosternal region. Covered with fine granules and a few setae, except for the coxapophyses I–II which are smooth. Sternum with posterior depression, outer ridge, and apical button well-marked. Genital operculum longitudinally divided and composed of two subtriangular plates.

Pectines. Basal piece sexually dimorphic (e.g., Figure 2 ) sub rectangular-shaped with an anterior elevated glandular area (female) ( Figure 2A ) or flat, shield-shaped with an anterior median notch well-marked (male) (e.g., Figure 2B ); pectinal tooth counts: 15–16 (female) and 17–17 (male). Intermediate plate, marginal plate, and fulcra moderately covered with setae ( Figure 14A, C ). Pectinal teeth peg sensillae flattened with a narrow apex ( Figure 14B, D ).

Legs. Carinae present; intercarinal areas with a few fine granules; telotarsus covered with numerous ventrosubmedian setae; claws short and symmetrical.

Mesosoma . Tergites I–VI moderately covered with fine granules and a few coarse granules; pre-tergite wellmarked with the median carina visible on the posterior margin of the post-tergite; tergite VII with DSM and DL carinae complete and crenulate; median carina of the tergite VII composed of an anterior median crenulate eminence located on the anterior half of the post-tergite. Sternites densely covered with fine granules; sternites III–VI each with a longitudinal median hyaline suture and a pair of elliptic spiracles on the posterior half; spiracles progressively larger towards sternite VI; posterior margin of the sternite V with a subtriangular glandular; sternite VI with VSM carinae crenulate occupying the posterior half of the sternite; sternite VII with VSM carinae crenulate occupying the two posterior thirds of the sternite and VL carinae crenulate occupying the anterior half of the sternite.

Hemispermatophore. General morphological characteristics as in other Tityus (Archaeotityus) species. Capsule ( Figure 13 A–C) with a straight distal carina ending in an 80° angle; sperm duct reduced not overpassing the basal carina level and with a translucent area between the base of the basal hook and the base of the basal carina; translucent area basally wide but progressively narrower towards the distal region ( Figure 13 B–C); basal carina with rounded apex ( Figure 13C ); basal hook semi ovate-shaped with anterior margin straight in lateral view ( Figure 13B ) and slightly curved in anterior and posterior views; basal hook forming a ‘’U’’-shaped curve with the stem ( Figure 13A, C ); basal hook slightly elongated with a rounded apex and approximately as long as its basal width in anterior view ( Figure 13A ).

Metasoma. Segments II–V short and of a medium-width in both sexes (Msl/Msw ratios= female/ male: II= 1.4/ 1.3; III= 1.6/ 1.5; IV= 1.76/ 1.68; V= 2.0/ 1.9). Segment V short and of a medium-width (female) ( Figure 15A ) or incrassate (male) ( Figure 16A ). Segments I–II ( Figures 15A , 16A, B ) with 10 complete, parallel, and crenulate carinae (paired DL, ML, LIM, VL, and VSM; DL carinae serratocrenulate on segment II); LIM on segment II composed of fine granules on the two posterior thirds and coarse granules on the first anterior third; intercarinal areas densely covered with fine granules. Segments III–IV ( Figures 15A , 16A, B ) with eight complete, parallel, and crenulate carinae (paired DL, ML, VL and VSM; DL carinae serratocrenulate); intercarinal areas densely covered with fine granules. Segment V with five complete and crenulate carinae (VM, paired DL and VL); lateral intercarinal areas densely covered only with fine granules (males) ( Figure 16A, B ) or densely covered with fine granules and moderately covered with coarse granules (female) ( Figure 15A ); ventral intercarinal areas with a few and feebly fine granules (male) ( Figure 16A, B ) or densely covered with fine granules (female) ( Figure 15A ); intercarinal areas with granules irregularly distributed on the middle of each area. Segment II–IV with DL carinae composed of granules which slightly increase in size towards the distal region of each segment; DL carinae ending in a distoterminal medium-sized coarse, and acute granule which increases in size towards the segment IV ( Figures 15A , 16A, B ).

Metasomal macrosetation. Segments I–IV each with two pairs of VSM macrosetae (2/2) and two pairs of VL macrosetae (2/2); pair VSM1 is located on the anterior third, and pair VSM2 is located near the posterior margin of the segment; pair VL1 is located near the anterior margin of the segment, and pair VL2 is located on the posterior third of the segment (e.g., Figure 1 ). Segment V ( Figures 17A , 18A ) with three pairs of VSM macrosetae (3/3), three pairs of VL macrosetae (3/3), and a single pair of DL macrosetae; pair VSM1 is located near the anterior margin of the segment; pair VSM2 is located slightly distal or at same level of the pair VL2 ( Figures 17A , 18A ), and pair VSM3 is located on the anal arch; pair VL1 is located near the anterior margin of the segment; pair VL2 is located on the posterior second third of the segment, and pair VL3 is located on the anal arch; pair DL1 is located near the posterior margin of the segment.

Telson . Vesicle not elongated and suboval (Vl/Vh= 2.2) ( Figures 19A , 20A ) with dorsal surface smooth and with a lateral longitudinal furrow on each side; VM, paired VSM, VL, and DL carinae composed of coarse granules; carinae well-marked (female) ( Figure 19 A ) or moderately marked (male) ( Figure 20A ); paired DL carinae located dorsally with respect to the lateral longitudinal furrow; VM carina basally marked and distally feebly marked ( Figures 19A , 20A ). Subaculear tubercle large-sized pyramidal, flat, and crest-like with a blunt apex in lateral view ( Figures 19A , 20A ); dorsal margin of the subaculear tubercle with a pair of ventrodistal small rounded granules with apices pointing to the basal portion of the aculeus; VM carina composed of fine granules and feebly crenulate in lateral view ( Figures 19A , 20A ); aculeus strongly curved shorter than vesicle and with a ventral groove.

Variability. Total length (including telson ): males 33.12–40.33 mm (n= 7; mean= 36.39; SD= 3.23; females 35.42–41.17 mm (n= 11; mean= 38.50; SD= 1.56) ( Table 9 ). Pectinal tooth counts: males 15–17 (n= 21; mode= 15 13–16 (n= 21; mode= 14) ( Table 2). Number of movable finger oblique rows of denticles: males 13–14 (n= 13; mode= 13); females 12–14 (n= 22; mode= 14) ( Table 3). Metasomal macrosetae counts: segments I–IV: males (n= 8) and females (n= 11): 2/2 VSM and 2/2 VL macrosetae on each segment; segment V: males (n= 8) and females (n= 11): 3/3 VSM; 3/3 VL macrosetae, and 1/1 DL macrosetae on the segment. Relative position of the VSM2 macrosetae on the metasomal segment V: males (n= 8): distal to VL2 (n= 1) or at the same level of VL2 (n= 7); females (n= 11): distal to VL2 (n= 2); at the same level of VL2 (n= 8) or asymmetric with one seta at the same level of VL2 (n= 1).

Hemispermatophore capsule: the basal hook of the capsule exhibited different shapes in the anterior view: (1) slightly elongated with a rounded apex, curved lateral margin, and forming a ‘’U’’-shaped curve with the stem ( Figure 13C ); (2) elongated with a rounded apex, straight lateral margin, and forming a narrow ‘’U’’-shaped curve with the stem ( Figure 13D ); and (3) elongated with a rounded apex, strongly curved lateral margin, and forming a ‘’U’’-shaped curve with the stem ( Figure 13E ).

Male intrasexual variations: adult males exhibited different body sizes ranging between 33.12–40.33 mm, but despite this they did not show discrete size-classes but rather a progressive increase in their total lengths ( Figure 4 C–F, Table 9 ). However, as the total body size increases among males the following: (1) Carapace and tergites: the number of coarse granules on intercarinal surfaces increases; (2) Chela : the manus is more incrassate without apparently exhibiting isometric growth (n= 6; Cl/Cw= 3.4–4.4; Cl/Ch= 3.2–4.0) ( Figure 12 A–F); (3) Metasomal segments II–IV: the DL carinae exhibit coarser distoterminal granules ( Figure 16A, B ); and (4) Metasomal segment V: segment is more incrassate without apparently exhibiting isometric growth (n= 6; Msl-V/Msw-V= 1.8–2.1; Msl- V/Msh-V= 1.7–2.1) ( Figure 16A, B ), and the granules associated with carinae and intercarinal areas are less marked ( Figure 16A, B ).

Distribution ( Figure 29 ). BRAZIL: Amazonas state: São Paulo de Olivença and Tefé. COLOMBIA: Amazonas department: La Pedrera. Boyacá department: Otanche, Puerto Boyacá, and Santa María. Caquetá department: La Montañita and Solano (National Natural Park Chiribiquete). Casanare department: Sácama. Cundinamarca department: Guayabetal. Guaviare department: El Retorno (Natural reserve Nukak Maku). Meta department: Cubarral, Fuente de Oro, Puerto Gaitán, Puerto López, Restrepo, San Juan de Arama, Serranía de la Macarena, and Villavicencio. Putumayo department: Orito. ECUADOR: Morona Santiago province: Los Tayos. Napo province: Estación Biológica Jatun Sacha. Sucumbíos province: San Pablo de Kantesiya. PERU: Loreto province: Iquitos and Pebas. San Martín province: Tarapoto. New records. COLOMBIA: Casanare department: Yopal. Meta department: San Martín and Villavicencio.

Figures

FIGURE 1. Tityus (Archaeotityus) Lourenço, 2006, metasomal segments I–V showing distribution of macrosetae.A. Segments I–V, ventral aspect. B. Segment V, lateral aspect. Abbreviations: ML, median lateral; VL, ventrolateral; VSM, ventrosubmedian.

FIGURE 2. Tityus (Archaeotityus) guane sp. nov., sternum and pectines, showing the shape of the basal pectinal piece. A. Female paratype (ICN-As-1003). B. Male holotype (ICN-As-1003). Abbreviations: A, anterior region; P, posterior region.

FIGURE 3. Tityus (Archaeotityus) Lourenço, 2006, right pedipalp, showing trichobothrial pattern. A–C. Pedipalp chela. A. External aspect. B. Dorsal aspect. C. Ventral aspect. D. Femur, dorsal aspect. E. Patella, dorsal aspect. Abbreviations: d, dorsal; db, dorsal basal; dt, dorsal terminal; e, external; eb, Eb, external basal; esb, Esb, external subbasal; est, Est, external subterminal; et, Et, external terminal; i, internal; it, internal terminal; v, ventral. Scale bars= A–C= 2.0 mm; D, E= 1.0 mm.

FIGURE 4. Tityus (Archaeotityus) bastosi Lourenço, 1984, habitus. A, B. Female (ICN-As-740). A. Dorsal aspect. B. Ventral aspect. C, E. Male (San Martín, Meta) (ICN-As-738). C. Dorsal aspect. E. Ventral aspect. D, F. Male (Puerto López, Meta) (ICN-As-741). D. Dorsal aspect. F. Ventral aspect. Scale bars= 10 mm.

FIGURE 5. Tityus (Archaeotityus) Lourenço, 2006, carapace coloration, dorsal aspect. A, B. Tityus bastosi Lourenço, 1984 (ICN-As-740, 741). A. Female. B. Male. C, D. Tityus columbianus (Thorell, 1876) (ICN-As-739, 524). C. Female. D. Male. E, F. Tityus guane sp. nov. (ICN-As-1003). E. Female paratype. F. Male holotype.

FIGURE 6. Tityus (Archaeotityus) Lourenço, 2006, metasoma coloration pattern, dorsal (left) and ventral (right). A, D. Tityus bastosi Lourenço, 1984 (ICN-As-740, 741). A. Female. D. Male. B, E. Tityus columbianus (Thorell, 1876) (ICN-As-739, 524). B. Female. E. Male. C, F. Tityus guane sp. nov. (ICN-As-1003). C. Female paratype. F. Male holotype.

FIGURE 7. Tityus (Archaeotityus) Lourenço, 2006, chelicerae, dorsal aspect, showing reticulation pattern. A, B. Tityus bastosi Lourenço, 1984 (ICN-As-740, 741). A. Female. B. Male. C, D. Tityus columbianus (Thorell, 1876) (ICN-As-739, 524). C. Female. D. Male. E, F. Tityus guane sp. nov. (ICN-As-1003). E. Female paratype. F. Male holotype.

FIGURE 8. Tityus (Archaeotityus) Lourenço, 2006, carapace sculpturing, dorsal aspect. A, B. Tityus bastosi Lourenço, 1984. A. Female (ICN-As-740). B. Male (ICN-As-741). C, D. Tityus columbianus (Thorell, 1876). C. Female (ICN-As-739). D. Male (ICN-As-524). E, F. Tityus guane sp. nov. (ICN-As-1003). E. Female paratype. F. Male holotype. Scale bars= 1.5 mm.

FIGURE 9. Tityus (Archaeotityus) Lourenço, 2006, female pedipalp femur and patella sculpturing, dorsal aspect. A, B. Tityus bastosi Lourenço, 1984 (ICN-As-740). A. Femur. B. Patella. C, D. Tityus columbianus (Thorell, 1876) (ICN-As-739). C. Femur. D. Patella. E, F. Tityus guane sp. nov. (ICN-As-1003). E. Femur. F. Patella. G, H. Tityus parvulus Kraepelin, 1914 (ICN- As-1004). G. Femur. H. Patella. I, J. Tityus tayrona Lourenço, 1991 (ICN-As-181). I. Femur. J. Patella. Scale bars= 1.0 mm.

FIGURE 10. Tityus (Archaeotityus) Lourenço, 2006, male pedipalp femur and patella sculpturing, dorsal aspect. A, B. Tityus bastosi Lourenço, 1984 (ICN-As-741). A. Femur. B. Patella. C, D. Tityus columbianus (Thorell, 1876) (ICN-As-524). C. Femur. D. Patella. E, F. Tityus guane sp. nov. (ICN-As-1003). E. Femur. F. Patella. G, H. Tityus parvulus Kraepelin, 1914 (ICN-As- 1004). G. Femur. H. Patella. I, J. Tityus tayrona Lourenço, 1991 (ICN-As-181). I. Femur. J. Patella. Scale bars= 1.0 mm.

FIGURE 11. Tityus (Archaeotityus) Lourenço, 2006, right female pedipalp sculpturing. A–C. Tityus bastosi Lourenço, 1984 (ICN-As-740). A. External aspect. B. Dorsal aspect. C. Ventral aspect. D–F. Tityus columbianus (Thorell, 1876) (ICN-As-739). D. External aspect. E. Dorsal aspect. F. Ventral aspect. Scale bars= 2.0 mm.

FIGURE 12. Tityus (Archaeotityus) bastosi Lourenço, 1984, male pedipalp chela. A–C. Male (Puerto López, Meta) (ICN-As- 741). A. External aspect. B. Dorsal aspect. C. Ventral aspect. D–F. Male (San Martín, Meta) (ICN-As-738). D. External aspect. E. Dorsal aspect. F. Ventral aspect. Scale bars= 2.0 mm.

FIGURE 13. Tityus (Archaeotityus) bastosi Lourenço, 1984, dextral hemispermatophore, showing capsule details, male (ICN- As-741). A. Posterior view. B. Lateral view. C. Anterior view. D, E. Capsule variations, anterior view. D. Male (San Martín, Meta) (ICN-As-738). E. Male (Villavicencio, Meta) (MZUSP 71740). Abbreviations: Bc, basal carina; Bh, basal hook; Dc, distal carina; Sdm, sperm duct membrane. Scale bar= 1.0 mm.

FIGURE 14. Tityus (Archaeotityus) Lourenço, 2006, left pectine with peg sensillae details, ventral aspect. A–D. Tityus bastosi Lourenço, 1984.A, B. Female (MZSP 71740). A. Pectine. B. Peg sensillae shape. C, D. Male (MZSP 71740). C. Pectine. D. Peg sensillae shape. E, F. Tityus columbianus (Thorell, 1876) (ICN-As-739). E. Female pectine. F. Female peg sensillae shape. G–J. Tityus guane sp. nov.. G, H. Female (MZSP 71736). G. Pectine. H. Peg sensillae shape. I, J. Male (MZSP 71736). I. Pectine. J. Peg sensillae shape. Scale bars= A, C, E, G, I= 400 µm; B, D, F, H, J= 10 µm.

FIGURE 15. Tityus (Archaeotityus) Lourenço, 2006, female metasomal segments I–V sculpturing, lateral aspect. A. Tityus bastosi Lourenço, 1984 (ICN-As-740). B. Tityus columbianus (Thorell, 1876) (ICN-As-739). C. Tityus guane sp. nov. (paratype) (ICN-As-1003). Scale bars= 2.0 mm.

FIGURE 16. Tityus (Archaeotityus) Lourenço, 2006, male metasomal segments I–V sculpturing, lateral aspect. A, B. Tityus bastosi Lourenço, 1984. A. Male (Puerto López, Meta) (ICN-As-741). B. Male (San Martín, Meta) (ICN-As-738). C, D. Tityus columbianus (Thorell, 1876). C. Male (Villa de Leyva, Boyacá) (ICN-As-524). D. Male (Villa de Leyva, Boyacá) (ICN-As- 739). Scale bars= 3.0 mm.

FIGURE 17. Tityus (Archaeotityus) Lourenço, 2006, female metasomal segment V sculpturing, showing macrosetae, ventral aspect. A. Tityus bastosi Lourenço, 1984 (ICN-As-740). B. Tityus columbianus (Thorell, 1876) (ICN-As-739). C. Tityus guane sp. nov. (paratype) (ICN-As-1003). D. Tityus parvulus Kraepelin, 1914 (ICN-As-1004). E. Tityus tayrona Lourenço, 1991 (ICN-As-181). Abbreviations: ML, median lateral; VL, ventrolateral; VSM, ventrosubmedian. Scale bars= 1.0 mm.

FIGURE 18. Tityus (Archaeotityus) Lourenço, 2006, male metasomal segment V sculpturing, showing macrosetae, ventral aspect. A. Tityus bastosi Lourenço, 1984 (ICN-As-741). B. Tityus columbianus (Thorell, 1876) (ICN-As-524). C. Tityus guane (holotype) (ICN-As-1003). D. Tityus parvulus Kraepelin, 1914 (ICN-As-1004). E. Tityus tayrona Lourenço, 1991 (ICN-As- 181). Abbreviations: ML, median lateral; VL, ventrolateral; VSM, ventrosubmedian. Scale bars= 1.5 mm.

FIGURE 19. Tityus (Archaeotityus) Lourenço, 2006, female telson sculpturing, lateral aspect. A. Tityus bastosi Lourenço, 1984 (ICN-As-740). B. Tityus columbianus (Thorell, 1876) (ICN-As-739). C. Tityus guane sp. nov. (paratype) (ICN-As-1003). D. Tityus parvulus Kraepelin, 1914 (ICN-As-1004). E. Tityus tayrona Lourenço, 1991 (ICN-As-181). Scale bars= 1.0 mm.

FIGURE 20. Tityus (Archaeotityus) Lourenço, 2006, male telson sculpturing, lateral aspect. A. Tityus bastosi Lourenço, 1984 (ICN-As-741). B. Tityus columbianus (Thorell, 1876) (ICN-As-524). C. Tityus guane sp. nov. (holotype) (ICN-As-1003). D. Tityus parvulus Kraepelin, 1914 (ICN-As-1004). E. Tityus tayrona Lourenço, 1991 (ICN-As-181). Scale bars= 1.5 mm.

FIGURE 29. Distribution of Tityus (Archaeotityus) bastosi Lourenço, 1984, showing known records and new locality records.

FIGURE 33. Tityus (Archaeotityus) Lourenço, 2006, metasoma coloration pattern, dorsal (left) and ventral (right). A, C. Tityus parvulus Kraepelin, 1914 (ICN-As-1004). A. Female. C. Male. B, D. Tityus tayrona Lourenço, 1991 (ICN-As-181). B. Female. D. Male.

FIGURE 39. Tityus (Archaeotityus) Lourenço, 2006, female metasomal segments I–V sculpturing, lateral aspect. A. Tityus parvulus Kraepelin, 1914 (ICN-As-104). B. Tityus tayrona Lourenço, 1991 (ICN-As-181). Scale bar= 3.0 mm.

FIGURE 40. Tityus (Archaeotityus) Lourenço, 2006, male metasomal segments I–V sculpturing, lateral aspect. A, B. Tityus parvulus Kraepelin, 1914. A. Male (Chinchiná, Caldas) (ICN-As-1004). B. Male (Chinchiná, Caldas) (ICN-As-1004). C, D. Tityus tayrona Lourenço, 1991. C. Male (Zambrano, Bolívar) (ICN-As-181). D. Male (Zambrano, Bolívar) (ICN-As-181). Scale bars= 3.0 mm.

FIGURE 41. Tityus (Archaeotityus) tayrona Lourenço, 1991, habitus. A, B. Female (ICN-As-181). A. Dorsal aspect. B. Ventral aspect. C, E. Male (Zambrano, Bolívar) (ICN-As-181). C. Dorsal aspect. E. Ventral aspect. D, F. Male (Zambrano, Bolívar). D. Dorsal aspect. F. Ventral aspect. Scale bars= 10 mm.

FIGURE 46. Tityus (Archaeotityus) Lourenço, 2006, habitus in life. A. Tityus bastosi Lourenço, 1984, female from El Carmen, Villavicencio, Meta, Colombia. B. Tityus columbianus (Thorell, 1876), female from Soacha, Cundinamarca, Colombia. C. Tityus guane sp. nov., male from Vereda Pasochico, Finca La Ceiba, Los Santos, Santander. D, E. Tityus parvulus Kraepelin, 1914, females. D. Female from Regional Natural Reserve El Vínculo [Natural Regional Park El Vínculo], Buga, Valle del Cauca, Colombia. E. Female from the Botanical Garden Maria Cespedes [Jardín Botánico Maria Cespedes], Tulúa, Valle del Cauca, Colombia. F. Tityus tayrona Lourenço, 1991, female from National Natural Park Tayrona [Parque Nacional Natural Tayrona], Santa Marta, Magdalena, Colombia. Photo B, courtesy of N. Herreño.