Tityus columbianus ( Thorell, 1876 )

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 : 29-38

Tityus columbianus ( Thorell, 1876)

Figures 5C, D , 6B, E , 7C, D , 8C, D , 9C, D , 10C, D , 11 D–F, 14E, F, 15B, 16C, D, 17B, 18B, 19B, 20B, 21–23, 30, 46B, 47B, C, 48, 49; Tables 1–3, 6–9.

Phassus columbianus Thorell, 1876: 8 .

References after Fet & Lowe (2000: 239). Tityus columbianus: Lourenço 2000: 456 , 458, figure 13; Flórez 2001a: 58, 59; Flórez 2001b: 28; Lourenço 2002a: 133; Lourenço 2006: 56, 58, 60; Botero-Trujillo & Fagua 2006: 130; Kovarik 2007: 3; Lourenço 2008: 197, 199; Teruel & García 2008: 1, 2–10, figures 1–6, table 1–4; Teruel & Kovarik 2011: 7; Francke et al. 2014: 232.

Tityus erikae (misidentification): Lourenço 2000: 456, figure 13.

Type material. Tityus columbianus : Holotype: COLOMBIA: Cundinamarca department: one adult female, Bogotá (‘’ America merid., Columbia , Bogotá?’’) ( NMG-Scorp-42 ) ( Not examined).

Examined material. COLOMBIA: Bolívar department: one adult male, Zambrano, Santuario de Flora y Fauna Los Colorados , 9°57’24’’ N 75°5’12’’ W, 390 masl, collected inside a dry forest, vii.1996, F. Escobar ( ICN-As- 188 ) (misidentified as T. erikae by W. R. Lourenço, xii.1998) . Boyacá department: three adult females, eight juvenile females and one juvenile male, Arcabuco, Vereda Montesuárez , 5°45’0’’ N 73°26’0’’ W, 2500 masl, under rock near an oak forest, 1–3.viii.2000, A. Sabogal ( ICN-As-364 ) ; one adult male, Chita, Vereda Chipa , 6°11’22’’ N 72°28’36’’ W, 2700 masl, under rock inside a crop, iv.1991, L. G. Baptiste ( ICN-As-130 ) ; three adult females and one adult male, Firavitoba, Vereda El Alcaparral , 6°11’22’’ N 72°28’36’’ W, 2500 masl, 7.i.2000, R. Alvarado ( ICN-As-288 ) ; two adult females, five juvenile males and three juvenile females, Iza , 5°36’48’’ N 72°58’51.93’’ W, 2590 masl, 10.vii.2000, E. Flórez ( ICN-As-388 ) ; two adult females, Susacón, Vereda San Ignacio , 6°13’58.2’’ N 72°42’0.23’’ W, 2315 masl, inside a rotten log, 21.v.2010, H. Gasca ( ICN-As-846 ) ; three adult females and one adult male, Villa de Leyva , 5°38’0’’ N 73°31’59.99’’ W, 2128 masl, 25.viii.2002, E. González ( ICN-As-739 ) ; two adult females, two adult males and one juvenile female, Villa de Leyva, Santuario de Flora y Fauna de Iguaque , Sector de Chaina , 5°41’0’’ N 73°29’0’’ W, 2600 masl, under rocks and logs inside a paddock, 9.vi.1991, F. Quevedo ( ICN- As-524 ) ; 24 adult females and eight adult males, Villa de Leyva , El Roble , 5°38’19’’ N 73°31’42’’ W, 2200 masl, x.1985, R. González ( ICN-As-352 ) ; one adult female, Villa de Leyva , near river Cebada and river Cene , 5°38’19’’ N 73°31’42’’ W, 2200 masl, collected inside a pastureland, iv.2000, E. Hernández ( ICN-As-314 ) ; one adult male, Villa de Leyva , 5°38’1.9’’ N 73°31’24.67’’ W, 2200 masl, inside a house, 1.viii.2000, H. Schultz ( ICN-As-419 ) ; one juvenile male, Villa de Leyva , 5°38’0’’ N 73°31’59.99’’ W, 2200 masl, manual capture inside leaf litter, 19.ii.2004, M. Porras ( ICN-As-570 ) ; one adult female, Chiquinquirá, Monte Batallón Sucre , 5°37’0.81’’ N 73°48’59.47’’ W, 2580 masl, 24.v.1979, C. Hernández ( ICN-As-129 ) ; one adult male, Tinjacá , 5°34’0.13’’ N 73°38’59.89’’ W, 2100 masl, 20.viii.2010, J. Iglesias ( ICN-As-851 ) . Cundinamarca department: three adult females and one juvenile female, Bogotá, Suba , 4°45’13.13’’ N 74°5’19.36’’ W, 2600 masl, 7.v.1961, J. M. Idroba ( ICN-As-088 ) ; one adult female, Bogotá, Parque de San Rafael , Cerros Orientales , 4°37’5.9’’ N 74°3’20.36’’ W, 2850 masl, under a rock, ix.2000, D. Tobar ( ICN-As-432 ) ; two adult females, Bogotá, rural zone Usme , 4°28’23.05’’ N 74°6’57.93’’ W, 2600 masl, under rocks, 27.iv.2001, M. Ramírez ( ICN-As-409 ) ; one adult female, Bogotá, Cerro Juan Rey , near Barrio La Fiscala in Usme , 4°28’23.05’’ N 74°6’57.93’’ W, 2600 masl, night manual capture, 4.v.2003, T. Vivas ( ICN-As- 525 ) ; four adult females and one adult male, Carmen de Carupa, Vereda San José , 5°21’01.01’’ N 73°54’19.06’’ W, 2971 masl, diurnal manual capture, under rocks, 25.vi.2011, J. Cabra ( ICN-As-995 ) ; six adult females, Carmen de Carupa, Vereda San José , 5°21’01.01’’ N 73°54’19.06’’ W, 2971 masl, diurnal manual capture, under rocks, iv.2010, J. Cabra ( ICN-As-996 ) ; six adult females, Facatativá, highway from El Rosal-Faca , 4°48’53.95’’ N 74°21’23.31’’ W, 2700 masl, collected on a hill with bushes, 24.viii.2003, F. Fernández ( ICN-As-512 ) ; one adult female, Guasca , 4°51’56.87’’ N 73°52’59.78’’ W, 2600 masl, without any other data available ( ICN-As-134 ) ; one adult male, Medina, Vereda Miralindo , 4°36’33.54’’ N 73°25’18.65’’ W, 1500 masl, manual capture, vi.1997, F. Escobar ( ICN- As-194 ) (misidentified as T. erikae by W. R. Lourenço, xii.1998) ; two adult females and 11 juvenile females, Mosquera, Mondoñedo , Zabrinsky desert, 4°40’4.96’’ N 74°15’45.95’’ W, 2600 masl, pitfall, ii-iv.2002, C. Niño, G. Mora & J. Martínez ( ICN-As-499 ) ; one adult female, Mosquera, Km 7- road to Cota , 4°35’56’’ N 74°4’51’’ W, 2600 masl, 19.ii.1988, P. Ruiz ( ICN-As-016 ) ; one adult female, Nariño , 4°24’2’’ N 74°50’9’’ W, 263 masl, inside a house, 1.xi.1998, R. Jaramillo ( ICN-As-347 ) ; five adult females, Soacha, Hacienda Terrenos , Hoya de la Quebrada , Tibanica , 4°32’59.66’’ N 74°10’59.83’’ W, 2600 masl, 13.v.1961, J. Hernández ( ICN-As-083 ) ; two adult females and one adult male, Susa, Finca Bermejales , Vereda La Fragua , 5°25’0’’ N 73°49’0’’ W, 2710 masl, 30.xii.2006, J. Pinzón & G. Alarcón ( ICN-As-727 ) ; two juvenile females, Tabio , 4°45’13.13’’ N 74°5’19.36’’ W, 2690 masl, 1.vii.2002, G. Stiles ( ICN-As-498 ) ; two adult females and one adult male, Ubaté , 5°18’42.86’’ N 73°49’5.56’’ W, 2600 masl, xii.2001, A. Uribe ( ICN-As-100 ) . Nariño department: seven adult females, two adult males and one juvenile female, Reserva Natural La Planada , 1°13’10.52’’ N 77°59’2.6’’ W, 1800 masl, vi.1992, A. Calixto ( ICN- As-712 ) . Tolima department: one adult female, Ibagué, Boquerón , Tigre Cave , 4°24’26.83’’ N 75°15’53.94’’ W, 1490 masl, 6.vii.1972, P. García ( ICN-As-131 ) ; two adult females, Ibagué, Botanical Garden San José , 4°26’26.36’’ N 75°14’38.91’’ W, 1200 masl, 20.v.2007, C. Samudio ( ICN-As-743 ) .

Diagnosis. This species can be readily recognized from other Tityus (Archaeotityus) species by the following combination of character states: Total length: 28.16–39.11 mm (males) and 28.16–39.11 mm (females). General body coloration ( Figure 21 ) dark yellow background moderately covered with dark brown variegated spots; metasomal segment V completely dark reddish-brown with three ventral black stripes and with a dorsomedian area without spots ( Figure 6B, E ). Metasomal segments II–III ( Figures 15B , 16C, D ) each has a distoterminal small acute granule on the DL carinae. Metasomal segment IV ( Figures 15B , 16C, D ) with a distoterminal medium-sized blunt, and coarse granule on the DL carinae. Metasomal segment V with the VSM2 macrosetae slightly distal or at the same level of the VL2 macrosetae ( Figures 17B , 18B ). Subaculear tubercle medium-sized pyramidal, flat, and triangle-like with an acute and coarse apex in lateral view ( Figures 19B , 20B ). Dorsal margin of the subaculear tubercle with a pair of ventrodistal small rounded granules with apices pointing to the middle portion of the aculeus. Vesicle and subaculear tubercle with VM carina composed of coarse granules and strongly crenulate in lateral view ( Figures 19B , 20B ). Telson with well-marked (female) ( Figure 19B ) or moderately marked (male) carinae ( Figure 20B ). Metasomal intercarinal areas densely covered with coarse granules ( Figures 15B , 16C, D ). Pectinal tooth counts and movable finger oblique rows of denticles: 11–15/ 11–13 (females) and 13–15/ 12–14 (males) ( Table 2, 3). Metasomal segments II–V ( Figures 15B , 16C, D ), female pedipalp ( Figure 11 D–F), and male pedipalp femur and patella ( Figure 10C, D ) short and of a medium-width. Chela ( Figure 22 A–C) and metasomal segment V ( Figure 16C, D ) incrassate in large males.

Affinities with other Colombian species. Tityus columbianus is most similar to T. parvulus , sharing with it the presence of a subaculear tubercle that is medium-sized pyramidal, flat, and triangle-like with an acute and coarse apex in lateral view ( Figures 19B , 20B ). Tityus columbianus can be readily distinguished by the presence of a general body coloration with a dark yellow background moderately covered with dark brown spots ( Figure 21 ); metasomal segment V completely dark reddish-brown with three ventral black stripes, a dorsomedian area without spots ( Figure 6B, E ) and with the VSM2 macrosetae; metasomal segments II–III each with a distoterminal small and acute granule on the DL carinae ( Figure 15B , 16C, D ) and metasomal segment IV with a medium-sized blunt, and coarse granule on the DL carinae ( Figures 15B , 16C, D ); metasomal intercarinal areas densely covered with coarse granules ( Figures 15B , 16C, D ); subaculear tubercle with VM carina composed of coarse granules and strongly crenulate in lateral view ( Figures 19B , 20B ); metasomal segments ( Figures 15B , 16C, D ) and male pedipalp femur and patella short and of a medium-width ( Figure 10C, D ).

Conversely, in T. parvulus the general body coloration is with a dark yellow background slightly covered with dark brown variegated spots ( Figure 31 ); the metasomal segment V is dark yellow slightly covered with dark brown variegated spots and a pair of distal VL dark brown spots, but does not have dorsal spots neither the VSM macrosetae ( Figure 33A, C ); the metasomal segments II–IV do not have conspicuous distoterminal granules on the DL carinae ( Figures 39A , 40A, B ); the metasomal intercarinal areas are densely covered with fine granules ( Figures 39A , 40A ); the subaculear tubercle has a VM carina composed of fine granules and is feebly crenulate in lateral view ( Figures 19D , 20D ); the metasomal segments are slender in both sexes ( Figures 39A , 40A, B ) and the male pedipalp femur and patella are slightly elongated and slender ( Figure 10G, H ).

Remarks. Lourenço (2000) provided records for Tityus erikae from Vereda Miralindo, Medina, Cundinamarca department (ICN-As-194) and Santuario de Flora y Fauna Los Colorados, Bolívar department (ICN-As-188). These specimens were examined and we consider that they correspond to adult males of Tityus columbianus . Also, we examined two adult males, seven adult females and one juvenile female from Reserva Natural La Planada, Nariño department (ICN-As-712); and they also correspond to Tityus columbianus .

The records from Santuario de Flora y Fauna Los Colorados, Bolívar, and Reserva Natural La Planada, Nariño are unusually far away from the known distribution range of T. columbianus which is in the Eastern Cordillera in Cundinamarca and Boyacá departments. However, only new collections from these localities could explain whether these records are due to mislabeling errors or introduced populations.

Redescription. Based on an adult female (ICN-As-739) and an adult male (ICN-As-524). For female live habitus see Figure 46B .

Total length. Female: 35.22 mm; male: 40.29 mm.

Coloration. General body coloration (in ethanol 70%) ( Figure 21 ) with a dark yellow background moderately covered with dark brown variegated spots. Carapace ( Figure 5C, D ) moderately covered with dark 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 7C, D ) coxa and hand with a light yellow background; hand with dark brown reticulated spots on the anterior half and a small areas without spots on the adjacent area to the movable finger articulation; movable and fixed fingers with dark brown spots along their lengths; teeth dark reddish-brown. Pedipalps ( Figure 21 ) moderately covered with dark 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 21B, E, F ) coxae I–IV, sternum, genital operculum, and pectines (teeth) light yellow; basal pectinal piece with an anterior white area (female); coxaphophyses I–II, pectines (marginal lamellae, middle lamellae, fulcra), and basal pectinal piece with a light yellow background moderately covered with dark brown variegated spots. Mesosoma ( Figure 21 ) tergites I–VII each with a dark yellow background moderately covered with dark brown variegated spots and with four posterior dark brown spots (two submedian and two lateral); sternites III–VII each densely covered with dark reddish-brown variegated spots; spiracles yellow. Metasoma ( Figure 6B, E ) with a dark yellow background moderately covered with dark brown variegated spots and with ventral yellow spots; each segment progressively darker towards segment V; segment V completely dark reddish-brown with three ventral black stripes and with a dorsomedian area without spots ( Figure 6B, E ). 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 21 ) almost completely dark reddish-brown with a lateral light reddish-brown stripe; subaculear tubercle and aculeus dark reddish-brown.

Morphology. Carapace ( Figure 8C, D ) densely covered with coarse granules and a few fine 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 (female) or feebly marked (male); median ocular tubercle well-marked and located on the anterior half of the carapace; median eyes separated by about 1.6 (female) and 1.3 (male) 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 7C, D ) dentition characteristic of the family Buthidae ( Vachon 1963) ; hand and fingers densely covered with setae on internal and ventral surfaces.

Pedipalps. Chela short and of a medium-width (female) (Cl/Cw= 4.9) or incrassate (male) (Cl/Cw= 3.1). 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 9C , 10C ) with five well-marked and complete carinae: VI, DI, and DE crenulate; VE and IM serratocrenulate; intercarinal areas moderately covered with coarse granules and densely covered with fine granules ( Figures 9C , 10C ). Patella ( Figures 9D , 10D ) 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 D–F, 22A–C) with eight carinae: VI, VE, D, DS, DMA, and ES complete and crenulate; DI complete and feebly crenulate (female) ( Figure 11 D–F) or obsolete (male) ( Figure 22 A–C); 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 22A ) or without a basal lobe (female) ( Figure 11D ); dorsal surface of the movable finger with 13–13 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) (e.g., Figure 2A ) or flat, shield-shaped with an anterior median notch well-marked (male) (e.g., Figure 2B ); pectinal tooth counts: 13–14 (female) and 14–14 (male). Intermediate plate, marginal plate, and fulcra moderately covered with setae ( Figure 14E ). Pectinal teeth peg sensillae flattened with a slightly spatulate apex ( Figure 14F ).

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

Mesosoma . Tergites I–VI densely covered with coarse granules and a few fine granules; pre-tergite well-marked 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 coarse granules and a few 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 area; sternite VI with VSM carinae crenulate occupying the posterior half of the sternite; sternite VII with VSM carinae crenulate occupying the posterior 2/3 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 23 A–C) with a straight distal carina ending in a 70° 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; trans- lucent area basally wide but progressively narrower towards the distal region ( Figure 23B, C ); basal carina with a rounded apex ( Figure 23C ); basal hook semi ovate-shaped with anterior margin straight in lateral view ( Figure 23B ), and slightly curved in anterior and posterior views; basal hook forming a ‘’U’’-shaped curve with the stem ( Figure 23A, C ); basal hook elongated with a thick and rounded apex and approximately 1.2 longer than its basal width in anterior view ( Figure 23C ).

Metasoma. Segments II–V short and of a medium-width in both sexes (Msl/Msw ratio= female/ male: II=1.4/ 1.5; III= 1.6/ 1.7; IV= 1.8/ 1.9; V= 2.0/ 1.7). Segment V short and of a medium-width (female) ( Figure 15B ) or incrassate (male) ( Figure 16C ). Segments I–II ( Figures 15B , 16C ) 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 coarse granules almost along all its length; intercarinal areas densely covered with coarse granules. Segment III–IV ( Figures 15B , 16C ) with eight complete, parallel, and crenulate carinae (paired DL, ML, VL, and VSM; DL carinae serratocrenulate); intercarinal areas densely covered with coarse granules. Segment V with five complete and crenulate carinae (VM, paired DL, and VL); lateral and ventral intercarinal areas densely covered with coarse granules and a few fine granules ( Figures 15B , 16C ); 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 acute and small granule (II–III) or a blunt and coarse granule (IV) which increase in size towards the segment IV ( Figures 15B , 16C ).

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 in 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 17B , 18B ) 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 distal or at same level of pair VL2 ( Figures 17B , 18B ), 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= 2.0) ( Figures 19B , 20B ) 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 19B ) or feebly marked (male) ( Figure 20B ); paired DL carinae located dorsally with respect to the lateral longitudinal furrow; VM carina well-marked along its length ( Figures 19B , 20B ). Subaculear tubercle medium-sized pyramidal, flat, and triangle-like with an acute and coarse apex in lateral view ( Figures 19B , 20B ); dorsal margin of the subaculear tubercle with a pair of ventrodistal small rounded granules with apices pointing to the middle portion of the aculeus; VM carina composed of coarse granules and strongly crenulate in lateral view ( Figures 19B , 20B ); aculeus strongly curved shorter than vesicle and with a ventral groove.

Variability. Total length (including telson ): males 28.16–39.11 mm (n= 14; mean= 31.83; SD= 3.91); females 25.64–37.54 mm (n= 57; mean= 33.46; SD= 2.08) ( Table 9 ). Pectinal tooth counts: males 13–15 (n= 42; mode= 14); females 11–15 (n= 208; mode= 12) ( Table 2). Number of movable finger oblique rows of denticles: males 12–14 (n= 39; mode= 13); females 11–13 (n= 203; mode= 13) ( Table 3). Metasomal macrosetae counts: segment I: males (n= 21): 2/2 (n= 10), 2/3 (n= 1), 3/2 (n= 2), 3/3 (n= 7), 3/4 (n= 1) VSM macrosetae on the segment; segments II–IV: males (n= 21) 2/2 VSM on each segment; segment V: males (n= 21): 3/3 VSM and 3/3 VL macrosetae on the segment. Segment I: females (n= 116): 2/2 (n= 57), 2/3 (n= 9), 3/2 (n= 2), 3/3 (n= 46), 3/4 (n= 1), 4/2 (n= 1) VSM macrosetae on the segment; segment II–III: females (n= 116): 2/2 (n= 115), 2/3 (n= 1) VSM macrosetae on each segment; segment IV: females (n= 116): 2/2 (n= 106), 3/2 (n= 2), 3/3 (n= 4) VSM macrosetae on the segment; segment V: females (n= 116): 3/2 (n= 1), 3/3 (n= 114), 3/4 (n= 1) VSM macrosetae; 3/3 VL macrosetae, and 1/1 DL macrosetae on the segment. Segments I–IV: males (n= 21) and females (n= 116): 2/2 VL macrosetae on each segment. Relative position of the VSM2 macrosetae on the metasomal segment V: males (n= 21): distal to VL2 (n= 7) or at the same level of VL2 (n= 14); females (n= 116): distal to VL2 (n= 27); at the same level of VL2 (n= 78); slightly proximal to the level of VL2 (n= 1) or asymmetric with just one seta at the same level of VL2 (n= 10).

Hemispermatophore capsule: the basal hook of the capsule exhibited different shapes in anterior view: (1) elongated with a thick and rounded apex, curved lateral margin, and forming ‘’U’’-shaped curve with the stem ( Figure 23C ) and (2) short with an acute apex, curved lateral margin, and forming a ‘’U’’-shaped curve with the stem ( Figure 23D ).

Male intrasexual variations: adult males exhibited different body sizes ranging between 28.16–39.11 mm, but despite this they did not show discrete size-classes, but rather a progressive increase in their total lengths ( Figure 21 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) Pedipalp femur: intercarinal areas have coarser and finest granules ( Figure 9C ); (3) Chela : the DI carina is more obsolete ( Figure 22 A–F); the manus is more incrassate without apparently exhibiting isometric growth (n= 14; Cl/Cw= 3.1–4.7; Cl/Ch= 3.0–4.0) ( Figure 22 A–F); (4) Metasomal segments II–IV: the DL carinae exhibit coarser distoterminal granules ( Figure 16C, D ); and (5) Metasomal segment V: segment is more incrassate without apparently exhibiting an isometric growth (n= 14; Msl-V/Msw-V= 1.8–2.1; Msl-V/Msh-V= 1.6–2.1) ( Figure 16C, D ), and the granules associated with carinae and intercarinal areas are less marked ( Figure 16C, D ).

Distribution ( Figure 30 ). COLOMBIA: Boyacá department:Arcabuco, Chiquinquirá, Chita, Cuítiva, Duitama, Iza, Firavitoba, La Uvita, Mongui, Úmbita and Villa de Leyva. Cundinamarca department: Bogotá, Cajíca, Chocontá, Gachancipá, Guasca, Guatavita, La Calera, Mosquera, Pacho, Ubaté, and Soacha ( Figure 47B, C ). New records. COLOMBIA: Bolívar department: Santuario de Flora y Fauna Los Colorados. Boyacá department: Susacón and Tinjáca. Cundinamarca department: Carmen de Carupa, Facatativá, Medina, Nariño, Susa, and Tabio. Nariño department: Natural Reserve La Planada. Tolima department: Ibagué.

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 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 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 21. Tityus (Archaeotityus) columbianus (Thorell, 1876), habitus. A, B. Female (ICN-As-739). A. Dorsal aspect. B. Ventral aspect. C, E. Male (Villa de Leyva, Boyacá) (ICN-As-739). C. Dorsal aspect. E. Ventral aspect. D, F. Male (Villa de Leyva, Boyacá) (ICN-As-524). D. Dorsal aspect. F. Ventral aspect. Scale bars= 10 mm.

FIGURE 22. Tityus (Archaeotityus) columbianus (Thorell, 1876), male pedipalp chela. A–C. Male (Villa de Leyva, Boyacá) (ICN-As-524). A. External aspect. B. Dorsal aspect. C. Ventral aspect. D–F. Male (Villa de Leyva, Boyacá) (ICN-As-739). D. External aspect. E. Dorsal aspect. F. Ventral aspect. Scale bars= 2.0 mm.

FIGURE 23. Tityus (Archaeotityus) columbianus (Thorell, 1876), dextral hemispermatophore, showing capsule details, male (ICN-As-739). A. Posterior view. B. Lateral view. C. Anterior view. D. Capsule variation, anterior view, male (Villa de Leyva, Boyacá) (ICN-As-524). Abbreviations: Bc, basal carina; Bh, basal hook; Dc, distal carina; Sdm, sperm duct membrane. Scale bar= 1.0 mm.

FIGURE 30. Distribution of Tityus (Archaeotityus) columbianus (Thorell, 1876) and Tityus (Archaeotityus) guane sp. nov., showing known records and new locality records.

FIGURE 31. Tityus (Archaeotityus) parvulus Kraepelin, 1914, habitus. A, B. Female (ICN-As-1004). A. Dorsal aspect. B. Ventral aspect. C, E. Male (Chinchiná, Caldas) (ICN-As-1004). C. Dorsal aspect. E. Ventral aspect. D, F. Male (Chinchiná, Caldas) (ICN-As-1004). D. Dorsal aspect. F. Ventral aspect. Scale bars= 10 mm.

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 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.

FIGURE 47. Tityus (Archaeotityus) Lourenço, 2006, habitat. A. Vereda El Carmen, Villavicencio, Meta, Colombia, habitat of Tityus bastosi Lourenço, 1984. B, C. Soacha, Cundinamarca, Colombia, habitat of Tityus columbianus (Thorell, 1876). D. Botanical Garden Maria Cespedes [Jardín Botánico Maria Cespedes], Tulúa, Valle, Colombia, habitat of Tityus parvulus Kraepelin, 1914. E. Vereda Pasochico, Finca La Ceiba, Los Santos, Santander, habitat of Tityus guane sp. nov. F. Natural National Park Tayrona [Parque Natural Nacional Tayrona], Santa Marta, Magdalena, Colombia, habitat of Tityus tayrona Lourenço, 1991. Photos B and C, courtesy of N. Herreño.