Tityus parvulus Kraepelin, 1914

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 : 53-66

Tityus parvulus Kraepelin, 1914

Figures 9 G–H, 10G–H, 17D, 18D, 19D, 20D, 25D–F, 31, 32A, B, 33A, C, 34A, B, 35A, B, 36, 37, 38A–D, 39A, 40A, B, 45, 46D, E, 47D, 48, 49; Tables 1–3, 6–8, 10 .

Tityus parvulus Kraepelin 1914: 19–21 .

References after Fet & Lowe (2000: 255). Tityus parvulus: Flórez 2001a: 60 , 61; Flórez 2001b: 28; Botero-Trujillo & Fagua 2006: 130; Lourenço 2006: 60; Teruel & García 2008: 12; Teruel & Cozijn 2011: 4.

Tityus betschi syn. nov.: Lourenço 1992: 474–476, figures 1–4, 5, table I; Lourenço 2000: 456, 458, figure 13; Flórez 2001a: 57; Flórez 2001b: 28; Kovarik 2007: 3; Lourenço 2008: 196–198, 205, figures 1, 2, 3–9, 10, table 1; Botero-Trujillo & Fagua 2006: 129, 130, 131–132, figures 9–10; Lourenço 2006: 60; Teruel & García 2008: 10, 12; Kovarik et al. 2015: 36.

Type material. Tityus betschi : Holotype: COLOMBIA: Valle del Cauca department: one adult female, Parque Nacional Farallones de Cali, Cerca a Pance , ix.1988, E. Flórez ( MNHN-RS-8552 ) (Examined). Tityus parvulus : Syntypes: COLOMBIA: Antioquia department: one adult female and one adult male, Angelópolis, 1911, Fuhrmann ( ZMH) ( Examined ).

Examined material. COLOMBIA: Antioquia department: one adult female, Tarso, Finca Las Camelias , Vereda El Cedrón , 5°52’17.21’’ N 75°49’36.25’’ W, 1200 masl, inside a secondary forest, 3.iv.2007, J. M. Palacios ( MPUJ-SCO-369 ) ; one adult female, Tarso, Finca Las Camelias , Vereda El Cedrón , 5°52’17.21’’ N 75°49’36.25’’ W, 1200 masl, inside a Secondary forest, 3.iv.2007, J. M. Palacios ( MPUJ-SCO-370 ) . Caldas department: two adult males and one adult female, Chinchiná, 4°59’28.97’’ N 75°35’38.33’’ W, 1400 masl, vii.2012, manual capture, Coll. Indet . ( ICN-As-1004 ) ; four adult males, Villamaría, 5°2’06.98’’ N 75°31’35.14’’ W, 1450 masl, v.2012, Coll. Indet . ( MZSP 71738 ) . Cauca department: four adult females, Santander de Quilichao, Vereda El Carbonero , 2°59’30.4’’ N 76°28’29.1’’ W, night manual capture with UV light, 18.iii.2012, N. Hazzi ( ICN-As-991 ) . Quindío department: four adult females, Armenia, forest inside of Universidad del Quindío , 4°33’14.14’’ N 75°39’35.4’’ W, 1532 masl, xii.2011, night manual capture with UV light, R. Botero-Trujillo ( MZSP 71737 ) . Risaralda department: one juvenile male, La Virginia, Finca Alejandría , 4°53'47.22'' N 75°52'59.41'' W, 900 masl, 28.iv.2005, C. Bermúdez ( MUSENUV 27573 ) . Valle del Cauca department: one adult female, Cali, Vereda Los Limones , 3°28’41.46’’ N 76°33’51.49’’ W, 1214 masl, night manual capture, viii.2012, J. A. Moreno ( MZSP 71739 ) ; one adult female, Roldanillo, Vereda Cajamarca , 4°28’22.59’’ N 76°12’28.75’’ W, 1496 masl, night manual capture, viii.2013, O. Saenz ( ICN-As) ; three adult females, Yotoco, Natural Reserve Bosque de Yotoco , 3°52’32.2’’ N 76°26’07.8’’ W, 1573 masl, night manual capture with UV light, 5.ii.2010, J. A. Moreno ( ICN-As-843 ) ; six adult females, Yotoco, Natural Reserve Bosque de Yotoco , 3°52’32.2’’ N 76°26’07.8’’ W, 1573 masl, night manual capture with UV light, 5.ii.2010, J. A. Moreno ( ICN-As-992 ) ; one adult female, Calima- Darién , 3°55’54.03’’ N 76°29’0.63’’ W, 1520 masl, diurnal manual capture inside a house, vi.2011, N. Hazzi ( ICN-As-994 ) ; one adult female, Zarzal, Finca Las Pilas , 4°26’27.63’’ N 75°59’24.7’’ W, 1000 masl, 10.xi.2005, C. Bermúdez ( MUSENUV-27572 ) ; eight adult females, Tuluá, Vereda Mateguadua , Botanical Garden José María Céspedes , 4°1’33.0’’ N 76°9’44.0’’ W, 1100 masl, L. Báez ( ICN-As-508 ) ; two adult females and one subadult female, Tuluá, Vereda Mateguadua , Botanical Garden José María Céspedes , 4°1’33.0’’ N 76°9’44.0’’ W, 1100 masl, S. Jaramillo ( ICN-As-509 ) ; one adult female, Tuluá, Vereda Mateguadua , Botanical Garden José María Céspedes , 4°1’33.0’’ N 76°9’44.0’’ W, 1100 masl, Dry Tropical Forest , x.1998, ( MPUJ-SCO-295 ) ; one juvenile female, Tuluá, Vereda Mateguadua , Botanical Garden José María Céspedes , 4°1’33.0’’ N 76°9’44.0’’ W, 1100 masl, Dry Tropical Forest , x.1998, L.A.S ( MPUJ-SCO-296 ) ; one adult female, Tuluá, Vereda Mateguadua , Botanical Garden José María Céspedes , 4°1’33.0’’ N 76°9’44.0’’ W, 1100 masl, Dry Tropical Forest , x.1998, N. Contreras ( MPUJ-SCO-297 ) ; one adult female, Tuluá, Vereda Mateguadua , Botanical Garden José María Céspedes , 4°1’33.0’’ N 76°9’44.0’’ W, 1100 masl, Dry Tropical Forest , 6.x.1998, N. Contreras ( MPUJ-SCO-298 ) ; seven adult females, Buga, Natural Regional Park El Vínculo , 3°50’04.3’’ N 76°17’54.8’’ W, 1022 masl, Dry Tropical Forest , night manual capture with UV light, 16.v.2010, J. A. Moreno ( ICN-As-993 ) .

Diagnosis. This species can be readily recognized from other Tityus (Archaeotityus) species by the following combination of character states: Total length: 35.07–41.42 mm (males) and 34.59–39.15 mm (females). General body coloration ( Figure 30 ) dark yellow background slightly covered with dark brown variegated spots; metasomal segment V dark yellow slightly covered with dark brown variegated spots, with a distal pair VL dark brown spots and without dorsal spots ( Figure 33A, C ). Metasomal segments II–IV ( Figures 39A , 40A, B ) without conspicuous distoterminal granules on the DL carinae. Metasomal segment V without the VSM2 macrosetae ( Figures 17D , 18D ). Subaculear tubercle medium-sized pyramidal, flat, and triangle-like with an acute and coarse ( Figures 19D , 20D ). 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 coarse granules and feebly crenulate in lateral view ( Figures 19D , 20D ). Telson with well-marked (female) ( Figure 19D ) or obsolete (male) carinae ( Figure 20D ). Metasomal intercarinal areas densely covered with fine granules ( Figures 39A , 40A, B ). Pectinal tooth counts and movable finger oblique rows of denticles: 12–14/ 12–13 (females) and 13–15/ 13–14 (males) ( Table 2, 3). Metasomal segments II–V ( Figures 39A , 40A, B ) slightly elongated and slender in both sexes; female pedipalp ( Figure 25 D–F) short and of a medium-width, and male pedipalp femur and patella ( Figure 10G, H ) slightly elongated and slender. Chela ( Figure 36 ) incrassate in large males; metasomal segment V more slender ( Figure 40A ) in large males.

Affinities with other Colombian species. Tityus parvulus is most similar to T. columbianus , 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 19D , 20D ). Tityus parvulus can be readily distinguished because the metasomal segment V is dark yellow, slightly covered with dark brown variegated spots, with a pair of distal VL dark brown spots and without dorsal spots ( Figure 33A, C ); metasomal segments II–IV without conspicuous distoterminal granules on the DL carinae ( Figures 39A , 40A, B ); metasomal intercarinal areas densely covered with fine granules ( Figures 39A , 40A, B ); subaculear tubercle with VM carina composed of fine granules and feebly crenulate in lateral view ( Figure 19D , 20D ); metasomal segments ( Figures 39A , 40A, B ) and male pedipalp femur and patella ( Figure 10G, H ) slightly elongated and slender.

Conversely, in T. columbianus the metasomal segment V is completely dark reddish-brown with three ventral black stripes and a dorsomedian area without spots ( Figure 6B, E ); the metasomal segments II–III each has a distoterminal small and acute granule on the DL carinae ( Figures 15B , 16C, D ); the metasomal segment IV has a medium-sized blunt and coarse granule on the DL carinae ( Figures 15B , 16C, D ); the metasomal intercarinal areas are covered with coarse granules ( Figures 15B , 16C, D ); the subaculear tubercle has a VM carina composed of coarse granules and is strongly crenulate in lateral view ( Figures 19B , 20B ); the metasomal segments of both sexes ( Figures 15B , 16C, D ) and the male pedipalp femur and patella ( Figure 10C, D ) are short and of a medium-width.

Remarks. Lourenço (1999b) describes the male of Tityus parvulus and provides a contradictory identification key. In this key, Tityus parvulus is separated from T. columbianus based on the presence of a subaculear tooth that is “strong and markedly rhomboidal” (Lourenço 1999b: 124). However, in the same paper Lourenço presents a draw- ing of the male metasoma of Tityus parvulus ( Lourenço 1999b: figure 3) which clearly shows a subaculear tubercle that is medium-sized pyramidal, and triangle-like in lateral view. A subaculear tubercle size and shape which are also found in T. columbianus ( Figures 19B , 20B ) and Tityus betschi (Lourenço 1992: figure 4; Lourenço, 2008: figure 2). Later on, Botero-Trujillo & Fagua (2006: 131–132, figure 4) presents a new record for Tityus betschi from Antioquia department. However, after it Teruel & Garcia (2008) criticizes this record arguing (italics added): ‘’... their specimens clearly belong to Tityus parvulus Kraepelin, 1914 …’’.

Therefore, to contribute to the solution of this taxonomic confusion, we examined the type material and specimens from localities between the type localities and near to the type localities of T. bestchi and T. parvulus . It is important to remark that both species are the only known members of Tityus (Archaeotityus) inhabiting the Andean valley of the Cauca river. The following examined material from this region is listed: Cali, Valle del Cauca department (near the type locality of T. bestchi ) ( MZSP 71739); Tarso- La Camelia, Antioquia (near the type locality of T. parvulus ) (MPUJ-SCO-369, MPUJ-SCO-370), and from other intermediate localities in Valle del Cauca, Cauca, Quindío, and Risaralda departments (see examined material) ( Figure 45 ).

After the examination of these specimens from these different populations, we concluded that they all share the following: (1) similar total sizes ( Table 10 ); (2) similar coloration patterns (e.g., chelicerae, carapace, mesosoma , and metasoma); (3) a subaculear tubercle medium-sized pyramidal, flat, and triangle-like with an acute and coarse apex in lateral view ( Figures 19D , 20D ); (4) metasomal intercarinal areas covered with fine granules ( Figures 39A , 40A, B ); (5) metasomal segments slightly elongated and slender with almost identical ratios ( Table 7 , 8 ); (6) the absence of the VSM2 macrosetae on the metasomal segment V ( Figures 17D , 18D ); and (7) the same ranges for pectinal tooth counts and the number of movable finger oblique rows of denticles ( Tables 2, 3). Therefore, given the morphological evidence and the shared geographical distribution, there are no reasons to support both species as valid entities; thus, we propose Tityus betschi Lourenço 1992 as a synonym of Tityus parvulus Kraepelin 1914 .

Redescription. Based on an adult female (ICN-As-1004) and an adult male (ICN-As-1004). For female live habitus see Figure 46D, E .

Total length. Female: 39.15 mm; male 41.24 mm.

Coloration. General body coloration (in ethanol 70%) ( Figure 31 ) with a dark yellow background slightly covered with brown variegated spots. Carapace ( Figure 32A, B ) moderately covered with dark brown variegated spots; lateral and median eyes surrounded by black spots; posterior area to the median ocular tubercle with an inverted ‘’Y’’-shaped spot. Chelicerae ( Figure 34A, B ) coxa and hand with a light yellow background; hand with a few brown reticulated spots on the anterior half and with a large area without spots on the anterior margin; movable and fixed fingers with dark brown spots on their posterior halves; teeth dark reddish-brown. Pedipalps ( Figure 31 ) 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 31B, E, F ) coxae I–IV, coxapophyses I–II, sternum, genital operculum, pectines, and basal pectinal piece light yellow; basal pectinal piece with an anterior white area (female). Mesosoma ( Figure 31 ) tergites I–VII each with a dark yellow background slightly covered with dark brown spots and with four posterior dark reddish-brown spots (two submedian and two lateral); sternites III–VII each moderately covered with dark brown variegated spots; spiracles yellow. Metasoma ( Figure 33A, C ) with a dark yellow background slightly covered with dark brown variegated spots and with ventral yellow spots; segment V dark yellow slightly covered with dark brown variegated spots, with a pair of distal VL dark brown spots and without dorsal spots ( Figure 33A, C ). DL intercarinal areas of segments I–IV each with a posterior median brown spot; ML and VSM intercarinal areas of segments I–IV each with a pair of distal brown spots. Telson ( Figure 31 ) almost completely dark yellow slightly covered with dark brown spots and with a lateral light reddish-brown stripe; subaculear tubercle and aculeus dark reddish-brown.

Morphology. Carapace ( Figure 35A, 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; mediun ocular tubercle well-marked and located on the anterior half of the carapace; median eyes separated by about 1.2 (female) and 1.1 (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 34A, B ) 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= 5.1) or incrassate (male, Cl/Cw= 3.9). 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 9G , 10G ) with five well-marked and complete carinae: VI, DI, and DE crenulate; VE and IM serratocrenulate; intercarinal areas moderately covered with fine granules. Patella ( Figures 9H , 10H ) 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. Chela (Tibia) ( Figures 25 D–F, 36) with eight carinae: VI, VE, D, DS, DMA, and ES complete and crenulate; DI complete and feebly crenulate ( Figures 25 D–F, 36); SA incomplete and crenulate, only present on the anterior half of the hand. Pedipalp movable and fixed fingers with an obsolete basal lobe (male) ( Figures 36A ) or without a basal lobe (female) ( Figure 25D ); dorsal surface of the movable finger with 13–13 (female) and 14–14 (male) oblique rows of denticles.

Coxosternal region. Covered with fine granules and a few setae, except for the coxaphophyses 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) or flat, shield-shaped with an anteriomedian notch well-marked (male) (e.g., Figure 2 ); pectinal tooth counts: 13–13 (female) and 13–14 (male). Intermediate plate, marginal plate, and fulcra moderately covered with setae ( Figure 38A, C ). Pectinal teeth peg sensillae flattened with a slightly spatulate apex ( Figure 38B, 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 densely covered with fine granules and a few coarse 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 moderately 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 37 A–C) with a straight distal carina ending in a 90° 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 37 B–C); basal carina with rounded apex ( Figure 37C ); basal hook semi ovate-shaped with anterior margin straight in lateral view ( Figure 37B ) and slightly curved in anterior and posterior views; basal hook forming a narrow ‘’U’’-shaped curve with the stem ( Figure 37A, C ); basal hook slightly elongated with an obtuse apex and approximately as long as its basal width in anterior view ( Figure 37A, C ).

Metasoma. Segments II–V slightly elongated and slender in both sexes (Msl/Msw ratio= female/ male: II= 1.7/ 2.2; III= 2.0/ 2.5; IV= 2.2/ 2.8; V= 2.5/ 3.0). Segments I–II ( Figures 39A , 40A ) with 10 complete, parallel and crenulate carinae (paired DL, ML, LIM, VL, and VSM); LIM on segment II composed of coarse granules on the two posterior thirds and fine granules on the first anterior third; intercarinal areas densely covered fine granules. Segment III–IV ( Figures 39A , 40A ) 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); intercarinal lateral and ventral areas densely covered with coarse granules and a few fine granules ( Figures 39A , 40A ); intercarinal areas with granules irregularly distributed on the middle of each area. Segment I–IV with DL carinae without a conspicuous distoterminal granule ( Figures 39A , 40A ).

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 near the posterior third of the segment (e.g., Figure 1 ). Segment V ( Figures 17D , 18D ) with two pairs of VSM macrosetae (2/2), 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 absent, 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 VL2 is located on the anal arch; pair DL1 is located near the posterior margin of the segment.

Telson . Vesicle not elongated and suboval (Vl/Vw= 2.0 (female) ( Figure 19D ) and Vl/Vh= 2.1 (male) ( Figure 20D )) 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 19D ) or obsolete (male) ( Figure 20D ); paired DL carinae located dorsally with respect to the lateral longitudinal furrow; VM carina feebly marked, basally crenulate, but fading progressively towards the subaculear tubercle apex ( Figures 19D , 20D ). Subaculear tubercle medium-sized pyramidal, flat, and triangle-like with an acute and coarse apex in lateral view ( Figures 19D , 20D ); 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 19D , 20D ); aculeus strongly curved shorter than vesicle and with a ventral groove.

Variability. Total length (including telson ): males 35.07–41.42 mm (n= 6; mean= 38.46; SD= 2.88); females 34.59–39.15 mm (n= 31; mean= 36.88; SD= 1.17) ( Table 10 ). Pectinal tooth counts: males 13–15 (n= 29; mode= 14); females 12–14 (n= 88; mode= 13) ( Table 2). Number of movable finger oblique rows of denticles: males 13–14 (n= 12; mode= 14); females 12–14 (n= 85; mode= 14) ( Table 3). Metasomal macrosetae counts: segments I–IV: males (n= 6) and females (n= 49): 2/2 VSM and 2/2 VL macrosetae on each segment; segment V: 1/1 VSM and 2/2 VL macrosetae on the segment. Relative position of the VSM2 macrosetae on the metasomal segment V: VSM2 macrosetae pair absent.

Male intrasexual variations: adult males exhibited different body sizes ranging between 35.07–41.42 mm, but despite this they did not show discrete size-classes but rather a progressive increase in their total lengths ( Figure 31 C–F, Table 10 ). However, as the total body size increases among males the following: (1) Carapace and tergites: the number of coarse and fine granules on all intercarinal surfaces increases; (2) Chela : the manus is more incrassate without apparently exhibiting isometric growth (n= 6; Cl/Cw= 3.8–5.1; Cl/Ch= 3.8–4.9) ( Figure 36 ); (3) Metasomal segment V: segment is more slender without apparently exhibiting isometric growth (n= 6; Msl-V/Msw-V= 2.9–3.0; Msl-V/Msh-V= 2.7–2.9) ( Figure 40A, B ), and the granules associated with carinae and intercarinal areas are less marked ( Figure 40A, B ).

Distribution ( Figure 45 ). COLOMBIA: Antioquia department: Angelópolis. Quindío department: Armenia. Valle del Cauca department: Cali (P.N.N. Farallones) and Tuluá ( Figure 46E ). New records. COLOMBIA: Antioquia department: Tarso. Caldas department: Chinchiná and Villamaría. Cauca department: Santander de Quilichao. Risaralda department: La Virginia. Valle del Cauca department: Cali (Vereda Los Limones), Buga ( Figure 46D ), Darién (Calima), Roldanillo, Zarzal, and Yotoco.

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 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 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 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 25. Tityus (Archaeotityus) Lourenço, 2006, right female pedipalp sculpturing. A–C. Tityus guane sp. nov. (paratype) (ICN-As-1003). A. External aspect. B. Dorsal aspect. C. Ventral aspect. D–F. Tityus parvulus Kraepelin, 1914 (ICN-As-1004). D. External aspect. E. Dorsal aspect. F. Ventral aspect. Scale bars= 2.0 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 32. Tityus (Archaeotityus) Lourenço, 2006, carapace coloration, dorsal aspect. A, B. Tityus parvulus Kraepelin, 1914 (ICN-As-1004). A. Female. B. Male. C, D. Tityus tayrona Lourenço, 1991 (ICN-As-181). C. Female. D. Male.

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 34. Tityus (Archaeotityus) Lourenço, 2006, chelicerae, dorsal aspect, showing reticulation pattern. A, B. Tityus parvulus Kraepelin, 1914 (ICN-As-1004). A. Female. B. Male. C, D. Tityus tayrona Lourenço, 1991 (ICN-As-181). C. Female. D. Male.

FIGURE 35. Tityus (Archaeotityus) Lourenço, 2006, carapace sculpturing, dorsal aspect. A, B. Tityus parvulus Kraepelin, 1914 (ICN-As-1004). A. Female. B. Male. C, D. Tityus tayrona Lourenço, 1991 (ICN-As-181). C. Female. D. Male. Scale bars= 1.5 mm.

FIGURE 36. Tityus (Archaeotityus) parvulus Kraepelin, 1914, male pedipalp chela. A–C. Male (Chinchiná, Caldas) (ICN-As- 1004). A. External aspect. B. Dorsal aspect. C. Ventral aspect. Scale bar= 2.0 mm.

FIGURE 37. Tityus (Archaeotityus) parvulus Kraepelin, 1914, dextral hemispermatophore, showing capsule details, male (Villamaría, Caldas) (MZSP 71738). A. Posterior view. B. Lateral view. C. Anterior view. D. Capsule variation, anterior view, male (Chinchiná, Caldas) (ICN-As-1004). Abbreviations: Bc, basal carina; Bh, basal hook; Dc, distal carina; Sdm, sperm duct membrane. Scale bar= 1.0 mm.

FIGURE 38. Tityus (Archaeotityus) Lourenço, 2006, left pectine with peg sensillae details, ventral aspect.A–D. Tityus parvulus Kraepelin, 1914. A, B. Female (ICN-As-1004). A. Pectine. B. Peg sensillae shape. C, D. Male (ICN-As-1004). C. Pectine. D. Peg sensillae shape. E–H. Tityus tayrona Lourenço, 1991. E, F. Female (MZSP 71735). E. Pectine. F. Peg sensillae shape. G, H. Male (MZSP 71735). G. Pectine. H. Peg sensillae shape. Scale bars= A, C, E, G= 400 µm; B, D, F, H= 10 µm.

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 45. Distribution of Tityus (Archaeotityus) Lourenço, 2006, showing known and new locality records. Tityus parvulus Kraepelin, 1914 (top). Tityus tayrona Lourenço, 1991 (bottom).

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.