Tityus tayrona Lourenço, 1991 (Lourenco, 1999)

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 : 66-77

Tityus tayrona Lourenço, 1991

Figures 9 I–J, 10I–J, 17E, 18E, 19E, 20E, 32C, D, 33B, D, 34C, D, 35C, D, 38E–H, 39B, 40C, D, 41–45, 46F, 47F, 48, 49; Tables 1–3, 5–8 , 10 .

Tityus tayrona Lourenço 1991: 279–281 , figures 3–4, tables 1, 2.

References after Fet & Lowe (2000: 263). Tityus erikae Lourenço 1999a (synonymized by Teruel & Roncallo 2010): 1–6, figures 1–7, table I; Lourenço 2000: 456, 458, figure 13; Flórez 2001a: 59, 60; Flórez 2001b: 28; Botero-Trujillo & Fagua 2006: 131; Kovarik 2007: 3; Teruel & García 2008: 12; Teruel & Roncallo 2010: 1, 6, 8.

Tityus tayrona: Lourenço 2000: 456 , 458, figure 13; Flórez 2001a: 61; Flórez 2001b: 28; Botero-Trujillo & Fagua 2006: 129– 130, 132–133, figures 12–15; Lourenço 2006: 60, figures 1–2; Rojas-Runjaic & Armas 2007: 63; Botero-Trujillo & Noriega 2008: 263; Teruel & García 2008: 9, 12; Teruel & Roncallo 2010: 1 , 4–8, figures 3–7, table 1; Teruel & Cozijn 2011: 1, 3, figure 2; Gómez et al. 2013: 151, figure 2c.

Tityus wayuu syn. nov. Rojas-Runjaic & Armas 2007: 58–63, figures 4–8, tables 4–5.

Type material. Tityus erikae : Holotype: COLOMBIA: Cesar department: one juvenile male, Chimichagua, 1997, E. B. Vargas ( ICN-As) (Examined).

Tityus tayrona : Holotype: COLOMBIA: Magdalena department: one adult male, Parque Nacional Natural Tayrona, Bahía de Guairaca , 13.iv.1985 (aus Fallaub) ( SMF 37019 ) (Examined) . Paratypes: COLOMBIA: Magdalena department: one adult female, Parque Nacional Natural Tayrona, Bahía de Neguangue , 8.x.1985 (aus Fallaub) ( SMF 37024 ) (Not examined) ; one adult male, same locality as the holotype, 6.xi.1985 (aus Fallaub) ( SMF 37020 ) (Not examined) ; one adult male and one adult female, Parque Nacional Natural Tayrona, same locality as the holotype, 31.x.1985 (aus Fallaub) ( SMF 37021 ) (Not examined) ; one adult male and two adult females, same locality as the holotype, 12.vii.1985 (aus Fallaub) ( SMF 37022 ) (Examined) ; one juvenile, same locality as the holotype, 19.vi.1985 (aus Fallaub) ( SMF 37023 ) (Not examined) ; three adult males and one adult female, Parque Nacional Natural Tayrona, Bahía de Neguangue , 30.ix.1985 (aus Fallaub) ( SMF 37025 ) (Examined) ; five adult males and six adult females, Villa Culebra near Bonda ( Barber-Falle in Trokkengebit ), vii.1985 ( SMF 37026 ) (Examined) .

Tityus wayuu : Holotype: VENEZUELA: Zulia state: one adult male, Sierra de Perijá, Mara , Parroquia Monseñor Marcos Sergio Godoy , Fundo La Orchila , Riecito Maché , 18.x.2003, F. Rojas-Runjaic ( MBLUZ 251 ) (Not examined) . Paratypes: VENEZUELA: Zulia state: four adult females, same data as the holotype ( MBLUZ 250 , MBLUZ 253 , IES, MHNLS 1210 ) (Not examined) ; one adult female, same locality as the holotype, 13.xii.2003, F. Rojas-Runjaic ( MBLUZ 259 ) (Not examined) ; one adult male, same locality as the holotype, 10.ix.2000, F. Rojas- Runjaic ( MBLUZ 169 ) (Not examined) ; three adult males ( MNHLS 1211 , MBLUZ 091 , MBLUZ 092 ) and three adult females ( MBLUZ 093 , MBLUZ 094 , IES), same locality as the holotype, 29.vi.2003, F. Rojas-Runjaic & C. Portillo (Not examined) ; one adult female, same locality as the holotype, 25.xi.2002, F. Rojas-Runjaic & C. Portillo ( MBLUZ 055 ) (Not examined) ; two adult males ( MNHLS 1212 , MNHLS 1213 ), four adult males ( IES) and one adult female ( IES), same locality as the holotype, 5-6.iv.2006, P. A. Colmenares, L. F. Armas & L. Morán (Not examined) .

Examined material. COLOMBIA: Antioquia department: three adult females, Santa Fé de Antioquia, Vereda El Espinal , agricultural station Cotové , 6°31’54.26’’ N 75°49’56.07’’ W, 513 masl, under rotten logs, 19.xi.2001, E. Cataño ( MZSP 71734 ) . Atlántico department: one adult male, Barranquilla, Pajonal , 10°59’9.5’’ N 74°46’13.06’’ W, 40 masl, collected with a pitfall tramp for dung-beetles, 15–20.vii.2006, G. Fagua ( MPUJ-SCO-335 ) ; one adult female, Barranquilla, Pajonal , 10°59’9.5’’ N 74°46’13.06’’ W, 40 masl, pitfall, 15–20.vii.2006, G. Fagua ( MPUJ- SCO-336 ) ; one adult female, Barranquilla, Pajonal , 10°59’9.5’’ N 74°46’13.06’’ W, 40 masl, collected with a pitfall tramp for dung-beetles, 15–20.vii.2006, G. Fagua ( MPUJ-SCO-337 ) ; one adult male, Barranquilla, Pajonal , 10°59’9.5’’ N 74°46’13.06’’ W, 40 masl, pitfall, 15–20.vii.2006, G. Fagua ( MPUJ-SCO-347 ) ; one adult female, Puerto Colombia, Loma China , 11°0’58.46’’ N 74°53’0.19’’ W, 40 masl, pitfall, 15–20.vii.2006, G. Fagua ( MPUJ- SCO-338 ) ; one adult male, Puerto Colombia, Costa Alta- El Nisperal , 11°0’58.46’’ N 74°53’0.19’’ W, 100 masl, 10–15.vii.2006, G. Fagua ( MPUJ-SCO-339 ) ; one adult female, Puerto Colombia, Contra Costa- El Nisperal , 11°0’58.46’’ N 74°53’0.19’’ W, 100 masl, pitfall, 10–15.vii.2006, G. Fagua ( MPUJ-SCO-340 ) ; one juvenile male, Puerto Colombia, Costa Alta- El Nisperal , 11°0’58.46’’ N 74°53’0.19’’ W, 100 masl, pitfall, 15–20.vii.2006, G. Fagua ( MPUJ-SCO-341 ) ; one adult female, Puerto Colombia, Costa Baja- El Nisperal , 11°0’58.46’’ N 74°53’0.19’’ W, 100 masl, pitfall, 10–15.vii.2006, G. Fagua ( MPUJ-SCO-342 ) ; one adult female, Puerto Colombia, Loma China , 11°0’58.46’’ N 74°53’0.19’’ W, 100 masl, pitfall, 15–20.vii.2006, G. Fagua ( MPUJ-SCO-343 ) ; one adult male, Puerto Colombia, Loma China , 11°0’58.46’’ N 74°53’0.19’’ W, 100 masl, pitfall, year 2000 (without an exact date), G. Fagua ( MPUJ-SCO-344 ) ; one adult female, Puerto Colombia, Loma China , 11°0’58.46’’ N 74°53’0.19’’ W, 100 masl, pitfall, 15–20.vii.2006, G. Fagua ( MPUJ-SCO-345 ) ; one adult female, Puerto Colombia, Loma China , 11°0’58.46’’ N 74°53’0.19’’ W, 100 masl, pitfall, year 2000 (without an exact date), G. Fagua ( MPUJ-SCO-346 ) ; one adult female and one juvenile male, Usiacurí , 10°44’58.83’’ N 74°59’0.16’’ W, 300 masl, 20.vi.2010, N. Galvis ( ICN-As-853 ) . Bolívar department: one juvenile, San Juan de Nepomuceno, Santuario de Flora y Fauna Los Colorados- La Suiris , 9°57’24’’ N 75°5’12’’ W, 126 masl, 16–31.i.2000, E. Deulofeut ( ICN-As-569 ) ; one adult female, San Juan de Nepomuceno, Santuario de Flora y Fauna Los Colorados , Diana , 9°45’0’’ N 75°7’0’’ W, 150 masl, Malaise, 2.iv.2001, E. Deulofeut ( ICN-As-613 ) ; one juvenile male, Santa Catalina, Hacienda El Ceibal , 10°36’25.20’’ N 75°17’20.12’’ W, 20 masl, x.1999, E. Flórez & Biology students- UN ( ICN-As-293 ) ; one adult female, Zambrano, Hacienda Monterrey , 9°37’48’’ N 74°54’44’’ W, 70 masl, pitfall, 16.viii.1993, F. Fernández ( ICN-As-173 ) ; one juvenile, Zambrano, Hacienda Monterrey , 9°37’48’’ N 74°54’44’’ W, 70 masl, 2.v.1997, F. Fernández ( ICN-As- 174 ) ; one adult female, Zambrano, Hacienda Monterrey , 9°37’48’’ N 74°54’44’’ W, 70 masl, pitfall, vi.1994, F. Fernández ( ICN-As-175 ) ; one adult female, Zambrano, Hacienda Monterrey , 9°37’48’’ N 74°54’44’’ W, 70 masl, pitfall, viii.1996, F. Fernández ( ICN-As-177 ) ; one adult female and one adult male, Zambrano, Hacienda Monterrey , 9°37’48’’ N 74°54’44’’ W, 70 masl, pitfall, 20.viii.1993, F. Fernández ( ICN-As-178 ) ; one adult female, Zambrano, Hacienda Monterrey , 9°37’48’’ N 74°54’44’’ W, 70 masl, Malaise , 2.ii.1994, F. Fernández ( ICN-As-179 ) ; one adult male, Zambrano, Hacienda Monterrey- Bosque el Totumito , 9°37’48’’ N 74°54’44’’ W, 70 masl, pitfall, 5.viii.1993, F. Fernández ( ICN-As-180 ) ; two adult females and two adult males, Zambrano, Hacienda Monterrey , 9°37’48’’ N 74°54’44’’ W, 70 masl, pitfall, 4.xii.1997, F. Fernández ( ICN-As-181 ) ; one adult male, Zambrano, Hacienda Monterrey- Casa Nueva , 9°37’48’’ N 74°54’44’’ W, 70 masl, pitfall, 2.xii.1993, F. Fernández ( ICN-As-182 ) ; one adult male, Zambrano, Hacienda Monterrey , 9°37’48’’ N 74°54’44’’ W, 70 masl, 5.xii.1997, C. Martínez ( ICN-As-183 ) ; one adult female, Zambrano, Hacienda Monterrey , 9°37’48’’ N 74°54’44’’ W, 70 masl, pitfall, 20.viii.1993, F. Fernández ( ICN-As-184 ) ; one adult female, Zambrano, Hacienda Monterrey , 9°37’48’’ N 74°54’44’’ W, 70 masl, pitfall, 12.iv.1994, F. Fernández ( ICN-As-185 ) ; three adult females, Zambrano, Hacienda Monterrey , 9°37’48’’ N 74°54’44’’ W, 70 masl, pitfall, 14.x.1993, F. Fernández ( ICN-As-186 ) ; one juvenile male, Zambrano, Hacienda Monterrey , 9°37’48’’ N 74°54’44’’ W, 70 masl, pitfall, 3.ii.1994, F. Fernández ( ICN-As-187 ) ; one adult female, Zambrano, Hacienda Monterrey , 9°37’48’’ N 74°54’44’’ W, 70 masl, 14.x.1993, F. Fernández ( ICN-As-23 ) ; one adult female, Zambrano, Socorro , 9°45’02.70’’ N 74°49’59.28’’ W, 26 masl, 19.viii.1993, F. Fernández ( ICN-As-416 ) ; one juvenile, Zambrano, 9°45’02.70’’ N 74°49’59.28’’ W, 50 masl, Malaise, 8.v.1993, G. Ulloa & F. Fernández (ICN-As-513) . Caldas department: four adult females and one adult male, La Dorada , 5°29’27.76’’ N 74°39’45.42’’ W, 160 masl, viii.2012, N. Hazzi ( ICN-As-1006 ) . Cesar department: one adult female and two adult males, Valledupar, Ecoparque Los Besotes , 10°34’15.3’’ N 73°16’15.2’’ W, 957 masl, night manual capture with UV light, 17.vii.2015, J. A. Moreno ( MZSP 71735 ) . Córdoba department: one adult female, Ayapel , 8°18’48.81’’ N 75°8’35.87’’ W, 25 masl, 2.iv.2004, O. Castaño ( ICN-As-599 ) ; one adult male, San Antero, Bahía Cispata , 9°23’42.77’’ N 75°46’36.41’’ W, 3 masl, manual capture, 2.ii.2000, G. Ulloa ( ICN-As-304 ) ; one adult male, San Antero, Caño Garzal , 9°22’41’’ N 75°45’22’’ W, 25 masl, collected in Mangrove, ii.2000, G. Ulloa ( ICN-As-332 ) ; four adult females, Montería, Vereda Jaraquiel , Finca Pensylvania , 08°40’15” N 75°55’30’’ W, 18 masl, Dry Tropical Forest, xi.2010, I. Rodiño & E. Bravo ( ICN-As-861 ) . Magdalena department: one adult female, Santa Marta, Parque Nacional Natural Tayrona- Sendero Arrieros , Cabo Pueblito , 11°17’17.45’’ N 74°9’09.62’’ W, 250 masl, pitfall, 20.vii.2001, P. Sánchez ( ICN-As-444 ) ; one adult male and three juvenile males, Santa Marta, Parque Nacional Natural Tayrona , 11°17’17.45’’ N 74°9’09.62’’ W, 0 masl, ii.2009, S. García ( ICN-As-780 ) . Sucre: one adult female, three adult males and one juvenile male, San Marcos , Hacienda Cocodrilla , Vereda La Florida , 8°39’39.99’’ N 75°8’05.03’’ W, 40 masl, 15–20.x.2004, G. Amat & Biology students- UN ( ICN-As-614 ) ; one adult female, Sincelejo, Barrio Bostón , 9°17’56.71’’ N 75°22’57.83’’ W, 203 masl, in a house backyard, 10.xi.2005, E. Hernández ( ICN-As-736 ) . VENEZUELA: Trujillo state: two adult females and two adult males, San Miguel, near the Water Dam AguaViva, 09°32.016’’ N 70°36.837’’ W, 195–270 masl, 27.ix.2008, S. Begó & J. A. Ochoa ( ICN- As-757 ) .

Diagnosis. This species can be readily recognized from other Tityus (Archaeotityus) species by the following combination of character states: Total length: 28.85–43.42 mm (males) and 31.88–42.81 mm (females). General body coloration ( Figure 40 ) dark yellow background moderately covered with dark brown variegated spots; metasomal segment V dark yellow moderately covered with dark brown variegated spots, with a pair of distal VL dark brown spots and with a dorsomedian area without spots ( Figure 33B, D ). Metasomal segments II–IV ( Figures 39B , 40C, D ) without conspicuous distoterminal granules on the DL carinae. Metasomal segment V with the VSM2 macrosetae proximal to the level of the VL2 macrosetae ( Figures 17E , 18E ). Subaculear tubercle large-sized pyramidal, flat, and crest-like with a blunt apex in lateral view ( Figures 19E , 20E ). Ddorsal 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 19E , 20E ). Telson with a well-marked (female) ( Figure 19E ) or feebly marked (male) carinae ( Figure 20E ). Metasomal intercarinal areas densely covered with fine granules ( Figures 39B , 40C, D ). Pectinal tooth counts and movable finger oblique rows of denticles: 14–20/ 13–15 (females) and 15–19/ 12–15 (males) ( Table 2–3). Metasomal segments II–V ( Figures 39B , 40C, D ), female pedipalp ( Figure 9I, J ), and male pedipalp femur and patella ( Figure 10I, J ) short and of a medium-width. Chela ( Figure 42 A–C) and metasomal segment V ( Figure 40C ) incrassate in large males.

Affinities with other Colombian species. Tityus tayrona is most similar to T. bastosi and T. guane , sharing with them the presence of a subaculear tubercle that is large-sized pyramidal, flat, and crest-like with a blunt apex in lateral view ( Figure 19E ). Tityus tayrona can be readily distinguished from both species by the presence of 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 without conspicuous distoterminal granules on the DL carinae ( Figures 39B , 40C, D ). Conversely, in T. bastosi and T. guane , the metasomal segment V has the VSM2 macrosetae slightly distal or at the same level of the VL2 macrosetae on the metasomal segment V ( Figures 17A, C , 18A, C ) and the metasomal segments II–IV have conspicuous distoterminal granules on the DL carinae ( Figures 15A, C , 16A, B ).

Remarks. Rojas-Runjaic & Armas (2007: 63) describes Tityus wayuu Rojas-Runjaic & Armas, 2007 and suggests it is closely related to T. tayrona . According to these authors, both species share similar total lengths, general coloration patterns, and pectinal tooth counts. However, they argue that T. tayrona differs from T. wayuu by the presence of a greater number of movable finger oblique rows of denticles; male metasomal segment V black in color and incrassate, and male chela incrassate with respect to the female chela . Nevertheless, to test the diagnostic morphological characters proposed for Tityus wayuu , we carried out an extensive examination of invididuals of Tityus tayrona from different Colombian populations (type and non-type material).

After comparing the morphology of Tityus tayrona against the original description of Tityus wayuu we could see that both species share several morphological features: (1) coloration pattern of the metasomal segment V with a dark yellow background moderately covered with reddish-brown variegated spots ( Rojas-Runjaic & Armas 2007: figures 6B, 7C; Figure 33B, D ); (2) pectinal tooth counts ( T. wayuu : females= 13–15 and males= 15–16; T. tayrona : females= 14–20 and males= 15–19) ( Rojas-Runjaic & Armas 2007: table 4; Table 2); (3) number of movable finger oblique rows of denticles ( T. wayuu : females and males= 13–14; T. tayrona : females= 13–15 and males 12–15) ( Rojas-Runjaic & Armas 2007: 60; Table 3); and (4) the presence of the VSM2 macrosetae proximal to the level of the VL2 macrosetae on the metasomal segment V ( Rojas-Runjaic & Armas 2007: figure 7C; Figures 17E , 18E ).

Additionally, we detected intrasexual variations related to the total size of Tityus tayrona males. For example, the male holotype of Tityus tayrona (SMF-37019) has an incrassate metasomal segment V and an incrassate pedipalp chela (Total length= 41.03 mm), but some male paratypes (SMF-37022 and SMF-37025) have a non-incrassate metasomal segment V and a non-incrassate pedipalp chela (SMF-37022: Total length= 31.03 mm; SMF-37025 (n= 5): Total length= 31.66–34.59 mm) ( Table 5 ). Identical variations were also observed in males of other non-type material of Tityus tayrona ( Figure 41 C–F, Table 10 ). We also saw that these intrasexual dimorphisms do not apparently follow an isometric growth pattern, thus, there are no morphometric ratios derived from the metasomal segment V and pedipalp chela that can be useful for species level distinction in Tityus tayrona .

As stated by Rojas-Runjaic & Armas (2007) the only available measurements for T. tayrona are those of its male holotype and one female allotype. So, only after the examination of some additional material of T. tayrona we corroborated the intrasexual dimorphism of this species. Finally, after critical examination of all available morphological evidence, we concluded that T. wayuu can no longer be supported as a diagnosable species, based on the character states proposed by Rojas-Runjaic & Armas (2007). For this reason, we formally propose Tityus wayuu Rojas-Runjaic & Armas, 2007 as a synonym of Tityus tayrona Lourenc ̧o, 1991.

Description. Based on an adult female and an adult male (ICN-As-181). For female live habitus see Figure 46F .

Total length. Female 40.47 mm; male 47.55 mm. Measurements: Non-type material ( Table 10 ) and type material ( Table 5 ).

Coloration. General body coloration (in ethanol 70%) ( Figure 41 ) with a dark yellow background moderately covered with dark brown variegated spots. Carapace ( Figure 32C, D ) moderately covered with dark brown varie- gated spots; lateral and median eyes surrounded by black spots; posterior area to the median ocular tubercle with an inverted ‘’Y’’-shaped spot. Chelicerae ( Figure 34C, D ) coxa and hand with a light yellow background; hand with dark brown reticulated spots on the anterior half and areas without spots on the median surface and a small area adjacent to the movable finger articulation; movable finger with dark brown spots on its posterior half, and fixed finger without spots; teeth dark reddish-brown. Pedipalps ( Figure 41 ) 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 41 ) coxae I, III, and IV, sternum, genital operculum, pectines, and basal pectinal piece light yellow; basal pectinal piece with an anterior white area (female); coxaphophyses I–II and coxa II with a light yellow background moderately covered with dark brown variegated spots. Mesosoma ( Figure 41 ) tergites I–VII and sternites III–VI each with a dark yellow background moderately covered with dark brown variegated spots; tergites I–VII each covered with four posterior dark brown spots (two submedian and two lateral); spiracles yellow. Metasoma ( Figure 32C, D ) 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 dark yellow moderately covered with dark brown variegated spots, with a pair of distal VL dark brown spots and with a dorsomedian area without spots. DL intercarinal areas of segments I–IV each with an anterior median arrowshaped brown spot and a posterior median brown spot; VSM intercarinal areas with a distal pair of dark brown spots; VM intercarinal areas without a distal spot. Telson ( Figure 41 ) almost completely dark reddish-brown with a lateral light reddish-brown stripe; subaculear tubercle and aculeus dark reddish-brown.

Morphology. Carapace ( Figure 35C, D ) 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; ocular tubercle well-marked and located on the anterior half of the carapace; median eyes separated by about 1.1 (female) or 1.2 (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 34C, 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= 3.3). 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 9I , 10I ) with five well-marked and complete carinae: VI, DI, and DE crenulate; VE and IM serratocrenulate; intercarinal areas densely covered with fine granules and a few coarse granules. Patella ( Figures 9J , 10J ) 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 42 A–C, 43A–C) with eight carinae: VI, VE, D, DS, DMA, and ES complete and crenulate; DI complete and serratocrenulate (female) ( Figure 42 A–C) or obsolete (male) ( Figure 43 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 43A ) or without a basal lobe (female) ( Figure 42A ); dorsal surface of the movable finger with 13–14 (female) and 13–14 (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 ) subrectangular-shaped with an anaterior elevated glandular area (female) (e.g., Figure 2A ) or flat, shield-shaped with an anteriomedian notch well-marked (male) (e.g., Figure 2B ); pectinal tooth counts: 18–19 (female) and 18–17 (male). Intermediate plate, marginal plate, and fulcra moderately covered with setae ( Figure 38E, G ). Pectinal teeth peg sensillae flattened with a slightly spatulate apex ( Figure 38F, H ).

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 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 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 44 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 44B ); basal carina with rounded apex ( Figure 44C ); basal hook semi ovate-shaped with anterior margin straight in lateral view ( Figure 44B ) and slightly curved in anterior and posterior views; basal hook forming a ‘’U’’-shaped curve with the stem ( Figure 44A, C ); basal hook elongated with a thick and rounded apex and approximately 1.0 longer than its basal width in anterior view ( Figure 44C ).

Metasoma. Segments II–V short and of a medium-width in both sexes (Msl/Msw ratio= female/ male: II= 1.5 /1.7; III=1.7/ 1.9; IV= 1.9/ 2.1; V= 2.3/ 2.0). Segment V short and of a medium-width (female) ( Figure 39B ) or incrassate (male) ( Figure 40C ). Segments I–II ( Figures 39B , 40C ) 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 first two thirds and composed of coarse granules in the anterior third; intercarinal areas densely covered with fine granules. Segment III–IV ( Figures 39B , 40C ), with eight complete, parallel, and crenulate carinae (paired DL, ML, VL and VSM; DL carinae serratocrenulate); intercarinal areas densely covered with fine granules ( Figures 39B , 40C ). Segment V with five complete and crenulate carinae (VM, paired DL and VL); lateral and ventral intercarinal densely covered with fine granules (female) ( Figure 39B ) or obsolete (male) ( Figure 40C ); intercarinal areas with granules irregularly distributed on the middle of each area. Segment I–IV with DL carinae slightly serratocrenulate, but without conspicuous distoterminal granules ( Figures 39B , 40C ).

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 ( Figure 1 ). Segment V ( Figures 17E , 18E ) 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, and pair VSM2 is located proximal to the level of the VL2 ( Figures 17E , 18E ), and pair VSM 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= 2.1 (female) ( Figure 19E ) and Vl/Vh= 2.3 (male) ( Figure 20E )) 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 19E ) or feebly marked (male) ( Figure 20E ); 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 19E , 20E ). Subaculear tubercle large-sized pyramidal, flat, and crest-like with a blunt apex ( Figures 19E , 20E ); 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 19E , 20E ); aculeus strongly curved shorter than vesicle and with a ventral groove.

Variability. Total length (including telson ): males 28.85–43.42 mm (n= 17; mean= 33.78; SD= 4.32); females 31.88–42.81 mm (n= 24; mean= 37.63; SD= 2.73) ( Tables 5 , 10 ). Pectinal tooth counts: males 15–19 (n= 40; mode= 18); females 14–20 (n= 68; mode= 18) ( Table 2). Number of movable finger oblique rows of denticles: males 12–15 (n= 37; mode= 14); females 13–15 (n= 68; mode= 14) ( Table 3). Metasomal macrosetae counts: segments I–V: males (n= 26) and females (n= 40): 2/2 VSM and 2/2 VL macrosetae on each segment. Relative position of the VSM2 macrosetae on the metasomal segment V: males (n= 26) and females (n= 40): proximal to VL2 (n= 7).

Hemispermatophore capsule: the basal hook of the capsule exhibited different shapes in the anterior view: (1) elongated with a rounded apex, curved lateral margin, and forming a ‘’U’’-shaped curve with the stem ( Figure 44C ); (2) elongated with an acute apex, curved lateral margin, and forming a ‘’U’’-shaped curve with the stem ( Figure 44D ); and (3) elongated with a rounded apex, strongly curved lateral margin, and forming a ‘’C’’-shaped curve with the stem ( Figure 44E ).

Male intrasexual variations: adult males exhibited different body sizes ranging between 28.85–43.42 mm, but despite this they did not show discrete size-classes but rather a progressive increase in their total lengths ( Figure 41 C–F, Tables 5 , 10 ). However, as the total body size increases among males the following: (1) Chela : the manus is more incrassate, without apparently exhibiting isometric growth (n= 22; Cl/Cw= 2.8–5.8; Cl/Ch= 3.1–4.4) ( Figure 43 ) and (2) Metasomal segment V: the segment is more incrassate without apparently exhibiting isometric growth (n= 22; Msl-V/Msw-V= 1.9–2.4; Msl-V/Msh-V= 1.6–2.4) ( Figure 40C, D ) and the granules associated with carinae and intercarinal areas are less marked ( Figure 40C, D ).

Distribution ( Figure 45 ). COLOMBIA. Atlántico department: Barranquilla and Puerto Colombia. Bolívar department: Santa Catalina, Turbaco, and Zambrano. Cesar department: Chimichagua. Córdoba department: San Antero. Magdalena department: Santa Marta (PNN Tayrona) and Santa Marta (Bonda). Sucre department: Colosó (Serranía de Coraza) ( Figure 46F ). PANAMA. Panama province: Panama city. New records. COLOMBIA: Antioquia department: Santa Fé de Antioquia. Atlántico department: Usiacurí. Bolívar department: San Juan de Nepomuceno (S.F.F. Los Colorados). Caldas department: La Dorada. Cesar department: Valledupar. Córdoba department: Ayapel and Montería. VENEZUELA: Trujillo state: San Miguel. Zulia state: Mara.

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 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 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 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 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 42. Tityus (Archaeotityus) tayrona Lourenço, 1991, female pedipalp chela (ICN-As-181).A. External aspect. B. Dorsal aspect. C. Ventral aspect. Scale bar= 2.0 mm.

FIGURE 43. Tityus (Archaeotityus) tayrona Lourenço, 1991, male pedipalp chela. A–C. Male (Zambrano, Bolívar) (ICN-As- 181). A. External aspect. B. Dorsal aspect. C. Ventral aspect. D–F. Male (Zambrano, Bolívar) (ICN-As-181). D. External aspect. E. Dorsal aspect. F. Ventral aspect. Scale bars= 2.0 mm.

FIGURE 44. Tityus (Archaeotityus) tayrona Lourenço, 1991, dextral hemispermatophore, showing capsule details (Zambrano, Bolívar) (ICN-As-181). A. Posterior view. B. Lateral view. C. Anterior view. D, E. Capsule variations, anterior view. D. Male (La Dorada, Caldas) (ICN-As-1006). E. Male (Valledupar, César) (MZUSP 71735). Abbreviations: Bc, basal carina; Bh, basal hook; Dc, distal carina; Sdm, sperm duct membrane. Scale bar= 1.0 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.

Tables

TABLE 5. Measurements (mm) of the type material of Tityus tayrona Lourenço, 1991.

Structure/ Measure		Male Holotype SMF-37019	Female Paratype SMF-37026	Male Paratype SMF-37022	Female Paratype SMF-37022	Female Paratype SMF-37025	Male Paratype SMF-37025	Male Paratype SMF-37025	Male Paratype SMF-37025	Male Paratype SMF-37025	Female Paratype SMF-37025	Male Paratype SMF-37025
Total length		41.03	40.96	31.03	36.96	37.29	31.66	34.49	32.14	34.59	33.01	32.13
Carapace	length	4.56	4.64	3.36	4.80	4.40	3.60	4.00	3.60	3.84	4.00	3.60
	anterior width	2.96	3.04	2.32	3.12	2.96	2.32	2.56	2.32	2.48	2.64	2.32
	posterior width	4.72	5.20	3.60	5.76	4.80	3.52	4.24	3.76	4.16	4.32	3.92
	eye diameter	0.42	0.39	0.36	0.39	0.36	0.30	0.36	0.30	0.33	0.36	0.30
	interocular distance	0.52	0.48	0.40	0.44	0.52	0.36	0.40	0.36	0.40	0.40	0.36
	ocular diada width	1.15	1.05	0.90	1.10	1.05	0.90	0.90	0.85	0.90	0.95	0.85
Tergite I	length	0.80	0.88	0.56	0.76	0.80	0.56	0.60	0.60	0.64	0.56	0.64
Tergite II	length	1.00	1.24	0.68	0.88	1.00	0.68	0.68	0.72	0.88	0.68	0.76
Tergite III	length	1.24	1.44	0.92	1.24	1.28	0.88	0.88	1.00	1.04	0.92	1.04
Tergite IV	length	1.48	1.72	1.04	1.52	1.44	1.12	1.08	1.20	1.28	1.12	1.20
Tergite V	length	1.76	1.92	1.24	1.68	1.64	1.36	1.20	1.28	1.40	1.32	1.32
Tergite VI	length	1.92	2.08	1.40	1.80	1.80	1.48	1.48	1.40	1.52	1.80	1.48
Tergite VII	length	2.88	2.96	2.32	2.40	3.00	2.40	2.20	2.20	2.44	2.28	2.32
Mesosoma	total length (tergites)	11.08	12.24	8.16	10.28	10.96	8.48	8.12	8.40	9.20	8.68	8.76
Metasoma I	length	2.96	2.76	2.12	2.40	2.60	2.20	2.48	2.28	2.48	2.24	2.28
	width	2.52	2.32	1.72	1.68	2.16	1.84	1.88	1.80	1.92	1.96	1.76
	height	2.52	2.52	1.84	1.88	2.40	1.96	2.20	1.92	2.08	2.24	2.00
Metasoma II	length	3.64	3.40	2.68	3.00	3.24	2.80	3.12	2.80	3.08	2.88	2.76
	width	2.28	2.28	1.88	1.72	2.16	1.84	1.96	1.76	1.96	1.92	1.76
	height	2.28	2.32	1.80	1.72	2.24	1.80	2.04	1.88	1.96	2.00	1.84
Metasoma III	length	3.68	3.92	3.08	3.40	3.44	3.20	3.72	3.16	3.44	3.28	3.20
	width	2.48	2.32	1.84	1.72	2.16	1.84	2.00	1.88	1.96	1.92	1.80
	height	2.40	2.36	1.80	1.76	2.20	1.80	2.08	1.88	2.00	2.00	1.88
Metasoma IV	length	4.96	4.60	3.68	4.08	3.60	3.68	4.20	3.80	4.20	3.88	3.68
	width	2.56	2.28	1.88	1.72	2.16	1.80	2.04	1.88	2.04	1.96	1.80
	height	2.52	2.40	1.84	1.72	2.24	1.88	2.12	1.88	2.04	2.00	1.88
Metasoma V	length	5.70	5.20	4.45	5.05	5.10	4.35	5.00	4.45	4.65	4.50	4.40