Taxonomic revision of the Colombian Tityus (Archaeotityus) (Scorpiones, Buthidae) species: a morphological and morphometric approach, with a description of a new species Author Moreno-González, Jairo A. Author O, Ranulfo González Author D, Eduardo Flórez text Zootaxa 2019 2019-08-26 4660 1 1 94 journal article 25919 10.11646/zootaxa.4660.1.1 2334efd2-e346-48ff-85ff-0bc8661538b2 1175-5326 3377134 DA70C158-7FB0-4B46-9A1D-EA32D24B19B6 Tityus guane sp. nov. Figures 2 , 5E, F , 6C, F , 7E, F , 8E, F , 9E, F , 10E, F , 14 G–J, 15C, 17C, 18C, 19C, 20C, 24, 25A–C, 26, 27, 28, 30, 46C, 47E, 48, 49; Tables 1–4 , 6–8 . http://zoobank.org/urn:lsid:zoobank.org:pub:urn:lsid:zoobank.org:act:DCB7088E-F939-44A0-945F- 3993772 E9E20 Tityus columbianus : Flórez, 2001b: 28 (misidentification); Teruel & García, 2008: 10 (misidentification). Type material. Holotype : COLOMBIA : Santander department : one adult male , Los Santos , Vereda Pasochico , Finca La Ceiba , 6°47'4.19'' N 73°8'31.25'' W , 1174 masl , night manual capture with UV light , 14–16.viii.2016 , J. A. Moreno ( ICN-As-1003 ) . Paratypes : COLOMBIA : Santander department : one adult female , same data as holotype ( ICN-As-1003 ) ; three adult males and five adult females , same data as holotype ( MZSP 71736 ) ; two adult males and three adult females , same data as holotype ( ICN-As-1003 ) ; two adult males and three adult females , Los Santos , Vereda La Mojarra , near Chicamocha’s canyon, 6°46.83’N 73°5.2’W , 18.ix.2008 , J. Ochoa & S. García ( ICN-As-735 ) ; three males and one female , Zapatoca , Vereda La Cacica , path to Natural Reserve Montaña Mágica , 6°48’48.94’’ N 73°16’29.81’’ W , 1704 masl , night manual capture with UV light , under rocks and leaf litter , 7.xi.2013 , E. Flórez , C. Perafán & D. Martínez ( ICN-As-1050 ) ; two adult males and one adult female , San Gil , 6°33’31.02’’ N 73°8’06.8’’ W , 1163 masl , diurnal manual capture under rocks , i.2004 , A. Otálora ( ICN-As-571 ) ; one adult male , San Gil , Barrio Altamira , 6°33’31.02’’ N 73°8’06.08’’ W , 1163 masl , inside a house , 13.iv.2001 , A. Otálora ( ICN-As-436 ) ; one adult male , San Gil , Vereda Resumidero , Path to Mogotes , 6°31’02.70’’ N 73°2’46.40’’ W , 1620 masl , A. García ( ICN-As-1048 ) . Etymology. This species is named after the Guane people (members of the Chibcha language family). They were a Pre-Columbian native tribe with considerable social advances that lived in what is now known as Santander department . They were exterminated during the Spanish colonization. The name is treated as a noun in apposition. Diagnosis. This species can be readily recognized from other Tityus ( Archaeotityus ) species by the following combination of character states: Total length: 27.63–38.28 mm (males) and 28.34–38.55 mm (females). General body coloration ( Figure 24 ) dark yellow background moderately covered with dark brown variegated spots; metasomal segment V completely dark reddish-brown with a few ventral yellow spots and a dorsomedian area without spots ( Figure 6C, F ). Metasomal segments II–III ( Figures 15C , 28 ) each has a distoterminal small acute granule on the DL carinae. Metasomal segment IV ( Figures 15C , 28 ) with a distoterminal small 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 17C , 18C ). Subaculear tubercle large-sized, pyramidal, flat, and crest-like with a blunt apex in lateral view ( Figures 19C , 20C ); dorsal margin of the subaculear tubercle with a pair of ventrodistal small rounded granules with apices pointing to the basal portion of the aculeus; vesicle and subaculear tubercle with VM carina composed of fine granules and feebly crenulate in lateral view ( Figures 19C , 20C ). Telson with well-marked (female) ( Figure 19C ) or feebly marked (male) carinae ( Figure 20C ). Metasomal intercarinal areas densely covered with coarse granules and a few fine granules ( Figures 15C , 28 ). Pectinal tooth counts and movable finger oblique rows of denticles: 15–18/ 13–14 (females) and 14–19/ 13–14 (males) ( Table 2 , 3 ). Metasomal segments II–V ( Figures 15C , 28 ), female pedipalp ( Figures 25 A–C, 26A–F), and male pedipalp femur and patella ( Figures 9E, F , 10E, F ) short and of a medium-width. Chela ( Figure 26 A–C) and metasomal segment V ( Figure 28A ) incrassate in large males. Affinities with other Colombian species. Tityus guane sp. nov. is most similar to T. bastosi , sharing with it the presence of a subaculear tubercle that is large-sized pyramidal, flat, and crest-like with a blunt apex in lateral view ( Figures 19C , 20C ); metasomal segments II–IV each with a conspicuous distoterminal granule on the DL carinae ( Figures 15C , 28 ), and metasomal segment V with the VSM2 macrosetae slightly distal or at the same level of the VL2 macrosetae ( Figures 17C , 18C ). Tityus guane can be readily distinguished by the presence of a general body coloration with a dark yellow background, moderately covered with dark brown variegated spots ( Figure 24 ); metasomal segment V dark reddish-brown with a few ventral yellow spots and a dorsomedian area without spots ( Figure 6C, F ); metasomal segments II–III each with a distoterminal small and acute granule on the DL carinae ( Figures 15C , 28 ), and metasomal segment IV with a small coarse and blunt granule on the DL carinae ( Figures 15C , 28 ). Conversely, in T. bastosi the general body coloration is with dark yellow background densely covered with dark reddish-brown spots ( Figure 4 ); the metasomal segment V is completely black ( Figure 6A, D ), and the metasomal segments II–IV each has a medium-sized acute distoterminal granule on the DL carinae ( Figures 15A , 16A, B ). Remarks. Teruel & García (2008) stated that the record of Tityus columbianus from Santander department presented by Flórez (2001b) appears to be feasible. Nevertheless, after an examination of this material from Santander department , we concluded that these populations correspond to Tityus guane sp. nov. Description. Based on an adult male ( holotype ) (ICN-As-1003) and an adult female ( paratype ) (ICN-As-1003). For male live habitus see Figure 46C . Total length. Male ( holotype ): 30.43 mm ; female ( paratype ): 36.29 mm . Measurements: Table 4 . Coloration. General body coloration (in ethanol 70%) ( Figure 24 ) with a dark yellow background moderately covered with dark brown variegated spots. Carapace ( Figure 5E, F ) 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 7E, F ) coxa and hand with light yellow background; hand with dark brown reticulated spots on the anterior half and areas without spots on the internal surface, median surface, and the adjacent area to the movable finger articulation; movable and fixed fingers with dark brown spots on their posterior halves; teeth dark reddish-brown. Pedipalps ( Figure 24 ) 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 24B, D ) coxae I–IV, sternum, genital operculum, pectines (middle lamellae, fulcra, and teeth), and basal pectinal piece light yellow; basal pectinal piece with an anterior white area (female); coxaphophyses I–II and pectines (marginal lamellae) with a light yellow background moderately covered with dark brown variegated spots. Mesosoma ( Figure 24 ) tergites I–VII and sternites III–VII each with a dark yellow background moderately covered with dark brown variegated spots; tergites I–VII with three posterior dark brown spots (two submedian and two lateral; spiracles yellow. Metasoma ( Figure 6C, F ) 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 a dorsomedian area without spots and a dorsomedian area without spots ( Figure 6C, F ). DL intercarinal areas of segments I–IV each with an anterior median arrow-shaped brown spot and a posterior median brown spot; ML and VSM intercarinal areas of segments I–IV each with a pair of distal brown spots. Telson ( Figure 24 ) almost completely dark reddish-brown with a lateral light reddish-brown stripe; subaculear tubercle and aculeus dark reddish-brown. Morphology. Carapace ( Figure 8E, F ) 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 1.3 (female paratype ) or 1.2 (male holotype ) 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 7E, F ) 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= 4.3; male: Cl/Cw=4.1). 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 9E , 10E ) 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 ( Figures 9E , 10E ). Patella ( Figures 9F , 10F ) with seven carinae: VI, VE, DI, DE, and EM complete and crenulate; DM incomplete and crenulate; IM complete and serratocrenulate with a spur near to the segment base; intercarinal areas densely covered with fine granules. Chela ( Tibia ) ( Figures 25 A–C, 26D–F) with eight carinae: VI, VE, D, DS, DMA, and ES complete and crenulate; DI complete and feebly crenulate ( Figures 25 A–C, 26D–F); SA incomplete and crenulate, only present on the anterior half of the hand. Pedipalp movable and fixed finger with an obsolete basal lobe ( Figures 25A , 26D ); dorsal surface of the movable finger with 14–14 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 longitudinal divided and composed of two subtriangular plates. Pectines . Basal piece sexually dimorphic ( Figure 2 ) sub rectangular-shaped with an anterior elevated glandular area (female paratype ) ( Figure 2A ) or flat, shield-shaped with an anterior median notch well-marked (male holotype ) ( Figure 2B ); pectinal tooth counts: 15–16 (female paratype ) and 18–17 (male holotype ). Intermediate plate, marginal plate, and fulcra densely covered with setae ( Figure 14G, I ). Pectinal teeth peg sensillae flattened with a slightly spatulate apex ( Figure 14H, J ). 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 fine granules and with a few coarse 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 the 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 occupying the anterior half of the sternite. Hemispermatophore. General morphological characteristics as in other Tityus ( Archaeotityus ) species. Capsule ( Figure 27 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 27B, C ); basal carina with rounded apex ( Figure 27C ); basal hook semi- ovate-shaped with anterior margin straight in lateral view ( Figure 27B ) and slightly curved in anterior and posterior views; basal hook forming a ‘’U’’-shaped with the stem ( Figure 27A, C ); basal hook short with an acute apex and approximately as long as its basal width in anterior view ( Figure 27C ). FIGURE 24. Tityus (Archaeotityus) guane sp. nov. , habitus. A, B. Male holotype (ICN-As-1003). A. Dorsal aspect. B. Ventral aspect. C, D. Female paratype (ICN-As-1003). C. Dorsal aspect. B. Ventral aspect. Scale bars= 10 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 26. Tityus (Archaeotityus) guane sp. nov. , male pedipalp chela . A–C. Male paratype (Vereda La Mojarra, Los Santos, Santander) (ICN-As-735). A. External aspect. B. Dorsal aspect. C. Ventral aspect. D–F. Male holotype (Vereda Pasochico, Los Santos, Santander) (ICN-As-1003). D. External aspect. E. Dorsal aspect. F. Ventral aspect. Scale bars= 2.0 mm. FIGURE 27. Tityus (Archaeotityus) guane sp. nov. , dextral hemispermatophore, showing capsule details, male (Los Santos, Santander) (MZSP 71736). A. Posterior view. B. Lateral view. C. Anterior view. D. Capsule variation, anterior view, male (Los Santos, Santander) (MZSP 71736). Abbreviations : Bc , basal carina; Bh , basal hook; Dc , distal carina; Sdm , sperm duct membrane. Scale bar= 1.0 mm. Metasoma. Segments II–V short and of a medium-width in both sexes (Msl/Msw ratio= female/ male: II= 1.2/ 1.4; III= 1.4/ 1.6; IV= 1.6/ 1.9; V= 1.9/ 2); segment V short and of a medium-width (male holotype and female paratype ) ( Figures 15C , 28B ). Segments I–II ( Figures 15C , 28B ) 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 (male holotype ) or coarse granules (female paratype ) almost along all its length; intercarinal areas densely covered with coarse granules. Segment III–IV ( Figures 15C , 28B ) 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 densely covered with coarse granules and a few fine granules ( Figures 15C , 28B ); 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 distally towards the distal region of each segment; DL carinae ending in a distoterminal granule acute and small granule which increases in size towards the segment IV ( Figures 15C , 28B ). Metasomal macrosetation. Segments I–IV each with two pairs of VSM macrosetae (2/2) and two pairs of VL macrosetae (2/2); pair VSM1 is located on the anterior third, and pair VSM2 is located near the posterior margin of the segment; pair VL1 is located near the anterior margin of the segment, and pair VL2 is located on the posterior third of the segment (e.g., Figure 1 ). Segment V ( Figures 17C , 18C ) with three pairs of VSM macrosetae (3/3), three pairs of VL macrosetae (3/3), and a single pair of DL macrosetae; pair VSM1 is located near the anterior margin of the segment; pair VSM2 is located slightly distal or at same level of pair VL2 ( Figures 17C , 18C ), 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. FIGURE 28. Tityus (Archaeotityus) guane sp. nov. , male metasomal segments I–V sculpturing, lateral aspect. A. Male paratype (Vereda La Mojarra, Los Santos, Santander) (ICN-As-735). B. Male holotype (Vereda Paso Chico, Los Santos, Santander) (ICN-As-1003). Scale bars= 2.0 mm. TABLE 4 . Measurements (mm) of Tityus guane sp. nov.

Structure/ Measure	Male Holotype ICN-As- 1003 #14	Female Paratype ICN-As- 1003 #13	Female Paratype ICN-As- 1003 #1	Male Paratype ICN-As- 1003 #2	Female Paratype ICN-As- 1003 #3	Male Paratype MZSP 71736 #4	Female Paratype ICN-As- 1003 #5	Male Paratype MZSP 71736 #6	Female Paratype MZSP 71736 #7	Male Paratype MZSP 71736 #8
Total length	30.43	36.29	33.14	28.37	35.44	28.69	34.52	27.63	35.72	-
Carapace	length	3.52	4.32	4.00	3.28	4.16	3.36	4.16	3.12	4.32	3.84
anterior width	2.32	2.88	2.80	2.24	2.88	2.16	2.80	2.08	2.80	2.56
posterior width	3.60	4.80	4.56	3.52	4.72	3.44	4.72	3.36	4.72	4.00
eye diameter	0.30	0.33	0.27	0.30	0.36	0.30	0.30	0.30	0.33	0.30
interocular distance	0.36	0.44	0.44	0.36	0.48	0.36	0.44	0.36	0.44	0.40
ocular diada width	0.80	0.90	0.90	0.75	0.90	0.70	0.90	0.70	0.90	0.85
Tergite I	length	0.60	0.76	0.68	0.56	0.68	0.56	0.72	0.48	0.72	0.68
Tergite II	length	0.72	1.00	0.88	0.72	1.00	0.68	0.96	0.68	1.00	0.84
Tergite III	length	0.92	1.24	1.16	0.92	1.24	0.88	1.20	0.88	1.28	1.04
Tergite IV	length	1.12	1.52	1.40	1.08	1.48	1.08	1.44	1.08	1.44	1.28
Tergite V	length	1.24	1.68	1.48	1.24	1.68	1.12	1.60	1.20	1.52	1.40
Tergite VI	length	1.40	1.80	1.60	1.32	1.88	1.24	1.72	1.28	1.76	1.48
Tergite VII	length	2.08	2.64	2.52	2.08	2.68	2.00	2.48	2.00	2.64	2.40
Mesosoma	total length (tergites)	8.08	10.64	9.72	7.92	10.64	7.56	10.12	7.60	10.36	9.12
Metasoma I	length	2.16	2.36	2.12	1.92	2.28	1.96	2.20	1.88	2.40	2.40
width	1.68	2.20	2.04	1.64	2.08	1.64	2.12	1.64	2.40	1.92
height	1.96	2.40	2.12	1.80	2.40	1.88	2.40	1.80	2.60	2.16
Metasoma II length		2.56	2.84	2.60	2.36	2.80	2.40	2.72	2.28	2.88	2.96
width	1.64	2.20	1.92	1.64	2.08	1.60	2.08	1.60	2.36	1.96
height	1.80	2.24	2.08	1.68	2.20	1.80	2.24	1.64	2.40	2.00
Metasoma III	length	3.00	3.32	3.00	2.68	3.16	2.80	3.12	2.64	3.32	3.32
width	1.72	2.28	2.00	1.64	2.08	1.64	2.20	1.64	2.36	2.00
height	1.80	2.28	2.12	1.68	2.20	1.80	2.28	1.68	2.40	2.00

...Continued on the next page TABLE 4. (Continued)

Structure/ Measure	Male Holotype ICN-As- 1003 #14	Female Paratype ICN-As- 1003 #13	Female Paratype ICN-As- 1003 #1	Male Paratype ICN-As- 1003 #2	Female Paratype ICN-As- 1003 #3	Male Paratype MZSP 71736 #4	Female Paratype ICN-As- 1003 #5	Male Paratype MZSP 71736 #6	Female Paratype MZSP 71736 #7	Male Paratype MZSP 71736 #8
Metasoma	length	3.56	3.96	3.60	3.16	3.80	3.36	3.80	3.16	3.84	3.96
IV
width	1.76	2.28	2.00	1.72	2.24	1.76	2.20	1.64	2.40	2.00
height	1.84	2.40	2.16	1.72	2.24	1.84	2.36	1.68	2.44	2.08
Metasoma V length		4.15	4.70	4.20	3.80	4.55	3.95	4.45	3.75	4.50	4.65
width	2.08	2.32	2.12	1.92	2.24	1.80	2.32	1.84	2.48	2.44
height	2.00	2.44	2.28	1.84	2.32	2.00	2.44	1.80	2.52	2.24
Telson	vesicle length	2.50	3.00	2.85	2.40	2.90	2.40	2.90	2.35	2.95	2.90
vesicle width	1.15	1.50	1.30	1.15	1.45	1.05	1.40	1.05	1.45	1.35
vesicle height	1.15	1.50	1.35	1.10	1.45	1.05	1.40	1.05	1.45	1.35
aculeus length	1.45	1.90	1.60	1.40	1.75	1.40	1.75	1.45	1.70	-
total length	3.40	4.15	3.90	3.25	4.05	3.30	3.95	3.20	4.10	-
Metasoma	total length	15.43	17.18	15.52	13.92	16.59	14.47	16.29	13.71	16.94	17.29
Femur	length	3.32	3.90	3.71	3.12	3.90	3.12	3.90	2.99	3.84	3.77
width	1.11	1.37	1.24	1.04	1.37	1.04	1.30	0.98	1.30	1.17
Patella	length	3.90	4.62	4.36	3.58	4.62	3.64	4.49	3.51	4.49	4.29
width	1.50	1.95	1.76	1.43	1.89	1.37	1.95	1.37	1.89	1.69
Chela	length	6.16	7.60	6.96	5.92	7.28	5.84	7.20	5.68	7.28	7.28
width	1.50	1.75	1.55	1.40	1.70	1.30	1.75	1.25	1.65	1.75
height	1.50	1.70	1.60	1.45	1.70	1.40	1.70	1.35	1.65	1.85
length movable finger	3.75	5.10	4.60	3.75	4.75	3.65	4.70	3.70	4.40	4.55
length fixed finger	3.20	4.40	4.10	3.30	4.25	3.25	4.20	3.30	3.85	3.95
length palm	2.45	2.70	2.50	2.30	2.60	2.30	2.65	2.15	2.65	2.85

TABLE 4 (Continuation). Measurements (mm) of Tityus guane sp. nov.

Structure/ Measure	Female Paratype MZSP 71736 #9	Female Paratype MZSP 71736 #10	Female Paratype MZSP 71736 #11	Female Paratype MZSP 71736 #12	Female Paratype ICN-As-571 #1	Male Paratype ICN-As-571 #2	Male Paratype ICN-As- 571 #3	Male Paratype ICN-As-436	Male Paratype ICN-As- 1048
Total length	31.58	28.34	36.02	33.73	38.20	33.73	32.30	33.71	32.77
Carapace	length	3.84	3.52	4.40	4.08	4.88	3.60	3.52	3.68	3.52
anterior width	2.48	2.32	2.80	2.56	3.28	2.32	2.40	2.40	2.32
posterior width	4.16	3.76	4.72	4.48	5.60	4.00	3.92	4.08	3.92
eye diameter	0.30	0.27	0.30	0.30	0.32	0.26	0.32	0.34	0.32
interocular distance	0.40	0.36	0.44	0.44	0.45	0.36	0.36	0.36	0.33
ocular diada width	0.85	0.75	0.95	0.85	0.96	0.75	0.72	0.81	0.78
Tergite I	length	0.60	0.56	0.68	0.72	0.84	0.60	0.64	0.64	0.60
Tergite II	length	0.80	0.76	0.96	0.92	1.00	0.72	0.76	0.80	0.80
Tergite III	length	1.00	0.92	1.28	1.16	1.28	1.12	0.92	1.00	0.96
Tergite IV	length	1.40	1.16	1.56	1.44	1.60	1.32	1.16	1.20	1.24
Tergite V	length	1.56	1.28	1.68	1.52	1.60	1.56	1.28	1.40	1.36
Tergite VI	length	1.60	1.36	1.76	1.68	1.88	1.60	1.36	1.48	1.48
Tergite VII	length	2.32	2.08	2.64	2.40	2.84	2.40	2.12	2.40	2.32
Mesosoma	total length (tergites)	9.28	8.12	10.56	9.84	11.04	9.32	8.24	8.92	8.76
Metasoma I	length	2.12	1.92	2.36	2.16	2.52	2.20	2.16	2.24	2.24
width	1.80	1.76	2.16	2.00	2.10	1.75	1.75	1.75	1.80
height	2.00	2.00	2.40	2.28	2.35	1.75	1.85	1.85	1.90
Metasoma II	length	2.48	2.32	2.96	2.72	2.84	2.72	2.72	2.80	2.72
width	1.84	1.76	2.16	2.08	2.05	1.75	1.75	1.80	1.85
height	1.88	1.80	2.20	2.08	2.20	1.80	1.75	1.95	1.80

...Continued on the next page TABLE 4. (Continued)

Structure/ Measure	Female Paratype MZSP 71736 #9	Female Paratype MZSP 71736 #10	Female Paratype MZSP 71736 #11	Female Paratype MZSP 71736 #12	Female Paratype ICN-As-571 #1	Male Paratype ICN-As-571 #2	Male Paratype ICN-As-571 #3	Male Paratype ICN-As-436	Male Paratype ICN-As- 1048
Metasoma III	length	2.92	2.68	3.32	3.04	3.52	3.08	3.16	3.20	3.12
width	1.88	1.76	2.24	2.08	2.10	1.80	1.75	1.85	1.80
height	1.92	1.80	2.24	2.08	2.20	1.80	1.80	1.85	1.90
Metasoma IV	length	3.44	3.08	3.92	3.64	4.20	3.76	3.60	3.72	3.76
width	1.88	1.80	2.28	2.08	2.10	1.80	1.95	2.00	1.95
height	1.92	1.84	2.28	2.12	2.25	1.85	1.80	1.95	1.85
Metasoma V	length	4.15	3.60	4.55	4.35	4.90	5.55	5.30	5.55	4.90
width	1.88	1.80	1.84	2.24	2.30	2.00	2.05	2.25	2.10
height	2.00	1.84	2.40	2.24	2.30	1.95	1.95	2.25	2.00
Telson	vesicle length	2.50	2.30	2.95	2.75	3.25	2.75	2.70	2.75	2.80
vesicle width	1.15	1.15	1.45	1.35	1.55	1.20	1.25	1.25	1.25
vesicle height	1.25	1.10	1.30	1.35	1.55	1.25	1.25	1.30	1.20
aculeus length	1.55	1.35	1.70	1.75	1.75	1.45	1.45	1.45	1.50
total length	3.35	3.10	3.95	3.90	4.30	3.50	3.60	3.60	3.75
Metasoma	total length	15.11	13.60	17.11	15.91	17.98	17.31	16.94	17.51	16.74
Femur	length	3.51	3.12	3.90	3.64	4.20	3.35	3.30	3.45	3.35
width	1.17	1.11	1.30	1.24	1.40	1.05	1.05	1.10	1.10
Patella	length	4.23	3.71	4.68	4.29	4.75	3.65	3.75	3.90	3.70
width	1.63	1.50	1.82	1.76	1.90	1.45	1.45	1.55	1.50
Chela	length	6.64	5.68	7.44	7.04	7.92	6.32	6.48	6.64	6.40
width	1.45	1.30	1.60	1.50	1.75	1.45	1.55	1.70	1.40
height	1.45	1.30	1.60	1.60	1.80	1.55	1.65	1.80	1.55
length movable finger	4.45	3.80	4.85	4.75	5.46	4.16	4.23	4.29	4.23
length fixed finger	3.90	3.40	4.25	4.10	4.81	3.64	3.58	3.64	3.64
length palm	2.35	2.05	2.65	2.45	2.90	2.40	2.65	2.65	2.55

TABLE 4 (Continuation). Measurements (mm) of Tityus guane sp. nov.

Structure/ Measure	Male Paratype ICN-As-1050 #1	Female Paratype ICN-As- 1050 #2	Male Paratype ICN-As-1050 #3	Male Paratype ICN-As- 1050 #4	Male Paratype ICN-As-735 #1	Female Paratype ICN-As- 735 #2	Female Paratype ICN-As- 735 #3	Male Paratype ICN-As-735 #4	Female Paratype ICN-As-735 #5
Total length	38.28	33.42	-	-	37.17	38.55	36.11	35.05	36.51
Carapace	length	4.32	4.08	3.50	3.50	4.08	4.56	4.32	3.84	4.24
anterior width	2.80	2.64	2.25	2.30	2.72	2.96	2.80	2.48	2.80
posterior width	4.80	4.48	3.90	3.90	4.24	4.96	4.64	4.00	4.72
eye diameter	0.34	0.30	0.26	0.30	0.36	0.36	0.36	0.33	0.33
interocular distance	0.36	0.36	0.40	0.40	0.40	0.48	0.48	0.40	0.44
ocular diada width	0.87	0.78	0.80	0.85	0.90	0.95	0.95	0.85	0.90
Tergite I	length	0.76	0.72	0.60	0.65	0.64	0.80	0.80	0.68	0.72
Tergite II	length	0.88	0.96	0.75	0.75	0.92	1.04	1.00	0.88	0.96
Tergite III	length	1.24	1.16	1.00	1.00	1.12	1.60	1.32	1.08	1.28
Tergite IV	length	1.44	1.36	1.20	1.20	1.40	1.84	1.44	1.40	1.48
Tergite V	length	1.48	1.52	1.35	1.35	1.52	1.84	1.68	1.48	1.68
Tergite VI	length	1.72	1.60	1.45	1.40	1.56	1.92	1.76	1.60	1.80
Tergite VII	length	2.68	2.48	2.35	2.35	2.68	2.84	2.60	2.48	2.64
Mesosoma	total length (tergites)	10.20	9.80	8.70	8.70	9.84	11.88	10.60	9.60	10.56
Metasoma I	length	2.60	2.24	2.10	2.15	2.64	2.48	2.36	2.48	2.40
width	2.15	2.00	1.65	1.75	2.04	2.20	2.12	1.84	2.12
height	2.20	2.05	1.70	1.75	2.24	2.40	2.32	2.08	2.28
Metasoma II	length	3.28	2.68	2.65	2.65	3.16	2.92	2.88	3.00	2.96
width	2.05	1.95	1.65	1.75	2.00	2.16	2.04	1.88	2.04
height	2.15	2.00	1.70	1.75	2.08	2.24	2.16	1.92	2.08
Metasoma III	length	3.88	3.08	3.35	2.95	3.68	3.44	3.28	3.44	3.40
width	2.10	2.00	1.70	1.75	2.04	2.24	2.12	1.96	2.16
height	2.20	2.05	1.75	1.85	2.12	2.28	2.20	1.96	2.08

...Continued on the next page TABLE 4. (Continued)

Structure/ Measure	Male Paratype ICN-As-1050 #1	Female Paratype ICN-As-1050 #2	Male Paratype ICN-As-1050 #3	Male Paratype ICN-As- 1050 #4	Male Paratype ICN-As-735 #1	Female Paratype ICN-As- 735 #2	Female Paratype ICN-As- 735 #3	Male Paratype ICN-As-735 #4	Female Paratype ICN-As-735 #5
Metasoma IV	length	4.40	3.64	3.55	3.45	4.32	4.12	3.92	4.04	4.00
width	2.20	2.00	1.75	1.75	2.16	2.24	2.24	2.08	2.16
height	2.30	2.00	1.80	1.80	2.24	2.36	2.28	2.00	2.16
Metasoma V	length	5.10	4.15	4.05	3.95	5.00	4.70	4.60	4.60	4.75
width	3.05	2.00	2.20	2.20	2.72	2.36	2.40	2.56	2.28
height	2.75	2.10	2.05	1.95	2.48	2.48	2.52	2.32	2.36
Telson	vesicle length	3.35	2.90	2.70	2.55	3.30	3.20	3.10	2.90	2.95
vesicle width	1.45	1.35	1.35	1.35	1.45	1.50	1.45	1.30	1.45
vesicle height	1.55	1.45	1.20	1.10	1.50	1.50	1.55	1.20	1.50
aculeus length	1.80	1.50	-	-	1.85	2.00	1.70	1.70	1.85
total length	4.50	3.75	-	-	4.45	4.45	4.15	4.05	4.20
Metasoma	total length	19.26	15.79	15.70	15.15	18.80	17.66	17.04	17.56	17.51
Femur	length	4.20	3.70	3.35	3.25	4.23	4.16	3.97	3.77	4.16
width	1.30	1.25	0.99	0.99	1.37	1.43	1.37	1.24	1.37
Patella	length	4.65	4.00	3.75	3.70	4.88	5.01	4.68	4.36	4.88
width	1.85	1.75	1.42	1.52	1.89	1.95	1.89	1.76	1.82
Chela	length	8.16	6.32	6.40	6.20	7.68	7.92	7.36	6.96	7.44
width	2.50	1.50	1.55	1.60	2.05	1.75	1.65	1.85	1.55
height	2.70	1.55	1.70	1.65	2.20	1.75	1.60	1.95	1.70
length movable finger	5.27	4.94	4.00	3.90	4.75	5.30	5.00	4.35	5.25
length fixed finger	4.42	4.29	3.50	3.40	4.10	4.60	4.35	3.75	4.55
length palm	3.40	2.70	2.70	2.50	3.25	2.80	2.60	2.85	2.50

Telson . Vesicle not elongated and suboval (Vl/Vh= 2.2 (male holotype ) ( Figure 20C ) and Vl/Vh= 2.0 (female paratype ) ( Figure 19C )) 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 19C ) or feebly marked (male) ( Figure 20C ); paired DL carinae located dorsally with respect to the lateral longitudinal furrow; VM carina well-marked along its length ( Figures 19C , 20C ). Subaculear tubercle large-sized pyramidal, flat, and crestlike with a blunt apex ( Figures 19C , 20C ); 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 19C , 20C ); aculeus strongly curved shorter than vesicle and with a ventral groove. Variability. Total length ( including telson ): males 27.63–38.28 mm (n= 11; mean= 32.56; SD= 3.53); females 28.34–38.55 mm (n= 14; mean= 34.71; SD= 2.67) ( Table 4 ). Pectinal tooth counts : males 14–19 (n= 29; mode= 17); females 15–18 (n= 28; mode= 16) ( Table 2 ). Number of movable finger oblique rows of denticles : males 13–14 (n= 28; mode= 14); females 13–13 (n= 27; mode= 14) ( Table 3 ). Metasomal macrosetae counts : segments I–IV: males (n= 15) and females (n= 14): 2/2 VSM and 2/2 VL macrosetae on each segment; segment V: males (n= 15) and females (n= 14): 3/3 VSM; 3/3 VL macrosetae, and 1/1 DL macrosetae on the segment. Relative position of the VSM2 macrosetae on the metasomal segment V : males (n= 14): slightly proximal to VL2 (n= 5) or at the same level of VL2 (n= 9); females (n= 14): slightly proximal to VL2 (n= 2) or at the same level of VL2 (n= 12). Male intrasexual variations : adult males exhibited different body sizes ranging between 27.63–38.28, but despite this they did not show discrete size-classes but rather a progressive increase in their total lengths ( Table 4 ). However, as the total body size increases among males the following: (1) Carapace and tergites : the number of coarse granules on intercarinal surfaces increases; (2) Chela : the manus is more incrassate without apparently exhibiting isometric growth (n= 13; Cl/Cw= 3.3–4.6; Cl/Ch= 3.0–4.2) ( Figure 26 A–F); (3) Metasomal segments II–IV : the DL carinae exhibit coarser distoterminal granules ( Figure 28 ); and (4) Metasomal segment V : segment is more incrassate without apparently exhibiting isometric growth (n= 13; Msl-V/Msw-V= 1.9–2.8; Msl-V/Msh-V= 1.7–2.8) ( Figure 28 ), and the granules associated with carinae and intercarinal areas are less marked ( Figure 28 ), Distribution ( Figure 30 ). COLOMBIA : Santander department : Los Santos, San Gil, and Zapatoca.