Tityus smithii Pocock, 1893

Teruel, Rolando, 2011, Redescription of Tityus pictus Pocock, 1893 and Tityus smithii Pocock, 1893, with notes on the Tityus species from the Lesser Antilles (Scorpiones: Buthidae), Euscorpius 125 (125), pp. 1-26 : 8-24

Tityus smithii Pocock, 1893 , stat. rest.

Figs. 4–15 , Tabs. 3–7

Tityus smithii Pocock, 1893: 377 , 383–384, 409, pl. XXX, figs. 9–9a; Lourenço, 1984: 91–92, 97–102, 104, figs. 20–27; tab. I; Kovařík, 1998: 122 (in synonymy).

Tityus pictus smithii : Kraepelin, 1899: 75, 86; Mello-Leitão, 1931: 126, 143; Mello-Leitão, 1939: 60–61, 65, 74, tab. IIc; Mello-Leitão, 1945: 310; Lourenço, 1984: 97–99, figs. 20–27, tab. I; Armas, 1988: 81, 93; Fet & Lowe, 2000: 256; Lourenço, 2006: 60.

Tityus smithii microdon Pocock, 1893: 384 ; Armas, 1988: 81. New synonym.

Tityus pictus microdon : Lourenço, 1984: 92, 97, 99; Fet & Lowe, 2000: 256; Lourenço, 2006: 60.

Tityus pictus : Mello-Leitão, 1931: 126, 143 (misidentification); Armas, 1982: 6, tab. 2 (misidentification: records from Grenada and the Grenadines); Lourenço, 1984: 97–99, figs. 20–27, tab. I (misidentification: specimens and records from Grenada and the Grenadines); Armas, 1988: 81, 93 (misidentification: specimens and records from Grenada and the Grenadines); Lourenço, 1992: 45, fig. 1 (misidentification: records from Grenada and the Grenadines); Lourenço, 1995: 28, fig. 1 (misidentification: records from Grenada and the Grenadines); Kovařík, 1998: 122 (misidentification); Fet & Lowe, 2000: 256 (misidentification).

Diagnosis: species of moderately large size (males 58–83 mm, females 57–72 mm) for the genus. Body yellowish brown, with a moderately dense pattern of dark brown spots; tergites with three poorly to well defined, irregular dark stripes; pedipalp fingers, metasomal segments IV–V and telson blackish. Pedipalp chela inflate and globular in larger males, incrassate in smaller males and subtriangular in females. Sternite V with a large smooth patch in males. Metasoma slightly attenuate and conspicuously enlarged distally in males, with all carinae weak to moderate and coarsely granulose to crenulate; dorsolateral carinae of metasomal segments II–IV with distal tooth not enlarged. Telson not inflate and coarsely granulose; subaculear tubercle obsolete to small. Pedipalp fixed finger with 13–14 principal rows of granules, movable finger with 14–15; basal lobe/notch combination very strong in males, moderate in females. Pectines with 19–22 teeth in males, and 19–21 in females; basal middle lamella obtusely angulose and slightly dilated in males, pyriform and strongly dilated in females.

Type data: 3 adult ♂♂ and 3 adult ♀♀ syntypes ( BMNH; not examined): Grenada; R. V. Sherring coll. Note: see Remarks for a detailed discussion on this topic .

Distribution ( Fig. 14 ): this species is widely distributed over the Lesser Antilles south of St. Vincent, where it has been collected at the islands of Bequia, Mustique, Union and Grenada. It probably occurs also in most (if not all) of the remaining Grenadines and satellite islets of Grenada.

Redescription (large adult male from Union; see also high-resolution color photos of an adult male topotype in Figs. 4–5 ): Coloration ( Figs. 6–7 , 13b–c ) basically yellowish brown, irregularly spotted with medium to dark brown all over the body and appendages; spots much lighter (pale to light brown) on pedipalps, metasomal segments I–III and venter. Chelicerae densely reticulated with blackish brown on distal half of manus only; fingers blackish. Pedipalps very faintly spotted with pale brown; fingers blackish. Carapace with interocular area irregularly infuscate; tergites with three well-defined dark stripes, composed of irregular spots; venter essentially immaculate; pectines light yellow, immaculate. Legs irregularly spotted with medium brown. Metasomal segments I–III very faintly spotted with pale brown on all surfaces; segment IV with dorsal surface reddish-yellow, lateral and ventral surfaces blackish; segment V and telson blackish. Carapace ( Fig. 7a ) trapezoidal, anterior margin very widely V-shaped; anterior median, lateral ocular, central median and posterior median carinae coarsely granulose (the latter fused into irregular rows), superciliary carinae very coarsely subgranulose, other carinae obsolete to absent; tegument finely and densely granulose, with coarser granules scattered; median eyes separated by about one ocular diameter; three pairs of lateral eyes, which are all much smaller than median eyes. Tergites ( Fig. 6a ) with the same granular sculpture as on carapace; longitudinal carina well defined and coarsely granulose; VII with two pairs of coarsely serrate lateral carinae. Chelicerae ( Fig. 7a ) with dentition typical for the genus; tegument smooth and shiny. Pedipalps ( Figs. 6a–b , 7b ) orthobothriotaxic A-α. Femur with all carinae coarsely granulose; intercarinal tegument densely granulose, with many coarser granules scattered. Patella with all carinae coarsely granulose to costate; intercarinal tegument with the same granular sculpture as on femur, internal surface with a few conical granules. Chela inflate and globular, much wider than patella ( Tabs. 3–5); hand with nine carinae, all moderate and finely granulose to subgranulose, intercarinal tegument with the same granular sculpture as on femur and patella; fingers with basal lobe/notch combination very strong, fixed finger with 14/14 principal rows of granules, movable finger with 15/15, apical subrow composed by four granules aligned similar to principal rows. Legs ( Figs. 6a–b ) with all carinae serrate to granulose; intercarinal tegument finely and densely granulose. Sternum ( Fig. 7c ) type 1, subtriangular. Pectines ( Fig. 7c ) somewhat small, just reaching the coxa-trochanter joint of leg IV; pectinal tooth count 21/21; basal middle lamella obtusely angulose and vestigially dilated. Sternites ( Figs. 7c–d ) with slit-like spiracles; sternite III with the lateral areas slightly depressed and finely granulose, mesially coriaceous, IV–VI coriaceous, VII finely and densely granulose; posterior margin of sternite V bilobed, with a large and smooth patch, which is yellowish, subtriangular, much wider than long and moderately bulky; sternites VI–VII with two pairs of granulose lateral carinae. Metasoma ( Figs. 7e–g ) slightly elongate and markedly enlarged distally; intercarinal tegument very fine and densely granulose, with coarser granules scattered; segment I with ten complete carinae, II–IV with eight (even though the lateral inframedian carinae are present on distal half of II), V with five, all moderately developed and coarsely granulose to crenulate; dorsolateral carinae on II–IV with the distal tooth vestigially enlarged; telson oval in lateral view, vesicle not inflate and sparsely granulose, with a subgranulose ventromedian carina progressively elevated towards the subaculear tubercle, which is very small, conical, placed adjacent to the base of aculeus and lacks any granules; aculeus long, sharp and evenly curved.

Female (Union Island, Figs. 10–11 , 13d ; Tabs. 3–7): in general is similar to the male, but there is a strong sexual dimorphism evidenced by: (1) mesosoma relatively wider; (2) metasoma shorter and stouter; (3) pedipalp chela much smaller, obtusely subtriangular in shape and narrower to slightly wider than patella; (4) pedipalp fingers with basal lobe/notch combination weaker; (5) genital papillae absent; (6) pectines with basal middle lamella pyriform and strongly dilated; (7) sternite V with smooth patch slightly smaller and less bulky; (8) metasomal segments IV–V and telson dark reddish-brown.

Variation: the adult size of T. smithii varies from 58.0– 82.7 mm in males, and 57.3–71.8 mm in females ( Tab. 3). Pocock (1893) recorded 62.5 mm for a male and 59–63 mm for two females, and Lourenço (1984) measured two females of 57.3 and 69.3 mm. The topotype male from FKPC matches Pocock's syntype in size, but in the sample from Union the total length is 58. 0–82.7 mm in males and 68.6–71.8 in females. These data suggest that there are two different size-classes in each sex.

The degree of expression of sexual dimorphism is positively correlated to the size in both sexes ( Figs. 4–11 , 13 ; Tabs. 3–4): the maximum robustness of pedipalp chela, metasomal segments and finger basal lobe/notch combination invariably corresponds to the largest males and females, and vice versa; the dimorphism is wellevident in all size-classes. This progressive size-related gradation on the expression of dimorphism has been documented also for T. pictus (see above), as well as many other species of this genus (Lourenço, 1983; Armas et al., 2002; Montoya & Armas, 2002; Kovařík, 2007; Teruel & Armas, 2006; Rojas-Runjaic & Armas, 2007; Teruel, 2000, 2011; Teruel & García, 2008a –b; Teruel & Sánchez, 2009, 2010; Teruel & Kovařík, 2011).

The coloration of the adults shows slight variations in the examined sample. In most specimens from Union, the three dark stripes of the mesosoma are completely separated by light areas in a perfectly defined pattern ( Figs. 6a , 10a , 13b ); nevertheless, in the male from Grenada and one female from Union the dark spots are interconnected by reticulations, and the three-striped pattern is diffuse ( Figs. 4a , 13a ). Also, two males and one female from Union have the dark pattern all over the body and appendages much fainter ( Fig. 8–9a ) than the remaining specimens.

The posterior margin of sternite V does show important variations among specimens from Union: it exhibits the complete gradation from essentially straight to deeply bilobed ( Figs. 6b , 7d , 10b , 11d ).

Similar variation is present in the size of the subaculear tubercle among the adults of both sexes: it varies from obsolete (a few specimens from Union) through small (most specimens, including the topotype male).

The variation of pectinal tooth counts known for T. smithii is compiled in Table 6. Pocock (1893) recorded 20–21 for males and 18–21 for females, and Lourenço (1984) recorded 19–21 teeth for three females. Among the additional specimens herein studied, the pectinal tooth counts varied from 19–22 in males and 18–21 in females; no differences were detected among different insular populations.

♂♂ ♀♀

Ratio T. pictus T. smithii T. pictus T. smithii 1. Chela (L/W) 4.14–4.41 3.19–3.45 4.48 3.66–4.38 2. Hand (W) / Patela (W) 1.16–1.26 1.46–1.56 0.96 0.96–1.12 3. Movable finger (L) / Hand (W) 2.41–2.59 1.83–2.03 2.96 2.34–2.77 4. Metasoma (L) / Carapace (L) 7.00–7.63 6.63–6.95 5.36 6.00–6.09 5. Metasomal segment I (L/W) 1.75–2.28 1.45–1.52 1.37 1.21–1.31 6. Metasomal segment II (L/W) 2.56–2.92 1.77–1.88 1.81 1.59–1.67 7. Metasomal segment III (L/W) 2.74–3.25 1.94–2.06 1.96 1.67–1.71 8. Metasomal segment IV (L/W) 2.57–3.08 1.77–1.97 2.04 1.71–1.87 9. Metasomal segment V (L/W) 2.73–2.96 1.74–1.84 2.35 1.86–1.88 10. Telson (L) / Aculeus (L) 2.69–3.00 2.08–2.09 3.09 2.00–2.54

The variation of the number of principal rows of granules on pedipalp fingers known for T. smithii is compiled in Table 7. Pocock (1893) did not present any data in the original description, and Lourenço (1984) recorded 15 rows for the movable finger of three females. Among the additional specimens herein studied, the number of principal rows of granules showed an interesting sex-related bias: all males have 14/14 on fixed finger and 15/15 on movable finger, but among females count varied from 13–14 on fixed finger and 14–15 on movable finger. Similar examples of sex-related variation in this character have been already documented for other species of the genus (Teruel & Armas, 2006; Teruel & García, 2008b; Teruel & Sánchez, 2010).

Juveniles ( Fig. 12 ) are very similar to adults, but can be distinguished by the somewhat different coloration (basically paler, with metasomal segments IV–V and telson of the same color as the rest of metasoma), a less sclerotized cuticle (especially in metasoma and pedipalps), the pedipalp chelae relatively smaller and narrower than patella, metasoma relatively more slender and parallel-sided, and a totally different subaculear tubercle (much stronger, sharp and with two dorsal granules).

Ecological notes: according to the information kindly provided by one of the collectors (MJR), in Union Island this species is common and shows some ecological plasticity: it lives under rocks and rotten logs, as well as inside termite nests in dry coastal forest ( Figs. 13 , 15 ), syntopically with Tityus atriventer Pocock, 1897 ; one female was observed carrying a litter of about 15 larvae on her back ( Fig. 13d ).

Material examined: GRENADA (no other data), 1 adult ♂ topotype ( FKPC) . SAINT VINCENT AND THE GRENADINES, Parish of the Grenadines, Union Island (new record), Chatham Bay Trail , 12°35.800 N 61°26.700 W, 31–23 4 m above sea level, 4–22 June 2010, M. J. Rivera & E. Bentz leg., 1 adult ♂ ( IES), 3 adult ♂♂, 2 adult ♀♀, 3 juvenile ♂♂, 2 juvenile ♀♀ (Sco-0472) .

Remarks: for a detailed comparison between this species and T. smithii , see General Remarks.

The status and composition of the name-bearing types of T. smithii is currently confused. Pocock (1893) did not designate a holotype, but declared to have based the original description on three adult males and three adult females from Grenada; two adult males and two adult females from the Grenadines (Mustique and Bequia) were also mentioned, but described “... as [...] a variety [...] called microdon...” [italics herein added].

Lourenço (1984) examined two adult females from Grenada deposited at BMNH (catalogue number 1894.10.20. 7–16), in page 99 he affirmed that both specimens belonged “... to the type-series ...” [original text in French, translation and italics herein added], and in pages 101–102 he listed them first as “ types ” [italics herein added], and then as “ lectotype ” and “ paratype ” [italics herein added] .

On the other hand, Fet & Lowe (20 00: 255–25 6) overlooked this apparently valid designation by Lourenço (1984), and listed as syntypes all specimens from Grenada and Grenadines originally mentioned by Pocock (1893).

Nevertheless, such lectotype designation by Lourenço (1984) is not valid according to the Article 72.4.1.1 of the Code (CINZ, 2000: 84), because there is enough evidence that it was not based upon an actual syntype of T. smithii . First of all, Pocock (1893) explicitly mentioned R. V. Sherring as the collector of the types, but the two females studied by Lourenço (19 84) were reportedly collected by H. H. Smith. Second, according to the numbering system applied by the BMNH the sample studied by Lourenço (1 984) was originally composed of 10 specimens (this is what the range “7–16” means in the catalogue number), but the original description was based only in six specimens as explicitly stated by Pocock (1 893). And third, in the original description of Tityus atriventer, Pocock (1897: 519) himself declared that its types (coincidently also from Grenada and collected by H. H. Smith) were “... received too late for notice in my paper upon the West Indian Scorpions ...”; this implies that the sample collected by H. H. Smith at Grenada was available to him only after the description of T. smithii was either submitted or published, as well as the fact that the types of T. atriventer and the specimens of T. smithii studied by Lourenço (1984) bear BMNH catalogue numbers (1894.10.20.29–30 and 1894.10.20.7–16, respectively) which indicate that all of them came from a single original sample which was later taxonomically divided. Thus, this case exactly fulfills the Article 74.2 of the Code (CINZ, 2000: 85), which renders invalid any designation of lectotype which has been demonstrated to be based upon a non-syntype specimen; this applies also to the wrong designation by Lourenço (1984: 101) of these two females first as syntypes, and then as lectotype and paralectotype of T. pictus (see above, in the Remarks section of this species).

It is highly recommended here that any other author who eventually decides to designate a lectotype, must select the adult male from Grenada measured and illustrated by Pocock (1893: 383–384; pl. XXX, fig. 9; the size match between the table and figure demonstrate that both refer to the same specimen about 62.5 mm long), following the Recommendation 74B of the Code that gives preference to a syntype which has been illustrated (CINZ, 2000: 86). It also concurs with the fact that all diagnostic characters of T. smithii are best expressed in this sex.

Apart from this, Fet & Lowe (2000: 255–256) wrongly listed as syntypes of T. smithii the four specimens from the Grenadines originally mentioned by Pocock (1893). According to the Article 72.4.1 of the Code (CINZ, 2000: 78), these specimens are not syntypes because Pocock (1893) explicitly excluded them all from the type-series by assigning them to the new "variety" he named microdon (see below).

Pocock (1893) described the subspecies T. smithii microdon on the basis of two couples of adults from Mustique and Bequia, and it was diagnosed by three characters: (1) subaculear tubercle obsolete; (2) pedipalp chelae more robust; (3) pedipalp movable finger with the distal third paler. However, the sample from Union herein examined (11 specimens) renders this distinction invalid, as it contains some specimens which match the definition of each subspecies exactly, together with others that have the same characters either mixed or present in intermediate states (i.e., the width of pedipalp chela is directly correlated to the size of the specimens, but the size of the subaculear tubercle and the color pattern do vary randomly; see Variation above). This is further supported by the comparison of this sample to a series of high-resolution color photos of an adult male from Grenada (kindly provided by František Kovařík), which revealed that this topotype is morphologically identical to a same-sized male from Union. Last, in view of the geographical distribution of these populations (with the morphologically intermediate specimens located coincidently at the center of the area; see Fig. 14 ), it is evident that the recognition of subspecies is not justified. Thus, the present nomenclatural act is proposed: Tityus smithii Pocock, 1893 = Tityus smithii microdon Pocock, 1893 , new synonym.

General Remarks

The specific distinction between T. pictus and T. smithii is a very interesting case, because both species are identical in two important characters which are usually used as diagnostic at this taxonomic level (pectinal tooth count and number of principal rows of granules on pedipalp fingers), but they are in turn very easy to distinguish from each other even to unaided eye on the basis of other characters. The most important of these features is the sexual dimorphism of the adults, which show a completely opposite trend in both taxa: the metasomal segments and pedipalp chelae of the males are much more elongate and slender than those of females in T. pictus , but much more robust in T. smithii ; such different is so obvious that it was already highlighted by Pocock (1893) in the original description of both species. Lourenço (1984) observed this dissimilitude, but incorrectly dismissed it as “ a case of polytypism [...] of interbreeding insular populations ” [original text in French, translation and italics herein added], obviously because he could not examine any adult males of T. smithii (which otherwise have remained either undiscovered or unavailable apart from the syntypes).

The additional samples herein studied have confirmed that Pocock (1893) was correct in most points of his original distinction between T. pictus and T. smithii . With the addition of further characters, adults of both taxa can be distinguished by: (a) a very different shape of pedipalps and metasoma, most remarkably in males; (b) a very different shape of telson, with the vesicle bulbous and the aculeus very short in T. pictus , but with the vesicle flattened and the aculeus remarkably longer in T. smithii ; (c) differences in pedipalpal and metasomal carination, with carinae composed of fine granules in T. pictus , but coarse granules in T. smithii ; (d) size conspicuously larger in T. smithii . Pocock (1893) also addressed slight differences in coloration, but the variation herein observed renders this character useless; the only exception is the base color, which is more yellowish in T. pictus and more brownish in T. pictus . All the above mentioned differences can be clearly seen in the figures and tables of this paper.

Their geographical distribution also shows an interesting pattern: T. pictus appears to be endemic from the island of St. Vincent, while T. smithii is widely distributed across Grenada and Grenadines. Despite the finding of the latter at Bequia (just 8 km south of St. Vincent), both species have never been collected together in the same island.

On the other hand, the present contribution sets the number of Tityus known from the Lesser Antilles to five. It is worth to make here some comments on the remaining three species:

1. Tityus atriventer Pocock, 1897 : originally described from Grenada, it was recently redescribed and recorded from the Grenadines ( Teruel & Kovařík, 2011). It has been long recognized as a member of the “ clathratus ” species-group (Pocock, 1897; Mello-Leitão, 1939; Lourenço, 1984, 2006; Fet & Lowe, 2000; Teruel & Kovařík, 2011).

2. Tityus exstinctus Lourenço, 1995 : described from a single adult male, collected in 1884 in Martinique. Lourenço (1995: 29–31) stated that its closest relative was Tityus trinitatis Pocock, 1897 (an endemic species from Trinidad & Tobago), but this association is clearly wrong in view of the deep differences shown by both species in almost all taxonomically important characters (i.e., T. trinitatis is a member of the “ androcottoides ” species-group which is characterized by the partial to complete fusion of the ventrosubmedian carinae of the metasoma, but in T. exstinctus these carinae are not fused, which is the condition present in all other species-groups of the genus); ironically, this incongruence was even illustrated and mentioned on the same paper (Lourenço, 1995: 29–31; figs. 8–9). The original description and figures of Lourenço (1995) clearly demonstrate that its closest relatives are in fact T. pictus and T. smithii , as T. exstinctus matches both species perfectly in three key characters (pedipalp movable finger with 14 principal rows of granules, pectines with 22 teeth, and telson with subaculear tubercle minute and aculeus shorter than vesicle), apart from coloration (yellowish brown with a pattern of dark spots, pedipalp fingers, metasomal segments IV–V and telson darker) and general shape of the pedipalps and metasoma.

3. Tityus insignis (Pocock, 1889) : it is endemic from St. Lucia and its satellite islets ( Pocock, 1893; Lourenço, 1984; Lourenço & Cloudsley-Thompson, 1999). It was associated by Pocock (1889, 1893) to T. americanus , which is a Buthidae incertae sedis according to Fet & Lowe (2000). Mello-Leitão (1939) placed it first in his “Group C” (which was actually a “catch-all” where most species of the genus were placed despite how dissimilar they could be), and then in the “ asthenes ” group (Mello-Leitão (1945); this second placement was implicitly followed by Lourenço (1984), who definitely later (Lourenço, 2006) placed it in the subgenus Tityus (Atreus) . Nevertheless, such association is incorrect: its closest relatives clearly are T. exstinctus , T. pictus and T. smithii , on the basis of the general shape and sculpture of the pedipalps and metasoma, and especially the shape of the telson (with subaculear tubercle minute to obsolete, and aculeus shorter than vesicle; see Pocock [1893: fig. 1a], Lourenço [1984: figs. 1, 10–11, 13], and Figs. 16a–c herein). Even the few characters which could disassociate T. insignis from these three species are not so important when examined in more detail: (a) its coloration is much darker to blackish, but in lighter specimens the typical spotted pattern of T. exstinctus , T. pictus and T. smithii is still traceable ( Fig. 16a ); (b) the higher numbers of pectinal teeth (21–25) and principal rows of granules on pedipalp movable finger (17) are congruent with the much larger size of this species (90–110 mm), a trend that has already been documented in other species-groups of this genus (Teruel & Armas, 2006; Rojas-Runjaic & Armas, 2007).

With the present contribution, the taxonomy of the Lesser Antilles members of Tityus has changed. Previously considered as a heterogeneous assemblage (Lourenço, 1984; Armas, 1988), it is herein demonstrated to be a much more homogeneous composite of five species which belong to only two well-defined lineages: ( T. atriventer ) and ( T. exstinctus + T. insignis + T. pictus + T. smithii ).

The first species belong to the “ clathratus ” species-group, which is widespread and well-diversified across South-America and adjacent Central America ( Costa Rica and Panama). It appears to be a recent immigrant into the Antilles, on the basis of its geographical distribution: T. atriventer is its only member present in (and endemic from) the Lesser Antilles ( Teruel & Kovařík, 2011), while about 20 other species occur in the continental part of its range, including Trinidad & Tobago (Lourenço, 2006, 2008; Rojas-Runjaic & Armas, 200 7; Teruel & Roncallo, 2010).

The remaining four species form a morphologically compact group, but it has been recognized as such only in this paper because in a similar way to T. exstinctus and T. insignis , the relationships of both T. pictus and T. smithii to the other members of the genus have been largely controversial. Pocock (1893) originally associated them to Tityus melanostictus Pocock, 1893 , which is currently placed in its own species-group (Fet & Lowe, 2000). Mello-Leitão (1939) placed them in his “Group C” (already commented on above), and finally, Fet & Lowe (2000) listed them in the “ clathratus ” group, a placement implicitly retained by Lourenço (2006) as the subgenus Tityus (Archaeotityus) . These four species ( T. exstinctus , T. insignis , T. pictus and T. smithii ) are much more closely related to each other than to any other member the genus, and also cannot be satisfactorily placed either in any of the species-groups of Fet & Lowe (2000), nor in the subgenera of Lourenço (2006). The only potential matches are first the “ crassimanus ” group as defined by Teruel & Armas (2006), and second the “ quisqueyanus ” group as defined by Armas & Abud (2004), but this relationship still needs to be investigated in detail.

Figures

Figure 4: Adult male topotype of Tityus smithii: a. entire dorsal view; b. entire ventral view. Photos courtesy František Kovařík.

Figure 5: Adult maletopotype of Tityus smithii: a. carapace; b. pedipalp, dorsal view; c. venter; d. metasoma, lateral view. Photos courtesy František Kovařík.

Figure 6: Large adult male of Tityus smithii from Union: a. entire dorsal view; b. entire ventral view.

Figure 7: Large adult male of Tityus smithii from Union: a. carapace; b. pedipalp, dorsal view; c. sternopectinal region; d. sternite V; e. metasomal segments I–III, lateral view; f. metasomal segments IV–V and telson, lateral view; g. telson, lateral close-up.

Figure 8: Small adult male of Tityus smithii from Union: a. entire dorsal view; b. entire ventral view.

Figure 9: Small adult male of Tityus smithii from Union: (a) pedipalp, dorsal view; (b) metasomal segments IV–V and telson, lateral view; (c) telson, lateral close-up.

Figure 10: Adult female of Tityus smithii from Union: a. entire dorsal view; b. entire ventral view.

Figure 11: Adult female ofTityus smithii from Union:a. carapace;b. pedipalp, dorsal view; c. sternopectinal region;d. sternite V; e. metasomal segments I–III, lateral view; f. metasomal segments IV–V and telson, lateral view; g. telson, lateral close-up.

Figure 12: Juvenile female of Tityus smithii from Union: (a) entire dorsal view; (b) metasomal segments III–V and telson, lateral view.

Figure 13: Live individuals of Tityus smithii, in their natural habitat: a. large adult male topotype; b–c. large adult male from Union;d. adult female from Union, with litter. Photos courtesy František Kovaříka., Alejandro J. Sánchez b–c.and Mel J. Rivera d.

Figure 14: Geographical distribution of Tityus pictus (white squares) and Tityus smithii (black squares).

Figure 15: Habitat of Tityus smithii at Chatham Bay Trail, Union: a. general view from seashore; b. closer view from hilltop. Photos courtesy Mel J. Rivera.

Figure 16: Adult female topotype of Tityus insignis (RMNH): a. entire dorsal view; b. entire ventral view; c. metasoma, lateral view. Photos courtesy František Kovařík.