Tityopsis Armas, 1974

Teruel, Rolando & Rodríguez-Cabrera, Tomás M., 2020, Revision of the genus Tityopsis Armas 1974 (Scorpiones Buthidae) Part 1 General updates and description of four new species, Euscorpius 304, pp. 1-40 : 6-7

Genus Tityopsis Armas, 1974

( Figures 1–40 , Tables 1–4)

Tityus (Tityopsis) Armas, 1974: 8 ; Armas, 1982: 11, 13; Francke, 1985: 16, 19.

Tityopsis: Armas, 1984a: 29 ; tab. 3; Francke, 1985: 14; Armas, 1988: 10–12, 14, 17, 62, 71, 91, 95, figs. 2h, 21; Sissom, 1990: 94, 102; Lourenço & Vachon, 1996: 33; fig. 2; Kovařík, 1998: 119; Fet & Lowe, 2000: 58, 227–228; Armas, 2001: 247–248, fig. 1 (in part, records from Cuba only); Armas, 2006: 1, 8; Vidal-Acosta & Francke, 2009: 333; Teruel & Kovařík, 2012: 143–151, figs. 7, 30, 42– 43, 49, 306–322, 570–577; Francke et al., 2014: 220–224.

Tityus: Moreno, 1940a: 161–164 ; pls. 22–23, figs. 1–5 (misidentification); Moreno, 1940b: 108–110, pls. XXXVI–XXXVII (misidentification); Mello-Leitão, 1945: 300–309 (misidentification, references to T. inespectatus [sic]); Jaume, 1954: 1092 (misidentification, references to T. inexpectatus ); Armas, 1973: 7, 17–18; tab. 5 (misidentification, references to T. inexpectatus ).

Nec Tityopsis: Armas & Martín Frías, 1998: 45–49 (misidentification); Armas, 2001: 247–248, fig. 1 (misidentification, record from Mexico).

TYPE SPECIES. Tityus inexpectatus Moreno, 1940 [currently Tityopsis inexpectata (Moreno, 1940) ], by original designation ( Armas, 1974: 8).

DIAGNOSIS (emended). Adult size small for the family (15– 30 mm in males, 20–40 mm in females), with males slightly smaller than females within the same size-class. Coloration light yellowish to blackish, immaculate to densely patterned all over, especially on carapace, legs and metasoma. Entire body and appendages covered with modified macrosetae: brightly UV-fluorescent, short to medium-sized, thick, translucent, with tip truncate to crown-shaped. Cheliceral fixed finger with one ventral denticle, movable finger with two. Pedipalp very similar in both sexes: not especially robust nor attenuate and strongly carinate; trichobothrial pattern A-α orthobothriotaxic, without clearly defined petite trichobothria (e.g., chela Eb 3 and Esb and fixed finger esb are somewhat smaller, but not as disproportionately reduced as a typical petite); femur with d 1 -d 3 -d 4 -d 5 regularly spaced along basal half, e 1 and e 2 located on subbasal and median position, respectively; patella with d 3 external to dorsal median carina and i slightly distal to midpoint; manus with V 2 clearly closer to ventroexternal carina than V 1; fixed finger with est-db-et-dt-it located on distal half, with it midway between dt and fingertip. Pedipalp chelae robust, wider than patella and strongly carinate; fixed and movable fingers with 11–13 principal rows of denticles which are short, oblique, subequal and flanked by very large internal and external accessory denticles, apical subrow oblique and composed of four denticles, basal lobe/notch combination absent. Carapace trapezoidal, essentially flat and with carinae distinct but variably fused and poorly defined from intercarinal granulation, which is variable but always well developed; 3–6 pairs of lateral eyes, not concealed below anterolateral margin. Legs without tibial spurs, both pedal spurs present; telotarsi ventrally with two well-defined, parallel, longitudinal rows of thin spiniform setae. Male genital papillae lip-like, not protruding from genital operculum and each with a distinct fleshy point, hardly visible by being located extremely basal and almost entirely concealed beneath pre-pectinal plate. Pre-pectinal plate well developed, heavily sclerotized and medially invaginate, not well visible by being largely concealed beneath genital operculum. Pectines with 11–16 teeth in males, 10–15 in females, fulcra well developed; basal middle lamella modified, slightly to moderately enlarged and angulose in males, moderately to remarkably enlarged and oval to round in females; basal plate unmodified, but in females with a large, whitish discal area of presumably glandular function. Tergites heavily granulose, monocarinate, with median carina and coarse granulation projecting over posterior margin. Sternites with small, round to short slitlike spiracles; III without stridulatory organ or deep furrows but with a raised, granulose, median triangular area flanked by two lateral depressions that fit pectines for protection or rest, V–VII with paired submedian and lateral keels, V with three smooth patches (one median and two laterals, all conspicuously larger, bulkier and also usually paler in males). Metasoma very similar in both sexes: short, robust and very strongly carinate, segment V with lateral median carinae irregularly defined but clearly present. Telson vesicle globose; subaculear tubercle vestigial to moderate, blunt, irregular to conical and widely separate from aculeus, which is long, very sharp and strongly curved.

SUBORDINATE TAXA. After the present revision, the genus includes six nominal species: Tityopsis canizaresorum sp. n., T. inaequalis ( Armas, 1974) , T. inexpectata (Moreno, 1940) , T. mulata sp. n., T. pumila sp. n., and T. sheylae sp. n.

ECOLOGICAL NOTES.All species of Tityopsis are forest scorpions that need shady, humid places with plenty of refuges to survive e.g., rocks, leaf litter and rotten logs, usually in karstic soil ( Figs. 37–39 ). Nevertheless, at least one species occurs also in open vegetation (sparse pine forest with grass understorey, on shale sandy soil) and another has endured heavy, longstanding urbanization and still can be found in relict forested patches (from groves to tiny house gardens and courtyards) in Havana City. One species is a presumed troglobite known only from a single cave ( Fig. 40 ), but others have been found sporadically or frequently in this subterranean habitat and are typical troglophiles. With a couple of noteworthy exceptions, population densities are extremely low and without a visible cause, individuals tend to concentrate (but never aggregate) in reduced spots within much larger, homogeneous areas.

It is important to note here that the most widely used standard technique to sample or study scorpions in situ, i.e., nocturnal search with ultraviolet light, is essentially useless for epigean populations of Tityopsis . Except for three scattered successful exceptions, we found the vast majority of our specimens during diurnal searches under rocks, logs and leaf litter, in exactly the same spots that were intensively sampled in vain the night before (all moon phases attempted).

The adult sex ratio of the studied samples usually biased towards female, nevertheless, in juveniles it was more balanced. Males (either adult or juvenile) have been found in most populations, thus, sexual reproduction is assumed for them. Nevertheless, thelytokous parthenogenesis (allfemale broods) was confirmed by us to occur in one species and suspected from strong factual evidence for at least two, maybe three others (R. Teruel and T. M. Rodríguez-Cabrera, unpublished data).

Pregnant females have been found all around the year, but more frequently during the warmer, rainy season (May through October). Litter size is small for buthids: 1–20 newborn ( Fig. 36d ). Sexually reproducing females give birth only once a year in captivity and apparently every parturition requires a mating, i.e., iteroparity does not seem to occur. In one species, captive females raised to adulthood in absolute isolation gave birth by parthenogenesis 7–10 weeks after reaching maturity. Postembryonic development has never been studied by the direct method, but we raised to adulthood in captivity many wild-caught juveniles (indirect method) and found that maturity is attained in about one year at instars 5–7 (i.e., after four to six ecdyses), with larger species such as T. inaequalis being more prone to mature at different instars, than smaller ones.

The humidity requirements of all Tityopsis spp. are extremely high: all individuals dehydrate critically and die within less than 36 hours, if air humidity in their enclosure is not continuously kept above 85%. This is independent from the habitat where the scorpions lived in nature (humid, mesic or xeric), as well as from the origin of the individuals (wildcaught or captive-born).

COMMENTS. Despite intensive and extensive searches, Tityopsis remains undetected from any of the many islets adjacent to the main island and especially from the large Isla de Pinos, where its apparent absence is enigmatic and represents a subject of debate among scorpiologists. This question became even more interesting after we (Teruel & Rodríguez-Cabrera, 2017) finally accomplished ourselves the long-awaited discovery of another scorpion genus in Isla de Pinos: the diplocentrid Heteronebo Pocock, 1899 .

DISTRIBUTION ( Figs. 1–2 ). This genus is endemic to western Cuba (Pinar del Río, Artemisa, La Habana, Mayabeque and Matanzas Provinces), with widespread but scattered occurrences mostly across mountainous and hilly areas of the main island only. See further details below, in the Comments section .

Figures

Figure 1. Western Cuba, showing the updated geographical distribution of the genus Tityopsis: a) Previous locality records from literature (red symbols); b) Same “a” map, with new locality records obtained during the present study added (yellow symbols); c) Same “a” map, with locality records personally confirmed during the present study added (white symbols). Image frame size = 500 x 250 km.

Figure 2. Western Cuba, showing the complete geographical distribution of the genus Tityopsis (white symbols) and overlaid, that of the four species of Tityopsis treated herein (red symbols, complemented with habitus of their holotypes): T. mulata sp. n. (star), T. canizaresorum sp. n. (rhombs, the southernmost is the type locality), T. pumila sp. n. (triangle) and T. sheylae sp. n. (square). Image frame size, 500 x 250 km.

Figures 3–4. Tityopsis mulata sp. n., male holotype (3) and female paratopotype (4), habitus: a) dorsal; b) ventral. Scale bar = 10 mm.

Figures 5–6. Tityopsis mulata sp. n., male holotype (5) and female paratopotype (6): a) pedipalp, dorsal view; b) pedipalp chela, external view; c) pedipalp chela, ventral view; d) telson, lateral view.

Figures 7–8. Tityopsis mulata sp. n., male holotype (7) and female paratopotype (8): a) chelicerae and carapace; b) tergites.

Figures 9–10. Tityopsis mulata sp. n., male holotype (9) and female paratopotype (10): a) sternopectinal region; b) sternite V.

Figures 11–12. Tityopsis mulata sp. n., male holotype (11) and female paratopotype (12), metasoma and telson: a) dorsal, b) lateral, and c) ventral views.

Figures 13– 14. Tityopsis pumila sp. n., male holotype (13) and T. canizaresorum sp. n., female holotype (14), habitus: a) dorsal; b) ventral. Scale bar = 10 mm.

Figures 15–16. Tityopsis pumila sp. n., male holotype (15) and T. canizaresorum sp. n., female holotype (16): a) pedipalp, dorsal view; b) pedipalp chela, external view; c) pedipalp chela, ventral view; d) telson, lateral view.

Figures 17–18. Tityopsis pumila sp. n., male holotype (17) and T. canizaresorum sp. n., female holotype (18): a) chelicerae and carapace; b) tergites.

Figures 19–20. Tityopsis pumila sp. n., male holotype (19) and T. canizaresorum sp. n., female holotype (20): a) sternopectinal region; b) sternite V.

Figures 21–22. Tityopsis pumila sp. n., male holotype (21) and T. canizaresorum sp. n., female holotype (22), metasoma and telson: a) dorsal, b) lateral, and c) ventral views.

Figures 23–24. Tityopsis sheylae sp. n., male holotype (23) and female paratopotype (24), habitus: a) dorsal; b) ventral. Scale bar = 10 mm.

Figures 25–26. Tityopsis sheylae sp. n., male holotype (25) and female paratopotype (26): a) pedipalp, dorsal view; b) pedipalp chela, external view; c) pedipalp chela, ventral view; d) telson, lateral view.

Figures 27–28. Tityopsis sheylae sp. n., male holotype (27) and female paratopotype (28): a) chelicerae and carapace; b) tergites.

Figures 29–30. Tityopsis sheylae sp. n., male holotype (29) and female paratopotype (30): a) sternopectinal region; b) sternite V.

Figures 31–32. Tityopsis sheylae sp. n., male holotype (31) and female paratopotype (32), metasoma and telson: a) dorsal, b) lateral, and c) ventral views.

Figure 33. Tityopsismulata sp. n., photographed alive inits naturalhabitat unless otherwise noted:a) male holotype;b–c) female paratopotypes; d–e) captive female paratopotype, with litter of instars 1 (d) and 2 (e); f) captive juvenile male paratopotype.

Figures 34–35: Figure 34. T.pumila sp. n., male holotype photographed alive in its natural habitat (two different shots, photos courtesy Raimundo López-Silvero).Figure 35. T. canizaresorum sp. n., photographed alive in its natural habitat:a) female holotype; b) juvenile female paratopotype.

Figure 36. Tityopsis sheylae sp. n., photographed alive in its natural habitat unless otherwise noted: a) male holotype; b) small-sized male paratopotype; c) female paratopotype; d) captive female paratopotype, with mixed litter of instars 1–2; e) juvenile female paratopotype; f) just-molted, small-sized adult male paratopotype, being predated upon by the pholcid Ciboneya nuriae (the spider is not visible because while taking the photographs, it fled from the camera’s flash and hid in the crevice behind the scorpion). Photos courtesy Sheyla Yong (a–b).

Figures 37–38: Type localities of Tityopsis mulata sp. n. (37) and T. pumila sp. n. (38), a) habitat, b) microhabitat.

Figures 39–40: Type localities of Tityopsis canizaresorum sp. n. (39) and T. sheylae sp. n. (40), a) habitat, b) microhabitat (photo courtesy Sheyla Yong).