Titanobochica, surprising discovery of a new cave-dwelling genus from southern Portugal (Arachnida: Pseudoscorpiones: Bochicidae) Author Reboleira, Ana Sofia P. S. Author Zaragoza, Juan A. Author Gonçalves, Fernando Author Oromí, Pedro text Zootaxa 2010 2681 1 19 journal article 10.5281/zenodo.199333 db8f4069-a3c7-4b2b-9d03-46f9826e17c0 1175-5326 199333 Titanobochica magna Zaragoza & Reboleira sp. nov. ( Figs 2–31 ) Type material. Male holotype : Portugal , Algarve, Algarão do Remexido, 23 May 2009 , collected by A.S.P.S. Reboleira, deposited in DEUA . Paratypes : Portugal , Algarve, Algarão do Remexido, 15 March 2009 : 1D ( MCNB ), 2T ( MCNB , MHNG ), 2 males ( MCNB , MNHNP ), 23 May 2009 : 1D ( DEUA ), 1T ( DEUA ), 2 females ( MCNB , MHNG ), 5 September 2009 : 1T ( MNHNP ), 29 December 2009 : 1 female ( DEUA ); Gruta de Ibne Ammar, 23 May 2009 : 1 male ( DEUA ); Gruta do Vale Telheiro, 30 January 2009 : 2D ( DZUL , SR ), 1T ( DEUA ), 13 March 2009 : 1T ( DEUA ), 2 males ( DEUA , MHNG ), 2 females ( DZUL , SR ), 24 May 2009 : 2D ( DEUA , MHNG ), 2T ( DZUL , SR ), 2 males ( MNCNM , SR ), 6 September 2009 : 1 male ( SR ); Gruta da Senhora, 14 March 2009 : 1 female ( DEUA ), 29 December 2009 : 1 male ( DZUL ); all collected by A.S.P.S. Reboleira. FIGURE 2. Titanobochica magna gen. nov. , sp. nov. , general appearance of living specimen from Algarão do Remexido. Diagnosis. As for the genus. Etymology. From the Latin adjective magnus,-a,-um , meaning big, in reference to its large size. Description. The data correspond to the male holotype , followed by paratype males and females in parentheses. Measurements and ratios of adults and nymphs in Table 2 . A large pseudoscorpion (habitus: Fig. 2 ), body length about 6 mm ( Table 2 ). Desclerotized and depigmented. Opisthosoma and legs pale. Carapace, chelicerae and pedipalps reddish. Carapace ( Figs 3 , 24 ). Elongate, maximum width at median portion. Without eyes or eyes spots. Epistome lacking ( Fig. 4 ). Chaetotaxy: 40 (M: 33–41; F: 34–40) setae, formula (total number of setae in each row followed by number of lateral setae in parentheses): 6:20(8):8:6 [M: 6:16–20(4–7):5–8:4–7; F: 6:18–21(4– 9):4–7:6–7]. One pair of lyrifissures on each side of pre-ocular zone. Numerous diminutive pores over entire carapace. TABLE 2. Titanobochica magna gen. nov. , sp. nov. : measurements and ratios. M: male; F: female; T: tritonymph., D: deutonymph. M holotype M paratypes F paratypes T paratypes D paratypes Body 6.16 4.83–5.76 5.26–6.06 4.56–4.82 3.06–3.10 Carapace 1.36× 1.27–1.37× 1.24–1.37× 1.16–1.20× 1.19–1.32× (1.66/1.22) (1.64–1.70/1.20–1.34) (1.66–1.78/1.24–1.40) (1.20–1.34/1.02–1.12) (0.80–1.00/0.67–0.76) Chelicera Palm 2.33× 2.27–2.32× 2.04–2.17× 1.95–2.12× 2.00× (0.98/0.42) (0.96–1.05/0.42–0.45) (1.00–1.08/0.46–0.50) (0.74–0.89/0.38–0.42) (0.57–0.60/0.29–0.30) Finger 0.59 0.58–0.61 0.58–0.68 0.45–0.52 0.30–0.36 Pedipalp Trochanter 2.00× 1.87–2.10× 1.89–1.96× 1.88–1.95× 1.78–1.94× (1.00/0.50) (1.00–1.13/0.48–0.56) (1.02–1.12/0.54–0.57) (0.75–0.84/0.40–0.43) (0.52–0.55/0.28–0.29) Femur 11.04× 10.18–11.24× 10.72–10.98× 7.52–8.19× 7.58–8.38× (4.09/0.37) (4.07–4.28/0.37–0.42) (4.21–4.40/0.38–0.40) (2.58–3.12/0.32–0.41) (1.76–1.82/0.21–0.24) Patella 9.00× 8.60–9.13× 8.81–9.18× 6.96–7.56× 5.93–6.42× (3.94/0.44) (3.94–4.20/0.43–0.47) (4.04–4.32/0.44–0.49) (2.46–2.84/0.35–0.41) (1.54–1.60/0.24–0.27) Patella pedicel 0.62 0.60–0.67 0.65–0.70 0.48–0.57 0.30–0.32 Hand 2.92× 2.55–2.81× 2.68–2.82× 2.26–2.43× 2.45–2.55× (2.10/0.72) (2.11–2.19/0.76–0.84) (2.08–2.31/0.75/0.82) (1.43–1.78/0.63–0.73) (1.08–1.16/0.44–0.46) Coxal area ( Fig. 18 ). Apex of pedipalpal coxa long and acute, with 2 distal setae (one female with 3 setae on one apex lobe and 2 on the other). Pedipalpal coxa with 15–19 setae (M: 15–19; F: 14–19), pedal coxa I: 21–24 (M: 18–26; F: 19–24), II: 18–19 (M: 15–21; F: 16–20), III: 16 (M: 12–20; F: 13–17), IV: 25 (M: 19–26; F: 17–26). Anterolateral process of coxa I rounded and moderately prominent; medial process absent.

Hand pedicel	0.30	0.30–0.32	0.31–0.32	0.18–0.22	0.15
Fixed finger Movable finger Chela	3.72 3.25 8.00×	3.66–3.82 3.30–3.42 6.98–7.53×	3.83–4.08 3.46–3.68 7.37–7.73×	2.69–3.09 2.33–2.70 6.29–6.51×	1.83–1.92 1.65–1.66 6.61–6.73×
Chela/carapace Chela/femur	(5.76/0.72) 3.47× 1.42×	(5.72–5.96/0.76–0.84) 3.45–3.59× 1.37–1.44×	(5.80–6.29/0.75–0.82) 3.41–3.64× 1.38–1.43×	(4.09–4.77/0.63–0.73) 3.20–3.67× 1.50–1.59×	(2.91–3.06/0.44–0.46) 3.06–3.64× 1.65–1.68×
Femur/carapace Femur/fixed finger Femur/patella	2.45× 1.09× 1.03×	2.45–2.55× 1.08–1.13× 1.02–1.05×	2.47–2.58× 1.08–1.10× 1.02–1.04×	2.02–2.45× 0.96–1.01× 1.05–1.10×	1.82–2.20× 0.95–0.96× 1.14×
Patella/hand Fixed Finger/hand Leg I	1.81× 1.77×	1.82–1.93× 1.72–1.79×	1.87–1.94× 1.77–1.84×	1.60–1.72× 1.73–1.88×	1.38–1.43× 1.66–1.70×
Trochanter	1.68× (0.61/0.36)	1.63–1.84× (0.61–0.68/0.37–0.38)	1.68–1.76× (0.62–0.70/0.37–0.40)	1.59–1.64× (0.46–0.50/0.28–0.34)	1.64–1.65× (0.34/0.21) continued next page

TABLE 2. (continued)
M holotype	M paratypes	F paratypes	T paratypes	D paratypes
Femur 7.56×	7.48–8.14×	7.57–7.77×	6.20–6.93×	5.90×
(1.97/0.26)	(1.98–2.10/0.25–0.27)	(2.02–2.14/0.26–0.28)	(1.26–1.49/0.20–0.24)	(0.83–0.92/0.14–0.16)
Patella 3.23×	3.19–3.36×	3.20–3.36×	2.81–3.06×	1.83–1.96×
(0.71/0.22)	(0.74–0.79/0.22–0.25)	(0.74–0.77/0.22–0.24)	(0.50–0.59/0.17–0.21)	(0.33–0.37/0.18–0.19)
Tibia 13.51×	12.46–14.10×	12.93–13.55×	9.33–10.69×	8.95–9.46×
(2.00/0.15)	(1.97–2.02/0.14–0.17)	(1.94–2.10/0.15–0.16)	(1.12–1.46/0.12–0.15)	(0.80–0.85/0.08–0.10)
Metatarsus 1.68×	1.63–1.79×	1.57–1.80×	1.55–1.67×	1.67–1.75×
(0.22/0.13)	(0.21–0.24/0.13–0.14)	(0.24/0.13–0.16)	(0.17–0.21/0.11–0.13)	(0.14–0.15/0.08–0.09)
Tarsus 14.17×	13.23–14.96×	13.60–14.07×	9.47–10.00×	10.33–10.50×
(1.70/0.12)	(1.68–1.75/0.12–0.13)	(1.74–1.90/0.13–0.14)	(1.12–1.42/0.12–0.15)	(0.84–0.93/0.08–0.09)
Femur/patella 2.77×	2.64–2.73×	2.73–2.83×	2.42–2.55×	2.50–2.51×
Tarsus/Metatarsus 7.80×	7.28–8.25×	7.25–8.09×	6.59–6.76×	6.00–6.20×
Leg IV
Trochanter 2.65×	2.43–2.63×	2.55–2.58×	2.62–2.75×	2.05–2.50×
(0.90/0.34)	(0.82–0.94/0.34–0.36)	(0.90–0.98/0.35–0.38)	(0.64–0.80/0.26–0.30)	(0.38–0.43/0.17–0.19)
Femur 1.86×	1.67–1.86×	1.86–1.93×	1.52–1.73×	1.75–1.77×
(0.54/0.29)	(0.50–0.57/0.29–0.31)	(0.56–0.60/0.30–0.32)	(0.37–0.46/0.23–0.27)	(0.27–0.32/0.15–0.18)
Patella 10.17×	8.94–9.87×	9.22–9.98×	7.21–7.91×	6.79–6.84×
(3.05/0.30)	(2.95–3.12/0.31–0.33)	(3.05–3.21/0.32–0.35)	(1.86–2.24/0.24–0.29)	(1.16–1.30/0.17–0.19)
Tibia 15.78×	15.23–16.71×	14.62–15.98×	11.44–13.09×	10.70–10.83×
(2.84/0.18)	(2.81–2.88/0.17–0.18)	(2.84–2.97/0.18–0.20)	(1.67–2.12/0.15–0.16)	(1.12–1.22/0.10–0.11)
Metatarsus 1.70×	1.77–1.94×	1.81–1.94×	1.89–2.02×	2.00×
(0.27/0.16)	(0.28–0.33/0.15–0.17)	(0.30–0.31/0.16–0.17)	(0.23–0.26/0.12–0.14)	(0.18–0.19/0.09–0.10)
Tarsus 17.29×	17.36–18.66×	17.97–18.37×	12.60–13.53×	12.56–13.37×
(2.42/0.14)	(2.46–2.54/0.13–0.14)	(2.57–2.75/0.14–0.15)	(1.55–2.01/0.12–0.15)	(0.18–0.19/0.09–0.10)
Tarsus/Metatarsus 8.64×	7.64–9.20×	8.43–8.96×	6.46–7.62×	6.28–6.68×

Opisthosoma ( Figs 22–23 , 25 ). Elongate. Pleural membranes smoothly, longitudinally striate. Some tergites with one or two median subdiscal microsetae; tergal chaetotaxy I–X (totals, followed by number of median microsetae in parentheses): 6: 6: 6: 10(1): 9(2): 9(1): 9(1): 9(1): 10(1): 7 [M: 4–6: 5–7: 5–6: 7–8(0–1): 7–10(1–2): 8–9(1–2): 7–10(1–2): 6–9(0–1): 7–9(0–1): 6–9; F: 4–7: 6–7: 6: 8–9(0–1): 8–12(1–2): 8–12(1–2): 7–12(1–2): 7–10(1–2): 7–9(0–1): 7–8]. Male genital area ( Fig. 19 ): 18 (22–26) setae on sternite II plus 15 (19–23) small setae on anterior genital opening, posterior genital opening with 15 (12–17) small setae, sternite III with 10 (7–12) discal and 11 (9–12) marginal setae. Female paratypes : genital area 18–21 very small setae on sternite II, 13–17 marginal setae and 8–10 small discal setae on sternite III. Sternites IV-VII, in addition to the marginal setae, with one or two rows of discal setae, most reduced to microsetae, resulting sternal plates biseriate or triseriate; sternites VIII-IX with some dispersed discal setae; sternal chaetotaxy IV-X (expressed as discal setae/posterior setae): 9/10: 8/14: 10/12: 9/12: 5/9: 3/11: 10; [M: 9/10: 8/14: 10/12: 9/12: 5/9: 3/11: 10; [M: 9–11/8–14: 7–10/8–12: 7–8/9–14: 5–7/7–12: 4–7/8–10: 3–6/7–11: 0–1/7–10; F: 6–11/6–11: 6–8/9– 15: 6–9/10–14: 6–10/8–13: 5–9/8–13: 2–5/8–13: 0–1/7–10]; segment XI 7 setae (7–8), segment XII (anal cone) bears 2 tergal and 3 sternal setae (teratology) (2–2). Tergites and sternites without tactile setae. Stigmata on sternite III ( Fig. 20 ) with 12-14 (M: 9–10; F: 11–14) setae, stigmata on sternite IV ( Fig. 21 ) with 9-10 (M: 5–8; F: 5–9) setae. Two lyrifissures in medial zone, one more on each side of anterolateral zone. Chelicera ( Figs 5–6 , 26–27 ). Four setae on hand (4, seldom 5–6; when variation occurs one chelicera always with 4) and one seta on movable finger, 0.77 (M: 0.69–075; F: 0.74–0.79) from base. Galea diminutive and simple, dagger-blade shaped, tip distinctly proximal of apex of finger, length: 0.03 (M: 0.03–0.05; F: 0.03) mm ( Fig. 6 , ga). Fixed finger with 8 teeth, distal tooth larger than the rest and two reduced to tiny protuberances (8–10). Movable finger with an extremely large distal tooth, turned slightly laterally to lodge the fixed finger, moreover with a ridge joining 4 (3–6) teeth and 3 (3–5) separate teeth up to beginning of basal third, total 8 (8–13) teeth. Rallum ( Fig. 7 ) with five externally serrate blades (5, one teratological male with only 4 in one rallum and 5 in the other), basal blade slightly shorter than others. Serrula exterior with 35 (36–40) blades, serrula interior with 34 (33–37) blades. Pedipalps ( Figs 13–17 , 28–30 ). Articles smooth, except fingers of chela; some micropores dispersed over whole pedipalp and in groups of 9–10 at bases of femur, patella and hand. Trochanter short. Femur and patella very slender, patella only slightly shorter than femur; femur distinctly longer than fixed chelal finger and without glandular tubercle at base of antiaxial face. Chela smooth, except distal portion of hand and the fingers, which are weakly granulated; fixed finger with irregular outline at the distal portion and distinctly longer than movable finger. Hand with oval outline at paraxial face and flattened at antiaxial face; trichobothrium ib located in basal half of dorsum of hand, eb in distal portion of hand; 10 (9–14) thin and small chemosensory setae ( Fig. 13 , cs) on dorsum of hand, in a row from ib to base of finger ( Figs 13 , 30 ). Fixed finger with tip rounded ( Figs 15–16 ), venedens and venom apparatus completely absent; trichobothria esb and isb at base of finger, medial portion of finger without trichobothria, ist slightly proximal of est , it and et at tip of finger; with one apical sublateral tooth and 112 (M: 107–121; F: 111–116) teeth up to base of finger, apically uniseriate, basally decreasing in size ( Fig. 14 ); 26th to 45th teeth from the apex are heterodentate, alternately weakly different in shape and with a slightly different orientation ( Fig. 14 ). Movable finger ( Figs 14, 17 ) with functional venom apparatus, nodus ramosus (nr) proximal of trichobothria st and not expanded ( Fig. 14 ); distance trichobothria sb–st 1.82× (1.17–1.92) longer than distance b–sb ; with 77 (M: 68– 81; F: 74–77) teeth, reaching to distal of trichobothrium b, rounded and progressively flattened in basal portion, last 6 (7–8) basal teeth without dental canal; one male paratype with teratological movable finger of one chela shorter than normal, length 1.40 mm , only trichobothria b and sb present, dental row reduced to 47 teeth. Lyrifissures of pedipalps as shown in Figs 13–14 . Legs smooth, tarsi not swollen basally, claws smooth and apically blunt, subterminal setae apically feathery. Leg I ( Fig. 8 ): metatarsus extremely short and apparently fused with tarsus, together composing a rigid article. Leg IV ( Figs 9–10 , 31 ): femur(F)/patella(P) suture strongly oblique, femur evidently shorter than ¼ length of combined segments, according to Muchmore’s formula (1998): F/F+P: 0.15 (0.14–0.16); articles without tactile setae. Tritonymph. Carapacal chaetotaxy: 31–39 setae, formula (total number of setae in each row followed by number of lateral setae in parentheses) 4–6:17–19(6–7):4–6:5–8. Coxal area: apex of the pedipalp with 2 setae, pedipalpal coxa 10–14 setae, pedal coxa I: 13–15, II: 11–16, III: 9–14, IV: 12–16. Opisthosomal tergites I-X chaetotaxy (totals, followed by number of median setae in parentheses): 4–6: 6: 6: 6–8: 6–9(1): 6–8(1): 6– 8(1): 6–8(1): 6–7: 4; sternite II: 4–5 setae, III-X (discal/posterior): 5/11–12: 6–8/7–8: 3–4/9–12: 3–5/8–12:2– 5/8–11: 2–4/8–10: 1–3/6–8: 6, segment XI: 6–8 setae, anal cone with 2 tergal and 2 sternal setae, stigmata sternite III: 6–7 setae, IV: 4–5. Cheliceral palm with 4 setae, some specimens with 4 setae on one chelicera and 5 on other, subgaleal seta on movable finger 0.79–0.80 from base; galea short and simple, length: 0.04 mm ; fixed finger with 8–9 teeth, the distal one larger than the rest; movable finger with 7–12 teeth, the distal one very large and sublateral; rallum with 4 externally serrate blades, with the same shape as in adults; serrula exterior with 30–34 blades, serrula interior with 25–31 blades. Pedipalp: chelal fixed finger about as long as femur; trichobothrium ist proximal of est ; fixed finger with 93–100 teeth; movable finger with 64–68 teeth, some basal ones without dental canal. Trichobothria isb and sb lacking. Legs: tarsi of all four legs basally swollen, maximum width of leg I at 0.18–0.22 of total length from base, leg IV ( Fig. 12 ) 0.29–0.32; metatarsus leg I apparently fused with the tarsus; leg IV: F/F+P 0.16–0.17. FIGURES 3–12. Titanobochica magna gen. nov. , sp. nov. , male holotype, unless otherwise noted. 3. carapace; 4. anterior margin of carapace; 5. right chelicera; 6. distal portion of fingers of right chelicera; 7. cheliceral rallum; 8. left leg I, lateral view; 9. left leg IV, lateral view; 10. distal portion of tarsus and apotele of left leg IV, lateral view. Deutonymph paratype: 11. metatarsus and tarsus of right leg I, lateral view; 12. tarsus of left leg IV, lateral view, tritonymph paratype. Scale bars (mm): 0.10 (Fig. 7), 0.20 (Figs 6, 10); 0.25 (Fig. 5), 0.50 (Figs 3, 4, 11, 12), 1.00 (Figs 8, 9). Abbreviations: ga : cheliceral galea. FIGURES 13–17. Titanobochica magna gen. nov. , sp. nov. , male holotype. 13. left pedipalp, dorsal view; 14. left chela, antiaxial view, some teeth magnified; 15. tip of fixed finger of left chela, ventral view; 16. tip of fixed finger of left chela, antiaxial view; 17. tip of movable finger of left chela, antiaxial view. Scale bars (mm): 0.20 (Figs 15–17), 1.00 (Figs 13, 14). Abbreviations: trichobothrial terminology following Chamberlin (1931); cs: chemosensory setae; nr: nodus ramosus. FIGURES 18–23. Titanobochica magna gen. nov. , sp. nov. , male holotype. 18. coxae of pedipalp and leg I; 19. genital sternites, ventral; 20. left stigma of sternite III; 21. left stigma of sternite IV; 22. sternites II – VII, ventral view; 23. tergites I – VII, dorsal view. Scale bars (mm): 0.20 (Figs 19 – 21), 0.50 (Figs 18, 22, 23). FIGURES 24–27. Titanobochica magna gen. nov. , sp. nov. , male paratype from Gruta do Vale Telheiro. SEM images. 24. carapace; 25. opisthosoma, tergite VII partially teratological; 26. chelicerae; 27. distal portion of fingers of left chelicera. Scale bars (mm): 0.12 (Fig. 27), 0.30 (Fig 26), 1.00 (Figs 24, 25). FIGURES 28–31. Titanobochica magna gen. nov. , sp. nov. , male paratype from Gruta do Vale Telheiro. SEM images. 28. fingers of the left chela, dorsal view, A: glandular pore magnified; 29. tip of the chelal movable finger, with venedens, A: magnification of a tooth; 30. right chelal hand, partial dorsal view; 31. leg IV, detail of claws and subterminal feathery setae. Scale bars (mm): 0.009 (Fig. 28A), 0.01 (Fig. 29A), 0.075 (Figs 29, 31), 0.75 (Fig. 30), 1.00 (Fig. 28). Deutonymph. Carapacal chaetotaxy: 25–27 setae, formula (total number of setae in each row, followed by number of lateral setae in parentheses) 4:13–14(3–4):4:4–5. Coxal area: pedipalpal apex with 2 setae; pedipalpal coxa 7–8 setae, pedal coxa I: 8–10, II: 7–8, III: 6–8, IV: 8. Tergites I-X chaetotaxy: 4–5: 4: 6: 6: 6: 6: 6: 6: 6: 4–6, without median setae; sternite II without setae, sternites III–X uniseriate, some with median subdiscal setae: 6–7: 2/5–6: 1–2/8–10: 2/8–10: 2/8: 2/8: 0–1/7: 6, segment XI: 6–7 setae, anal cone with 2+2 setae, stigmata sternite III: 4–5 setae, IV: 3–4. Cheliceral palm with 4 setae, subgaleal seta on movable finger 0.74–0.75; galea short and simple, length 0.02–0.04 mm ; fixed finger with 7–10 teeth, distal tooth large; movable finger with 8–10 teeth, distal tooth large and sublateral; rallum with 3 unilaterally serrate blades; serrula exterior with 27 blades, serrula interior with 22. Pedipalp: chelal fixed finger longer than the femur; trichobothrium ist proximal of est ; fixed finger with 83–84 teeth; movable with 55–58 teeth, basal 12–17 flattened, all with dental canal. Trichobothria esb , isb , sb and st lacking. Tarsi of legs basally swollen, leg I maximum width 0.20 from base, leg IV 0.29; suture between metatarsus and tarsus leg I almost indistinguishable ( Fig. 11 ); leg IV F/F+P: 0.19–0.20. Remarks. No significant differences are found in the form and length of the cheliceral galea between males, females and nymphs. The development of the chelal femur length and slenderness relative to the other pedipalpal articles is progressive from the nymphal to adult stages, as shown in the ratios of the femur, femur/fixed finger and chela/femur ( Table 2 ). Heterodentate marginal teeth of fixed chelal finger have been previously reported in Bochicidae for Vachonium boneti Chamberlin, 1947 ( Chamberlin 1947 ) . Presence of chemosensory setae on the palm of the chela was reported for Bochicidae by Judson (2007) , who assumed the chemosensory function of these setae because they are smaller, thinner and more strongly inclined than ordinary setae. Tritonymphs and deutonymphs show tarsi swollen basally, which does not occur in adults. Basally swollen tarsi have been described for nymphs and, exceptionally, for adults of some genera of the family Syarinidae : Pseudoblothrus Beier ( Vachon 1954 ) , Hadoblothrus Beier and Microcreagrella Beier ( Mahnert 1980 ) . In the family Bochicidae this characteristic has also been reported for nymphal stages of the genera Mexobisium Muchmore, 1972 , Paravachonium and Vachonium ( Muchmore 1973a , 1973b , 1982c ), to which Titanobochica is now added; apparently in correlation with this characteristic, the nymphal stages of these bochicid genera also usually show fused metatarsi and tarsi of legs I and II ( Muchmore 1973a , 1973b , 1982c ). Titanobochica shows a rigid suture between metatarsi and tarsi of legs I–II in adults and tritonymphs, whereas in deutonymphs they have a very indistinct, almost unrecognizable suture.