Hyidae Chamberlin, 1930

Harvey, Mark S., Burger, Mieke A. A., Abrams, Kym M., Finston, Terrie L., Huey, Joel A. & Perina, Giulia, 2023, The systematics of the pseudoscorpion genus Indohya (Pseudoscorpiones: Hyidae) in Australia, Zootaxa 5342 (1), pp. 1-119 : 19-21

Family Hyidae Chamberlin, 1930

Diagnosis. The family Hyidae currently includes those pseudoscorpions in which the femora of legs I and II have a basi-dorsal mound surmounted by a small seta and slit sensilla, the setae of the female anterior genital operculum are extremely small, and the pedipalpal femur has 2–3 stout setae on the retrolateral-basal margin ( Harvey 1992, 1993; Harvey & Volschenk 2007).

Genus Indohya Beier, 1974

urn:lsid:zoobank.org:act:41313090-4004-46AE-92DF-994574CC67AC

Indohya Beier 1974: 1003

Hyella Harvey 1993: 27–28 (synonymised by Harvey and Volschenk 2007: 386). urn:lsid:zoobank.org:act:CE45A219-6717-4E3C-8E6E-FE65773E0B72

Type species. Indohya : Indohya besucheti Beier, 1974 , by original designation.

Hyella : Hyella humphreysi Harvey, 1993 , by original designation.

Diagnosis. Species of Indohya differ from the only other genus of the family, Hya , by the absence of venom glands in the fixed chelal finger (e.g. Figs. 8C , 40C ) and the juxtadentate teeth of the chelal fingers (e.g. Figs. 8C , 40C ).

Remarks. The genus Indohya has a remarkable disjunct distribution in the southern hemisphere with species known from southern India, Madagascar and north-western Australia ( Harvey & Volschenk 2007). This is highly suggestive of an ancient Gondwanan origin for the genus. The presence of an extensive fauna in the Pilbara—comprising epigean and troglobitic species—extends the distribution of the family into a new Australian bioregion.

The ratio sb -st / b -sb has been recently introduced in pseudoscorpion taxonomy to denote the relative position of trichobothrium sb on the movable chelal finger ( Zaragoza 2017). This value was calculated for all of the new species described in this paper. Its utility, however, was found to be less useful than for chthoniid pseudoscorpions, with considerable intraspecific variation found in some species. For example, males of I. boltoni vary from 1.26 to 2.26 and females vary from 2.00 to 2.45, and males of I. lynbeazleyae vary from 1.49 to 1.89, and females vary from 0.41 to 1.87. The discrepancies within I. lynbeazleyae were evident in individuals, with significant bilateral variation. The adult specimens possess ratios as follows: male holotype (WAM T91735) with 1.49 on the left chela (with the right chela missing); male paratype (WAM T91742) with 1.71 (left) and 1.89 (right); female paratype (WAM T138709) with 0.41 (left) and 1.23 (right); female paratype (WAM T138707) with 1.10 (left) and 1.55 (right); and female paratype (WAM T138728) with 1.87 (left), but the right chela is missing from the specimen.

Key to adults of species of Indohya

1. Carapace with 18 setae arranged 4: 2: 4: 4: 4, with 4 setae in posterior row........... Indohya haroldi Harvey & Volschenk Carapace with 12–16 setae arranged 4: 2: 4: 4: 2 or 4: 2: 4: 2: 2 or 4: 0: 4: 4: 2 or 4: 0: 4: 2: 2, with 2 setae in posterior row.. ................................................................................................... 2

2. Carapace with 16 setae arranged 4: 2: 4: 4: 2 ( Fig. 55A ), with 4 setae in intermediate row............................ 3 Carapace with 12–14 setae arranged 4: 2: 4: 2: 2 (e.g. Figs. 10A , 23A , 26A , 47A ) or 4: 0: 4: 2: 2 (e.g. Figs. 8A , 12A , 16A , 20A , 30A , 32A , 34A ), with 2 setae in intermediate row........................................................... 12

3. Carapace with 14 setae arranged 4: 0: 4: 4: 2, with 4 setae in intermediate row..................................... 4 Carapace with 12 setae arranged 4: 0: 4: 2: 2, with 2 setae in intermediate row (e.g. Figs. 8A , 16A ).................... 5

4. Cheliceral hand with 5 setae; fixed chelal finger with 20–21 very flat teeth...................... Indohya besucheti Beier Cheliceral hand with 6 setae; fixed chelal finger with 29 moderately flat teeth.................... Indohya caecata Beier

5. Larger species, e.g. chela (with pedicel) 1.90–2.01 (Ô), 1.84–1.855 (♀) mm long................................... 6 Smaller species, e.g. chela (with pedicel) less than 1.0 mm long................................................ 7

6. Prolateral margin of chelal hand smooth; chelal hand (without pedicel) 1.79–1.96 (Ô), 1.84 (♀) × longer than broad; setae on pedipalpal femur of moderate length, not as long as width of femur; cheliceral hand with 6 setae............................................................................................. Indohya gollum Harvey & Volschenk Prolateral margin of chelal hand finely granulate; chelal hand (without pedicel) 1.71 (♀) × longer than broad; most setae on pedipalpal femur long, about as long as width of femur; cheliceral hand with 7 setae.................................................................................................... Indohya napierensis Harvey & Volschenk

7. Eyes present......................................................................................... 8 Eyes absent (e.g. Fig. 55A )............................................................................ 11

8. With two pairs of eyes................................................................................. 9 With one pair of eyes................................................................................. 10

9. Pedipalpal femur 0.41–0.425 mm in length; pedipalpal femur with granulations on basal half....... Indohya panops Harvey Pedipalpal femur 0.32–0.33 mm in length; pedipalpal femur smooth........................... Indohya pusilla Harvey

10. Pedipalpal femur 0.365 (♀) mm in length; carapace with lateral setae of ocular row located on same level as median ocular setae, and median setae of intermediate row located slightly anterior to lateral setae of intermediate row................................................................................................... Indohya beieri Harvey Pedipalpal femur 0.402 (Ô), 0.467 (♀) mm in length; carapace with lateral setae of ocular row located anterior to median ocular setae, and median setae of intermediate row located anterior to lateral setae of intermediate row........................................................................................... Indohya jacquelinae Harvey & Volschenk

11. Chela (with pedicel) 3.21 (♀) × longer than broad; fixed chelal finger with 41 teeth (♀).......... Indohya typhlops Harvey Chela (with pedicel) 3.98 (♀) × longer than broad; fixed chelal finger with 46 teeth (♀) ( Fig. 55C )......................................................................................... Indohya currani Harvey & Burger , n. sp.

12. Carapace with 14 setae arranged 4: 2: 4: 2: 2, with 2 setae in ocular row (e.g. Figs. 10A , 23A , 26A , 47A , 59A , 62A )...... 13 Carapace with 12–14 setae arranged 4: 0: 4: 2: 2 or 4: 0: 4: 4: 2 (e.g. Figs. 8A , 16A ), without setae in ocular row........ 18

13. Eyes present (e.g. Figs. 23A , 26B , 59A ).................................................................. 14 Eyes absent (e.g. Figs. 10A , 47A , 62A )................................................................... 16

14. Basal teeth of fixed chelal finger prominent and reclined ( Fig. 26F )............... Indohya cardo Harvey & Burger , n. sp. Basal teeth of fixed chelal finger small and upright ( Figs. 23C , 59D )............................................ 15

15. Larger species, e.g. chela (with pedicel) greater than 1.0 mm.................... Indohya boltoni Harvey & Burger , n. sp. Smaller species, e.g. chela (with pedicel) less than 0.8 mm .................. Indohya julianneae Harvey & Burger , n. sp.

16. Larger species, e.g. chela (with pedicel) greater than 1.5 mm ................... Indohya adlardi Harvey & Burger , n. sp. Smaller species, e.g. chela (with pedicel) less than 1.3 mm ................................................... 17

17. Trichobothrial ratio sb–st / sb–b = 2.47 (♀) ( Fig. 62C ); larger species, e.g. chela (with pedicel) 1.22–1.27 (♀) mm............................................................................. Indohya sachsei Harvey & Burger , n. sp. Trichobothrial ratio sb–st / sb–b = 1.81 (Ô) ( Fig. 47C ); larger species, e.g. chela (with pedicel) 1.055 (Ô) mm.................................................................................... Indohya rixi Harvey & Burger , n. sp.

18. Eyes present ( Fig. 60A )................................................ Indohya karenae Harvey & Burger , n. sp. Eyes absent (e.g. Figs. 8A , 20A , 30A , 57A )............................................................... 19

19. Tergite I with 2 setae................................................................................. 20 Tergite I with 4 (or rarely 3) setae....................................................................... 22

20. Smaller species, e.g. chela (with pedicel) 1.12 mm (Ô) in length................. Indohya aquila Harvey & Burger , n. sp. Larger species, e.g. chela (with pedicel) 1.52–1.65 mm (Ô), 2.09–2.18 mm (♀) in length............................ 21

21. Trichobothrium isb midway between esb and ist ...................................... Indohya humphreysi (Harvey) Trichobothrium isb closer to esb than to ist ( Fig. 8C )...................... Indohya anastomosa Harvey & Burger , n. sp.

22. Distal teeth of the fixed chelal finger rounded.............................................................. 23 At least 2 distal teeth of the fixed chelal finger pointed....................................................... 24

23. All teeth of the fixed chelal finger rounded ( Fig. 20E )................... Indohya arnoldstrongi Harvey & Burger , n. sp. At least some teeth of the fixed chelal finger pointed ( Fig. 57E )................. Indohya finitima Harvey & Burger , n. sp.

24. Teeth of the fixed chelal finger rounded except for 5–7 distal teeth and occasionally the basal-most tooth ( Fig. 30E ).......................................................................... Indohya cockingi Harvey & Burger , n. sp. Teeth of the fixed chelal finger including many pointed teeth (e.g. Fig. 12D , 34F )................................. 25

25. Smaller species, e.g. chela (with pedicel) less than 1.20 mm; fixed chelal finger with 46–49 teeth ( Figs. 12C , 34D )....... 26 Larger species, e.g. chela (with pedicel) at least 1.39 mm; fixed chelal finger with 69–85 teeth (e.g. Figs. 32C , 42D , 49C ). 27

26. Chelal hand slightly narrowed distally ( Fig. 12B ).......................... Indohya alexanderi Harvey & Burger , n. sp. Chelal hand rounded ( Fig. 34C )......................................... Indohya draconis Harvey & Burger , n. sp.

27. Pedipalpal femur and patella mostly granulate ( Figs. 32B , 42C )................................................ 28 Pedipalpal femur and patella smooth or only partially granulate ( Figs. 40B , 49B , 51B )............................. 29

28. Carapaceal setae of the median row slightly recurved ( Fig. 42A )............ Indohya lynbeazleyae Harvey & Burger , n. sp. Carapaceal setae of the median row strongly recurved ( Fig. 32A )................ Indohya cribbi Harvey & Burger , n. sp.

29. Pedipalpal segments narrow, e.g. chela (with pedicel) at least 5.0 × longer than broad; femur at least 6.5 × longer than broad ( Fig. 49B ).......................................................................................... 30 Pedipalpal segments less narrow, e.g. chela (with pedicel) less than 4.5 × longer than broad; femur less than 6.0 × longer than broad ( Figs. 40B , 51B )................................................................................ 31

30. Larger species, e.g. chela (with pedicel) 2.22–2.32 (Ô), 2.22–2.55 (♀) mm......... Indohya damocles Harvey & Volschenk Smaller species, e.g. chela (with pedicel) 1.50–1.66 (Ô) mm.................. Indohya sagmata Harvey & Burger , n. sp.

31. Larger species, e.g. chela (with pedicel) 1.81 (♀) mm......................... Indohya jessicae Harvey & Burger , n. sp. Smaller species, e.g. chela (with pedicel) 1.395 (Ô), 1.505 (♀)................ Indohya scanloni Harvey & Burger , n. sp.

Cape Range fauna

Figures

FIGURE 8. Indohya anastomosa Harvey & Burger, n. sp., holotype female (WAM T141292): A, carapace, dorsal; B, right pedipalp, dorsal; C, left chela, retrolateral; D, teeth, left fixed finger. Scale lines = 1.0 mm (Fig. B), 0.5 mm (Figs. A, C), 0.1 mm (Fig. D).

FIGURE 10. Indohya adlardi Harvey & Burger, n. sp., holotype female (WAM T127062):A, carapace, dorsal; B, right pedipalp, dorsal; C, left chela, retrolateral; D, fixed finger, chelal teeth, retrolateral. Scale lines = 0.2 mm (Figs. A, C), 0.5 mm (Fig. B), 0.1 mm (Fig. D).

FIGURE 12. Indohya alexanderi Harvey & Burger, n. sp., holotype male (WAM T138460): A, carapace, dorsal; B, right pedipalp, dorsal; C, left chela, retrolateral; D, left movable finger, teeth, retrolateral; E, left fixed finger, teeth, retrolateral. Scale lines = 0.5 mm (Figs. B, C), 0.2 mm (Fig. A), 0.1 mm (Figs. D, E).

FIGURE 16. Indohya aquila Harvey & Burger, n. sp., holotype male (WAM T130511): A, carapace, dorsal; B, left pedipalp, dorsal; C, left chela, retrolateral; D, left movable finger, chelal teeth, retrolateral; E, left fixed finger, chelal teeth, retrolateral. Scale lines = 0.5 mm (Figs. A, C), 0.2 mm (Fig. A), 0.1 mm (Figs. D, E).

FIGURE 20. Indohya arnoldstrongi Harvey & Burger, n. sp., holotype male (WAM T111408): A, carapace, dorsal; B, right chelicera, dorsal; C, right pedipalp, dorsal; D, left chela, retrolateral; E, fixed finger, chelal teeth, retrolateral; F, male genitalia, ventral. Scale lines = 0.5 mm (Figs. C, D), 0.2 mm (Fig. A), 0.1 mm (Figs. B, E, F).

FIGURE 23. Indohya boltoni Harvey & Burger, n. sp., holotype male (WAM T112020): A, carapace, dorsal; B, right pedipalp, dorsal; C, left chela, retrolateral; D, left fixed finger, chelal teeth, retrolateral. Scale lines = 0.5 mm (Fig. B), 0.2 mm (Figs. A, C), 0.1 mm (Fig. D).

FIGURE 26. Indohya cardo Harvey & Burger, n. sp., holotype male (WAM T113253): A, carapace, dorsal; B, right pedipalp, dorsal; C, left chela, retrolateral; D, left movable finger, chelal teeth, retrolateral; E, left fixed finger, chelal teeth, retrolateral; F, left fixed finger, chelal teeth, retrolateral. Scale lines = 0.5 (Figs. B, C), 0.25 mm (Fig. A), 0.1 mm (Figs. D, E).

FIGURE 30. Indohya cockingi Harvey & Burger, n. sp., holotype male (WAM T111708), unless stated otherwise: A, carapace, dorsal; B, right chelicera, dorsal; C, right pedipalp, dorsal; D, right chela, retrolateral; E, right fixed finger, chelal teeth, retrolateral; F, left chelal fingers, retrolateral, tritonymph (WAM T111712). Scale lines = 0.5 mm (Figs. C, D), 0.2 mm (Fig. A), 0.1 mm (Figs. B, E).

FIGURE 32. Indohya cribbi Harvey & Burger, n. sp., holotype male (WAM T138721), unless stated otherwise: A, carapace, dorsal; B, right pedipalp, dorsal; C, left chela, retrolateral; D, left chelal teeth, retrolateral; E, left chela, retrolateral, tritonymph (WAM T150970); F, left chela, retrolateral, protonymph (WAM T138974). Scale lines = 0.5 mm (Figs. B, C), 0.25 mm (Figs. E, F), 0.2 mm (Fig. D), 0.1 mm (Fig. A).

FIGURE 34. Indohya draconis Harvey & Burger, n. sp., holotype female (WAM T129681), unless stated otherwise: A, carapace, dorsal; B, left chelicera, dorsal; C, right pedipalp, dorsal; D, left chela, retrolateral; E, left movable finger, chelal teeth, retrolateral; F, left fixed finger, chelal teeth, retrolateral; G, left chela, retrolateral, protonymph (WAM T140482). Scale lines = 0.5 mm (Figs. C, D), 0.2 mm (Figs. A, B, G), 0.1 mm (Figs. E, F).

FIGURE 40. Indohya jessicae Harvey & Burger, n. sp., holotype female (WAM T133678), unless stated otherwise:A, carapace, dorsal; B, left pedipalp, dorsal; C, left chela, retrolateral; D, right movable finger, chelal teeth, retrolateral; E, right fixed finger, chelal teeth, retrolateral; F, right chela, retrolateral, tritonymph (WAM T133676); G, left chela, retrolateral, deutonymph (WAM T133677); H, left chela, retrolateral, protonymph (WAM T139291). Scale lines = 0.5 mm (Figs. A–C, F–H), 0.2 mm (Figs. D, E).

FIGURE 42. Indohya lynbeazleyae Harvey & Burger, n. sp., holotype male (WAM T91735): A, carapace, dorsal; B, right chelicera, dorsal; C, right pedipalp, dorsal; D, left chela, retrolateral; E, movable finger, chelal teeth, retrolateral; F, fixed finger, chelal teeth, retrolateral. Scale lines = 0.5 mm (Figs. C, D), 0.2 mm (Figs. A, B), 0.1 mm (Figs. E, F).

FIGURE 47. Indohya rixi Harvey & Burger, n. sp., holotype male (WAM T144193): A, carapace, dorsal; B, right pedipalp, dorsal; C, left chela, retrolateral; D, left fixed finger, chelal teeth, retrolateral; E, left movable finger, chelal teeth, retrolateral. Scale lines = 0.5 mm (Figs. B, C), 0.2 mm (Fig. A), 0.1 mm (Figs. D, E).

FIGURE 49. Indohya sagmata Harvey & Burger, n. sp., holotype male (WAM T138967), unless stated otherwise: A, carapace, dorsal; B, right pedipalp, dorsal; C, right chela, retrolateral; D, right fixed finger, chelal teeth, retrolateral; E, right movable finger, chelal teeth, retrolateral; F, left chela, retrolateral, tritonymph (WAM T111712). Scale lines = 0.5 mm (Figs. B, C, F), 0.2 mm (Fig. A). 0.1 mm (Figs. D, E).

FIGURE 51. Indohya scanloni Harvey & Burger, n. sp., holotype male (WAM T140479):A, carapace, dorsal; B, right pedipalp, dorsal; B, left chela, retrolateral; C, fixed chelal finger teeth, retrolateral; D, left fixed finger, chelal teeth, retrolateral. Scale lines = 0.5 mm (Figs. B, C), 0.2 mm (Figs. A, D).

FIGURE 55. Indohya currani Harvey & Burger, n. sp., holotype female (WAM T134915):A, carapace; B, right pedipalp, dorsal; C, right chela, retrolateral; D, right fixed finger, chelal teeth, retrolateral; E, right movable finger, chelal teeth, retrolateral. Scale lines = 0.2 mm (Figs. A–C), 0.1 mm (Figs. D, E).

FIGURE 57. Indohya finitima Harvey & Burger, n. sp., holotype adult (WAM T120197):A, carapace, dorsal; B, right pedipalp, dorsal; C, left chela, retrolateral; D, movable finger, chelal teeth, retrolateral; E, fixed finger, chelal teeth, retrolateral. Scale lines = 0.5 mm (Figs. A–C), 0.1 mm (Figs. D, E).

FIGURE 59. Indohya julianneae Harvey & Burger, n. sp., holotype female (WAM T127193), unless stated otherwise: A, carapace, dorsal, paratype male (WAM T142931); B, right pedipalp, dorsal; C, left chela, retrolateral; D, left fixed finger, chelal teeth, retrolateral; E, right chela, retrolateral, paratype tritonymph (WAM T152602). Scale lines = 0.2 mm (Figs. A–C, E), 0.1 mm (Fig. D).

FIGURE 60. Indohya karenae Harvey & Burger, n. sp., holotype female (WAM T126382), unless stated otherwise:A, carapace, dorsal (distorted); B, right pedipalp, dorsal; C, left chela, retrolateral; D, left movable finger, chelal teeth, retrolateral; E, left fixed finger, chelal teeth, retrolateral; F, right chela, retrolateral, paratype tritonymph (WAM T82510); G, left chela, retrolateral, paratype protonymph (WAM T153117). Scale lines = 0.2 mm (Figs. A–C, F, G), 0.1 mm (Figs. D, E).

FIGURE 62. Indohya sachsei Harvey & Burger, n. sp., holotype female (WAM T152600), unless stated otherwise:A, carapace, dorsal; B, right pedipalp, dorsal; C, left chela, retrolateral; D, left fixed finger, chelal teeth, retrolateral; E, left chela, retrolateral, protonymph (WAM T152601). Scale lines = 0.5 mm (Fig. B, C), 0.2 mm (Fig. A), 0.1 mm (Figs. D, E).