Psechrus Thorell, 1878

Psechrus Thorell 1878: 170 . [Type species: Tegenaria argentata Doleschall, 1857, transferred to Psechrus by Thorell (1878)]. Simon 1890: 80 (Transfer to Psechridae). Simon 1892: 226. Dalmas 1917: 324. Homann 1950: 66. Lehtinen 1967: 260, 383. Levi 1982: 118. Coddington 1990: 7. Griswold 1993: 539. Murphy and Murphy 2000: 264. Griswold et al. 2005: 37.

Lancaria Karsch 1879: 557 . [Type species: Tegenaria torva O. Pickard-Cambridge, 1869, transferred to Lancaria by Karsch (1879)]. Simon 1887: 194 (Syn., formal transfer of Lancaria torva to Psechrus).

Species transferred to other genera:

Psechrus nicobarensis Tikader, 1977 to Fecenia (Levi 1982: 138) .

Note: Fecenia nicobarensis (Tikader, 1977) is junior synonym of Fecenia protensa Thorell, 1891 (Bayer 2011) .

Diagnosis. Psechrus is distinguished from Fecenia by the following characters: AME at most equal to other eyes, mostly smaller; white longitudinal line ventrally on opisthosoma; clypeus (relatively) high, 2–3.5 times diameter of AME, consequently cephalic region of carapace higher than in Fecenia; length of leg IV more or less equal to leg II; males (mostly) without RTA or apophyses on other palp limbs (except femoral extension, see below) and always without median apophysis on tegulum; females with rather simple median septum of epigyne; vulva mostly with spherical spermathecal heads.

Description. Large Psechridae, body length in males: 8.5–23.8 mm; females: 11.0– 31.1 mm. Width of anterior part of carapace smaller than broadest section of carapace. Both anterior and posterior eye row recurved. Chelicerae long and strong, basal limb at least 2.5 times longer than broad. Cheliceral furrow with three teeth anteriorly, four to five posteriorly and a longitudinal row of 5–10 small denticles in between both rows of teeth (Fig. 66e). Their number varies intraspecifically. Basal limb of chelicerae ventrally with long field of short, transversal striae. Labium longer than broad (Fig. 2e). Gnathocoxae ca. 2.5 times longer than broad (Fig. 2e). Sternum longer than broad, with pointed posterior ending and broad-angled (160°) anterior ending (Fig. 2f). Pedipalp in females with single, toothed claw (Fig. 21g), which is similarly shaped as the pair of distal tarsal claws of the legs. The tarsi of the legs long, gracile, elastic, and apically additionally with a third, small, toothed and short tarsal claw (median hook). Legs very long in males (metatarsus I ca. 2.5–3.5x carapace length, Fig. 81c), long in females (metatarsus I ca. 2x carapace length, Fig. 81d). Leg formula 1243 or 1423. Coxae of legs I and II slightly broader than of IV and especially III, which is smallest. Calamistrum dorso-retrolaterally on metatarsus IV consisting of 4–5 rows of setae, inner rows generally with irregular arrangement. In adult males calamistrum rudimentary, if not completely reduced.

Spination of palp and legs: Variable within each species. Mostly no species-specific spination pattern recognisable. The spination pattern is in parts useful for the characterisation of the different species groups (see below). At the following positions spines are always absent: All patellae, tarsi, dorsal surface of all metatarsi, ventral surface of all femora. Ventral spines on tibiae and metatarsi generally paired, except for most distal one (in the case of odd numbers).

Males of many species possess macrosetae ventrally on coxae of legs I (MC-I) and/or II (MC-II) as well as an apical row of macrosetae ventrally on trochanter I (MT-I).

Colouration: Chelicerae brown to dark red-brown. Sternum mostly yellowish brown at lateral margins and with (dark) brown, tapered patch centrally (Fig. 82h). Rarely unicoloured light brown (Fig. 82j) and even more rarely brown with light longitudinal line (Fig. 82k). Carapace yellowish brown, always with two dark brown median bands (Figs 82a–g), which may be serrated laterally. Most distal lateral margins often also with dark bands (Figs 82a,f–g), however these are (much) narrower than median ones. Palpal femur ventrally with a longitudinal row of 6–10 long bristles, which may be reduced in adult males. Legs from yellowish brown or light brown to brown, often dark brown annulated (Figs 81a–d). Opisthosoma dorsally yellowish-brown with (dark) brown patches (Figs 81a–b), ventrally brown with a straight, light, longitudinal line (Figs 81e–i). On each side next to that line with a longitudinal row of small patches of muscle sigillae (Figs 81h–i). Opisthosoma ventrally very rarely unicoloured brown (Fig. 81j).

Anterior lateral spinnerets are short (broader than long) and more or less conical, posterior median spinnerets distinctly smaller, slender and cylindrical. Posterior lateral spinnerets also cylindrical and bipartite, similar size to anterior lateral spinnerets. Cribellum divided into two very narrow parts (Fig. 81h).

Copulatory organs: Male palp with more or less oval tegulum (T). Embolus (E) filiform (Figs 2b–c, 25b–c), broad and strong (Figs 10a–c, 54b), or of intermediate shape. E arising in retrolateral half of T or centrally. Conductor (C) mostly membranous, rarely sclerotised, rudimentary or completely reduced. C arising medially in upper half of T (Figs 2b, 70b). Expanded bulb clearly showing the large basal haematodocha (BH), surrounding subtegulum partly (Figs 1a–c). A median haematodocha, like present e.g. in Araneidae (Grasshoff 1968), Oecobius Lucas, Uroctea Dufour (Baum 1972), Liphistius Schiödte (Kraus 1978) or in Pisauridae (Sierwald 1987), is absent in Psechrus . Bulbal ligament (BL) and bulbal petiolus (BP) are visible through the BH (Figs 1a–b). Cymbium (slightly) broader than palpal tibia and patella or more or less equal in width. RTA usually absent, but a few species belonging to the mulu -group (see below) with a tibial process (TP) with a bunch of long setae (Figs 10b–c). It is not clarified if this process represents a RTA, if it is homologous to a RTA or if it is a completely different structure. A “regular” RTA usually lacks setae and is generally strongly sclerotised. Palpal femur may be modified (e.g. Figs 2d, 10d, 15d or 55d). The respective modifications may be species-specific. Cymbium dorsally mostly with scopula (CS) (Figs 83a–b, d–g). There are differences in density among the different species: very dense (Figs 83d–g) or moderate dense (Figs 83a–b). CS can be of different length (covering cymbium from 1/4 up to 6/7, Figs 83d–g). In a few species CS is absent (Fig. 83c).

Female epigyne mostly with simple septum (e.g. Fig. 29e). Anterior to septum mostly lots of curved wrinkles (Figs 29e, 32a). Vulva specifically shaped (see each species description) with internal duct system generally divided into three sections: copulatory duct, receptaculum with spermathecal head and fertilisation duct (Figs 2h, 29d).

Biology. The lace-sheet-weavers generally live in shady habitats near the ground. They can be found in forests (e.g. between tree roots, in holes of tree trunks or underneath dead wood), between rocks, boulder, at rock walls or at clay escarpments. Sometimes they appear in untended barracks or huts, too. Several species, especially those that are often found at rock walls or between boulders, can be found in the entrance areas of caves, too. According to colleagues’ and my own observations in the field in Laos, there are several species, Psechrus laos sp. nov., P. ancoralis Bayer & Jäger, 2010, P. antraeus Bayer & Jäger, 2010, P. khammouan Jäger, 2007 and P. steineri Bayer & Jäger, 2010, which prefer these habitats (rock walls etc., see above). Others, P. luangprabang Jäger, 2007, P. ghecuanus Thorell, 1897 and P. jaegeri sp. nov., are found mainly in forests, between roots or in tree holes or at escarpments. There are a few Psechrus species reported only from caves, e.g. P. mulu Levi, 1982, P. cebu Murphy, 1986 or P. steineri . However, they had mostly been reported from the entrance areas of caves (rarely from the aphotic zone). Presumably they can be found on rock walls or boulders outside of the caves, too. I would not assume that there is any Psechrus species that is restricted to caves. The lace-sheet weavers build a horizontal sheet web, which reaches up to 1.2 m length (in rare cases even 2 m [Robinson and Lubin 1979]). At one side the web turns into a long tube, which leads into a narrow crevice or hole, where the spider is safe from invaders. This tuberetreat is generally located in a rigid environment, e.g. rocks, stones, rigid soil or wood. It never appears between leaves and only very rarely between grass. Psechrus moves upside down underneath the sheet web like representatives of Linyphiidae do. Psechrus behaves extremely shyly and careful in its sheet web. At the slightest disturbance it runs back to its retreat with extreme speed. This explains why Psechrus, though it is abundant in many regions, is not easy to catch. Females carry their egg sacs, which can be up to 25 mm in diameter, in their chelicerae (Fig. 93b). According to my own observations in P. jaegeri sp. nov., P. argentatus (Doleschall, 1857), P. mulu Levi, 1982, P. ghecuanus Thorell, 1897 and according to Jäger (2007) for P. khammouan Jäger, 2007, egg sacs contain 70– 96 eggs. Yoshida (2009) counted 174 spiderlings in an egg sac of P. clavis sp. nov. (sub P. taiwanensis).

Robinson and Lubin (1979) observed and described the predatory behaviour of Psechrus argentatus (Doleschall, 1857) . Most of the behavioural units described therein, were also observed in my own trials using several P. laos sp. nov., P. luangprabang Jäger, 2007 and P. torvus (O. Pickard-Cambridge, 1869) specimens. Each spider was transferred into a glass terrarium with a piece of wood in one corner. A sheet web was built by the second day at the latest, with the retreat situated between the piece of wood and the corner of the terrarium. Crickets or large flies were used as prey, which were placed in the centre of the web. After a fly was put into the web, it took at most one second until the spider moved slightly forward in its retreat. A few seconds later it moved to the mouth of the retreat and pulled slowly at the web with the forelegs. Finally it ran out very fast to the respective site of the sheet web, grasped the fly with its forelegs and bit it. Immediately it ripped the fly out of the web with the chelicerae and ran back with it forward into the retreat. Later, it turned its direction within the retreat to be ready for the next prey attack (in some cases not before finishing up eating the fly). When larger prey items, e.g. large crickets, were offered, the behaviour was the same up to the point before grasping the prey. In this case the spider moved more slowly before directly encountering the prey. After a few attempts of stretching out and drawing back the forelegs over the cricket, it was bitten. The bite was mostly located between head and thorax or at the base of antennae and lasted for ca. 5 seconds. Then shorter bites at other sites of the cricket’s body (e.g. legs, antennae) followed. After ca. 1 minute the prey seemed to be paralysed and Psechrus began to bind its prey with threads produced from the spinnerets but also with threads taken from the web. Subsequently the prey was cut out of the web and carried within the chelicerae back to the retreat. The binding behaviour was not executed in every trial, but in ca. 50 % of prey attacks upon large prey items. The approaching behaviour in the terrarium is rapid and does not pass off stepwise as has been observed in the natural environment by Robinson and Lubin (1979) and myself. In the wild the webs are far larger in size and consequently prey are much more difficult to localise. The three Psechrus species examined showed no significant differences in web structure and predatory behaviour.

Mating behaviour was observed only once. In the respective trial a male of Psechrus luangprabang was released into the terrarium of a conspecific female (ca. 11:00 o’ clock a.m). It moved to an upper corner of the terrarium and stayed there for 4 ½ hours. At ca. 3: 30 p. m. the male slowly approached from a peripheral section of the web. The approach was interrupted by stops, from about 5–15 minutes duration. At each resting position the web was gently pulled rhythmically with the two pairs of forelegs. Meanwhile the female appeared at the mouth of the tube retreat. Finally the male reached the females legs and stroked them with his tarsi for about 5–10 minutes. Then he crawled underneath the female, both specimens facing the same direction. He surrounded her body with his long legs. They both swung violently up and down. Suddenly the male turned underneath the female and once again surrounded the latter with his legs. The spines on his legs, especially the femora, were erected to ca. 45°. The male turned again and pulled down the two first legs of the female with the metatarsus of his right leg I. Once again he turned and both spiders showed trembling movements. Then the male approached very closely to the female and a few seconds after that he departed again. This sequence was repeated about three times. Finally he moved his body perpendicular to the female and cleaned his two first legs and the palps. Both specimens trembled, even more intensely than before and the male’s spines erected to almost 90°. The male changed his position to a 45° angle towards the female and in this position he pulled her towards him and copulation took place. It was very rapid, lasting about 20 seconds. During copulation the first two pairs of the female’s legs and the second pair of the male’s legs were stretched straight forward. The expansion of the bulb and the exact position of insertion could not be observed with the naked eye. After copulation the male was chased away by the female and moved to an upper corner of the terrarium (Fig. 93a shows a Psechrus couple in the field observed by a colleague).

Species groups. The species groups are defined mainly by the basic structures of the copulatory organs of their representatives (see diagnoses of each species group below). A few somatic characters are in parts useful for such a “classification”, but often only by trend as there are exceptions in character patterns. These useful characters are: 1) the shape of median and lateral bands on carapace, 2) the shape of the light, longitudinal line ventrally on opisthosoma, 3) the relative leg length (measured as ratio between femur I + metatarsus I / carapace length), 4) dorsal spines on tibia III and IV. The length of legs is variable among different specimens of the same sex, which is the case for every species. It will be noted as an approximate ratio for males and females in the description of each species-group, with the following convention: Ratio between femur + metatarsus of leg I / carapace length (FEM-I + MTT-I / CL). Presently, eight groups, the argentatus-, mulu-, annulatus-, singaporensis-, ancoralis-, himalayanus-, sinensis-, and the torvus- group are differentiated.

Key to species of Psechrus:

1 Male [unknown in: borneo, annulatus, aluco sp. nov., norops sp. nov., arcuatus sp. nov., jinggangensis, fuscai sp. nov., kenting, taiwanensis, tauricornis sp. nov.; identification not absolutely certain in: demiror sp. nov., zygon sp. nov.; those of demiror sp. nov. and zygon sp. nov. are included in the present key, but with question mark].......................... 2

- Female [unknown in: ulcus sp. nov., kinabalu, schwendingeri sp. nov.]......................................... 37

2 Harpago (Figs 70b–c, 72b–c, 74b–c) present................................................................ 3

- Harpago absent....................................................................................... 5

3 C longer than width of T........................................................................... torvus

- C shorter than width of T............................................................................... 4

4 C centrally as broad as in distal fourth and located medially in upper half of T....................... hartmanni sp. nov.

- C broadest in distal fourth and located in retrolateral half of T..................................... zygon sp. nov. (?)

5 C absent or strongly reduced............................................................................ 6

- C well developed...................................................................................... 9

6 C absent............................................................................................ 7

- C rudimentary (a very short and stout structure still recognisable), E quite broad and strongly sclerotised, resting in CA (Figs 78a–c)............................................................................ schwendingeri sp. nov.

7 With three apophyses close to E (Figs 7a–b)............................................................ mulu

- With less than three apophyses close to E.................................................................. 8

8 E retrolaterad; E and EB constitute an extremely bulky structure (Fig. 10b).............................. ulcus sp. nov.

- E rather slim, prolatero-apicad..................................................................... kinabalu

9 C with numerous small or very small, short spines or tubercles (Figs 52a–b, 54a–b, 57a–b, 66a–b).................... 31

- C without such structures.............................................................................. 10

10 Palpal femur modified with ventral bulge (the latter may be flat) (Figs 14d, 15d) or with pointed, ventral extension (Fig. 2d). ................................................................................................... 11

- Palpal femur without modification....................................................................... 16

11 E dorsally with one distinct, pointed apophysis (Fig. 79b)................................................... cebu

- E different.......................................................................................... 12

12 C strongly sclerotised and narrow, its distal half just as broad as E (Figs 2b, 6b)................................... 13

- C membranous and/or fleshy, its distal half (distinctly) broader than E........................................... 14

13 C more than half as long as E........................................................................ libelti

- C less than half as long as E..................................................................... argentatus

14 C ca. as long as T....................................................................... decollatus sp. nov.

- C far shorter than T................................................................................... 15

15 EB in ventral view in alignment with upper retrolateral margin of T (Fig. 15b)........................... singaporensis

- EB protruding beyond upper retrolateral margin of T (Fig. 17b)..................................... elachys sp. nov.

16 Bulb with elongated EB possesing a distinct ventral protrusion basally (Figs 76a–c).................... crepido sp. nov.

- Bulb different....................................................................................... 17

17 E arising medially on upper half of T, coxa of leg I (Figs 82l,r) or proximal part of palpal femur (Fig. 35d) ventrally with distinct field of macrosetae............................................................................... 23

- E arising retrolaterally on T, neither coxa of leg I nor proximal part of palpal femur ventrally with distinct field of macrosetae (may be with few unconspicuous macrosetae subdistally in addition to an apical row of macrosetae)................... 18

18 C with distinct, broadened base (Figs 23b, 25b)............................................................ 19

- C without distinct, broadened base....................................................................... 20

19 E longer than T.............................................................................. laos sp. nov.

- E shorter than T..................................................................................... rani

20 C broader than 1/3 the width of T................................................................. ancoralis

- C narrower than 1/3 the width of T....................................................................... 21

21 E longer than width of T......................................................................... antraeus

- E shorter than width of T............................................................................... 22

22 E longer than half the width of palpal tibia........................................................ khammouan

- E shorter than half the width of palpal tibia............................................................ steineri

23 E (almost) straight.................................................................................... 26

- E curved........................................................................................... 24

24 EB with particular flat, elongated and proximally curved extension (Fig. 43b)............................ luangprabang

- EB different......................................................................................... 25

25 Distal section of E curved and distinctly narrower than central section (Fig. 50f)....................... jaegeri sp. nov.

- Distal section of E hardly curved and not significantly narrower than central section (Fig. 50e).............. vivax sp. nov.

26 C as long as T......................................................................... demiror sp. nov. (?)

- C shorter than T...................................................................................... 27

27 T apically with quite strongly sclerotised, semicircular extension (Figs 37b, 39g, 41b).............................. 28

- T apically without such an extension..................................................................... 29

28 E arising distinctly further distally than C.......................................................... ghecuanus

- E arising at most at the same level as C, but mostly further proximally.............................. pakawini sp. nov.

29 Sperm duct simply U-shaped................................................................... himalayanus

- Sperm duct with at least two loops....................................................................... 30

30 C with broad and large proximal section and small and narrow distal section (Fig. 35b).................. inflatus sp. nov.

- C broadest distally............................................................................ marsyandi

31 C apically (at least slightly) bifid (Figs 56a–c, 57a–c)........................................................ 32

- C apically not bifid................................................................................... 33

32 EB with apophyses only in its distal half (Figs 55b–c, 56b–c)......................................... tingpingensis

- EB with apophyses only in its proximal half (Figs 57b–c)........................................ obtectus sp. nov.

33 C in ventral view proximally with striking, hemispherical bulge (Fig. 62b)............................... senoculatus

- C without such a bulge................................................................................ 34

34 Sperm duct with distinctly U-shaped section in prolateral half of T (Figs 66b)........................... clavis sp. nov.

- Sperm duct different (e.g. as in Figs 52b, 54b, 86m,o)....................................................... 35

35 EB with long, apically bifurcated apophysis (Fig. 86n).............................................. kunmingensis

- EB without long, apically bifurcated apophysis............................................................. 36

36 E in ventral view short, broad and blunt (Fig. 54b)................................................... triangulus

- E in ventral view with filiform distal section (Fig. 52b)................................................... sinensis

37 Epigyne complex, tegimentum (TM) present (Figs 69a,d, 71a,d, 73a,e, 75a,e, 89m –p).............................. 38

- Epigyne without TM................................................................................. 41

38 Branches of TM distally rounded (Figs 69a, 73a)............................................................ 39

- Branches of TM distally pointed, resembling the horns of a bull (Figs 75a,e)........................ tauricornis sp. nov.

39 Branches of TM directed anterio-mediad............................................................... torvus

- Branches of TM directed (anterio-) laterad................................................................ 40

40 Distal section of CD located medial to SH....................................................... zygon sp. nov.

- Distal section of CD located lateral to SH.................................................... hartmanni sp. nov.

41 Epigyne with flat, large-area bulge in front of MS (Figs 77a,e, 89k)................................. crepido sp. nov.

- Epigyne without such a bulge in front of MS............................................................... 42

42 MS longer than broad................................................................................. 43

- MS broader than long................................................................................. 51

43 Anterior half of MS broader than posterior half (Figs 61a, 63a, 65a, 67a)......................................... 44

- Posterior half of MS broader than anterior half (Figs 53a, 56d, 60a,e)........................................... 48

44 CD with twisted section (Fig. 61b), the latter narrower than spermatheca............................... jinggangensis

- CD with bulbous section, the latter broader than spermatheca................................................. 45

45 MS distinctly longer than broad (more than 1.6 times, Fig. 63a), initial section of CD straight (Fig. 63b)........ senoculatus

- MS less than 1.6 times longer than broad, initial section of CD either integrated within bulbous section (Fig. 67b) or anteriorly curved mediad (Figs 64b, 65b).......................................................................... 46

46 Initial section of CD integrated in kidney-shaped, bulbous section.................................... clavis sp. nov.

- Initial section of CD anteriorly curved mediad.............................................................. 47

47 Posterior half of MS just slightly narrower than anterior half (Fig. 65a), bulbous section of CD anteriorly with small bulge (Fig. 65b)....................................................................................... taiwanensis

- Posterior half of MS clearly narrower than anterior half (Figs 64a,d), bulbous section of CD anteriorly without small bulge .................................................................................................. kenting

48 CD with twisted section (Figs 55f, 58b, 59b)............................................................... 51

- CD without twisted section (Figs 53b, 54f, 60b,d)........................................................... 49

49 MS and LL complicatedly folded interleaved (Figs 60a,e)............................................ kunmingensis

- MS and LL different.................................................................................. 50

50 Posterior half of MS anteriorly rounded (Figs 53a,d)..................................................... sinensis

- Posterior half of MS triangular (Fig. 54e)............................................................ triangulus

51 Epigyne anteriorly with two rounded, flattened, glossy fields near CO (Figs 59a, 89e).................... fuscai sp. nov.

- Epigyne without rounded, flattened, glossy fields near CO.................................................... 52

52 CD at least 4x longer, but less than 2x broader than diameter of receptaculum.................................... 53

- CD different........................................................................................ 54

53 Twisted sections of CD, located anteriorly beyond transversal, initial sections of CD, shorter than half the diameter of one receptaculum (Figs 58b,g).................................................................. obtectus sp. nov.

- Twisted sections of CD, located anteriorly beyond transversal, initial sections of CD, almost as long as diameter of one receptaculum (Figs 55f, 56f)....................................................................... tingpingensis

54 CD with voluminous, spherical/bulbous section, the latter distinctly larger than receptaculum (Figs 64b,e, 65b).......... 47

- CD may be large and broad (Figs 2h, 5c), but without voluminous, spherical/bulbous section......................... 55

55 Epigyne and EF strongly sclerotised, with dark red-brown colour (Figs 87c,d).................................... 56

- At least EF different.................................................................................. 57

56 CD running transversally and straight (Figs 7e, 8f)........................................................ mulu

- CD curved and flowing into receptaculum from anterior (Fig. 9b).......................................... borneo

57 Epigyne protruding, CO large (Figs 12a,b, 13a, 14f, 87e–g)................................................... 58

- Epigyne not or hardly protruding, CO rather small.......................................................... 60

58 Vulva with spherical SH............................................................................... 59

- Vulva without spherical SH (Fig. 14h)....................................................... decollatus sp. nov.

59 CD distinctly larger than receptaculum (Fig. 12c)..................................................... annulatus

- CD not larger than receptaculum (Fig. 13b, 90f).................................................. aluco sp. nov.

60 CD very large, broad and flat (CD at least 5x larger than receptaculum, Figs 2h, 5c, 20b)............................ 61

- CD not or not distinctly larger than receptaculum........................................................... 63

61 CO located anterior to SH, receptacula round................................................... arcuatus sp. nov.

- CO located posterior to SH, receptacula cross-oval.......................................................... 62

62 MS at most half as long as broad and with notches laterally (Figs 2g, 3i).................................. argentatus

- MS not distinctly broader than long, with continuous lateral margins (Figs 5a–b)............................... libelti

63 Lateral, dark bands on carapace clearly broader than diameter of PME (Fig. 82a, f–g), receptacula remarkably round (Figs 15f, 17e), CD curved medially (Figs 15h,f, 19b)................................................................ 64

- At least one of the features listed in item 63 different........................................................ 68

64 CD ca. 4–5x larger than receptaculum........................................................ arcuatus sp. nov.

- CD at most 2x larger than receptaculum................................................................... 65

65 Anterior margins of LL strongly diverging (Fig. 80a; Attention: anterior margins of LL may be confound with the anterior margins of MS, but the latter are short and pointing more or less anteriad)......................................... cebu

- Anterior margins of LL not or hardly diverging............................................................. 66

66 SH located upon receptacula................................................................. norops sp. nov.

- SH (still) located upon CD............................................................................. 67

67 CO small, pointing anteriorly (Fig. 17d), penetration of E happens frontally.......................... elachys sp. nov.

- CO elongated (Fig. 15e,i), penetration of E happens laterally......................................... singaporensis

68 Lateral bands on carapace quite broad (1.5–2.5x diameter PME)............................................. cebu

- Lateral bands on carapace narrow to medium-sized (at most 1.2x diameter PME).................................. 69

69 MS mushroom-like shaped (Figs 49a, 51a)................................................................ 70

- MS differently shaped................................................................................. 71

70 Anterior half of MS more than 2.5x broader than posterior section................................... jaegeri sp. nov.

- Anterior half of MS less than 2.5x broader than posterior section...................................... vivax sp. nov.

71 Lateral margins of MS anteriorly diverging................................................................ 72

- Lateral margins of MS anteriorly converging............................................................... 73

72 Lateral margins of MS strongly diverging anteriorly (Fig. 30e), CD at most 1.5x longer than receptaculum.......... steineri

- Lateral margins of MS moderately diverging anteriorly (Figs 29a,e), CD 2.5–3x longer than receptaculum...... khammouan

73 MS ca. 2x broader than long, its anterior margins running transversally mediad, parallel with anterior margins of LL (Figs 32a, 34a, 88h–i).......................................................................................... 74

- MS and/or anterior margins of LL different................................................................ 75

74 CD shorter than distance in between both CO....................................................... marsyandi

- CD longer than distance in between both CO....................................................... himalayanus

75 MS more than 2x broader than long................................................................ ancoralis

- MS less than 2x broader than long....................................................................... 76

76 Epigyne without EF................................................................................... 77

- Epigyne with EF..................................................................................... 79

77 Initial parts of CD clearly located anterior to spermathecae (Fig. 27f)....................................... antraeus

- Initial parts of CD located posterior to spermathecae or (at most) at the same level................................. 78

78 Helical section of spermatheca high, with more than three windings (Figs 25d, 26d)........................ laos sp. nov.

- Helical section of spermathecae not or just slightly higher than broad, with less than two windings (Fig. 24b).......... rani

79 Spermathecae high, with at least three helical windings (Figs 25d, 26d)................................. laos sp. nov.

- Spermathecae compact, with at most one helical winding..................................................... 80

80 Anterior margins of LL constitute strongly sclerotised clasps (Figs 44a, 47a, 88l–m), CD longer than diamter of one spermatheca 81

- Anterior margins of LL rather inconspicuous, CD shorter than diameter of one spermatheca......................... 82

81 CD ca. two times longer than diameter of one spermatheca (Fig. 47b)................................ demiror sp. nov.

- CD at most 1.5x longer than diameter of one spermatheca (Fig. 44b, 45b)............................... luangprabang

82 SH very flat, hardly protruding out of spermathecae.............................................. inflatus sp. nov.

- SH regularly spherical, protruding out of spermathecae....................................................... 83

83 CD extending medially (clearer in frontal view) (Figs 41f,h)...................................... pakawini sp. nov.

- CD constitutes one compact structure together with spermathecae (Fig. 37e,g)............................. ghecuanus argentatus- group