Troglokhammouanus steineri Lourenço, 2007

Prendini, Lorenzo, Ehrenthal, Valentin L. & Loria, Stephanie F., 2021, Systematics Of The Relictual Asian Scorpion Family Pseudochactidae Gromov, 1998, With A Review Of Cavernicolous, Troglobitic, And Troglomorphic Scorpions, Bulletin of the American Museum of Natural History 2021 (453), pp. 1-153 : 69-80

Troglokhammouanus steineri Lourenço, 2007

Figures 3, 5, 7–9, 11, 13B, D, 15B, 16C, D, 18C, D, 20C, D, 26–31, tables 1–5, 7, 8

Troglokhammouanus steineri Lourenço, 2007a: 771 , 775, 776, figs. 1, 3 (4–15), 4 (16–24); 2007b: 1–6, figs. 1–24; Lourenço and Pham, 2010: 2, 12; Botero-Trujillo and Noriega, 2011: 42; Jäger, 2012: 37; Lourenço, 2012a: 233, 236, 237, appendix A; Lourenço and Leguin, 2012: 71; Lourenço and Pham, 2012: 80, 84; Soleglad et al., 2012: 89; Steiner, 2013: 416; Loria and Prendini, 2014: appendix S1: 10, appendix S2: 7; Lourenço, 2014: 31; Beron, 2015: 184; Sharma et al., 2015a: 354; 2015b: 4, 7, figs. 1e, 2c, 3b; Lourenço, 2017a: 19–21, 23, fig. 1; Beron, 2018: 834; Lourenço et al., 2018: 265; Ballesteros et al., 2019: table S1; Francke, 2019: 32; Howard et al., 2019: 76, table 1; Santibáñez-López et al., 2019b: 26, table 3.

Troglokhammouanus louisanneorum Lourenço, 2017a: 20–23 , figs. 1, 4–16; syn. nov.

TYPE MATERIAL: LAOS: Khammouane Prov.:

Boualapha Distr.: Hin Namno National Biodiver-

sity Conservation Area: Holotype ♀ ( MNHN RS 9098) [examined], juv. ♂ paratype ( MNHN RS 9099) Boualapha, Tham Xe Bangfai (cave), Ban Nong Ping [17°22′23.6″N 105°50′11.8″E], 14.ii.2007, H. Steiner, midsection of cave [2–3 km from entrance], on sand [alluvial soil and sand bank next to river edge]. Gnommalath Distr.: Troglokhammouanus louisanneorum : Holotype ♀ ( MNHN), Ban Naden, Tham Nam Lot cave (17°30′17.9″N 105°23′08.2″E), 176 m, 8.xi.2011, L. Deharveng and A. Bedos, 350–450 m from main cave entrance.

DIAGNOSIS: As for subfamily.

DESCRIPTION: The following description, based on the holotype female of T. steineri , and material examined, including 10 adult males and 61 adult females, supplements the original descriptions of Lourenço (2007a, 2017a) and supplemental data in Lourenço (2007b).

Total length: Small, 32.2 mm (28.4–36.4 mm, n = 9) (♂) or 38.5 mm (35.8–41.3 mm, n = 13) (♀) (tables 3, 7, 8).

Color: Tegument base coloration reddish yellow to reddish brown. Carapace diffusely infuscate, with brownish marbled pattern; tergites with paired submedial and sublateral stripes of brownish infuscation longitudinally, or immaculate; other surfaces immaculate. Carapace, pedipalps, tergites, and metasoma reddish yellow to reddish brown. Chela fingers, pedipalpal and metasomal carinae dark reddish brown. Chelicerae, legs, sternum, genital operculum, and sternites pale yellow. Pectines pale, yellowish white. Telson paler than metasomal segment V. Aculeus black.

Chelicerae: Fixed finger, dorsal margin with four teeth (basal, median, subdistal, distal); basal and median teeth fused into bicuspid (“conjoined on trunk”); space between median and subdistal teeth U-shaped; ventral margin with five or six small denticles (ventral accessory denticles). Movable finger, dorsal margin with three teeth (median, subdistal, retrolateral distal), without basal teeth; ventral margin with five or six small denticles (ventral accessory denticles) and serrula in distal third; retrolat- eral (dorsal) distal and prolateral (ventral) distal teeth subequal, retrolateral (dorsal) distal tooth smaller than prolateral (ventral) distal tooth, and opposable. Ventral surface of fingers and manus with numerous long, dense macrosetae.

Carapace: Carapace slightly longer than wide, anterior width 51% (43%–55%, n = 9) (♂) or 50% (45%–57%, n = 13) (♀) of length, posterior width 90% (85%–94%, n = 9) (♂) or 95% (79%–102%, n = 13) (♀) of length (tables 3, 7, 8). Anterior margin convex (procurved), protruding medially; anterolateral margins entire; posteromedian margin shallowly concave (recurved); posterolateral margins gently curved (fig. 11). One pair of very small anterolateral major (ALMa) ocelli situated close to anterolateral margins of carapace, other lateral ocelli absent. Anterosubmedial depressions, medial to lateral ocelli, well developed. Median ocular tubercle situated anteromedially on carapace, distance from anterior margin 32% (26%– 37%, n = 9) (♂) or 33% (30%–35%, n = 11) (♀) of carapace length (tables 7, 8); pair of median ocelli considerably larger than lateral ocelli; pair of weakly developed, smooth superciliary carinae, not protruding above ocelli; interocular sulcus shallow, obsolete. Circumocular sutures complete but disconnected posteriorly; circumocular triangle subtriangular (broad V-shape); median ocular curvatures present. Anteromedian depression narrow, shallow; posteromedian sulcus very shallow anteriorly, becoming slightly deeper posteriorly; posterolateral sulci very shallow, wide, weakly curved; posteromarginal sulcus narrow, shallow. Carapace acarinate; surfaces finely to moderately granular, except on anteromedian depression and along lateral and posterior margins.

Pedipalps: Pedipalps relatively robust; segments almost apilose, sparsely covered in short microsetae and occasional macrosetae. Pedipalp femur length 68% (65%–71%, n = 9) (♂) or 64% (59%–68%, n = 13) (♀) greater than width (tables 3, 7, 8). Femur with seven carinae evident (fig. 28A–D); promedian carina obsolete, reduced to few spiniform granules in proximal two-thirds of segment; prodorsal, proventral and retrodorsal carinae well developed, costate granular; dorsomedian carina moderately developed, granular; retromedian and retroventral carinae vestigial, reduced to few granules proximally; ventromedian carina absent; intercarinal surfaces smooth. Pedipalp patella length 62% (59%–70%, n = 9) (♂) or 61% (58%–62%, n = 13) (♀) greater than width (tables 3, 7, 8). Patella with six carinae evident (fig. 28E–G); retrodorsal, retroventral, and proventral carinae well developed, costate granular to costate; prodorsal carina well developed, granular to costate granular; prolateral surface, dorsoventral “vaulted” projection (“anterior process”) moderately developed, with prominent pair of dorsal and ventral spiniform granules (“patellar spurs”) proximally, the dorsal more strongly developed, demarcating pair of obsolete, granular prolateral dorsal and ventral carinae (“patellar spur carinae”); ventromedian carina vestigial, reduced to proximal granules; dorsomedian and retromedian carinae absent; intercarinal surfaces smooth. Pedipalp chela relatively short and broad; manus globose, width 28% (21%– 32%, n = 9) (♂) or 25% (18%–33%, n = 13) (♀) greater than height and length 14% (8%–19%, n = 9) (♂) or 21% (17%–26%, n = 11) (♀) greater than width (tables 3, 7, 8); length of movable finger 16% (8%–24%, n = 9) (♂) or 24% (16%–35%, n = 11) (♀) greater than length of manus. Chela with 10 carinae evident (figs. 29, 30); dorsomedian carina complete, granular; dorsal secondary and subdigital carinae vestigial, reduced to granules proximally on manus; digital carina well developed, costate granular; retromedian (retrolateral secondary) carina partial, comprising granular row medially; secondary accessory and retroventral carinae incompletely fused, slight disjunction evident in proximal third; retroventral carina well developed, granular, aligned parallel to longitudinal axis of chela, with distal margin connected to retrolateral movable finger condyle; ventromedian carina partial, comprising granular row in proximal quarter; prodorsal, promedian, and proventral carinae obsolete, each reduced to series of isolated granules; intercarinal surfaces smooth except for coarse, scattered granules on prolateral surface of manus, near base of fixed and movable fingers. Fixed and movable fingers, dentate margins sublinear (♀) or markedly sinuous, movable finger with shallow lobe, proximally and fixed finger with correspondingly shallow notch, proximally (♂), no proximal “gap” evident when closed; median denticle rows comprising seven (6 or 7, n = 143) (fixed finger) or eight (6–8, n = 141) (movable finger) oblique and slightly imbricated subrows (tables 3, 4, 7, 8), decreasing in length distally; each subrow comprising large retrolateral denticle proximally (absent from proximal subrow, for total of seven retrolateral denticles on fixed and movable fingers), several small median denticles, and large prolateral denticle distally, slightly offset (total of seven and eight prolateral denticles on fixed and movable fingers, respectively); terminal (prolateral) denticle of first subrow enlarged; accessory denticles absent.

Trichobothria: Orthobothriotaxic, Type D, β configuration, trichobothrium d 2 situated on femur dorsal surface, d 3 and d 4 in same axis, parallel and closer to retrodorsal carina than d 1, angle formed by d 1, d 3 and d 4 opening toward prolateral surface, with the following segment totals (figs. 28–30): femur, 12 (6 dorsal, 3 prolateral, 3 retrolateral); patella, 10 (3 dorsal, 1 prolateral, 6 retrolateral); chela, 13 (5 manus, 8 fixed finger). Total number of trichobothria per pedipalp, 35. Five trichobothria on femur, i 4, d 1, d 2, d 4, and d 6, one on patella, est 1, and one on chela fixed finger, ib 2, noticeably smaller than others (“petite”). Trichobothrium Et 1 situated distally on manus, aligned with or proximal to movable finger condyle; eb situated on fixed finger, slightly distal to movable finger condyle; esb 2 situated in proximal third of fixed finger, approximately aligned with proximal retrolateral denticle of median denticle row.

Legs: Leg I maxillary lobes (coxapophyses), distal margins rounded, unmodified (not spatulate or dilate) anteriorly, terminating flush with lobes of leg II (figs. 26B, 27B, D). Legs I and II tibiae, retrolateral margins each with scattered macrosetae, without spurs (fig. 15B); III and IV with spurs, smooth, without spinelets. Basitarsi with few scattered macrosetae, pro- and retrolateral rows of long spinules, and pair of pro- and retrolateral pedal spurs. Telotarsi with pair of ventrosubmedian rows of long spinules; proventral and retroventral rows of macrosetae absent, only few scattered macrosetae laterally; laterodistal lobes reduced and truncated; median dorsal lobe very short; ungues short, distinctly curved, equal in length; unguicular spur (dactyl) pronounced, pointed.

Sternum: Sternum, Type 1, pentagonal, posterior width 89% (80%–104%, n = 9) (♂) or 86% (79%–97%, n = 11) (♀) of length (tables 7, 8), lateral margins recurved medially, ventral surface shallowly concave, posteromedian depression round (fig. 13B, D).

Pectines: Three marginal (anterior) lamellae, proximal sclerite considerably longer, distal sclerite short (fig. 13B, D); 15/14 (13–16, n = 92) (♂) or 13/13 (12–14/12–15, n = 190) (♀) median lamellae (tables 3, 5, 7, 8); fulcra present, small. Proximal median lamella and basal pectinal tooth unmodified. Pectinal teeth present along entire posterior margin of each pecten (♂, ♀); 16/15 (14–17, n = 94) (♂) or 14/14 (13–15/13– 16, n = 190) (♀) teeth. Pectinal peg sensillae long, cylindrical, tubular or bottle shaped, and rounded distally, without pair of laterodistal processes; sensillar sockets smooth.

Genital operculum: Opercular sclerites completely separated, prominent genital papillae visible along entire length (♂) or absent (♀) (fig. 13B, D). Genital plugs observed in some females.

Male reproductive organs: Paraxial organ without accessory glands. Hemispermatophore lamelliform, pars recta absent (fig. 31); basal lobe present.

Female reproductive organs: Ovariuterus comprising reticulate mesh of six cells. Oocytes contained in sessile follicles directly contacting ovaruterine tubes.

Mesosoma: Pretergites surfaces smooth and shiny. Posttergites I–VI surfaces moderately to coarsely granular, granulation becoming slightly coarser near posterior margins, acarinate, each with pair of shallow submedian depressions (figs. 26A, 27A, C); VII surface sparsely granular, with pair of costate granular dorsosubmedian and dorsolateral carinae reaching posterior margin of segment. Sternites III–VII surfaces entirely smooth, acarinate (figs. 26B, 27B, D), with scattered macrosetae, posterior margins with sparse row of macrosetae; III–VI, respiratory spiracles (stigmata) small, oval (fig. 13B, D); V posteroventral margin with pale, raised, triangular surface; VII width 41% (34%–47%, n = 9) (♂) or 41% (36%– 46%, n = 11) (♀) greater than length (tables 7, 8).

Metasoma: Metasoma relatively short, total length 24% (16%–28%, n = 9) (♂) or 18% (13%–32%, n = 11) (♀) greater than combined length of prosoma and mesosoma (tables 7, 8); segments I–V progressively increasing in length, I and II wider than long, width of length for segment I, 126% (118%–136%, n = 9) (♂) or 138% (124%–156%, n = 13) (♀); II, 101% (92%–109%) (♂) or 107% (96%–118%) (♀); III, 88% (76%–97%) (♂) or 93% (88%–100%) (♀); IV, 65% (59%–68%) (♂) or 68% (64%–74%) (♀); and V, 35% (32%–37%) (♂) or 36% (34%– 39%) (♀). Metasoma almost apilose, sparsely covered in short microsetae. Eight carinae on segment I, 10 on II–IV, and nine on V (figs. 16C, D, 18C, D, 20C, D). Dorsosubmedian carinae well developed, costate granular throughout length of segments I–IV, absent on V; converging posteriorly. Dorsolateral carinae well developed, costate granular throughout length of segments I–V; converging posteriorly on I and V, subparallel on II–IV. Dorsosubmedian and dorsolateral carinae of segments I–IV each terminating posteriorly with enlarged, spiniform granule. Median lateral carinae complete, costate granular throughout length of segments I–V. Ventrolateral carinae well developed, costate granular, but becoming obsolete anteriorly on segments I and II, continuous throughout length of III–V; converging posteriorly on I and V, subparallel on II–IV. Ventrosubmedian carinae absent on segment I, obsolete, costate on II, well developed, costate granular throughout length of III–V; subparallel on II and III, converging posteriorly on IV and V. Ventromedian carina moderately developed, granular, on segment V. Intercarinal surfaces smooth.

Telson: Telson large, suboval (figs. 16C, D, 18C, D, 20C, D). Vesicle elongate, width 105% (101%–112%, n = 9) (♂) or 106% (97%–117%, n = 13) (♀) of metasomal segment V width (tables 3, 7, 8), with flattened dorsal surface and rounded ventral surface, height 37% (32%–44%, n = 9) (♂) or 39% (34%–47%, n = 13) (♀) of length; dorsal surface smooth, lateral surfaces with three pairs of obsolete, granular carinae, ventral surfaces with two pairs of obsolete, granular carinae; anterodorsal lateral lobes (“vesicular tabs”) distinct, each with two spiniform granules; subaculear tubercle absent. Aculeus short, shallowly curved, 23% (21%–27%, n = 9) (♂) or 24% (21%–32%, n = 13) (♀) of telson length. Venom glands thin walled, simple and unfolded.

Intraspecific variation: One juvenile was completely depigmented and one adult female partially depigmented, including the ocelli (figs. 11E, 27A, B). No other significant variation was noted.

Ontogenetic variation: Immature stages of T. steineri are paler in color and less sclerotized than adults, and the posteroventral margin of mesosomal sternite V does not exhibit a pale, raised, triangular surface as in the adults of both sexes. Males and females closely resemble one another externally until the final instar. However, immature stages are easily sexed by examination of the genital aperture.

Sexual dimorphism: The genital papillae, visible the entire length and thus completely separating the two sclerites of the genital operculum (fig. 13B), are the characters of primary external sexual dimorphism in the male. The opercular sclerites are also completely separated in the female, although genital papillae are absent (fig. 13D). Adult males may be further separated from adult females and immature stages based on several secondary sexual characters. The most obvious are the sexually dimorphic pedipalp chelae, the fixed and movable fingers of which are markedly sinuous in the adult male (figs. 29, 30), as well as the more developed pectines, the median lamellae of which are more numerous and the teeth somewhat larger, and more numerous (fig. 13B, D; tables 3, 5, 7, 8). In addition, the adult male is proportionally more slender (fig. 26), with a slightly longer metasoma and telson (tables 7, 8), and slightly more pronounced granulation on the carapace, pedipalps, and tergites than the adult female (fig. 27). The pale, raised, triangular surface on the posteroventral margin of sternite V is more pronounced in the adult male (fig. 26B) than the adult female.

DISTRIBUTION: Endemic to the Boualpha District of Khammouane Province, Laos, where it has been recorded from a single cave, Tham Xe Bangfai (also known as Xe Bangfai River Cave or Tham Khoun Xe), in the Hin Namno National Biodiversity Conservation Area, near Nong Ping (fig. 7). Two expeditions to the putative type locality of T. louisanneorum, Tham Nam Lot (Lod) , near Ban Naden, in the Gnommalath District, failed to confirm the presence of this species.

ECOLOGY: Specimens collected by the first author at Tham Xe Bangfai were found with UV light detection, from the twilight zone to approximately 3 km from the downstream entrance (resurgence) (fig. 3B–D). Whereas specimens were observed in the dark zone during daytime, specimens were only observed in the twilight zone at night. It is not known how far the species extends into the river cave, which is approximately 11 km long, but the greatest abundance of individuals was observed between 300–500 m from the entrance, and their abundance appeared to decrease with increasing distance (more than 500 m upstream). Although a few specimens were collected on flowstone rock faces, the vast majority were collected close to the water, on moist muddy riverbanks at several “beaches” within the cave (fig. 3E). Most were collected in cracks between the drying mud slabs or among sticks and other vegetative debris trapped between rocks, almost always on moist soil (fig. 3F–H). None were taken on the dry sand banks or dry rock faces higher upslope. Although these moist areas presumably harbor a greater abundance of prey, the apparent preference of T. steineri for moist substrates, also observed in the epigean P. mischi and P. ovchinnikovi (Fet et al., 2004; Prendini et al., 2006; Soleglad et al., 2012), strongly suggests an ecological requirement for moisture.

Although no specimens of T. steineri have thus far been collected outside the cave, the morphology of this species suggests it is troglophilous rather than troglobitic ( Prendini, 2001b; Volschenk and Prendini, 2008). The ocelli, especially the median pair, are well developed, as in some epigean scorpions from low light environments, e.g., rainforests ( Loria and Prendini, 2021), the tegument of most individuals does not lack pigmentation (indeed, the base coloration is darker than in the two epigean species of Pseudochactas ), and there is no obvious attenuation of the pedipalps or legs, as in Vietbocapinae . It is conceivable that T. steineri is restricted to the cave interior due to its requirement for high humidity rather than for darkness per se.

Two buthids, Lychas aberlenci Lourenço, 2013 , and Lychas mucronatus (Fabricius, 1798) were collected in the deciduous forest surrounding the cave entrance. The remains of a scorpionid, Heterometrus laoticus Couzijn, 1981 , presumably washed in by the Xe Bang Fai River, were collected inside.

CONSERVATION STATUS: Troglokhammouanus steineri is very abundant in Tham Xe Bang Fai, which is very large (ca. 11 km long), relatively undisturbed, and situated within a protected area, the Hin Namno National Biodiversity Conservation Area. Due to difficult access, the surrounding forest is relatively intact, except for seasonal subsistence cultivation. Aside from an increase in disturbance associated with recreational ecotourism (e.g., caving and kayaking) in the cave, which could become a threat if unchecked, there appear to be few immediate threats to the survival of this species. Nevertheless, it is characterized by an acute restriction in both its area of occupancy and number of known localities: it is known to exist at one or perhaps two locations, the extent of occurrence is less than 5000 km 2, and the area of occupancy less than 500 km 2. Therefore, while presently of Least Concern, this species is prone to the effects of human activities (or stochastic events, the impact of which may be increased by human activities, e.g., climate change) within a very short period of time in an unforeseeable future, and poten- tially faces a high risk of extinction in the wild, warranting its assignment to the Endangered category of the IUCN Red List in the future.

REMARKS: In the description of T. louisanneorum, Lourenço (2017a: 23) noted the following:

The new species is very similar to the only other known species of the genus, Troglokhammouanus steineri … Nevertheless , a few differences can be observed: (i) a generally paler coloration in the new species, which is yellow to reddish-yellow, with pale yellow legs, (ii) the granulation on pedipalps and tergites is stronger in the new species, (iii) dorsal and ventral spinoid tubercles of pedipalp patella weaker than on T. steineri , (iv) some morphometric values differ between the two species, although their overall size is almost identical. Furthermore, the two species were found in separate caves that are not connected.

No other morphological differences between T. louisanneorum and T. steineri can be discerned from the diagnosis or description, in the absence of which the minor differences noted by Lourenço (2017a) cannot be considered diagnostic for the following reasons. Coloration is known to vary considerably within and between conspe- cific populations of scorpions and the variation described falls well within the variation among the material examined from the type locality, a single cave, during the present study (e.g., although immatures of T. steineri are generally more infuscate than adults, a few are depigmented; figs. 11E, 27A, B). Similarly, the unspecified differences in granulation of the pedipalps and tergites between the two species, which includes the “weaker” tubercles on the pedipalp patella of T. louisanneorum , fall within the range of variation among the material examined from the type locality. Any possible morphometric differences offered by Lourenço (2017a) may be discounted on the grounds that two individuals (i.e., the holotypes of T. louisanneorum and T. steineri ) do not comprise a statistical sample. Moreover, multivariate statistical analysis of the morphometric data presented by Lourenço (2007a, 2017a), together with new data (tables 3–5, 7, 8; appendices 4, 5, 7), was unable to separate the holotype of T. louisanneorum from the holotype of T. steineri or other individuals from the type locality, all of which clustered together across a morphospace comparable to that of other pseudochactid species (fig. 8).

It would therefore appear that the distinction between T. louisanneorum and T. steineri rests solely on their occurrence in two caves “that are not connected,” also known as the “caves as islands” paradigm ( Mammola, 2019). The assumption that species occurring in different caves must be heterospecific, promulgated by others (e.g., Francke, 2009a), is problematic for three reasons. Firstly, geography is not heritable and cannot comprise part of a taxon diagnosis, regardless of the presence or absence of morphological (or genomic) differences. Secondly, no evidence has been presented that the caves are disconnected or, if true, that the scorpion (which appears to be troglophilous) might not occur in other caves, rock shelters or deep crevices in the 50 km between them. The type locality of T. steineri, Tham Xe Bang Fai , is 11 km long, but many other caves and cave networks are more than 50 km in length, e.g., the Clearwater Cave System, Malaysia (over 48 km long), Lechuguilla Cave, New Mexico (over 221 km long), the Shuanghedong Cave Network, China (over 240 km long), Optymistychna Cave, Ukraine (over 258 km long), the Sistema Ox Bel Ha (over 268 km long) and Sistema Dos Ojos (over 320 km long), both in Mexico, and the Mammoth Cave Network, Kentucky (over 652 km long). In attributing islandlike properties to caves, researchers must consider the presence of subterranean connections, as well as the vagility of the species of interest ( Mammola, 2019). A network of fissures connecting two caves would make it difficult to consider the scorpion populations inhabiting them, to be isolated from one another. Lastly, there is doubt about the accuracy of the original collection data for T. louisanneorum . Two expeditions to Tham Nam Lot (Lod), in different years (June 2012, May 2018), failed to confirm the presence of this species, which is extremely abundant at the type locality, Tham Xe Bang Fai, although numerous other scorpions of several taxa (including the pseudochactid, A. lao ), were collected. Not a single specimen of this species was found during either visit to Tham Nam Lot (Lod), whereas ca. 140 specimens were collected during a single visit to Tham Xe Bang Fai, conducted during the same season.

In view of the evidence that the holotypes are conspecific, and absent any evidence to the contrary, the following new synonym is proposed: Troglokhammouanus louisanneorum Lourenço, 2017 = Troglokhammouanus steineri Lourenço, 2007 , syn. nov.

MATERIAL EXAMINED: LAOS: Khammouane Prov.: Boualapha Distr.: Hin Namno National Biodiversity Conservation Area: Tham Xe Bang Fai ( Xe Bang Fai River Cave ), 17°22′23.6″N 105°50′11.8″E, 159 m, left bank of Xe Bang Fai River (coming from downstream entrance): 19.ii.2012, L. Prendini and P. Kanyavong, very steep muddy beach on riverbank, with few cracks and no debris, UV detection, specimen on mud bank, 1 ♂ ( AMNH), muddy beach on riverbank, with moderate to steep mud bank with few rocks, UV detection, all specimens on moist mud bank, 2 ♀, 1 subad. ♂, 4 juv. ♂, 1 juv. ♀ ( AMNH), rocky beach of riverbank at base of “Stairway to Heaven,” with little mud except at base of rocks close to river, UV detection, 1 ♀, several imm. on moist mud, 1 imm. among vegetative debris caught between rocks, none on dry sand or rocks, 1 ♀, 1 subad. ♀, 3 juv. ♂ [1 juv. ♂ totally depigmented], 3 juv. ♀, pair of pedipalp chelae ( AMNH), 1 juv. ♀ ( AMCC [ LP 11275 ]), 19–21. ii.2012, L. Prendini, P. Kanyavong, and W. Phimmathong, rocky beach on riverbank, with extensive mud and sand banks in places, UV detection, specimens mostly on mud banks, almost all on moist soil, 5 ♂, 9 ♀, 4 subad. ♂, 9 juv. ♂, 4 juv. ♀, dextral pedipalp femur and patella ( AMNH), 1 juv. ♂ ( AMCC [ LP 11273 ]), 20.ii.2012, L. Prendini and P. Kanyavong, in twilight zone at cave entrance, muddy beach on riverbank, UV detection at night, specimens sitting on muddy banks (none collected at this site during daylight), 2 ♀ ( AMNH), 20–21.ii.2012, L. Prendini, P. Kanyavong, and W. Phimmathong, large sandy beach of riverbank, midway through cave [type locality], with muddy bank immediately along river and muddy patches in several depressions further from river edge, UV detection, all specimens collected on moist muddy soil, some among vegetative debris (sticks, etc.) trapped between stones, others sitting in open on mud, one in mud cracks, one below stone, 7 ♀, 1 juv. ♀ ( AMNH), 1 juv. ♀ ( AMCC [ LP 11272 ]) ; right bank of Xe Bang Fai River (coming from downstream entrance): 19–20.ii.2012, L. Prendini and P. Kanyavong, muddy beach of riverbank at bottom of “Balcony Passage,” ca. 300–500 m from downstream entrance of cave, UV detection during day: specimens very common on moist mud banks, mostly in cracks between drying mud slabs or among sticks and other vegetative debris caught between rocks, all collected on moist soil, none on dry sand banks higher upslope ; UV detection at night: several ♂♂ sitting on flowstone rock faces, several ♀♀ and juv. on mud banks and cracked clay, 2 ♂ (1 ♂ partly cannibalized by ♀), 22 ♀, 2 subad. ♂, 4 subad. ♀, 5 juv. ♂, 7 juv. ♀, carapace and sinistral pedipalp chela ( AMNH), 1 juv. ♀ ( AMCC [ LP 11271 ]), 19–21.ii.2012, L. Prendini, P. Kanyavong, and W. Phimmathong, muddy beach of riverbank between “Balcony Passage” and “Stairway to Heaven,” steep and dry with little cracked mud, relatively few rocks with trapped vegetative debris, UV detection, specimens uncommon, mostly on mud close to river edge, few higher upslope against cave walls, near moist flowstone, all specimens collected on moist mud surfaces, none on dry sand, 1 ♂, 7 ♀ ( AMNH), 20.ii.2012, L. Prendini and P. Kanyavong, rocky beach of riverbank before second rapids, UV detection, several adult ♀♀ and imm. on mud bank or in rock crevices close to river, none on dry soil higher upslope, 5 ♀, 1 subad. ♀, 1 juv. ♂ ( AMNH), 20–21.ii.2012, L. Prendini, P. Kanyavong, and W. Phimmathong, rocky/muddy beach of riverbank before “Stairway to Heaven,” UV detection, several ♀♀ and imm. on muddy banks or among trapped vegetative debris between rocks, none on dry sand, 1 ♂, 4 ♀ [1 ♀ depigmented ocelli], 4 subad. ♂, 1 subad. ♀, 2 juv. ♂, 8 juv. ♀ ( AMNH), 1 juv. ( AMCC [ LP 11274 ]), 21.ii.2012, L. Prendini and W. Phimmathong, Oxbow Lake , rocky beach on riverbank, UV detection, specimens on rocky shore, 2 ♀, 1 subad. ♂, dextral pedipalp chela ( AMNH) .