Revision of the genera Eutrecha and Xenotrecha (Solifugae: Ammotrechidae), taxonomic notes on Ammotrechinae, and description of a remarkable new Eutrecha from Colombia Author Botero-Trujillo, Ricardo https://orcid.org/0000-0002-6199-6572 Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79 th Street, New York, NY 10024 - 5192, USA & Division Aracnologia, Museo Argentino de Ciencias Naturales " Bernardino Rivadavia " - CONICET, Avenida Angel Gallardo 470, CP: 1405 DJR, C. A. B. A., Buenos Aires, Argentina Author Martinez, Leonel Division Aracnologia, Museo Argentino de Ciencias Naturales " Bernardino Rivadavia " - CONICET, Avenida Angel Gallardo 470, CP: 1405 DJR, C. A. B. A., Buenos Aires, Argentina Author Iuri, Hernan Augusto Division Aracnologia, Museo Argentino de Ciencias Naturales " Bernardino Rivadavia " - CONICET, Avenida Angel Gallardo 470, CP: 1405 DJR, C. A. B. A., Buenos Aires, Argentina Author Ojanguren-Affilastro, Andres Alejandro Division Aracnologia, Museo Argentino de Ciencias Naturales " Bernardino Rivadavia " - CONICET, Avenida Angel Gallardo 470, CP: 1405 DJR, C. A. B. A., Buenos Aires, Argentina Author Carvalho, Leonardo Sousa https://orcid.org/0000-0003-4700-5610 Universidade Federal do Piaui, Campus Amilcar Ferreira Sobral, BR 343, km 3.5, Bairro Meladao, s / no. CEP 64800 - 000, Floriano, PI, Brazil carvalho@ufpi.edu.br text Arthropod Systematics & amp; Phylogeny 2023 2023-04-04 81 317 344 http://dx.doi.org/10.3897/asp.81.e95181 journal article http://dx.doi.org/10.3897/asp.81.e95181 1864-8312-81-317 BE57FF40714740AC8BEA1706A95DA4ED C3B28C3FAD9250BC906A4A865EAA18B4 3.1.7. Xenotrecha huebneri (Kraepelin, 1899) Figs 1 , 3 , 11 , 12 , 13 , 14 , 15 , 16 , 17 , 18 , 19 Cleobis huebneri Kraepelin, 1899: 239-240; Weidner 1959 : 109; Maury 1982 : 124. Cleobis huebneri Kraepelin, 1899: Harms and Duperre 2018 : 12-13, figs 5a-c. Ammotrecha huebneri (Kraepelin, 1899): Kraepelin 1901 : 112-114. Ammotrechella huebneri (Kraepelin, 1899): Roewer,1934: 593, 594-595, 598, figs 336b, 338c. Ammotrechella hubneri (Kraepelin, 1899): Muma and Nazario 1971 : 506, 507; Muma 1976 : 25. Xenotrecha huebneri (Kraepelin, 1899): Maury 1982 : 125-127, 129, 134-138, figs 18-28; Maury 1984 : 75, fig. 11; Rocha and Cancello 2002a : 4; Rocha and Cancello 2002b : 2; Harvey 2003 : 210-211; Bird et al. 2015 : 123; Harms and Duperre 2018 : 13. Type material. Holotype. VENEZUELA • 1 ♀; "South Venezuela" [locality not specified]; 25 Nov 1898; G. Huebner & O. Schneider leg; ZMH. Examined by images from Harms and Duperre (2018) . Revised diagnosis. As for the genus. Redescription of male. Based on nontype male from Vila Tepequem (CHNUFPI 1247). - Measurements. Linear measurements in Table 1 . - Color. In 80% ethanol-preserved specimen. Prosomal dorsal shields and opisthosomal tergites with overall brown coloration (Fig. 12A, C ). Propeltidium with a design of pale brown areas in a darker contour (Fig. 12C ), one large that narrows anteriorly, and two small oval areas one on each side of the ocular tubercle, the latter of which is blackish; eyespots shiny white (Fig. 13A ). Meso-, metapeltidium, and opisthosomal tergites predominantly dark brown (Fig. 12A ), with scattered faded patches. Chelicerae, base color pale brown (same as pale propeltidial areas) (Figs 12C , 14C, E ), with three dark brown, narrow longitudinal stripes on prodorsal, retrolateral, and retroventral surfaces of manus, which fuse into a large dark brown retrolateral area on distal part of manus; stridulatory plate predominantly yellow, with brownish stridulatory ridges (Fig. 14D ). Asetose area of fixed and movable fingers red, with all teeth darkened (Fig. 14E ); movable finger setose area with ventral, brown-spotted area. Pedipalp coxae yellowish white, trochanter pale brown, femur and tibia dark brown, with faint paler areas (Fig. 17 ), as are the patella and tibia of legs. Pedipalps and legs, basitarsus proximal half dark brown, distal half yellowish brown, same color as telotarsus. Coxosternal region and trochanters of legs immaculately yellowish white (Fig. 13B ). Femora of legs I-II, basifemora and telofemora of legs III-IV yellowish white, with scattered darker patches, mostly on dorsal surface. Malleoli white. Opisthosomal pleural membranes with faded, dark brown color dorsally, paler towards the venter. Sternites yellowish white (Fig. 13C ), except for three posteriormost sternites which have some scattered dark brown patches. - Prosoma. Propeltidium longer than wide (Table 1 ); covered with small to medium-sized, spicule-like stout setae, straight and rigid (Fig. 12C ); these setae fall off and break easily; at least the larger macrosetae exhibit a bilaterally symmetrical distribution on propeltidium. Ocular tubercle slightly elevated, with abundant macrosetae. Anterolateral propeltidial lobes separated from the propeltidium principal shield by incomplete lateral groove (Fig. 13A ). Eyespots elongated, its length approximately half the length of the anterolateral propeltidial lobe ventral margin. Meso- and metapeltidium wider than long, with abundant macrosetae similar to those on propeltidium (Fig. 12A ). Coxae densely covered with abundant thin setae (Fig. 13B ). Sternum glabrous. - Chelicera-dentition and processes. Fixed finger with median teeth series comprising well-developed primary teeth (FP, FM, and FD) and very small FSM tooth (Figs 3B , 14E , 15A ); FSD tooth absent; FSM and FM contiguous to adjacent teeth (i.e., without medial notch or FMAD); retrofondal teeth series uninterrupted (i.e., without FRFD), with four teeth (RFSP, RFP, RFM, RFA) (Figs 3B , 15A ); basal retrofondal margin heavily sclerotized (Fig. 15A ); profondal teeth series consisting of four teeth (PFSP, PFP, PFSM, PFM). Fixed finger asetose area with dorsal and ventral margins notably curved; prodorsal carina sharp, not elevated in lateral aspect, without angular dorsal crest; proventral carina weakly pronounced on the mucron area; fixed finger retrolateral carina (FRLC) obsolete, represented by few granules on the proximal region of the asetose area (Figs 3B , 14E ). Fixed finger mucron without subterminal (FST) teeth; apex (FT tooth) curved. Movable finger with median teeth series comprising well-developed and similar-sized MP and MM primary teeth, and one MSM secondary tooth which is smaller than MP and MM (i.e., MP ≈ MM> MSM) (Figs 3B , 14E ); all three teeth of the median series adjacent to each other; MSM upright and triangular. Movable finger prolateral carina (MPLC) markedly developed, ending slightly basal to MP in a small but distinct prolateral (MPL) tooth, which is about half the size of MSM tooth (Fig. 14D ). Movable finger without subproximal (MSP) or subterminal (MST) teeth; movable finger retroventral longitudinal carina (MRVC) present on distal half, or third, of finger, forming a smooth elevated ridge (Figs 3B , 14E , 16D ); retrolateral longitudinal carina (MRLC) consisting of scattered conspicuous granules on the retrolateral surface of finger (Fig. 3B ). Movable finger mucron moderately long, with gnathal edge carina ordinary (not convex). Closure of FM tooth distal to MM, when fingers are closed. - Chelicera-setose areas and stridulatory plate. Retrolateral and dorsal surfaces with abundant retrolateral manus ( rlm ) and retrolateral finger ( rlf ) setae, of different sizes, which are predominantly straight and rigid (Figs 12C , 14C-F ); some of these setae are arranged in bilaterally symmetrical pattern, as are some principal retrolateral finger ( principal rlf ) setae that are more flexible than others; movable finger retrolateral proximal setal cluster ( rlpc ) dorsally with a single, long and markedly plumose seta (Figs 14E , 16D ). Prolateral surface with array of setal types (Fig. 14D, F ), as follows: row of plumose proventral distal ( pvd ) setae starting at level of the interdigital condyle ( pic ) and ending near level of FP tooth; proventral subdistal ( pvsd ) setae arranged in rather disorganized pattern, pvsd comb slightly differentiated; carpet-like field of bristle-like promedial ( pm ) setae narrow (Fig. 16C ). Stridulatory plate longer than high, occupying approximately two-thirds of the prolateral surface of manus (Fig. 14D ); stridulatory apparatus consisting of eleven distinct ridges approximately parallel to the manus ventral surface (Fig. 16C ); most ridges not reaching the limit with the pm setae field. Distal limit of the prolateral setose area of movable finger reaching the level of MSM tooth; movable finger prodorsal ( mpd ) setal series consisting of plumose setae (similar to the pvd setae), adjacent to abundant non-plumose setae of the movable finger promedial ( mpm ) and proventral ( mpv ) setal series. - Chelicera-flagellum. Of the composite type, without shaft. A thin, translucent, membranous structure immovably attached prodorsally to the fixed finger (Figs 14F , 15A ); flagellar base general aspect bowl-shaped, long and narrow, with apex reaching about two thirds of the mucron length; prolateral surface with plumose setiform organ arising from the center of the flagellar base (Figs 14F , 16A, B ); plumose setiform organ robust basally, long, progressively narrowing distally, its apex almost reaching the apex of the flagellar base, covered with acuminate fringes over distal two thirds of its length; other than fringes on the plumose setiform organ, the flagellum is predominantly smooth; flagellum dorsal margin visible over the prodorsal carina in retrolateral aspect; attachment point elliptical, horizontally elongated, placed at level of the PFM tooth. - Pedipalp. All segments coated with abundant short and delicate setae (Fig. 17A ); those on ventral surface of tibia, basitarsus, and telotarsus stouter and more distinct than those on other surfaces. Proventral surface of femur with some spicule-like, somewhat spiniform macrosetae similar to those on propeltidium (Fig. 12A ); tibia with proventral and retroventral rows of six spiniform setae each, which are short and stout, distributed along distal two thirds of tibia, in addition to a basal pair of conspicuous, thinner, and slightly longer setae (Fig. 17A ); basitarsus with proventral and retroventral rows of eight and eleven spiniform setae, respectively, similar to those on tibia; telotarsus without spiniform setae. Femur, tibia, basitarsus, and telotarsus with few long thin setae; clubbed setae apparently absent. Retroventral surface of femur proximally with a suture-like cleavage plane (Figs 17 , 18 ). Telotarsus retrodorsal pore area, if present, not visible under light stereomicroscopy. - Leg I. All segments coated with abundant short and delicate setae similar to those on pedipalps, without stout or spiniform setae; tibia and basitarsus with few long thin setae. Telotarsus without claws or spiniform setae; retrodorsal pore area, if present, not visible under light stereomicroscopy. - Walking legs. Covered with abundant short and delicate setae, in addition to a few long setae like those on pedipalps and legs I. Legs II and III: basitarsus with five spiniform setae: two proventral (distal and subdistal), one retroventral (distal), one retrolateral (subdistal), and one retrodorsal (distal); telotarsus bi-segmented (consisting of large basal and small distal segments), with proventral row of four spiniform setae and a retroventral row of three, in 2.2.2/1 pattern. Leg IV: basitarsus with row of three proventral and one distal retroventral spiniform setae, in 1.1.2 pattern; telotarsus 3-segmented (the two segmentation lines are complete), with proventral and retroventral rows of four spiniform setae each, in 2.2.2/2/0 pattern. - Opisthosoma. Tergites with abundant setae similar to those on propeltidium; setation of the sternites comparable to that of coxae. Ctenidia present on 1st and 2nd post-genital sternites (spiracular sternites I and II) (Fig. 13C ); ctenidia short, in the form of abundant, lanceolate setae irregularly distributed on the sternites; other sternites without ctenidia. Figure 11. Xenotrecha huebneri (Kraepelin, 1899), live habitus of adults at Serra do Tepequem , Roraima State, Brazil. A ♂ (CHNUFPI 1247). B ♀ (CHNUFPI 1249). Figure 12. Xenotrecha huebneri (Kraepelin, 1899), habitus, dorsal aspect ( A , B ), propeltidium ( C , D ). A , C ♂ (CHNUFPI 1247), Serra do Tepequem , Roraima State, Brazil. B , D ♀ (CHNUFPI 1248), same locality. Scale bars = 2 mm (A, B), 1 mm (C, D). Figure 13. Xenotrecha huebneri (Kraepelin, 1899), prosoma, anterolateral propeltidial lobe, lateral aspect ( A ), anterior part of coxosternal region, ventral aspect ( B ) [note the severed dextral pedipalp], ctenidia on 2nd post-genital sternite ( C ), genital plate, ventral aspect ( D ). A-C ♂ (CHNUFPI 1247), Serra do Tepequem , Roraima State, Brazil. D ♀ (CHNUFPI 1249), same locality. Scale bars = 0.5 mm (A, D), 1 mm (B), 0.2 mm (C). Figure 14. Xenotrecha huebneri (Kraepelin, 1899), dextral chelicera, retrolateral aspect ( A , C ) and close-up of fingers ( E ), prolateral aspect ( B , D ) and close-up of flagellum ( F ). A , B ♀ (CHNUFPI 1248), Serra do Tepequem , Roraima State, Brazil. C-F ♂ (CHNUFPI 1247), same locality. Scale bars = 1 mm (A-D), 0.5 mm (E), 0.2 mm (F). Arrows in C-F indicate the position of hole left by broken FD tooth. Figure 15. Xenotrecha huebneri (Kraepelin, 1899), sinistral ( A ) and dextral ( B ) chelicera, fixed finger, ventral aspect. A ♂ (CHNUFPI 1247), Serra do Tepequem , Roraima State, Brazil. B ♀ (CHNUFPI 1248), same locality. Scale bars = 0.5 mm. Arrow in A indicates hole left by broken FD tooth. Figure 16. Xenotrecha huebneri (Kraepelin, 1899), ♂ (CHNUFPI 1247), Serra do Tepequem , Roraima State, Brazil, sinistral chelicera, flagellum, prolateral aspect ( A ) and closeup of the plumose setiform organ ( B ), stridulatory apparatus, prolateral aspect ( C ), movable finger, retrolateral aspect ( D ). Scale bars = 0.2 mm (A), 0.1 mm (B), 0.5 mm (C, D). Figure 17. Xenotrecha huebneri (Kraepelin, 1899), ♂ (CHNUFPI 1247), Serra do Tepequem , Roraima State, Brazil, sinistral pedipalp, retrolateral aspect ( A ) and close-up of cleavage plane proximally on femur ( B ) [indicated by arrow], healed severed stump of dextral pedipalp femur (post-autotomy in nature), retrodorsal ( C ) and retrolateral ( D ) aspects. Scale bars = 1 mm (A), 0.5 mm (B-D). Figure 18. Xenotrecha huebneri (Kraepelin, 1899), ♂ (CHNUFPI 1247), Serra do Tepequem , Roraima State, Brazil, scanning electron micrographs of cleavage plane on healed severed stump (post-autotomy in nature) of dextral pedipalp femur, retrolateral aspect ( A ) and close-up of edge ( B ), inclined, anterior aspect ( C ) and close-up of pointy process on prolateral part of stump ( D ). Scale bars = 0.5 mm (A, C), 0.1 mm (B), 0.2 mm (D). Supplementary description of female. Based on nontype female from Vila Tepequem (CHNUFPI 1248). Measurements in Table 1 . Similar to the male in most aspects, including size and general appearance. Pedipalps with short and stout spiniform setae on the ventral surface of basitarsus only, arranged in proventral and retroventral rows of seven and nine spiniform setae, respectively. Tegument setation similar to that of male; setae on dorsal surfaces of prosomal and opisthosomal shields, and on dorsal and retrolateral surfaces of the chelicerae and legs, weaker and more flexible. Opisthosoma without ctenidia. Genital plate posterior margin with deep median indentation (Fig. 13D ); posteromedian region conspicuously glabrous and shiny, with a central pocket. Chelicera without the secondary sexual characteristics of males (Figs 14A, B , 15B ). Stridulatory apparatus with all the ridges parallel to each other and to the manus ventral surface, as in male (Fig. 14B ); ridges short, progressively occupying a more distal position the more dorsal they are. Fixed finger, lateral aspect with distinct and pronounced angular dorsal crest at level of the RFM tooth (Fig. 14A, B ); retrolateral carina (FRLC) evident, as in male (Fig. 15B ). Fixed finger without FSD tooth; mucron short and tooth-like (i.e., ventral margin sublinear), without subterminal teeth (FST). Movable finger with MP, MM, and MSM teeth, MP being largest and MSM smallest (Fig. 14A ); MM tooth not displaced distally. Movable finger prolateral carina (MPLC) ending in small but distinct prolateral (MPL) tooth, which is less than half the size of the MSM tooth. Retrolateral longitudinal carina (MRLC) consisting of abundant granules; gnathal edge carina and retroventral longitudinal carina (MRVC) evident; subproximal (MSP) and subterminal (MST) teeth absent. Variability. One female (CHNUFPI 1248) has nine spiniform setae on the retroventral series of the pedipalp basitarsi, whereas the other female (CHNUFPI 1249) has ten. Distribution. Originally described from an unspecified locality in southern Venezuela ( Kraepelin 1899 ), X. huebneri has also been recorded from the Henri Pittier National Park and Pardillar, respectively in the states of Aragua and Carabobo ( Maury 1982 ), and from El Rincon , in the state of Sucre ( Rocha and Cancello 2002a ), all in northern Venezuela (Fig. 1 ). In Brazil, a record of Xenotrecha (as " Xenotrecha sp.") has been presented from Furo do Firmino, southeastern Maraca Island in the state of Roraima ( Rocha and Cancello 2002a ), locality that is situated some 50 km south of Vila Tepequem (record here presented; Fig. 1 ). Records from Brazil are located some 350 km east of the headwaters of the Orinoco River, where the type specimen was most likely collected ( Harms and Duperre 2018 ). An additional record of X. huebneri from an unspecified locality in Suriname was identified in the Global Information Facility (GBIF) ( Goud et al. 2020 ), further extending the putative distribution of the genus far eastward. Natural history. Specimens from Serra do Tepequem were collected at night. All specimens were found on Curatella americana L. ( Dilleniaceae ) tree trunks. One female (CHNUFPI 1249) was observed foraging, moving upwards in the tree trunk while inspecting small holes and under the tree barks, using both pedipalps to sense the surface (Fig. 11B ). The sampling locality (Fig. 19 ) is a small tepui (reaching 1100 m), forested on its slopes and with savannas on the higher plateaus ( Almeida et al. 2009 ). Several other specimens of C. americana were inspected at other sampling localities in the municipalities of Boa Vista ( 02°52′08.4″S 60°43′13.1″W , at ca. 90 m.a.s.l.) and Bonfim ( 03°16′20.5″S 60°03′09.3″W , at 140 m.a.s.l.), but no additional specimens of X. huebneri were detected at these localities. However, an unidentified Ammotrecha species was found in the inspected trees at Bonfim, locality that is situated close to the Brazil-Guiana border. The specimen of X. huebneri reported by Rocha and Cancello (2002a) from El Rincon was found inside of a dead tree-trunk in a forest with many lianas, whereas the specimen from Maraca Island was collected in a forest inside of a termite mound of a possibly undescribed Araujotermes Fontes, 1982 ( Isoptera , Termitidae ) species. Figure 19. Landscape and habitat of Xenotrecha huebneri (Kraepelin, 1899) at Serra do Tepequem , Roraima State, Brazil. Note the dense vegetation ( A ), grasslands, and rocky outcrops ( B - D ) present in the area. Other material examined. BRAZIL • 1 ♂ ; Roraima , Amajari , Serra do Tepequem , Vila Tepequem , near Pousada PSJ; 03°47′10.4″S 61°43′15.3″W ; 640 m . a.s.l.; 17 Jul 2014 ; J. Cabra-Garcia leg.; CHNUFPI 1247; • 2 ♀♀ ; same data, except: L.S. Carvalho leg.; CHNUFPI 1248-1249 . Literature records (material not examined). BRAZIL • 1 sex not specified; Roraima , Alto Alegre , southeastern Maraca Island , Uraricoera River , Furo do Firmino ; 03°23′60″N 61°25′60″W ; 01 Nov 1986 , E.M. Cancello and C.R.F. Brandao leg.; MZUSP 14295; listed as " Xenotrecha sp." SURINAME • 1 sex not specified; 07 Aug 1959 ; RMNH.SOL.11; gbifID 2434367917 ; VENEZUELA • 1 sex not specified; Sucre , El Rincon ; 10°38′14″N 64°14′09″W ; 27 Sep 1987 ; O.F.F. Souza leg.; MZUSP 14296. These records were obtained from Rocha and Cancello (2002a) and Goud et al. (2020) .