PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3687, 56 pp., 28 figures, 2 tables June 25, 2010 The Genus Hadruroides Pocock, 1893 (Scorpiones: Iuridae), in Peru: New Records and Descriptions of Six New Species JOSE´ A. OCHOA1 AND LORENZO PRENDINI1 ABSTRACT WereviewthetaxonomyoftheHadruroidesPocock,1893(Iuridae:Caraboctoninae),scorpions of Peru, describe sixnew speciesfrom the north of the country, andreport new records of other poorlyknownspecies.Thedescriptionofthesespeciesraisesto16thenumberofdescribedspecies in the genus, 13 of which occur in Peru. Four species inhabit dry forest in northern Peru: H. charcasus (Karsch, 1879); H. chinchaysuyu, n. sp.; H. geckoi, n. sp.; H. leopardus Pocock, 1900. Three species occur in inter-Andean valleys along the Cordillera: H. bustamantei Ochoa and Chaparro, 2008; H. carinatus Pocock, 1900; H. mauryi Francke and Soleglad, 1980. Six species inhabitdesertalongthePacificcoast:H.aguilariFranckeandSoleglad,1980;H.graceae,n.sp.;H. juanchaparroi,n.sp.;H.lunatus(L.Koch,1867);H.tishqu,n.sp.;H.vichayitos,n.sp.Mostspecies of Hadruroides have restricted distributions, except H. charcasus and H. lunatus, which are apparentlymorewidelydistributed.Weconsideritnecessarytoreassessallpreviousrecordsofthe lattertwospecies, because we suspectseveral are basedon misidentifications. INTRODUCTION pelin, 1905, together with Caraboctonus Pocock, 1893, a monotypic genus endemic to Hadruroides Pocock, 1893, is a small genus Chile (Francke and Soleglad, 1981; Prendini of Neotropical iurid scorpions, currently and Wheeler, 2005; Francke and Prendini, comprising 10 species (Maury, 1975; Francke 2008). Hadruroides has been reported from and Soleglad, 1980; Lourenc¸o, 1995; Sissom Bolivia, Chile, Colombia, Ecuador, Peru, and and Fet, 2000; Ochoa and Chaparro, 2008), Venezuela (Karsch, 1879; Pocock, 1900; included in subfamily Caraboctoninae Krae- Mello-Leita˜o, 1945; Esquivel de Verde, 1968; 1Scorpion Systematics Research Group, Division of Invertebrate Zoology, American Museum of Natural History, CentralParkWestat79thStreet,NewYork,NY,10024-5192([email protected];[email protected]). CopyrightEAmericanMuseumofNaturalHistory2010 ISSN0003-0082 2 AMERICAN MUSEUMNOVITATES NO. 3687 TABLE1 Currently recognizedspecies of thescorpion genusHadruroides Pocock, 1893, with countries anddepartmentsor provinces from whichthey havebeenrecorded HadruroidesaguilariFranckeandSoleglad,1980 Peru:Lima HadruroidesbustamanteiOchoaandChaparro,2008 Peru:Ayacucho,Huancavelica HadruroidescarinatusPocock,1900 Peru:Cajamarca Hadruroidescharcasus(Karsch,1879) Peru:Lambayeque,Piura,Tumbes? Hadruroideschinchaysuyu,n.sp. Peru:Tumbes HadruroidesgalapagoensisMaury,1975 Ecuador:Gala´pagos Hadruroidesgeckoi,n.sp. Peru:Cajamarca Hadruroidesgraceae,n.sp. Peru:Ancash Hadruroidesjuanchararroi,n.sp. Peru:Trujillo,Ancash HadruroidesleopardusPocock,1900 Peru:Lambayeque Hadruroideslunatus(L.Koch,1867) Peru:Lima Hadruroidesmaculatus(Thorell,1876) Ecuador:Manab´ı,Guayas HadruroidesmauryiFranckeandSoleglad,1980 Peru:Cusco Hadruroidestishqu,n.sp. Peru:Ancash(IslaSanta) HadruroidesudvardyiLourenc¸o,1995 Ecuador:Azuay,Loja Hadruroidesvichayitos,n.sp. Peru:Piura Kinzelbach, 1973; Maury, 1975; Cekalovic, History Museum, London, U.K. (BMNH); 1983; Sissom and Fet, 2000), but is actually Ca´tedra de Diversidad Animal I, Facultad de restricted to Ecuador, Peru, northern Chile, Ciencias Exactas, F´ısicas y Naturales, and several offshore islands, including the Universidad Nacional de Co´rdoba, Argen- Gala´pagos (Cekalovic, 1966; Maury, 1975; tina (CDA); Field Museum of Natural Francke and Soleglad, 1981; Ochoa, 2005; History, Chicago (FMNH); Museo Civico di unpubl. data). Species of Hadruroides inhabit Storia Naturale ‘‘Giacomo Doria,’’ Genova, inter-Andean valleys, Pacific desert, and dry Italy (MCSNG); Museo de Historia Natural, forest habitats. Universidad Nacional de San Antonio Abad In the present contribution, we describe six del Cusco, Peru (MHNC); Museo de Historia new species of Hadruroides from northern Natural, Universidad Nacional San Agust´ın, Peru, collected during recent fieldwork, and Arequipa, Peru (MUSA); Museo de Historia report new records of other poorly known Natural, Universidad Nacional Mayor de species. The description of these species raises San Marcos, Lima, Peru (MUSM); U.S. to 16 the number of described species in the National Museum of Natural History, Smith- genus.ThirteenoftheseoccurinPeru(table 1; sonian Institution, Washington (USNM); figs. 1, 2). Museum fu¨r Naturkunde der Humboldt- Universita¨t, Berlin, Germany (ZMB); Zoolo- gisches Museum der Universita¨t Hamburg, MATERIAL AND METHODS Germany (ZMH). Most of the material reported here was MorphologicalterminologyfollowsVachon collected during the course of several field (1974) for trichobothrial nomenclature; expeditions to Peru in recent years, especially Vachon (1952) and Prendini (2000) for pedi- 2004 and 2008. Specimens were collected palp carinae, abbreviated as follows: inter- during the day by turning rocks or stones, nomedian (IM), dorsointernal (DI), dorsome- and at night by ultraviolet (UV) light detec- dian (DM), dorsoexternal (DE), externome- tion (Stahnke, 1972). Point-locality records dian (EM), ventroexternal (VE), ventrome- weregeoreferencedinthefieldwithaportable dian(VM),ventrointernal(VI),dorsalpatellar Garmin GPS V. process(DPP),ventralpatellarprocess(VPP); Material is deposited in the following Stahnke (1970) for other characters; and a collections: American Museum of Natural modified version of Prendini (2004) for History, New York (AMNH); Natural metasomal carinae including carinae on ster- 2010 OCHOA AND PRENDINI:HADRUROIDES POCOCK, 1893 3 Fig.1. MapofPeru,plottingknownlocalityrecordsofHadruroidesPocock,1893.Hadruroidesaguilari Francke and Soleglad, 1980, solid circle. Hadruroides bustamantei Ochoa and Chaparro, 2008, squares. Hadruroides geckoi, n. sp., open triangle. Hadruroides graceae, n. sp., open circle. Hadruroides leopardus Pocock,1900, diamonds.Hadruroides tishqu, n.sp., solid triangles.Hadruroides vichayitos, n. sp.,star. 4 AMERICAN MUSEUMNOVITATES NO. 3687 Fig.2. MapofPeru,plottingknownlocalityrecordsofHadruroidesPocock,1893.Hadruroidescarinatus Pocock,1900,opencircles.Hadruroidescharcasus(Karsch,1879),solidsquares.Hadruroideschinchaysuyu, n.sp.,diamonds.Hadruroidesjuanchaparroi,n.sp.,solidcircles.Hadruroideslunatus(L.Koch,1867),open squares. Hadruroides mauryi Francke and Soleglad, 1980, open triangles. Undetermined Hadruroides species, stars. 2010 OCHOA AND PRENDINI:HADRUROIDES POCOCK, 1893 5 nite VII and tergites, abbreviated as follows: DIAGNOSIS: The genus Hadruroides com- dorsolateral (DL), lateral supramedian prises small- to medium-sized scorpions, (LSM), median lateral (ML), lateral inframe- varying from ca. 30 mm, e.g., H. leopardus dian (LIM), ventrolateral (VL), ventrosubme- Pocock, 1900, and H. graceae, n. sp., to ca. dian(VSM),ventromedian(VM);terminology 80 mm in total length, e.g., H. charcasus for leg carinae follows that used for the (Karsch, 1879) (Maury, 1975). The most pedipalps. important diagnostic characters for the genus Measurements (in mm) were recorded and are as follows: cheliceral movable fingers, illustrations produced using a Nikon SMZ– internal surface with two subdistal teeth and 1500 stereomicroscope fitted with an ocular one prominent basal tooth; carapace anterior micrometer and camera lucida. Photographs margin slightly convex, with three pairs of under UV and visible light were taken using a lateral ocelli; sternum subpentagonal, with Y- Microptics ML-1000 digital-imaging system. shaped sulcus; pedipalp chela smooth, acar- Distribution maps were generated using inate in most species, except H. charcasus, in DIVA-GIS Version 5.4 (http://www.diva-gis. whichgranularcarinaearepresentoninternal org/) by superimposing georeferenced point- surface;chelamovablefinger,mediandenticle locality records on a digital elevation dataset row with six or seven median subrows of from the CGIAR Consortium for Spatial denticles and variable number of accessory Information (CGIAR-CSI; available at http:// denticles; neobothriotaxic major Type C srtm.csi.cgiar.org). trichobothrial pattern (Vachon, 1974): femur with three trichobothria (i, e, d); patella with 20: one internal (i), two dorsal (d ), three 1, 2 SYSTEMATICS ventral (v1–3), v3 situated on external surface, and14externaltrichobothria(et ,est,em , 1–3 1–3 esb , eb ); chela with 26 trichobothria: 14 1, 2 1–5 FAMILY IURIDAE THORELL, 1876 onmanus(Et ,Est,Esb,Eb ,V )and12 1–5 1–3 1–4 on fixed finger(Dt,Db,et,est,esb,eb,dt,dst, SUBFAMILY CARABOCTONINAE dsb, db, it, ib); leg telotarsus with ventrome- KRAEPELIN, 1905 dian row of spinule clusters (setaceous tufts); Hadruroides Pocock, 1893 metasomalsegmentVwithcomplete,granular VL and VM carinae; telson slightly concave Telegonus:L.Koch,1867(part):235. dorsally, with short aculeus; hemispermato- Hadrurus: Thorell,1876(part):186;Karsch,1879(part): phore lamelliform, with elongated crest, ac- 135,1881:290;Boeris,1889:125. companied externally by medial slit and one Caraboctonus:Pocock,1893a(part):92. Hadruroides Pocock, 1893b: 306, 309, 329, 330, type free lobe, truncal flexure absent, internobasal species by original designation: Hadrurus charcasus reflection of the sperm duct absent. Karsch, 1879 (5 Hadruroides charcasus [Karsch, Hadruroides appears to be most closely 1879]); Kraepelin, 1894: 182, 206; Laurie, 1896: 130; related to Caraboctonus. The two genera Kraepelin,1899:188;Pocock,1900:474;Birula,1917a: 163,189;Birula,1917b:44,48,51;Mello-Leita˜o,1945: exhibit several similarities, e.g., trichobothrial 119; Bu¨cherl,1964:61,1967:115; Williams,1970:31; pattern, ventromedian row of spinule clusters Bu¨cherl, 1971:328; Stahnke, 1974:122; Maury, 1975: on telotarsus, dentition of chelicerae and 10,11;Francke,1977:75;FranckeandSoleglad,1981: pedipalp chela fingers. They may be distin- 235,256,figs.9,13,27–33,58;Francke,1985:8,17,20; Sissom, 1990: 130, 131; Nenilin and Fet, 1992: 14; guished by means of the ventral carinae of Lourenc¸o, 1994: 157, 1995: 74–76, 1997a: 601, 602, metasomal segment V: in Caraboctonus, the 1998: 135; Kovarˇ´ık, 1998: 135; Lourenc¸o, 2000: 25; VL carinae are restricted to the posterior half SissomandFet,2000:411–413;Lourenc¸oandDastych, of the segment and the VM carina is absent, 2001: 55, 56; Soleglad and Sissom, 2001: 31, fig.34; whereas in Hadruroides, both VL and VM AcostaandOchoa,2002:19;SolegladandFet,2003a: 2, 5, 12, 2003b: 8, 111; Fet et al., 2004: 18, 23, 24; carinaearepresentandcomplete(exceptinH. Prendini and Wheeler, 2005: 482; Fet and Soleglad, tishqu,n.sp.,inwhichtheVLandVMcarinae 2005:12;Ochoa,2005:65,fig.17;Dupre´,2007:5,14; arerestrictedtotheposteriortwo-thirdsofthe OchoaandChaparro,2008:5;FranckeandPrendini, segment). The two genera may be further 2008:210;FetandSoleglad,2008:255;Solegladetal., 2009:1. distinguished on the basis of pedipalp chela 6 AMERICAN MUSEUMNOVITATES NO. 3687 finger dentition: internal and external acces- well developed and comprising strong gran- sory denticles, flanking the median denticle ules (fig. 4E). .. .. .. .. . .. . H. charcasus row in Hadruroides, are absent in – Pedipalp chela smooth, acarinate (figs. 8E, 11G).. . .. .. .. .. .. .. . .. .. .. .. .. . 2 Caraboctonus. Furthermore, the hemisperma- 2. Sternite VIIacarinate.. .. . .. .. .. .. .. . 3 tophore of Hadruroides exhibits an elongated – SterniteVIIwithtwoorfourcarinae(figs. 7C, crest and a medial slit, both absent from the D, 27C) .. .. .. .. .. .. . .. .. .. .. .. . 6 hemispermatophore of Caraboctonus, which 3. Metasomal segments I–IV elongated; segment instead exhibits a digit-shaped process. IIlongerthanwide;segmentV(-)approxi- DISTRIBUTION: The genus Hadruroides is mately three times longer than wide, endemic to Ecuador, Peru, northern Chile, length:width ratio: 3.18 . . .. .. H.aguilari and several offshore islands, including the – Metasomal segments I–IV relatively short Gala´pagos (table 1). Thirteen of the 16 (fig. 20E); segment II as wide as long; - segment V ( ) approximately twice longer described species occur in Peru (figs. 1, 2). than wide, length:width ratio: 2.04–2.59 Four species inhabit dry forest in northern (figs. 19G, 23B). .. .. .. . .. .. .. .. .. . 4 Peru: H. charcasus (fig. 4E), H. chinchaysuyu, 4. Pigmentation reduced to faint spots on car- n.sp.(fig. 4B);H.geckoi,n.sp.(figs. 3B,4C); apace and tergites (fig. 24B), absent on H. leopardus Pocock, 1900 (figs. 3A, 5B). metasomal segments; segment V, VM and Three species occur in inter-Andean valleys VLcarinaereducedtoposteriortwo-thirdsof along the Cordillera: H. bustamantei Ochoa segment (fig. 23B); telson, ventral surfaces - U and Chaparro, 2008 (fig. 3D); H. carinatus smooth ( , ) (fig. 23A,C). .. . H.tishqu Pocock, 1900 (fig. 4A); H. mauryi Francke – Pigmentation pronounced on carapace, ter- gites, and metasomal segments (fig.20C, and Soleglad, 1980. Six species inhabit desert D);segmentV,VMandVLcarinaecomplete along the Pacific coast: H. aguilari Francke (fig. 19G);telson,ventralsurfacessmoothor and Soleglad, 1980; H. graceae, n. sp. - sparsely granular anteriorly ( ), granular (fig. 4D); H. juanchaparroi, n. sp. (figs. 3C, (U). .. . .. .. .. .. .. .. . .. .. .. .. .. . 5 5A);H.lunatus(L.Koch,1867)(figs. 3G,5C); 5. Tergites I–VI with rectangular dorsosubmedian H.tishqu,n.sp.(figs. 3E,5E);H.vichayitos,n. spots of pigmentation (fig.20C); sternite VII sp. (figs. 3F, 5D). We exclude H. maculatus and metasomal segment I, ventral surfaces (Thorell, 1876), which is restricted to the unpigmented;hemispermatophorebroaderba- sallywithrelativelylongcrest ... H.lunatus central coastline of Ecuador, from the list of – Tergites I–VI with irregular dorsosubmedian Peruvian species. Two other Hadruroides spotsofpigmentation(fig.20D);sterniteVII species are endemic to Ecuador: H. galapa- and metasomal segment I, ventral surfaces goensis Maury, 1975; H. udvardyi Lourenco, with several spotsof pigmentationsurround- 1995 (table 1). Most species of Hadruroides inginsertionofsetae(fig.20E);hemisperma- have restricted distributions, except H. char- tophoreslenderwithshortcrest(fig.21) ... casus and H. lunatus, which are apparently ..................... H.juanchaparroi - more widely distributed. We consider it 6. Chela fixed finger (adult ) curved, creating necessary to reassess all previous records of distinctproximalgapwithmovablefingerwhen fingersclosed(figs.8C,11H,27H)....... 8 the latter two species, because we suspect - – Chela fixed and movable fingers (adult ) several are based on misidentifications. Other straight, no proximal gap evident when recordsofHadruroidesfromthecoastaldesert fingers closed (fig.15I).. . .. .. .. .. .. . 7 of southern Peru and northern Chile (Ochoa, 7. MetasomalsegmentIV,ventralsurfacedensely 2005;unpubl.data)correspondtonewspecies granular (fig.15C); segment V with 7–8 related to H. lunatus (fig. 2), the descriptions ventral setae and 6–8 lateral setae; pedipalp - of which are in preparation by the authors. chelalength:widthratio:5.00–5.41( ),4.46– U 5.12( )................... H.graceae – Metasomal segment IV, ventral surface PERUVIAN SPECIES OFHADRUROIDES smooth; segment V with 12–18 ventral setae and 3–5 lateral setae; pedipalp chela KEY TO IDENTIFICATION OF THE SPECIES length:width ratio: 3.93–4.54 (-), 4.23–4.64 U ( ). .. . .. .. .. .. .. .. . .. . H. leopardus 1. Adult pedipalp chela robust, internomedian, 8. Metasomal segments, VSM carinae absent on dorsointernal, and dorsal marginal carinae segments I–III,presenton IV.. .. .. .. 12 2010 OCHOA AND PRENDINI:HADRUROIDES POCOCK, 1893 7 Fig.3. HadruroidesPocock,1893,habitatsinPeru.A.PuchacaAlto(LambayequeDepartment),habitat ofHadruroidesleopardusPocock,1900.B.Balsas(CajamarcaDepartment),habitatofHadruroidesgeckoi, n. sp. C.Cerro Campana (La Libertad Department), habitat of Hadruroides juanchaparroi,n. sp. D.Wari (Ayacucho Department), habitat of Hadruroides bustamantei Ochoa and Chaparro, 2008. E. Isla Santa (AncashDepartment),habitatofHadruroidestishqu,n.sp.F.PlayaVichayitos(PiuraDepartment),habitat ofHadruroidesvichayitos,n.sp.G.LomasdeLachay(LimaDepartment),habitatofHadruroideslunatus(L. Koch,1867). 8 AMERICAN MUSEUMNOVITATES NO. 3687 - Fig. 4. Hadruroides Pocock, 1893, habitus in life. A. Hadruroides carinatus Pocock, 1900, . B. U - - Hadruroideschinchaysuyu,n.sp., .C.Hadruroidesgeckoi,n.sp., .D.Hadruroidesgraceae,n.sp., .E. - Hadruroides charcasus(Karsch, 1879), . 2010 OCHOA AND PRENDINI:HADRUROIDES POCOCK, 1893 9 - Fig.5. HadruroidesPocock,1893,habitusinlife.A.Hadruroidesjuanchaparroi,n.sp., .B.Hadruroides - U - leopardusPocock,1900, .C.Hadruroideslunatus(L.Koch,1867), .D.Hadruroidesvichayitos,n.sp., . - E.Hadruroides tishqu, n. sp., . 10 AMERICAN MUSEUMNOVITATES NO. 3687 – Metasomal segments, VSM carinae present on relatively slender, length:width ratio: 2.7– segmentIand,usually,IIandIII,absenton 2.8. .. . .. .. .. .. .. .. . .. .. . H.mauryi IV .. .. .. .. .. . .. .. .. .. .. .. .. . .. 9 9. Pigmentationpronouncedoncarapace,tergites, Hadruroides aguilari and metasomal segments (figs.8I, 9C, 12F,D);VSMcarinaepresentonmetasomal Francke and Soleglad, 1980 segments I–III; pedipalp chela fixed finger - Figure 1 (adult ) strongly curved, creating well developed proximal gap with movable finger whenfingersclosed(fig.11H) ......... 10 Hadruroides aguilari Francke and Soleglad, 1980: 2–8, – Pigmentation absent, except for small, faint figs.1, 3–20; Kovarˇ´ık, 1998: 135; Sissom and Fet, spots on carapace and tergites (fig.12G); 2000: 411; Fet et al., 2004: 24; Ochoa and Chaparro, 2008:5. VSMcarinaeobsoleteonmetasomalsegment Ifi,xaebdsefinntgoenrs(aegdmuletn-ts)IsIliagnhdtlIyIIc;uprevdeidp,aclrpecahtienlga meTnYt:PELiMmaATEPRrIoAvLi:nceP:ERHUol:otyLpimea-D, e2parUt- weakproximalgapwithmovablefingerwhen paratypes (AMNH), Cajamarquilla (11u579S fingersclosed(fig.27H)...... H.vichayitos 76u549W, ca. 800 m), 3.i.1976, O.F. Francke. 10. Sternite VII, VL and VSM carinae well developed; metasomal segment V, DL car- DIAGNOSIS: Hadruroides aguilari appears U to be most closely related to H. lunatus and inae smooth; telson, ventral surface ( ) smooth; hemispermatophore lamina apically H. juanchaparroi, based on the similar carina- rounded .. .. .. . .. .. .. .. . H.carinatus tion of sternite VII and metasomal segments – Sternite VII, VL carinae distinct, VSM carinae I–IV. Haduroides aguilari may be separated obsolete (fig.7C); metasomal segment V, DL from these species by the elongated metaso- carinaesparselytodenselygranular(figs.8A, mal segments, e.g., segment II is longer than D, 11A); telson, ventral surface (U) granular wide and segment V (-) approximately three (fig.8B); hemispermatophore lamina apically times longer than wide in H. aguilari, acuminate(figs.9E,13).............. 11 - compared with H. lunatus and H. juancha- 11. Metasomal segment V elongated ( ): parroi, in which segment II is as wide as long, length:width ratio: 2.40–2.89 (fig.11B); seg- - and the length:width ratio of segment V ( ) ment V, ventral surface sparsely granular, varies from 2.04 to 2.59. Additionally, in H. comprisingscatteredgranulesinposteriorthird (fig.11B);segmentVwith10–14DLsetaeand lunatus and H. juanchaparroi, the pedipalp 9–13 VL setae; pedipalp chela slender, chela fixed finger of the adult male is curved, - length:width ratio: 4.41–4.40 ( ), 4.53–4.74 creating a distinct proximal gap with the U ( )(figs.11E,12B);pectinaltoothcount:21– movable finger when the fingers are closed, - U 23( ),18–21( );hemispermatophorelamina whereas the fixed and movable fingers of the curvedindistalhalf(fig.13) ..... H.geckoi adultmale arestraight,suchthatno proximal - – Metasomal segment V short ( ): length:width gap is evident when the fingers are closed, in ratio: 1.97–2.14 (fig. 8F); segment V, ventral H. aguilari. surfacedenselygranular(fig. 8F);segmentV with 5–8 DL setae and 6–10 VL setae; DISTRIBUTION: This endemic Peruvian spe- cies is known only from the type series and, pedipalp chela relatively broad, length:width ratio: 3.66–4.02 (-), 4.0–4.18 (U) (fig. 8E); despite extensive searches, no additional ma- pectinal tooth count: 17–20 (-), 16 (U); terial has been collected. The type locality is hemispermatophore lamina straight in distal now an urban zone, close to the Peruvian half(fig. 9E–H). . .. .. .. H. chinchaysuyu capitalofLima(fig. 1),andnonaturalhabitat 12. TergitesI–IVeachwithpaireddorsosubmedian remains there. However, it is possible that the and dorsolateral spots of pigmentation, speciesstilloccursinadjacentareasofnatural forming four longitudinal stripes along me- habitat. sosoma; legs I–IV each with several prolat- ECOLOGY: According to Francke and eral spots of pigmentation; pedipalp chela - Soleglad(1980),H.aguilarihasbeencollected ( ) relatively broad, length:width ratio: atnightfromthetopsofterrestrialbromeliads 3.09–3.36 . .. .. . .. .. .. . H. bustamantei – TergitesI–IVunpigmented,mesosomawithout (Tillandsia sp., Bromeliaceae), and is sympa- longitudinal stripes; legs without prolateral tric with H. lunatus, found under stones and - spots of pigmentation; pedipalp chela ( ) bromeliad mats.