Systematic Revision Of The Andean Scorpion Genus Orobothriurus Maury, 1976 (Bothriuridae), With Discussion Of The Altitude Record For Scorpions

OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 AMNH BULLETIN 359 2011 BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY SYSTEMATIC REVISION OF THE ANDEAN SCORPION GENUS OROBOTHRIURUS MAURY, 1976 (BOTHRIURIDAE), WITH DISCUSSION OF THE ALTITUDE RECORD FOR SCORPIONS JOSÉ A. OCHOA, ANDRES A. OJANGUREN AFFIL ASTRO, C A MILO I. MATTONI, AND LORENZO PRENDINI Scientific Publications of the American Museum of Natural History American Museum Novitates Bulletin of the American Museum of Natural History Anthropological Papers of the American Museum of Natural History Publications Committee Robert S. Voss, Chair Board of Editors Jin Meng, Paleontology Lorenzo Prendini, Invertebrate Zoology Robert S. Voss, Vertebrate Zoology Peter M. Whiteley, Anthropology Managing Editor Mary Knight Submission procedures can be found at http://research.amnh.org/scipubs O n t h e c o v e r : Orobothriurus parvus Maury, 1976 (Bothriuridae), adult ♂ from La Oroya, Junín, Peru. Photo, José A. Ochoa. All issues of Novitates and Bulletin are available on the web from http://digitallibrary.amnh.org/dspace Order printed copies from http://www.amnhshop.com or via standard mail from: American Museum of Natural History—Scientific Publications Central Park West at 79th Street New York, NY 10024 This paper meets the requirements of ANSI/NISO Z39.48-1992 (permanence of paper). SYSTEMATIC REVISION OF THE ANDEAN SCORPION GENUS OROBOTHRIURUS MAURY, 1976 (BOTHRIURIDAE), WITH DISCUSSION OF THE ALTITUDE RECORD FOR SCORPIONS JOSE ´A. OCHOA 1,2 ANDRES A. OJANGUREN AFFILASTRO 3 CAMILO I. MATTONI 4 LORENZO PRENDINI 1 1 Scorpion Systematics Research Group, Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, NY, 10024-5192 2 Departamento de Zoologia, Instituto de Biocieˆncias, Universidade de Sa˜oPaulo,RuadoMata˜o, travessa 14, 101, Sa˜o Paulo, SP, 05508-900, Brazil 3 Museo Argentino de Ciencias Naturales ‘‘Bernardino Rivadavia,’’ Avenida A ´ngel Gallardo 470, CP: 1405DJR, Buenos Aires, Argentina 4 Laboratorio de Biologı ´aReproductivay Evolucio´n, Ca´tedra de Diversidad Animal I, Facultad de Ciencias Exactas, Fı ´sicas y Naturales, Universidad Nacional de Co´rdoba, Av. Ve´lez Sarsfield 299, 5000, Co´rdoba, Argentina BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Number 359, 90 pp., 54 figures, 3 tables Issued November 23, 2011 Copyright E American Museum of Natural History 2011 ISSN 0003-0090 CONTENTS Abstract .......................................................... 3 Introduction . ...................................................... 3 MaterialandMethods ................................................ 5 On the distribution of Orobothriurus ...................................... 8 On the altitude record for Scorpiones. . ................................... 11 Systematics ....................................................... 12 Orobothriurus (Maury,1976)........................................... 12 Key to identification of the species of Orobothriurus .......................... 14 Orobothriurus alticola (Pocock,1899)..................................... 15 Orobothriurus ampay OchoaandAcosta,2003 .............................. 20 Orobothriurus atiquipa OchoaandAcosta,2002 ............................. 25 Orobothriurus calchaqui,n.sp........................................... 25 Orobothriurus compagnuccii,n.sp........................................ 41 Orobothriurus curvidigitus (Kraepelin,1911) ................................ 48 Orobothriurus famatina Acosta,2001..................................... 52 Orobothriurus grismadoi Ojanguren Affilastro et al., 2009....................... 52 Orobothriurus huascaran,n.sp. ......................................... 56 Orobothriurus paessleri (Kraepelin,1911) .................................. 68 Orobothriurus parvus Maury,1976....................................... 69 Orobothriurus quewerukana,n.sp........................................ 70 Orobothriurus ramirezi,n.sp............................................ 74 Orobothriurus tamarugal,n.sp.......................................... 77 Orobothriurus wawita AcostaandOchoa,2000.............................. 81 Acknowledgments .................................................. 83 References........................................................ 83 Appendix1....................................................... 88 Appendix2....................................................... 88 2 ABSTRACT The systematics of the Andean scorpion genus, Orobothriurus Maury, 1976 (Bothriuridae Simon, 1880), is revised. New locality records, obtained during recent field expeditions, distribution maps, and a key to identification of the 15 known species, are provided. Six new species are described: Orobothriurus calchaqui, n. sp., from northwestern Argentina; Orobothriurus compagnuccii, n. sp., from the central Andes of Argentina; Orobothriurus huascaran, n. sp., from central Peru; Orobothriurus quewerukana, n. sp., from southern Peru and northern Chile; Orobothriurus ramirezi, n. sp., from central Chile; and Orobothriurus tamarugal, n. sp., from northern Chile. The known distribution of Orobothriurus and the altitude record for scorpions are discussed. The world’s altitude record for a scorpion, previously reported as 5550 m, is demonstrated to be 4910 m. INTRODUCTION The genus Orobothriurus Maury, 1976, comprises several fairly small scorpion species from the high altitudes and arid zones of central-western and southern South America (figs. 1–4). The genus is endemic to the Andes Cordillera, extending from central Peru (Cordillera Blanca in Ancash Department) to central-western Argentina, including western Bolivia, northern and central Chile, and northern and western Argentina (fig. 1). The southernmost record is the El Nevado mountain range in Mendoza Province, Argentina (Ojanguren Affilastro et al., 2009; figs. 1, 3D, 54). After its original description (Maury, 1976), this genus received little attention from taxonomists, mostly due to the scarcity of specimens in collections and the inaccessibility of the habitats where these scorpions occur, notably the high Andes Mountains, where each species is range restricted and microhabitat specific. This situation changed in recent years, as result of many expeditions conducted in the Andes of Argentina, Chile, and Peru, mostly by members of our research group. Orobothriurus initially included eight species (Maury, 1976): Orobothriurus alticola (Pocock, 1899); Orobothriurus curvidigitus (Kraepelin, 1911); Orobothriurus paessleri (Kraepelin, 1911); Orobothriurus peruvianus (Mello-Leita˜o, 1948); Orobothriurus dumayi (Cekalovic, 1974); Orobothriurus crassimanus Maury,1976; Orobothriurus inca Maury, 1976, and Orobothriurus parvus Maury, 1976. Five of these were originally placed in Bothriurus Peters, 1861. The following species were added subsequently: Orobothriurus wawita Acosta and Ochoa, 2000; Orobothriurus famatina Acosta, 2001; Orobothriurus iskay Acosta and Ochoa, 2001; Orobothriurus atiquipa Ochoa and Acosta, 2002, Orobothriurus ampay Ochoa and Acosta, 2003; Orobothriurus lourencoi Ojanguren Affilastro, 2003; Orobothriurus grismadoi Ojanguren Affilastro et al., 2009. Orobothriurus dumayi was returned to Bothriurus (Acosta and Ochoa, 2001; Mattoni and Acosta, 2006). Two species groups, alticola and inca, were also recognized in the genus (Maury, 1976; Acosta and Ochoa, 2001), but the monophyly of Orobothriurus remained doubtful (Prendini, 2000). Based on a cladistic analysis, Ochoa (2004a) described a new genus, Pachakutej Ochoa, 2004, to accommodate the four species of the inca group: Pachakutej crassimanus (Maury, 1976); Pachakutej inca (Maury, 1976); Pachakutej iskay (Acosta and Ochoa, 2001); Pachakutej peruvianus (Mello-Leita˜o, 1948). In the most recent taxonomic developments on this genus, Acosta (2002) transferred the enigmatic taxon, Cercophonius brachycentrus var. bbivittatus Thorell, 1877, to Orobothriurus, and subsequently (Acosta, 2005) recognized it as a valid species. Orobothriurus bivittatus (Thorell, 1877) was later synonymized with O. alticola (Ojanguren Affilastro et al., 2009). In the present contribution we describe six new species of Orobothriurus from Argentina, Chile and Peru, collected during recent field expeditions, raising to fifteen the number of species in the genus (table 1; appendix 1), and present new records and comprehensive distribution maps for the other 10 species, along with a key to the identification of all species in the genus. 3 Fig. 1. Orobothriurus Maury, 1976, map of the distribution of the genus plotting known locality records. 4 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 MATERIAL AND METHODS Scorpion specimens were collected by turning stones during the day, or by ultraviolet (UV) light detection at night (Stahnke, 1972). Portable UV lamps, comprising mercury vapor tubes attached to a chromium reflector, and powered by a 12V, 7 amp/hour battery, or MagliteHflashlights modified with UV LED attachments, were used in fieldwork. Material is deposited in the following collections: American Museum of Natural History, New York (AMNH); Natural History Museum, London, U.K. (BMNH); Ca´tedra de Diversidad Animal I, Facultad de Ciencias Exactas, Fı ´sicas y Naturales, Universidad Nacional de Co´rdoba, Argentina (CDA); Frantisˇek Kovar ˇı ´k Private Collection, Prague, Czech Republic (FKPC); Field Museum of Natural History, Chicago. (FMNH); Instituto Argentino de Investigaciones de las Zonas A ´ridas, Mendoza, Argentina (IADIZA); Instituto Miguel Lillo, Tucuma´n, Argentina (IML); Luı ´s Eduardo Acosta Private Collection, Co´rdoba, Argentina (LEA); Laboratorio de Biologı ´a Reproductiva y Evolucio´n, Universidad Nacional de Co´rdoba, Argentina (LBRE); Museo Argentino de Ciencias Naturales ‘‘Bernardino Rivadavia’’, Buenos Aires, Argentina (MACN-Ar); Museo de Historia Natural, Universidad Nacional de San Antonio Abad delCusco,Peru(MHNC);Muse´um d’Histoire Naturelle de Gene`ve, Switzerland (MHNG); Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru (MUSM); Museo Regionale di Scienze Naturali, Turin, Italy (MRSN); Museu de Zoologia, Universidade de Sa˜o Paulo, Brazil (MZSP); Museo de Zoologı ´adelaUniversidad de Concepcio´n,Chile(MZUC);Naturhistoriska Riksmuseet, Stockholm, Sweden (NRS); Zoologisches Museum, Universita¨t Hamburg, Germany (ZMH). Measurements (mm) were recorded with an ocular micrometer, and illustrations of external morphology produced using a Nikon SMZ 1500 stereomicroscope with a camera lucida. Ultraviolet fluorescence images were produced using a Microptics ML1000 digital imaging system, or a NikonH DS-Fi1-U2 digital camera, attached to a NikonHSMZ 1500 steremicrocope, under long-wave UV light. Habitus images were taken using the Microptics ML-1000 system, or a NikonHD100 digital camera, under visible light. Hemispermatophores where dissected in ethanol and manually extracted from the tissues of the paraxial organs. Illustrations of hemispermatophores were produced with a camera lucida fitted to a Leica M 165C stereomicroscope. Photographs of hemispermatophores were taken with a Leica DFC 290 camera fitted to a Leica M 165C stereomicroscope. Morphological terminology follows Vachon (1974) for trichobothrial nomenclature; Vachon (1952) and Prendini (2000) for pedipalp carinae, abbreviated as follows: internomedian (IM), dorsointernal (DI), dorsomedian (DM), dorsoexternal (DE), externomedian (EM), ventroexternal (VE), ventromedian (VM), ventrointernal (VI), dorsal patellar process (DPP), ventral patellar process (VPP), digital (D), dorsal secondary (DS), dorsomarginal (DMA), external (E); Mattoni and Acosta (2005) for macrosetae of the metasomal segments; Stahnke (1970) for other characters; a modified version of Prendini (2004) used by Ochoa et al. (2010) for carinae of the tergites, sternites, and metasoma, abbreviated as follows: dorsolateral (DL), lateral supramedian (LSM), lateral median (LM), lateral inframedian (LIM), ventrolateral (VL), ventrosubmedian (VSM), ventromedian (VM). Hemispermatophore terminology follows Ochoa (2004a) and Mattoni and Acosta (2005). Point-locality records were georeferenced in the field with a portable GPS (GarminH Etrex, GarminHVista or GarminHII), or retroactively using the GeoNet Names Server (GNS, http://earth-info.nga.mil/gns/html/). Distribution maps were generated using ArcMap 9.0 (Enviromental Systems Research Institute [ESRI], Redlands, California), by superimposing point locality records of species on coverages depicting political boundaries and topography. The topographic coverage (as a shaded relief) was generated from digital elevation model files (1 arc degree resolution) from the United States Geological Survey (USGS) (http://edc.usgs.gov/products/ elevation/gtopo30/gtopo30.html). 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 5 Fig. 2. Orobothriurus Maury, 1976, habitats in Peru. A. Polobaya, Arequipa (Arequipa Department), habitat of Orobothriurus curvidigitus (Kraepelin, 1911). B. Pacchac, Urubamba (Cusco Department), habitat of Orobothriurus wawita Acosta and Ochoa, 2000. C. Quiquijana, inter-Andean valley near Cusco (Cusco Department), habitat of O. wawita.D. Lomas Atiquipa (Arequipa Department), habitat of Orobothriurus atiquipa Ochoa and Acosta, 2002. E. Puna, central Peru (Junin Department), habitat of Orobothriurus parvus Maury, 1976. 6 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 Fig. 3. Orobothriurus Maury, 1976, habitats in Argentina, Chile and Peru. A. Pampa de Tamarugal (Tarapaca´ Region, Chile), habitat of Orobothriurus tamarugal, n. sp. B. Puente del Inca (Mendoza Province, Argentina), habitat of Orobothriurus alticola (Pocock, 1899). C. El Tontal (Mendoza Province, Argentina), habitat of O. alticola.D. Cerro Nevado (Mendoza Province, Argentina), habitat of Orobothriurus grismadoi Ojanguren-Affilastro et al., 2009. E. Ampay, Abancay (Apurimac Department, Peru), habitat of Orobothriurus ampay Ochoa and Acosta, 2003. F. Llanganuco (Ancash Department, Peru), habitat of Orobothriurus huascaran, n. sp. G. Sierra de Famatina (La Rioja Province, Argentina), habitat of Orobothriurus famatina Acosta, 2001. 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 7 ON THE DISTRIBUTION OF OROBOTHRIURUS Most species of Orobothriurus inhabit the Andes and orographic systems of Argentina, Bolivia, Chile and Peru, where they have been collected up to 4910 m. However, some species occur at low altitude near the Pacific coast (figs. 1, 53). All Orobothriurus species are allopatric (figs. 52–54). This pattern of Fig. 4. Orobothriurus Maury, 1976, habitus in life. A. Orobothriurus ampay Ochoa and Acosta, 2003, U .B. Orobothriurus grismadoi Ojanguren-Affilastro et al., 2009, - .C. Orobothriurus parvus Maury, 1976, U .D. Orobothriurus huascaran,n.sp., U . TABLE 1 Currently recognised species of the scorpion genus Orobothriurus Maury, 1976, with countries and provinces (Argentina), regions (Chile) or departments (Peru) from which they have been recorded. Orobothriurus lourencoi Ojanguren Affilastro, 2003, is excluded as this species will be placed in another genus (Mattoni et al., in litt.) Orobothriurus alticola (Pocock, 1899) Argentina: Mendoza, San Juan Orobothriurus ampay Ochoa and Acosta, 2003 Peru: Apurı ´mac Orobothriurus atiquipa Ochoa and Acosta, 2002 Peru: Arequipa Orobothriurus calchaqui, n. sp. Argentina: Tucuma´n Orobothriurus compagnuccii, n. sp. Argentina: La Rioja Orobothriurus curvidigitus (Kraepelin, 1911) Peru: Arequipa Orobothriurus famatina Acosta, 2001 Argentina: La Rioja Orobothriurus grismadoi Ojanguren Affilastro et al., 2009 Argentina: Mendoza Orobothriurus huascaran, n. sp. Peru: Ancash Orobothriurus paessleri (Kraepelin, 1911) Peru: Arequipa Orobothriurus parvus Maury, 1976 Peru: Junin, Lima Orobothriurus quewerukana, n. sp. Chile: Tarapaca´, Peru: Tacna Orobothriurus ramirezi, n. sp. Chile: Coquimbo Orobothriurus tamarugal, n. sp. Chile: Tarapaca´ Orobothriurus wawita Acosta and Ochoa, 2000 Peru: Ayacucho, Cusco 8 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 VSM carinae well developed, granular, occupying entire length of segment I and weakly developed but complete on II; hemispermatophore apex 66%–72%of lamina length, terminal process of basal lobe small (fig. 32A, B,36F–H,K–O).................. 9 9. Metasomal segment IV with two or three pairs of ML macrosetae; segment III usually with one pair of DL macrosetae . . . O. calchaqui – Metasomal segment IV with one pair of ML macrosetae; segment III usually without DL macrosetae............... O. famatina 10. Pedipalp chela manus ( - ), internal surface with apophysis (fig. 8B); pedipalp chela fixed finger ( - ) slightly curved, creating small gap with movable finger when fingers closed (fig. 22A); metasomal segment V, VL and VM carinae complete (fig. 22E); metasomal segments with distinct VM pigmentation stripe (at least on segments II–V); hemispermatophore, ventral border of apex slightly curved (fig. 27B) . . . . . . . . . . . . O. parvus – Pedipalp chela manus ( - ), internal surface without apophysis (fig. 25C); pedipalp chela fixed finger ( - ) straight, no gap evident when fingers closed (fig. 25B); metasomal segment V, VL and VM carinae absent ( - ; fig. 21C) or restricted to distal third ( U ; fig. 22F); metasomal segments irregularly pigmented, VM pigmentation stripe indistinct (fig. 13D); hemispermatophore, ventral border of apex straight (fig. 27C) . . . . . . . . . . . O. wawita 11. Metasoma segments I–III ( U ), VSM carinae present (fig. 19D); segment V, VL and VM carinae restricted to posterior two-thirds (fig. 21F); pedipalp chela manus, trichobothrium Esb situated between Eb 2 and Eb 3 ,Db slightly closer to Dt than to Eb 3 (fig. 39C); hemispermatophore, distal border of apex slightly rounded, frontal crest with internal lateral projection vestigial (fig. 40A–C) .... ...................... O. huascaran – Metasoma segments I–III ( U ), VSM carinae absent; metasomal segment V, VL, and VM carinae complete (fig. 20A, C, F); pedipalp chela manus, trichobothrium Esb situated dorsal to Eb 2 ,Db equidistant between Dt and Eb 3 (figs. 25D, 46B, C); hemispermatophore, distal border of apex subtriangular, frontal crest with internal lateral projection complete (figs. 28A, 47A, B) . . . . . . . . . 12 12. Sternite VII acarinate (fig. 18A); metasomal segment II ( - ), VSM carinae absent (fig. 18A); segment V, VSM and VM carinae not discernible from dense surface granulation (fig. 21A); legs III and IV, telotarsi each with 3/4 spiniform macrosetae in proand retroventral rows; metasomal segments I–III, paired VL stripes contiguous with VM stripe posteriorly (but not on segments IV and V) (fig. 11D); segment IV with four pairs of VSM macrosetae. . . . . . . . . . . O. atiquipa – Sternite VII with one or two pairs of carinae (fig. 18C, D); metasomal segment II ( - ), VSM carinae present; segment V, VM, and VL carinae well defined (figs. 20A, C, F, 21E); legs III and IV, telotarsi each with 3/3 spiniform macrosetae in proand retroventral rows; metasomal segments I–V, paired VL stripes not contiguous with VM stripe posteriorly (fig. 11C); segment IV with three pairs of VSM macrosetae ....................... 13 13. Hemispermatophore apex elongated, angle of inclination 135.18u–139.1u(fig. 36D, E); tergite VII entirely pigmented (fig. 13G); telson vesicle ( - ), dorsal surface concave (fig. 23E) ....... ........................ O. grismadoi – Hemispermatophore apex short, angle of inclination 138.99u–144.99u(figs. 36I, J, 47A, B); tergite VII with paired lateral spots of pigmentation delimiting unpigmented median longitudinal stripe (fig. 13F); telson vesicle ( - ),dorsalsurfaceflat(fig.23A).......14 14. Hemispermatophore, distal crest extremely developed, extending almost entire length of lamina (fig. 47A, B); telson length/width ratio ( - ), 2.65–2.94 (mean 52.78), ( U ), 2.13–2.37 (mean 52.27)............. O. ramirezi – Hemispermatophore, distal crest restricted to distal half of lamina (fig. 36I); telson length/ width ratio ( - ), 2.87–3.21 (mean 53.07), ( U ), 2.44–2.55 (mean 52.49) . . . O. alticola Orobothriurus alticola (Pocock, 1899) Figures 13F, 15B, 18D, 20A, 21A, 36I, J, 54 Cercophonius brachycentrus var. bbivittatus Thorell, 1877: 183 [synonymized by Ojanguren Affilastro et al., 2009: 35]. Bothriurus alticola Pocock 1899: 357–358, fig. 1; Kraepelin, 1911: 91, 97; Mello-Leita˜o, 1931: 82, 92; 1932: 34; 1934: 63, 65; 1935: 93; 1937: 103; 1945: 138, 144, 145; Bu¨cherl, 1959a: 23, 24, fig. 4; Abalos, 1959: 591; 1963: 113; Cekalovic, 1966: 3; Bu¨cherl, 1969: 769; Maury, 1973: 110; Roig Alsina, 1973: 198; Masnu´ de Moreno, 1991: 184, 189, map 1. Bothriurus (Andibothriurus)alticola:Bu¨cherl et al., 1963: 217, 224, fig. 8. Urophonius brachycentrus bivittatus: Mello-Leita˜o 1931: 99–100; 1932: 35; 1934: 48, 51; 1938: 94– 95; 1945: 213, 215; Acosta and Maury, 1998: 559; Lowe and Fet, 2000: 44. Urophonius brachicentrus bivittatus: Mello-Leita˜o, 1939: 612; Abalos, 1959: 592; 1963: 117; Roig Alsina, 1973: 200 [incorrect subsequent spelling]. 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 15 Fig. 5. Most parsimonious tree (length 5167; CI: 0.479; RI: 0.813), obtained by cladistic analysis of 65 morphological characters (appendix 2) scored for 15 species of Orobothriurus Maury, 1976, and 15 outgroup taxa with the implied weighting regime that maximized average support (Mattoni et al., in litt.). Synapomorphies optimized with accelerated transformation indicated with bars. Black bars indicate uniquely derived apomorphic states, white bars indicate parallel derivations of apomorphic states. Numbers above bars indicate characters, numbers below indicate states. 16 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 Orobothriurus bivitattus: Acosta, 2005: 3–12, figs. 1–8; Ojanguren Affilastro, 2005: 181–183, 220, 241; Acosta, 2006: 20, 21; ICZN, 2008: 69–70. Orobothriurus alticola: Maury, 1976: 17–18, figs. 1–10, table 1 (part); 1979: 717, map 10 (part); 1981: 98 (part); Acosta and Maury, 1998: 559 (part); Kovar ˇı ´k, 1998: 101; Acosta and Ochoa, 2000: 135, 136, 143; Lowe and Fet, 2000: 35; Acosta and Ochoa, 2001: 203–205, 208, 209; Acosta, 2002: 176, 177; Ochoa, 2004a: 44, 49, 50, 52, 55, figs. 1, 2, 21, table 1; Acosta, 2005: 1, 2, 8, 9, 12; Ojanguren Affilastro, 2005: 176, 178, 179, 180, 220, 241; Kamenz and Prendini, 2008: 39, pl. 111, table II; Ojanguren Affilastro et al., 2009: 35–40, figs. 2, 18, 19, 25–35, table 2. T YPE M ATERIAL : ARGENTINA: Mendoza Province: Las Heras Department:1 - , 1 U syntypes (BMNH), Puente del Inca [32u49909.150S69u55901.820W, 2721 m]. Cercophonius brachycentrus bivittatus: 1 juv. holotype (NRS), San Juan Province. N EW R ECORDS : ARGENTINA: Mendoza Province: Las Heras Department: Puente del Inca, 32u49.5689S69u54.6069W, 2759 m, 2.xi.2003, J.A. Ochoa, L. Prendini and C.I. Mattoni, alpine vegetation (low spiny bushes) on hard, rocky ground, under stone, 1 juv. (AMNH [LP 2386]). San Juan Province: Calingastra Department: Cerro el Tontal, 31u31924.70S69u12923.30W, 3600 m, 25.i. 2006, A. Ojanguren Affilastro, L. Compagnucci and L. Piacentini, 2 juv. (AMNH [LP 5848]); between Paso de Agua Negra and Aduana, vega and surrounds, 30u17933.10S 69u46945.60W, 4005 m, 27.i.2005, C.I. Mattoni and A. Ojanguren Affilastro, UV sampling, 1 subad. - , 1 subad. U , 1 juv. - (AMNH [LP 4309]). D IAGNOSIS : Orobothriurus alticola is similar to O. grismadoi in pigmentation pattern, external morphology, and hemispermatophore. The two species can be separated by the shape of the hemispermatophore: the apex is narrower, and the angle formed by the apex with the rest of the distal lamina smaller in O. grismadoi (fig. 36D, E) than in O. alticola (fig. 36I, J). Males can also be separated by the shape of the telson, which is more slender in O. grismadoi. Additionally, the dorsal surface of the telson vesicle is concave in males of O. grismadoi, whereas it is flat in males of O. alticola (fig. 23A, E). Tergite VII is almost entirely pigmented, without an unpigmented median stripe, in O. grismadoi (fig. 13G), but exhibits paired lateral spots of pigmentation, delimiting an unpigmented median stripe, in O. alticola (fig. 13F). On the other hand, O. alticola can be separated from O. compagnuccii by the shape of the lamina of the hemispermatophore: the apex comprises 52.17%–54.54% (n53; mean 552.98%) of the lamina in O. compagnuccii and 40.87%–46.08%(n520; mean 543.88%)inO. alticola.Orobothriurus alticola and O. ramirezi can be separated by Fig. 6. Orobothriurus Maury, 1976, metasomal segments I–V showing distribution of macrosetae. A. Lateral aspect. B. Ventral aspect. Annotations: dorsolateral (DL); median lateral (ML); ventrolateral (VL); ventrosubmedian (VSM). 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 17 the distal crest of the hemispermatophore, which is less developed, occupying only half the apex of the lamina, in O. alticola, than in O. ramirezi, in which it occupies almost the entire apex (figs. 36I, J, 47A, B). The telson vesicle of O. alticola is narrower, especially in females (telson length/width ratio: - , 2.87–3.21, mean 53.07; U , 2.44– 2.55, mean 52.49; fig. 23A) than that of O. ramirezi (telson length/width ratio: - , 2.65– 2.94, mean 52.78; U , 2.13–2.37, mean 5 2.27; figs. 23G, 24E). The dorsal surface of the femur and ventral surface of the metasoma are less pigmented, with narrower stripes, in O. alticola than O. ramirezi. The metasomal carinae, especially the VL and VSM carinae of segments I and V, and the DL carinae of segment V, are less developed in males of O. alticola than O. ramirezi (fig. 20A, C). Fig. 7. Orobothriurus Maury, 1976, dextral pedipalp femur (A) and patella (C–D), showing trichobothrial pattern. A. Femur, dorsal aspect. B. Patella, dorsal aspect. C. Patella, external aspect. D. Patella, ventral aspect. Annotations: dorsal (d); external (e); external basal (eb); external medial (em); external suprabasal (esb); external subterminal (est); external terminal (et); internal (i); ventral (v). Scale bar 51 mm. 18 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 R EMARKS : Acosta (2002) transferred Cercophonius brachycentrus bivittatus to Orobothriurus, and subsequently redescribed it as a valid species based on a single male specimen (Acosta, 2005). Based on an examination of additional material, however, Ojanguren Affilastro et al. (2009) were unable to identify any characters by means of which this species could be consistently separated from O. alticola, and it was therefore synonymized. D ISTRIBUTION : Orobothriurus alticola is endemic to central-western Argentina, from the central part of Mendoza Province (Laguna Diamante) to the central part of San Juan Province (Paso del Agua Negra; fig. 54). It occurs at 2700–4000 m in the Andes (fig. 3B) and nearby El Tontal mountain range of the Precordillera (fig. 3C, Ojanguren Affilastro et al., 2009). E COLOGY : The area inhabited by O. alticola belongs to the Puna and the Altoandina Fig. 8. Orobothriurus Maury, 1976, dextral pedipalp chela, showing trichobothrial pattern. A. External aspect, with dorsal aspect of fixed finger. B. Ventral aspect. C. Ventrointernal aspect. Annotations: dorsal basal (db,Db); dorsal suprabasal (dsb); dorsal subterminal (dst); dorsal terminal (dt,Dt); external basal (eb, Eb); external suprabasal (esb,Esb); external subterminal (est,Est); external terminal (et,Et); internal basal (ib); internal terminal (it); ventral (V). Scale bar 51 mm. 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 19 phytogeographic provinces (Cabrera and Willink, 1980). This species was collected in sympatry with another bothriurid, Brachistosternus montanus Roig Alsina, 1977. Orobothriurus ampay Ochoa and Acosta, 2003 Figures 4A, 11A, 12D, 17B, 22A, 25A, 27A, 52 Orobothriurus ampay Ochoa and Acosta, 2003: 2– 6, figs. 1–14; Ochoa, 2004a: 43, 52, 54, 73, figs. 1, 2, 21, table 1; 2005: 55, figs. 7, 9, table 2. T YPE M ATERIAL : PERU: Apurı ´mac Department: Abancay Province: Holotype - (MUSM), Tamburco, Sector Arapato, Santuario Nacional Ampay, 13u359S72u529W, 3580 m, 25.viii.1998, R. Aimituma. Paratypes: Santuario Nacional Ampay, Bosque Nativo del Ampay [13u359410S72u529480W], 3100–3300 m, 26.viii.1998, J.C. Chaparro andJ.A.Ochoa,2 U (MHNC), 1 U (CDA 088), 1 U (MACN-Ar 10039); Santuario Nacional Ampay, Ccorhuani [ca. 13u359S Fig. 9. Orobothriurus parvus Maury, 1976, sternum and pectines. A. - (MHNC). B. U (MHNC). Scale bars 51mm. 20 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 72u529W], 3280 m, 18.ix.1998, R. Aimituma, 1 U (MUSM). N EW R ECORDS : PERU: Apurı ´mac Department: Abancay Province: Santuario Nacional Ampay, Bosque Nativo del Ampay, 13u359S 72u529W, 3200 m, 17.vii.2000, J.A. Ochoa, in Podocarpus glomeratus forest, 1 U (AMNH [LP 1918]). D IAGNOSIS : Orobothriurus ampay differs from all other species in the genus by its welldeveloped pigmentation pattern. The tergites of this species are entirely, densely pigmented (fig. 12D) and the paired VL stripes contiguous with the VM stripe posteriorly on metasomal segments I–V (fig. 11A) whereas, in other species, tergites I–VI each possess two lateral spots delimiting an unpigmented median stripe along the mesosoma (fig. 13E– G) and the paired VL stripes of the metasomal segments are not contiguous with the VM Fig. 10. Orobothriurus Maury, 1976. Sinistral hemispermatophore showing morphology and measurements. A. Ental aspect. B. Dorsal aspect. C. Ectal aspect. Annotations: basal lobe (bl); capsular concavity (cc); constriction (cns); distal crest (Dc); external lobe (el); external lateral projection (elp); frontal crest (Fc); internal lobe (il); internal lateral projection (ilp); incision (in); Lamina (Lam); pedicel (Ped); terminal process (tp); trunk (Tr); tortuous stem (ts); apex length (x); length of basal portion of lamina (y). 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 21 stripe posteriorly, at least on segments IV and V (figs. 11B–D, 13A, D). Orobothriurus ampay may be further distinguished on the basis of metasomal carination, trichobothriotaxy and pedipalp chela ornamentation. The VSM carinae, present on metasomal segment IV in O. ampay (fig. 17B), are absent in other species of Orobothriurus (fig. 17A). The angle formed between pedipalp chela trichobothria eb–Et 5 –Et 4 is less than 90uin O. ampay Fig. 11. Orobothriurus Maury, 1976, sternite VII and metasomal segments I–V, ventral aspect, showing pigmentation pattern. A. Orobothriurus ampay Ochoa and Acosta, 2003, allotype U (MUSM). B. Orobothriurus curvidigitus (Kraepelin, 1911), U (MHNC). C. Orobothriurus huascaran, n. sp., paratype U (MHNC). D. Orobothriurus atiquipa Ochoa and Acosta, 2002, - paratype (MHNC). Scale bars 51 mm. 22 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 (fig. 25A), but greater than 90uin other species (figs. 25B, D, 26B, C). The acuminate conical apophysis on the internal surface of the male pedipalp chela manus, observed in all other Orobothriurus species (except O. wawita, in which it is absent altogether), is reduced to a few granules in O. ampay. The hemispermatophore of O. ampay resembles that of O. parvus and O. wawita in possessing an elongated apex and a short frontal crest, which is less than half the length of the lamina. However, the ventral margin of the apex is inclined to the dorsal surface and curved distally in O. ampay (fig. 27A) and O. parvus (fig. 27B), but straight in O wawita (fig. 27C). Finally, O. ampay possesses the lowest pectinal tooth count in the genus (12–15). D ISTRIBUTION : Orobothriurus ampay is known only from the Santuario Nacional Ampay, near Abancay in the Apurı ´mac Department of southeastern Peru, at 3100– 3580 m (figs. 3E, 52). E COLOGY : Orobothriurus ampay appears to be endemic to an inter-Andean forest of Podocarpus glomeratus Don (Podocarpaceae) (fig. 3E). All known records of this species were collected inside the forest under stones on moist soil. No other scorpion species are known from this forest. The closest records are two other bothriurids, Brachistosternus peruvianus Piza, 1974, and Pachakutej iskay Fig. 12. Orobothriurus Maury, 1976, tergite VII (A,C,Donly) and metasomal segments I–IV, dorsal aspect, showing pigmentation pattern. A. Orobothriurus huascaran, n. sp., paratype U (MHNC). B. Orobothriurus parvus Maury, 1976, U (MHNC). C. Orobothriurus curvidigitus (Kraepelin, 1911), U (MHNC). D. Orobothriurus ampay Ochoa and Acosta, 2003, allotype U (MUSM). Scale bars 51 mm. 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 23 Fig. 13. Orobothriurus Maury, 1976, tergites, sternites and metasomal segments I–V, showing pigmentation pattern. A, B. Orobothriurus paessleri (Kraepelin, 1911), U (MHNC), metasomal segments I– V, ventral aspect (A), tergite VII and metasomal segments I–IV, dorsal aspect (B). C, D. Orobothriurus wawita Acosta and Ochoa, 2000, U (MHNC), tergite VII and metasomal segments I and II, dorsal aspect (C), sternite VII and metasomal segments I–V, ventral aspect (D). E. Orobothriurus curvidigitus (Kraepelin, 1911), U (MHNC), tergite V, dorsal aspect. F. Orobothriurus alticola (Pocock, 1899), U (MACN-Ar), tergites VI and VII, dorsal aspect. G. Orobothriurus grismadoi Ojanguren-Affilastro et al., 2009, tergites VI and VII, dorsal aspect. Scale bars 51 mm. 24 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 Fig. 19. Orobothriurus Maury, 1976, sternite VII and metasomal segments I and II, ventral aspect. A. Orobothriurus curvidigitus (Kraepelin, 1911), U (MHNC). B. Orobothriurus paessleri (Kraepelin, 1911), U (MHNC). C. Orobothriurus quewerukana, n. sp., paratype - (MHNC). D. Orobothriurus huascaran,n. sp., paratype U (MACN-Ar). E. Orobothriurus wawita Acosta and Ochoa, 2000, paratype U (MHNC). F. Orobothriurus tamarugal, n. sp., holotype - (MACN-Ar). Scale bars 51 mm. 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 31 Fig. 20. Orobothriurus Maury, 1976, metasomal segment V, ventral aspect. A. Orobothriurus alticola (Pocock, 1899), U (MACN-Ar). B. Orobothriurus compagnuccii, n. sp., paratype - (MACN-Ar). C. Orobothriurus ramirezi, n. sp., paratype - (MACN-Ar). D. Orobothriurus quewerukana , n. sp., paratype - (MHNC). E. Orobothriurus tamarugal, n. sp., holotype - (MACN-Ar). F. O. ramirezi, n. sp., paratype U (MACN-Ar). Scale bars 51 mm. 32 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 Fig. 21. Orobothriurus Maury, 1976, metasomal segment V, ventral aspect. A. Orobothriurus atiquipa Ochoa and Acosta, 2002, paratype - (MHNC). B. Orobothriurus paessleri (Kraepelin, 1911), - (MACNAr). C. Orobothriurus wawita Acosta and Ochoa, 2000, paratype - (MHNC). D. Orobothriurus calchaqui, n. sp., paratype - (MACN-Ar). E. Orobothriurus grismadoi Ojanguren Afillastro et al., 2009, paratype - (MACN-Ar). F. Orobothriurus huascaran, n. sp., paratype - (MACN-Ar). Scale bars 51 mm. 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 33 Fig. 22. Orobothriurus Maury, 1976, metasomal segment V, ventral aspect. A. Orobothriurus ampay Ochoa and Acosta, 2003, paratype U (MHNC). B. Orobothriurus paessleri (Kraepelin, 1911), U (MHNC). C. Orobothriurus curvidigitus (Kraepelin, 1911), U (MHNC). D. Orobothriurus huascaran, n. sp., paratype U (MACN-Ar). E. Orobothriurus parvus Maury, 1976, U (MHNC). F. Orobothriurus wawita Acosta and Ochoa, 2000, paratype U (MHNC). Scale bars 51 mm. 34 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 sublaterally (fig. 29B, D). Metasomal segment I, dorsal surface unpigmented or with two small faint spots medially; lateral surfaces densely pigmented between ML and LIM carinae (fig. 29); ventral surface with two narrow VL stripes, becoming slightly broader in posterior half. Segment II, dorsal surface with two subtriangular spots medially; lateral surface as for segment I; ventral surface with one narrow VM and two narrow VL stripes, becoming slightly broader in posterior half. Segment III, dorsal surface as for segment II; lateral surfaces densely pigmented below ML carinae; ventral surface as for segment II. Segment IV, dorsal surface with reticulate pigmentation medially; lateral and ventral surfaces as for segment III. Segment V, dorsal and lateral surfaces as for segment IV; ventral surface with three dark narrow stripes (one VM and two VL), becoming slightly broader in posterior half but not contiguous with lateral pigmentation at posterior margin. Telson vesicle with narrow VM and two broad, dark VL stripes, separated by two narrow unpigmented stripes; aculeus sclerotized, dark reddish brown. Chelicerae: Movable finger with two subdistal teeth. Carapace: Surfaces smooth medially, finely granular near lateral margins. Anterior margin with shallow median notch (fig. 14A). Anteromedian longitudinal sulcus weakly developed; median ocular, posteromedian longitudinal and posterolateral sulci well developed. Median ocular tubercle raised, situated anteromedially; median ocelli two ocular diameters apart. Pedipalps: Femur, DI and VI carinae complete, finely granular; DE carina complete, finely granular proximally (fig. 30A); internal surface finely and sparsely granular medially; ventral surface finely granular proximally; other surfaces smooth. Patella, DI and VI carinae complete, finely granular (fig. 30B–D); DPP carina finely granular; VPP carina vestigial, comprising two or three small granules proximally; internal surface with prominent granule adjacent to trichobothrium inear DI carina. Chela manus slender, fingers relatively elongated (fig. 31); length/width ratio: - , 3.97–4.51 (n510; mean 54.17), U , 3.47–3.73 (n54, mean 5 3.63); length/height ratio: - , 3.40–4.03 (n5 10; mean 53.71), U , 3.07–3.31 (n54, mean 53.18); D, DS, DMA, DI, and VM carinae obsolete; intercarinal surfaces smooth; internal surface with acuminate apophysis ( - )or low bulge ( U ) near articulation of movable finger (fig. 31A, D); fingers, dentate margins each with median denticle row and 4–5 pairs of internal and external accessory denticles. Trichobothria: Femur with 3 trichobothria, patella with 19, chela with 27 (fig. 30, 31). Patella trichobothrium em 2 situated proximal to em 1 in 80%of specimens (fig. 30C). Chela trichobothrium Et 3 situated proximal to Est (fig. 31C). Tergites: Tergites I–VI, surfaces finely granular, becoming smooth posterolaterally. Tergite VII tetracarinate, paired DL carinae restricted to posterior two-thirds of segment, paired DSM carinae to posterior half; intercarinal surfaces coarsely granular, other surfaces finely granular. Legs: Femur and patella, prolateral surfaces finely granular, retrolateral surfaces smooth. Femur, ventral carinae weakly developed; other carinae absent. Patella acarinate. Telotarsi, proand retroventral rows of spiniform macrosetae with following counts on leg I, 1/1; II,2/2;IIIandIV,3/3. Pectines: Pectinal tooth count: - , 18–21 (n5 21, mode 518); U ,14(n54, mode 514). Sternites: Sternites III–VI, surfaces smooth; spiracles small, narrow. Sternite VII, surface smooth; VSM and VL carinae well developed ( U , fig. 17B) or obsolete ( - ). Metasoma: Segment I, DL carinae complete, moderately granular; ML carinae complete, moderately granular in posterior two-thirds; one pair of ML macrosetae; LIM carinae complete, moderately granular in posterior half; VL and VSM carinae complete, well developed (especially in U ), granular (fig. 17B); three pairs of VL and two pairs of VSM macrosetae. Segment II, DL carinae complete, granular; ML carinae complete, granular in posterior two-thirds; one pair of ML macrosetae; LIM carinae restricted to posterior third; VL and VSM carinae complete, well developed ( U )or obsolete ( - ); three pairs of VL and VSM macrosetae. Segment III, DL carinae complete, granular; ML carinae complete, granular in posterior third; one pair of ML 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 35 Fig. 23. Orobothriurus Maury, 1976, telson, lateral aspect. A. Orobothriurus alticola (Pocock, 1899), - (MACN-Ar). B. Orobothriurus calchaqui, n. sp., paratype - (MACN-Ar). C. Orobothriurus compagnuccii,n.sp.,paratype - (MACN-Ar). D. Orobothriurus huascaran,n.sp.,paratype - (MACN-Ar). E. Orobothriurus grismadoi Ojanguren Afillastro et al., 2009, paratype - (MACN-Ar). F. Orobothriurus quewerukana, n. sp., paratype - (MHNC). G. Orobothriurus ramirezi, n. sp., paratype - (MACN-Ar). H. Orobothriurus tamarugal, n. sp., holotype - (MACN-Ar). Scale bars 51 mm. 36 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 Fig. 24. Orobothriurus Maury, 1976, telson, lateral aspect. A. Orobothriurus calchaqui, n. sp., paratype U (MACN-Ar). B. Orobothriurus compagnuccii, n. sp., paratype U (MACN-Ar). C. Orobothriurus huascaran, n. sp., paratype U (MACN-Ar). D. Orobothriurus quewerukana, n. sp., paratype U (MHNC). E. Orobothriurus ramirezi, n. sp., paratype U (MACN-Ar). F. Orobothriurus tamarugal, n. sp., paratype U (MZUC). Scale bars 51 mm. 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 37 macrosetae; LIM carinae obsolete, restricted to posterior quarter; VL and VSM carinae complete, well developed ( U ) or absent ( - ); three pairs of VL and VSM macrosetae. Segment IV, DL carinae complete, granular; one pair of DL macrosetae; ML carinae obsolete, restricted to posterior margin; one pair of ML macrosetae; LIM carinae absent; VL and VSM carinae absent; three pairs of VL and VSM macrosetae. Segment V, length/width ratio, - , 2.33–2.73 (n510; mean 52.52), U , 1.93–1.98 (n54; mean 51.96); DL carinae granular, restricted to anterior half; ML carinae absent; four or five pairs of ML macrosetae; lateral margin smooth; VL carinae complete ( - ) or restricted to posterior three-quarters ( U ); VL and VSM carinae situated close together, fused in anterior and Fig. 25. Orobothriurus Maury, 1976, dextral pedipalp chela. A. Orobothriurus ampay Ochoa and Acosta, 2003, paratype U (MHNC), external aspect. B, C. Orobothriurus wawita Acosta and Ochoa, 2000, paratype - (MHNC). B. External aspect. C. Ventral aspect. D. Orobothriurus atiquipa Ochoa and Acosta, 2002, paratype - (MHNC), external aspect Scale bars 51 mm. 38 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 posterior thirds; four pairs of VL and VSM macrosetae; two pairs of macrosetae along posterior margin; VM carina complete, obscured by surface granulation in posterior half (fig. 21D). Telson: Length/height ratio: - , 3.52–3.89 (n510, mean 53.74); U , 2.93–3.13 (n54, mean 53.01). Vesicle elongated ( - , fig. 23B) or globose ( U , fig. 24A); dorsal surface slightly concave, gland not apparent ( - ); ventral surface smooth ( - ) or granular ( U ). Aculeus short and curved, more so in U . Hemispermatophore: Apex very well developed; distal crest curved like ventral margin. Frontal crest weakly developed, less than half length of lamina; basal part oblique; distal part short, parallel to ventral margin of lamina; lateral projections slightly undulated. Basal lobe, terminal process extending almost to distal end of frontal crest (fig. 32A, B). R EMARKS : We examined some Orobothriurus specimens from other Andean localities in Salta and Tucuma´n provinces of northern Argentina. However, specimens from Fig. 26. Orobothriurus Maury, 1976, dextral pedipalp chela, external aspect. A. Orobothriurus parvus Maury, 1976, - (MHNC). B. Orobothriurus paessleri (Kraepelin, 1911), - (MACN-Ar). C. Orobothriurus curvidigitus (Kraepelin, 1911), - (MHNC), external aspect. Scale bars 51 mm. 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 39 these localities are so scarce and the morphological differences between them and O. calchaqui so subtle that at this time it is unclear whether or not they are conspecific. More specimens from these Orobothriurus populations are required to establish their identity. D ISTRIBUTION : This species is known only from the type locality, Infiernillo Mountain (Tucuma´n Province, Argentina), belonging to the Cumbres Calchaquies, part of the Andes (fig. 54). Altitude records from Huaca Huasi (4250 m) and Vaca Huasi (4300 m) should be confirmed, because, at this latitude, the environment above 4000 m is usually too extreme for scorpions to survive. E COLOGY : All personally collected specimens were found at night in a rocky area near a small mountain river, at 2700 m. The area is a typical Puna habitat (Cabrera and Willink, 1980). Most specimens collected were on the ground, but some were taken from near-vertical rock walls besides the river, almost 2 m from its base. No specimens could be found in the surrounding area, which has a sandy soil without rocks. Most of the specimens collected were adult males, moving very rapidly, probably searching for females, suggesting that late November, the start of spring in the Southern Hemisphere, is the start of the active period for this species. Fig. 27. Orobothriurus Maury, 1976, sinistral hemispermatophore, ectal aspect. A. Orobothriurus ampay Ochoa and Acosta, 2003, holotype - (MUSM). B. Orobothriurus parvus Maury, 1976, - (MHNC). C. Orobothriurus wawita Acosta and Ochoa, 2000, paratype - (MHNC). Scale bar 50.5 mm. 40 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 Fig. 33. Orobothriurus compagnuccii, n. sp., habitus. A, B. Paratype - (MACN-Ar). C, D. Paratype U (MACN-Ar). A, C. Dorsal aspect. B, D. Ventral aspect. Scale bars 55 mm. 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 47 Hemispermatophore: Apex well developed, slightly longer than frontal crest, comprising 52.17–54.54%(n53; mean 552.98%)of lamina; distal crest curved like ventral margin. Frontal crest divided into two parts, basal part oblique, distal part parallel to ventral margin of lamina; lateral projections slightly undulated and larger than basal part. Basal lobe, terminal process extending to median part of frontal crest (fig. 32C, D). D ISTRIBUTION : Orobothriurus compagnuccii is known only from the type locality, Laguna Brava Provincial Park, in the western Andres of La Rioja Province, northwestern Argentina (fig. 54). All specimens were collected on the ascent to Pircas Negras international pass, on the border between Argentina and Chile, at 3200–3900 m. E COLOGY : The habitat at the type locality is intermediate between Puna and high Andean habitats (Cabrera and Willink, 1980), with very sparse vegetation. All specimens were collected in areas with very pronounced slopes, on high rocks or spiny shrubs. We conducted several expeditions to the type locality of this species, in different years and at different times during the summer, when Orobothriurus are presumed to be most active at this latitude. However, despite considerable effort, only a few specimens were collected each time. Most were very difficult to find, hiding in inaccessible places, e.g., in vertical rocks more than two or three meters above ground level. In the same area, we collected many Brachistosternus montanus, a bothriurid species more than twice the size of O. compagnuccii. The scarcity of O. compagnuccii during summer expeditions, and its habitat preference for places inaccessible to larger scorpions adapted to sandy soils like Brachistosternus, may be explained by the abundance of B. montanus. It is possible that the active period of O. compagnuccii starts in early spring, before that of B. montanus, to avoid competition and predation by the larger species. Orobothriurus curvidigitus (Kraepelin, 1911) Figures 11B, 12C, 13E, 14C, 19A, 22C, 26C, 28B, D, 53 Bothriurus curvidigitus Kraepelin, 1911: 91, 97–99, fig. 7a, 7b; Mello-Leita˜o, 1931: 82, 92; 1932: 34; 1934: 65; 1935: 93; 1937: 103; 1945: 161; Aguilar and Meneses, 1970: 2; Maury, 1973: 110; 1981: 104. Bothriurus lampei Werner, 1916: 92 [synonymized by Maury, 1973: 110]; Lampe, 1917: 203; MelloLeita˜o, 1931: 88; 1932: 34; 1945: 175; Aguilar and Meneses, 1970: 2; Maury, 1973: 110; 1981: 104. Bothriurus (Andibothriurus)curvidigitus:Bu¨cherl et al., 1963: 216. Bothriurus (Andibothriurus)lampei: Francke, 1974: 219. Orobothriurus curvidigitus: Maury, 1976: 18, 19, figs. 11–24, table I; 1980: 338; 1981: 101; Francke, 1977: 75; Kovar ˇik, 1998: 101; Acosta and Ochoa, 2000: 136, 143; Lowe and Fet, 2000: 35, 36; Acosta and Ochoa, 2001: 205; Ochoa and Acosta, 2002a: 102, fig. 9; Ochoa, 2004a: 52, 55, 73, figs. 1, 2, 13, 19, 21, table 1; 2005: 55, figs. 7, 10, table 2. T YPE M ATERIAL : PERU: Arequipa Department: Arequipa Province:1 - , 1 subad. Fig. 34. Orobothriurus compagnuccii, n. sp., paratype - (MACN-Ar), dextral pedipalp femur (A) and patella (B–D). A, B. Dorsal aspect. C. External aspect. D. Ventral aspect. Scale bar 5 1 mm. 48 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 Fig. 35. Orobothriurus compagnuccii, n. sp., dextral pedipalp chela. A, B. Paratype U (MACN-Ar). C–E. Paratype - (MACN-Ar). A. Dorsal aspect. B, C. External aspect. D. Ventral aspect. E. Ventrointernal aspect. Scale bars 51 mm. 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 49 U syntypes (ZMH), Yura [16u129530S71u429 330W, 2642 m], 20.vi.1909. N EW R ECORDS : PERU: Arequipa Department: Arequipa Province: Agua del Milagro, Characato [16u289S71u289W, 2460 m], 12.x.1983, E. Maury, 4 U , 8 juv. (MACN-Ar); Arequipa, 1937, P. Crawford, 1 U (MACNAr); Canchimayo, Chiguata [16u249130S 71u239340W, 2950 m], 18.x.1998, J.A. Ochoa, 4 juv. (MHNC); Charcani, Arequipa, 2800 m, xi.1947, W. Weyrauch, 3 U , 2 juv. (MACNAr); entrance to Charcani Quinto, near Arequipa city, Chili (Radio Azul) [16u199330S 71u319470W, 2550 m], 28.ii.1998, E. Ponce and J.A. Ochoa, 1 juv. (MHNC); Espı ´ritu Santo, Chiguata [16u249280S71u239070W, 2900 m], 7.iii.1998, E. Ponce and J.A. Ochoa, 2 U ,1juv.(MHNC);Miraflores,nearChiguata [16u239S71u219W,3295m],7.iii.1998,E.Ponce and J.A. Ochoa, 12 U , 3 juv. (MHNC), 3 U (CDA 166); near Mollebaya river [16u289560S 71u289010W, 2513 m], 3.iii.1998, E. Ponce, D. Mun˜iz and J.A. Ochoa, 2 - ,6 U , 2 juv. (MHNC), 1 - ,1 U (CDA 167); Ribera del Rio Mollebaya [ca. 16u289560S71u289010W, 2513 m], 8.ii.2004, R. Carden˜as and J.A. Ochoa, 2 juv. (AMNH [LP 3064]); Socabaya [16u289010S71u319180W, 2300 m], 1948, W. Weyrauch,1 - (MUSM),2329m,5.ix.1998,U. Zanabria, 1 U (MHNC); Yura [16u129530S 71u429330W, 2642 m], 7.viii.1939, K.P. Schmidt, 2 - , 1 juv. (FMNH), 8.viii.1939, K.P. Schmidt, 1 - ,1 U (FMNH), 10.viii.1939, K.P. Schmidt, 1 - , 1 juv. (AMNH), 9.v.1998, U. Zanabria, 1 U , 2 juv. (MHNC); Yura, Rı ´o Yura, 20 m del Cementerio, 16u139590S 71u429270W,2426m, 29.xii.2007,R. Gutie´rrez, A. Quiroz and J.A. Ochoa, Serrania Esteparia, collectedat nightwithUV light,3juv.(AMNH [LP 8357]). Cailloma Province: Calera Chivay [15u389340S71u369010W, 3640 m], 27.ix.1998, S. Rivera, 1 U (MHNC); Curin˜a, Yanque [15u399340S71u429150W, 3390 m], 20.xii.1996, J.A. Ochoa, 10 - , 3 juv. (MHNC); Curin˜a [3600 m], 21.xii.1996, J.A. Ochoa, 2 U , 4 juv. (MHNC). D IAGNOSIS : Orobothriurus curvidigitus is most closely related to O. paessleri,O. quewerukana,andO. tamarugal (fig. 5), all of which have the following characters: anterior margin of carapace with weak median projection (epistome; figs. 14C, 15A, C, D); pedipalp chela movable finger of male curved (figs. 26B, C, 43C, 51A); pedipalp femur and patella of male more elongated than that of female. Orobothriurus curvidigitus may be separated from O. paessleri and O. tamarugal by the shape of the lamina of the hemispermatophore (figs. 28B, D), the frontal crest of which is more developed and elongated in O. curvidigitus, but shorter in O. paessleri and O. tamarugal (fig. 28C, E, 47D). The hemispermatophore of O. curvidigitus differs from that of O. quewerukana in the slightly shorter frontal crest and absence of folds in the distal crest (figs. 28B, 40E). Orobothriurus curvidigitus also differs from O. quewerukana in the shape and granulation of the male telson vesicle: the length/height ratio is 3.72–4.23 (mean 53.96) and only the anterior third of the ventral surface is granular in O. curvidigitus, whereas the length/height ratio is 3.59– 4.00 (mean 53.76) and the ventral surface is entirely granular in O. quewerukana (fig. 23F). The male pedipalp chela carinae are finely and sparsely granular to smooth in O. curvidigitus, smooth in O. paessleri, and finely and densely granular in O. quererukana and O. tamarugal (figs. 43D, E, 51B, C). The pigmentation pattern is the most conspicuous difference between O. curvidigitus and the other three species. Orobothriurus curvidigitus exhibits well-developed pigmentation on the carapace, tergites, metasoma, and pedipalps (figs. 11B, 12C, 13E), compared to O. quewerukana, in which the pigmentation, especially that of the carapace and tergites, is faint or barely discernible (fig. 41) and O. tamarugal, which is unpigmented (fig. 48). Orobothriurus curvidigitus and O. paessleri may be further distinguished by the dimensions of metasomal segment V and the setation of metasomal segment I: segment V is relatively elongated, its length greater than twice its width, in O. curvidigitus (fig. 22C), but shorter, its length less than twice its width, in O. paessleri (fig. 22B); two pairs of VSM macrosetae are present on segment I in O. curvidigitus (fig. 19A), whereas three pairs are present in O. paessleri (fig. 19B). D ISTRIBUTION : Orobothriurus curvidigitus inhabits the western slopes of the Andes at 2300–3600 m (figs. 2A, 53), in the Arequipa Department, southern Peru. 50 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 Fig. 36. Orobothriurus Maury, 1976, sinistral hemispermatophore, ectal aspect showing lamina. A–C. Orobothriurus compagnuccii, n. sp., - holotype, - paratypes (MACN-Ar). D, E. Orobothriurus grismadoi Ojanguren Affilastro et al., 2009, - paratypes (MACN-Ar). F–H. Orobothriurus famatina Acosta, 2001, - (MACN-Ar). I, J. Orobothriurus alticola (Pocock, 1899), - (MACN-Ar). K–O. Orobothriurus calchaqui, n. sp., - paratypes (MACN-Ar). Scale bar 51 mm. 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 51 E COLOGY : The vegetation in the areas inhabited by this species falls within the Serrania Estaparia Ecoregion, characterized by dry zones with herbaceous vegetation, cacti, shrubs, and few trees (Brack, 1986; Ochoa, 2005; fig. 2A). Three other bothriurids, Brachistosternus ehrenbergii Gervais, 1841, and two undescribed species of Brachistosternus, also occur in this area. R EMARKS : We examined populations of an undetermined Orobothriurus species, apparently related to O. curvidigitus, from Cotahuasi and Chuquibamba in the Arequipa Department, Peru, and Lucanas in the Ayacucho Department. Lourenc¸o and Dastych (2001: 54) listed the record from Chuquibamba under ‘‘O. aff. curvidigitus’’ and stated that it is ‘‘probably a new species.’’ We cannot establish the identity of these specimens until more material becomes available, however. Orobothriurus famatina Acosta, 2001 Figures 36F–H, 54 Orobothriurus alticola: Maury 1976: 17 (part); Cei, 1982: 660, 669 [misidentification]. Orobothriurus famatina Acosta in Acosta and Ochoa, 2001: 205–211, figs. 4–14; Ochoa, 2004a: 43, 52, 55, 73, table 1; Ojanguren Affilastro, 2005: 177–181, figs. 458–466, 477, 488, 663, table 15. T YPE M ATERIAL : ARGENTINA: La Rioja Province: Famatina Department: Holotype - (MACN-Ar 9932), Sierra de Famatina, Rio Oro canyon, path to Mina El Oro, 29u069S 67u429W, 2450 m, 6.xii.1998, L. Acosta, M. Acosta and G. Repossi. Paratypes: same data, 1 - ,1 U (LEA 240), 1 - (CDA 051), 1 - (MHNC); Famatina, iv.1951, R. Gonza´lez Amorı ´o, 1 U (MACN-Ar 6844); Mina El Oro, Chilecito, ii.1956, M.E. Galiano, 1 U (MACN-Ar 6843); Sierra de Famatina, path to Mina La Mejicana, 3060 m, 5.xii.1998, L. Acosta, M. Acosta, and G. Repossi, 1 U (CDA 050), 1 U (MACN-Ar 9933), 1 U (MHNC). N EW R ECORDS : ARGENTINA: La Rioja Province: Famatina Department: Sierra de Famatina, Cuevas de Noronha (path to La Mejicana mine), 28u559310S67u409230W, 2846 m, 29.i.2006, L. Piacentini, L. Compagnucci, and A. Ojanguren Affilastro, 2 - ,6 U , 12 juv. (MACN-Ar), 2 U (AMNH [LP 5846]). D IAGNOSIS : Orobothriurus famatina is most closely related to O. calchaqui (fig. 5), from which it differs in the following respects. Orobothriurus famatina possesses one ML macroseta on metasomal segment IV and only 10%of the specimens possess one DL macroseta on segment III (absent in 90%), whereas O. calchaqui possesses two or three macrosetae on segment IV and 70%of the specimens possess one DL macroseta on segment III (absent in 30%). There are also differences in the trichobothrial pattern. Pedipalpal chela trichobothria db and Et 3 are located in the same axis as, or slightly distal to Est in 70%and 30%of the specimens of O. famatina, respectively, but always located proximal to Est in O. calchaqui. Pedipalp patella trichobothrium em 2 is located in the same axis as, or distal to em 1 in 70%of the specimens of O. famatina, but proximal to em 1 in 80%of the specimens of O. calchaqui.Orobothriurus famatina may be differentiated from O. compagnucci by the more elongated apex of the hemispermatophore lamina: the apex comprises 66.6– 68.48%(n53, mean 567.02%) of the lamina in O. famatina (fig. 36F–H) and 52.17%–54.54%(n53; mean 552.98%)in O. compagnuccii (fig. 36A–C). The two species can also be separated by the VL and VSM carinae of metasomal segment I and sternite VII in the male, which are well developed in O. famatina, and weakly developed to absent in O. compagnuccii (fig. 18F). D ISTRIBUTION : Orobothriurus famatina is known only from high altitudes (2500– 3200 m) of the Sierra Famatina, a mountain range close to the Andes, in La Rioja Province of central-western Argentina (figs. 3G, 54). E COLOGY : This species inhabits an area of grassland and shrub steppe that belongs to the Prepuna and Altoandina phytogeographic provinces (Cabrera and Willink, 1980; fig. 3G). It occurs in sympatry with another bothriurid, Bothriurus olaen Acosta, 1997 (Ojanguren Affilastro, 2005). Orobothriurus grismadoi Ojanguren Affilastro et al., 2009 Figures 4B, 13G, 21E, 23E, 36D, E, 54 Orobothriurus grismadoi Ojanguren Affilastro et al., 2009: 29–35, figs. 1, 3–13, 15–17, 20–24, 35, tables 1, 2. 52 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 Fig. 37. Orobothriurus huascaran, n. sp., habitus. A, B. Paratype - (MACN-Ar). C, D. Paratype U (MACN-Ar). A, C. Dorsal aspect. B, D. Ventral aspect. Scale bars 510 mm. 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 53 T YPE M ATERIAL : ARGENTINA: Mendoza Province: Malargu¨e Department: Holotype - (MACN-Ar 17986), Cerro Nevado, 35u35945.060S68u30924.120W, 3130 m, 25.ii.2006, F. Ferna´ndez Campo´n and S. Lagos Silnik. Paratypes: same data, 6 - ,2 U (MACN-Ar 17987), 2 - (CDA), 1 juv. (IADIZA); Cerro Nevado, 35u36904.080S 68u30944.280W, 2900 m, 16.xi.2004, G. Flores, 1 U (CDA), 2949 m, 7.i.2006, F. Ferna´ndez Campo´n and S. Lagos Silnik, 1 juv. (IADIZA); Cerro Nevado, 35u36902.460S 68u30940.920W, 2953 m, ii.2006, F. Ferna´ndez Campo´n and S. Lagos Silnik, 1 - (AMNH), same data except 25.ii.2006, 5 - (IADIZA), 1 - (AMNH). N EW R ECORDS : ARGENTINA: Mendoza Province: Malargu¨e Department: Cerro Nevado, road to antennas, SW face of mountain, 35u35958.60S68u30943.90W, 2984 m, 23.iv.2011, C.I. Mattoni and M. Vivanco, UV sampling on rocky slopes, with small bushes and shrubs, no moon, 3uC, winds up to 45 km/h, wind chill factor 210uC, specimens walking or sitting on rocks, 5 - , 6 U , 4 juv. (LBRE), 4 juv. (AMNH), 1 juv. (AMNH [LP 10968]). D IAGNOSIS : Orobothriurus grismadoi is similar to O. alticola in pigmentation pattern, external morphology, and hemispermatophore. The two species may be separated by the shape of the hemispermatophore: in O. grismadoi, the apex is more slender and the angle it forms with the rest of the distal lamina more acute (fig. 36D, E) than in O. alticola (fig. 36I, J). Additionally, the apex comprises 42.70%–45.69%(n57; mean 544.17%)of the lamina in O. grismadoi and 40.87–46.08% (n520; mean 543.88%)inO. alticola. Males of O. grismadoi may also be recognized by the slender telson vesicle, with a concave dorsal surface; the vesicle is broader, with a flat dorsal surface, in males of O. alticola (fig. 23A, E). Orobothriurus grismadoi is also more densely pigmented, with tergite VII entirely pigmented (fig. 13G), compared to O. alticola, in which tergite VII exhibits paired lateral spots of pigmentation delimiting an unpigmented median stripe (fig. 13F). D ISTRIBUTION : Orobothriurus grismadoi is the southernmost species of the genus. It is known only from the Cerro El Nevado, an isolated mountain range, 200 km east of the Andes, in central-western Mendoza Province, Argentina (fig. 54). The El Nevado range, with a maximum altitude of 3833 m, extends north to south between 34uSand36uS, parallel to the Andes, and is separated from it by a plateau of 1800 m (Ojanguren Affilastro et al., 2009). El Nevado is an extinct, eroded, stratovolcano, unrelated to and younger than the Andes. The age of these volcanoes is estimated at between 2.5 and 0.01 MA (Caminos, 1999). E COLOGY : Orobothriurus grismadoi was collected in a high-altitude shrub steppe at 2900–3130 m (fig. 3D), in an area belonging to the Altoandina and Puna phytogeographic provinces (Cabrera and Willink, 1980). Fig. 38. Orobothriurus huascaran,n.sp.,paratype - (MHNC), dextral pedipalp femur (A) and patella (B–D). A, B. Dorsal aspect. C. External aspect. D. Ventral aspect. Scale bar 51 mm. 54 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 Fig. 39. Orobothriurus huascaran, n. sp., dextral pedipalp chela. A, B. Paratype U (MACN-Ar). C–E. Paratype - (MACN-Ar). A. Dorsal aspect. B, C. External aspect. D. Ventral aspect. E. Ventrointernal aspect. Scale bars 51 mm. 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 55 Arthropod surveys conducted using pitfall traps in the summers of 2004 and 2006 did not collect other scorpion species. A recent expedition to the area confirmed that this species is the only scorpion present and active during the summer at this altitude on the El Nevado volcano. All specimens were collected at night in rocky areas (mostly basalt and pumice), and efforts to find the species on sandy slopes (with ash and pumice fragments) surrounding these rocky habitats were unsuccessful. Despite the harsh conditions, with very low temperatures and strong winds during the night, the species is apparently active on the surface from midNovember to the end of April. Orobothriurus huascaran, n. sp. Figures 4D, 11C, 12A, 14D, 19D, 21F, 22D, 23D, 24C, 37–39, 40A–C, 52; table 3 Orobothriurus crassimanus: Polis, 1990: 252; Lourenc¸o, 1997: 588 (part); Coddington and Colwell, 2001: 207; Lourenc¸o, 2002: 400; 2003: 227; Lourenc¸o and Qi, 2006: 290. [misidentification]. Orobothriurus paessleri: Lourenc¸o and Dastych, 2001: 54 (part) [misidentification: 1 U , from Quebrada Yanganuco]. Orobothriurus sp. 2: Ochoa, 2004a: 50, 73, figs. 1, 2, 14, 21, table 1. T YPE M ATERIAL : PERU: Ancash Department: Huaraz Province: Holotype - (MHNC), Cordillera Blanca, Huascara´n National Park, Fig. 40. Orobothriurus Maury, 1976, sinistral hemispermatophores. A–C. Orobothriurus huascaran,n. sp., paratype - (MACN-Ar). D, E. Orobothriurus quewerukana, n. sp., paratype - (MACN-Ar). A, D. Ental aspect. B, E. Ectal aspect. C. Dorsal aspect. Scale bar 51 mm. 56 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 huascaran (fig., 39C), but equidistant between Dt and Eb 3 in other species. The VM and VL carinae are restricted to the posterior two-thirds of metasomal segment V in O. huascaran (fig. 21F), absent in O. wawita and complete in all other species. Orobothriurus huascaran is similar to four other Peruvian species, O. atiquipa,O. curvidigitus,O. paessleri, and O. quewerukana, in several respects. Pedipalp chela trichobothrium Et 3 is situated proximal to Est, and Esb situated between Eb 2 and Eb 3 in all these species, except O atiquita, in which Esb is situated dorsal to Eb 2 . The hemispermatophore of these species exhibits an elongated frontal crest, with a strongly curved, S-shaped apex and a basal lobe with a well-developed terminal process. Orobothriurus huascaran may be distinguished by the presence of VM and VSM carinae on sternite VII and metasomal segment I (fig. 19D), which are absent in O. atiquipa,O. curvidigitus,O. paessleri,andO. quewerukana (figs. 18A, 19A, C). Additionally, the pedipalp chela movable finger of the male is straight in O. huascaran (fig. 39C) and curved in O. curvidigitus,O. paessleri,andO. quewerukana (fig. 26B, C, 43C). D ESCRIPTION : Based on holotype - and paratypes. Measurements of holotype - and paratype U recorded in table 3. Total length: - , 26.43–30.79 mm (n59, mean 527.97 mm); U , 30.05–33.21 mm (n5 10, mean 531.65 mm). Color: General color yellowish with dark brown spots (fig. 4D). Carapace densely pigmented, especially laterally and posteriorly; anterior margin with narrow stripe (fig. 37A, C); median lateral surfaces densely pigmented; posterolateral surfaces with dense reticulate pigmentation; median ocular tubercle and lateral ocelli dark brown to black; posterior half of anteromedian longitudinal sulcus, median ocular tubercle and anterior half of posteromedian longitudinal sulcus densely pigmented; posterolateral sulci unpigmented. Chelicerae, dorsal surfaces with fine reticulate pigmentation, contiguous distally near base of movable finger; movable finger external surface pigmented. Pedipalp coxa and trochanter faintly pigmented; femur densely pigmented along margins with small unpigmented areas in proximal half; patella densely and irregularly pigmented; chela manus with faint longitudinal stripes along carinae, contiguous at base of movable finger. Legs, femur, and patella densely pigmented prolaterally; tibia and basitarsus faintly pigmented. Tergites I–VI each with two dark spots sublaterally along entire margin and pretergites, delimiting broad, unpigmented median stripe; VII with similar pattern and additional reticulate pigmentation (fig. 12A). Sternum, genital opercula and pectines unpigmented. Sternites III–VI unpigmented, VII with two dark stripes sublaterally (fig. 11C). Metasomal segments I–III, dorsal surfaces each with two subtriangular spots medially, separated by narrow unpigmented line on I, contiguous or separated on II, contiguous on III, becoming broader posteriorly and often connected to median pigmentation by fine reticulate pigmentation, with additional pigmentation along DL carinae; lateral surfaces densely pigmented, especially in anterior half, posterior half with reticulate pigmentation (figs. 11C, 12A); ventral surface with narrow VM stripe, not contiguous with lateral pigmentation, and with two dark VL stripes, becoming broader in posterior half and extending to LIM carinae. Segment IV as for segment III, but with dorsal surface more densely reticulate. Segment V, dorsal surface faintly pigmented along DL carinae; lateral surfaces with dense reticulate pigmentation; ventral surface with three dark stripes, two VL and one VM, becoming slightly broader in posterior half but not contiguous with lateral pigmentation (fig. 11C). Telson vesicle with narrow VM and two broad VL stripes, separated by two narrow unpigmented stripes; aculeus sclerotized, dark reddish-brown. Chelicerae: Movable finger with two subdistal teeth. Carapace: Surfaces smooth in anterior third, finely granular elsewhere. Anterior margin linear, without median notch (fig. 14D). Anteromedian longitudinal sulcus complete, well developed, more so in - ; median ocular and posteromedian longitudinal sulci well developed; posterolateral sulci obsolete. Median ocular tubercle raised, situated anteromedially; median ocelli two ocular diameters apart. 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 63 Pedipalps: Femur, DI and VI carinae complete, finely granular; DE carina obsolete, smooth (fig. 38A); internal surface finely and sparsely granular medially; other intercarinal surfaces smooth. Patella, DI and VI carinae obsolete, finely granular ( - )or smooth ( U ) (fig. 38B–D); internal surface with two prominent granules distally, the larger adjacent to trichobothrium i; other intercarinal surfaces smooth. Chela manus slender, fingers relatively elongated (fig. 39); length/width ratio: - , 3.46–4.14 (n511, mean 53.76), U , 3.37–4.14 (n520, mean 5 3.74); D, DS, DMA, DI, and VM carinae obsolete; intercarinal surfaces smooth; internal surface with acuminate apophysis ( - )or low bulge ( U ) near articulation of movable finger (fig. 39A, D); fingers, dentate margins each with median denticle row and 4–5 pairs of internal and external accessory denticles. Trichobothria: Femur with 3 trichobothria, patella with 19, chela with 27 (figs. 38, 39). Chela trichobothrium Et 3 situated proximal to Est;Esb situated between Eb 2 and Eb 3 (fig. 39). Tergites: Tergites I–VI, surfaces finely granular. Tergite VII tetracarinate, paired DL carinae restricted to posterior two-thirds of segment, paired DSM carinae to posterior half; surfaces more coarsely and densely granular in posterior half. Legs: Femur and patella, prolateral surfaces finely granular, retrolateral surfaces smooth. Femur, ventral carinae weakly developed; other carinae absent. Patella acarinate. Telotarsi, proand retroventral rows of spiniform macrosetae with following counts on leg I, 1/1; II, 2/2; III and IV, 3/3. Pectines: Pectinal tooth count: - , 17–20 (n530, mode 518); U , 14–17 (n556, mode 515). Sternites: Sternites III–VI, surfaces smooth; spiracles small, narrow. Sternite VII, surface smooth in anterior half, granular in posterior half; VL and VSM carinae well developed ( U , fig. 19D) or obsolete (indistinct from granulation) to absent ( - ). Metasoma: Segment I, DL carinae complete, more developed in U ; usually one pair of DL macrosetae; ML carinae complete; two pairs of ML setae; LIM carinae obsolete, entirely smooth or with few granules in posterior third; VL and VSM carinae complete, obsolete, smooth ( - ) or granular ( U ); three pairs of VL and two or three pairs of VSM macrosetae (fig. 19D). Segments II and III, DL carinae complete; one pair of DL macrosetae; ML carinae restricted to posterior half ( - ) or complete but less developed than on segment I ( U ); two pairs of ML macrosetae; LIM carinae reduced to one or two granules posteriorly ( U ) or absent ( - )on segment II, absent on III; VL carinae as for segment I, but less developed ( U ) or absent ( - ) on II, obsolete ( U ) or absent ( - ) on III; VSM carinae as for segment I, but less developed on II, obsolete ( U ) or absent ( - ) on III; three pairs of VL and VSM macrosetae. Segment IV, DL carinae complete; one or (usually) two pairs of DL macrosetae; ML carinae reduced to few small granules posteFig. 45. Orobothriurus ramirezi, n. sp., paratype - (MACN-Ar), dextral pedipalp femur (A) and patella (B–D). A, B. Dorsal aspect. C. External aspect. D. Ventral aspect. Scale bar 51 mm. 64 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 Fig. 46. Orobothriurus ramirezi, n. sp., dextral pedipalp chela. A, B. Paratype U (MACN-Ar). C–E. Paratype - (MACN-Ar). A. Dorsal aspect. B, C. External aspect. D. Ventral aspect. E. Ventrointernal aspect. Scale bar 51 mm. 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 65 riorly; LIM carinae absent; VL and VSM carinae absent; three pairs of VL and four pairs of VSM macrosetae. Segment V, length/ width ratio: - , 2.07–2.40 (n55, mean 5 2.17), U , 1.71–2.02 (n57, mean 51.87); DL carinae reduced to three or four prominent granules anteriorly; lateral intercarinal surfaces smooth ( - ) or finely and sparsely granular medially near VL carinae ( U ); VL carinae usually restricted to posterior twothirds (but complete, weakly granular in anterior third in some U ), with terminal granules more developed; VSM and VM carinae usually obscured by surface granulation in posterior two-thirds (fig. 21F), but VM carinae faintly evident in some U (fig. 21D); four pairs of VL and VSM macrosetae; two pairs of macrosetae along posterior margin; ventral intercarinal surfaces densely granular in posterior two-thirds, more so in U , granules more acute in - . Telson: Length/height ratio: - , 2.86–3.52 (n511, mean 53.24); U , 2.61–3.39 (n519, mean 53.04). Vesicle slightly elongated ( - , fig. 23D) or globose ( U , fig. 24C); dorsal surface smooth, flat, gland not apparent ( - ); ventral surface smooth ( - ) or granular in anterior third ( U ). Aculeus short and curved. Hemispermatophore: Apex shorter than frontal crest, distal border slightly rounded; Fig. 47. Orobothriurus Maury, 1976, sinistral hemispermatophore. A, B. Orobothriurus ramirezi, n. sp., paratype - (MACN-Ar). C, D. Orobothriurus tamarugal, n. sp., holotype - (MACN-Ar). A, C. Ental aspect. B, D. Ectal aspect. Scale bar 51 mm. 66 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 Fig. 48. Orobothriurus tamarugal, n. sp., habitus. A, B. Holotype - (MACN-Ar). C, D. Paratype U (LBRE). A, C. Dorsal aspect. B, D. Ventral aspect. Scale bars 510 mm. 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 67 distal crest curved like ventral margin. Frontal crest elongated; basal part oblique; distal part parallel to ventral margin of lamina, lateral projections slightly undulated and larger than basal part, ental lateral projection vestigial, ectal lateral projection complete. Basal lobe, terminal process extending to constriction of frontal crest (fig. 40A–C). D ISTRIBUTION : Orobothriurus huascaran is endemic to the Ancash Department of central Peru (fig. 52). All known localities of this species occur in the Cordillera Blanca mountain range of the western Andes, the world’s highest tropical mountain range (fig. 3F). This species has been collected from 3730 m (Porganillo, trekking route to Aquilpo mountain) to 4910 m (Ishinca ravine). It is present above 4050 m in the Ishinca area, but more abundant at 4100 m; only four specimens were found above 4500 m. The record of this species from Laguna Paro´n is currently the northernmost record for the genus Orobothriurus. The record from the Ishinca ravine is the world’s highest record for a scorpion. E COLOGY : The area inhabited by O. huascaran represents typical Puna vegetation with shrub steppe, grasses and small patches of Polylepis spp. forest (‘‘quen˜ua’’ in Quechua), situated at 3800–4200 m (fig. 3F). All specimens were found under stones during the day. No other scorpion species have been recorded in the area. R EMARKS : ThefemalespecimenfromQuebrada Yanganuco, listed under O. paessleri by Lourenc¸o and Dastych (2001: 54), is probably conspecific with O. huascaran, n. sp., basedontheknowndistributionofthisspecies. Orobothriurus paessleri (Kraepelin, 1911) Figures 13A, B, 15A, 19B, 21B, 22B, 26B, 28C, E, 53 Bothriurus paessleri Kraepelin, 1911: 91, 98, 99, figs. 8, 9a, 9b; Mello-Leita˜o, 1931: 92; 1932: 34; 1934: 63; 1945: 181; Bu¨cherl, 1953: 117; Aguilar and Meneses, 1970: 2; Maury, 1973: 110. Bothriurus (Andibothriurus)paessleri:Bu¨cherl et al., 1963: 216. Orobothriurus paessleri: Maury, 1976: 19, 20, figs. 25–33, table I; 1981: 105; Francke, 1977: 75; Kovar ˇı ´k, 1998: 101; Acosta and Ochoa, 2000: 136, 143; Lowe and Fet, 2000: 36; Acosta and Ochoa, 2001: 205; Ochoa and Acosta, 2002a: 102, fig. 9; Ochoa, 2004a: 52, 55, 73, figs. 1, 2, 21, table 1; Ochoa, 2005: 55, figs. 7, 10, table 2. Orobothriurus curvidigitus: Lourenc¸o and Dastych, 2001: 54 [misidentification: 3 U from Mejı ´a]. T YPE M ATERIAL : PERU: Arequipa Department: Islay Province: Syntypes: 1 - ,3 U (ZMH), ‘‘Kataringo’’ [Catarindo], near Mollendo [17u009580S72u029030W, 34 m?], 25.ii.1907, R. Paessler. N EW R ECORDS : PERU: Arequipa Department: Camana´ Province: S of Camana´, 7.viii.1977, L. Pen˜a, 1 U (MACN-Ar). Islay Province: Lomas de Mejı ´a [17u029S71u519W, 530 m], 1–3.ii.2000, J.A. Ochoa, 1 - ,2 U ,3 juv. (MHNC), 1 - ,1 U (CDA 168); Lomas de Mejı ´a, Challascapa [17u029S71u519W, 550 m], 3.iii.2004, R. Gutierrez and J.A. Ochoa, 2 juv. (AMNH [LP 3057]); near Mejı ´a, 17u079S71u559W, 350–500 m, 15.i.1956, Koepcke, Park loma, Kp 1350, 1 - , 1 subad. - , 2 juv. - , 2 juv. U (ZMH A18/ 101) [misidentified by W.R. Lourenc¸o as ‘‘O. curvidigitus’’]; Lomas de Yuta, Matarani, 16u579160S72u049100W, 570 m, 12.iii.1998, E. Ponce and J.A. Ochoa, 1 U (MHNC). D IAGNOSIS : Orobothriurus paessleri is most closely related to O. curvidigitus,O. quewerukana and O. tamarugal (fig. 5), from which it may be separated as follows. Metasomal segment V is markedly shorter, its length less than twice its width, in the female of O. paessleri (fig. 22B), than in the females of other species, in which its length is greater than or equal to twice its width. Orobothriurus paeslleri possesses three pairs of VSM macrosetae on metasomal segment I (fig. 19B), whereas the other species possess two (fig. 19A). The pedipalp chela movable finger of the male is slightly curved in O. paessleri (fig. 26B), but strongly curved in the other species (figs. 26C, 43C, 51A). Orobothriurus paessleri may be further separated from O. curvidigitus and O. quewerukana by the shape of the lamina of the hemispermatophore (figs. 28B, C, 40E), the frontal crest of which is shorter and less developed in O. paessleri (fig. 28E) than in O. curvidigitus and O. quewerukana (fig. 28D). Additionally, the ventral surfaces of metasomal segments I–III are finely granular in O. paessleri (fig. 19B), whereas only the ventral surfaces of segment 68 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 I in males are finely granular in O. curvidigitus and O. quewerukana. D ISTRIBUTION : Orobothriurus paessleri appears to be endemic to the area near Mejı ´a and Yuta (Matarani) in the Arequipa Department of southern Peru (fig. 53). A single record from Toquepala (17u149S70u369W, 3000 m) in the Tacna Department (Maury, 1976), probably based on a misidentification, requires confirmation. E COLOGY : As with O. atiquipa, this species has only been collected in Lomas biotopes, isolated patches of green vegetation on hillsides below 1000 m, sustained by coastal fog, and surrounded by hyperarid desert. The Lomas vegetation near Mejı ´a and Yuta is dominated by herbaceous shrubs, e.g., Nolana L.f., Palaua Cav., Portulaca L., Weberbauerella Ulbr., and the tree Caesalpinia spinosa (Molina) Kuntze (Pe´faur, 1981; Da´vila Flores, 1982). R EMARKS : Specimens of O. paessleri from Mejı ´a deposited in the ZMH collection were examined, and found to be misidentified as O. curvidigitus by Lourenc¸o and Dastych (2001: 54). Currently two Orobothriurus species have been recorded from Lomas formations: O. atiquipa and O. paessleri. In addition, we examined a single female deposited at MHNC from Lomas de Sama, southern Peru. This specimen is poorly preserved and difficult to identify with certainty. Orobothriurus parvus Maury, 1976 Figures 4C, 9, 12B, 17A, 22E, 26A, 27B, 52 Orobothriurus parvus Maury, 1976: 17, 21–23, figs. 45–54, table I; Galiano and Maury, 1979: 327; Maury, 1980: 338, fig. 12; Kovar ˇı ´k, 1998: 101; Acosta and Ochoa, 2000: 136, 143; Lowe and Fet, 2000: 36; Acosta and Ochoa, 2001: 205; Lourenc¸o and Dastych, 2001: 54; Ochoa 2004a: 52, 55, 73, table 1, figs. 1, 2, 21. Bothriurus borellianus Mello-Leita˜o, 1934: Bu¨cherl, 1959b: 273 [misidentification]. Bothriurus chilensis (Molina, 1782): Aguilar and Meneses, 1970: 3 [misidentification]. Bothriurus (Andibothriurus)peruvianus MelloLeita˜o, 1948: Francke, 1974: 217, 218, figs. 1– 5, table 1 [misidentification]. T YPE M ATERIAL : PERU: Junı ´n Department: Yauli Province: Holotype - (MACN-Ar 6837), Abra de Anticona, Oroya [11u369310S 76u119480W], 4750 m, 28.xi.1974, A. Martı ´nez. Paratypes: same data, 1 - , 1 juv. (MACN-Ar 6838); Acaya, Rı ´oMantaro[11u499S75u369W], 3480 m, 1.iii.1957, W. Weyrauch, 1 U (IML 388). N EW R ECORDS : PERU: Junı ´n Department: Jauja Province: Acolla, near Jauja, [11u449170S 75u329350W, 3400 m], 8.viii.1953, F. Blancas, 4 U , 12 juv. (MUSM), ii.1965, F. Blancas, 2 U (MUSM). Junı ´n Province: Ondores [11u049060S76u099210W, 4300 m], 6.v.2000, E. Ponce, 2 - ,7 U (MHNC); Quebrada Antacocha, between Junı ´n and Carhuamayo [11u039S76u009W, 4090 m], viii.1952, F. Blancas, 1 juv. (MUSM). Tarma Province: Tarma, 2 km E [11u249580S75u399590W, 3400 m], 4.i.1973, N.F. Hadley and O.F. Francke, 1 U (AMNH). Yauli Province: Casaragra?, xi.1947, H. de Madedo, 1 U (MUSM); CerroQuinan, TupacAmaru,Yauli,La Oroya [11u329S75u549W, 3968 m], 26.viii.2005, W. Paredes, 1 U (MHNC), 1 juv. (AMNH [LP 6257B]). D IAGNOSIS : Orobothriurus parvus is most closely related to O. wawita (fig. 5). Both species share similar hemispermatophore morphology, including an elongated lamina apex and a short frontal crest (fig. 27B, C). The reticulate pigmentation along the metasomal DL carinae (figs. 12B, 13C) and the macrosetal counts on the ventral surface of the metasoma are also similar. The following characters separate the two species. The pedipalp chela manus of the male O. parvus possesses an apophysis on the internal surface, and the fixed finger is slightly curved, creating a small gap with the movable finger when the fingers are closed (fig. 26A), whereas the apophysis is absent and the fixed finger straight, such that no gap is evident when the fingers are closed, in the male of O. wawita (fig. 25B, C). The VL and VM carinae of metasomal segment V are complete in O. parvus (fig. 22E), but absent ( - ) or restricted to the distal third of the segment ( U )inO. wawita (figs. 21C, 22F). Metasomal segments II–V exhibit a distinct VM pigmentation stripe in O. parvus that is absent in O. wawita (fig. 13D). The two species also differ in the shape of the ventral margin of the hemispermatophore apex, which is inclined to the dorsal surface and curved distally in O. parvus (fig. 27B), but straight in O. wawita (fig. 27C). D ISTRIBUTION : Orobothriurus parvus is endemic to the central Andes, recorded from 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 69 3400–4750 m, in the Junı ´n and Lima departments, Peru (figs. 2E, 52). E COLOGY : The area inhabited by O. parvus corresponds to the Puna ecoregion or Puna biogeographical province (Brack, 1986; Ceballos Bendezu, 1976). The typical Puna vegetation of this area comprises shrub, steppe, and grasses (fig. 2E). No other scorpion species have been recorded in sympatry with O. parvus. R EMARKS : Orobothriurus parvus was misidentified in several papers as Bothriurus borellianus,Bothriurus chilensis or Bothriurus (Andibothriurus)peruvianus (Bu¨cherl, 1959b; Aguilar and Meneses, 1970; Francke, 1974). A record from Colcabamba (Lourenc¸o and Dastych, 2001: 54), based on a single female, requires confirmation. Orobothriurus quewerukana, n. sp. Figures 15C, 19C, 20D, 23F, 24D, 40D, E, 41–43, 53; table 3 Orobothriurus dumayi: Maury, 1976: 21, figs. 40– 44; Acosta and Ochoa, 2000: 136, 143. [misidentification]. T YPE M ATERIAL : CHILE: Regio´n I (Tarapaca´): Parinacota Province: Holotype - (AMNH), Putre, 6 km W, 18u13926.80S 69u32947.10W, 3732 m, 16.i.2005, A. Ojanguren Affilastro, C.I. Mattoni and J.A. Ochoa, UV and under stones. Paratypes: CHILE: Regio´n I (Tarapaca´): Iquique Province: Termas de Mamin˜a, 8.viii.1967, J.C. Ortiz, 4 juv. (MACN-Ar 6846); Mamin˜a, 20u04933.60S 69u12905.40W, 2822 m, 19.i.2005, A. Ojanguren Affilastro, C.I. Mattoni and J.A. Ochoa, UV and under stones, small wet valley with old terrace cultivation, 2 - ,2 U , 14 juv. (LBRE), 2 - ,2 U , 8 juv. (MACN-Ar), 2 - ,1 U , 1 subad. U (MHNC), 1 - ,1 U , 6 juv. (MZUC). Parinacota Province: same data as holotype,2 - ,2 U ,6juv.(MACN-Ar),2 - ,1 U , 3 juv. (LBRE), 1 - ,1 U , 3 juv. (MHNC), 1 subad. - (AMNH). PERU: Tacna Department: Tacna Province: Palca, 17u469450S 69u579040W, 3230 m, 26.ii.2000, J.A. Ochoa, 1 U (AMNH), 1 - ,2 U (MHNC). A DDITIONAL M ATERIAL : CHILE: Regio´n I (Tarapaca´): Iquique Province: Mamin˜a, 20u04933.60S69u12905.40W, 2822 m, 19.i.2005, A. Ojanguren Affilastro, C.I. Mattoni, and J.A. Ochoa, UV and under stones, small wet valleywitholdterracecultivation,1subad. - ,1 subad. U ,2juv. - ,2juv. U (AMNH[LP4307]), 5 juv. (AMNH), 5 juv. (MHNC). Parinacota Province: Putre, 6 km W, 18u13926.80S 69u32947.10W,3732m,16.i.2005,A.Ojanguren Affilastro, C.I. Mattoni and J.A. Ochoa, UV and under stones, 1 subad. - , 4 juv. - , 1 juv. U (AMNH [LP 4306]), 4 juv. (LBRE), 3 juv. (MHNC). E TYMOLOGY : The specific name is a noun in apposition composed of two Quechua words, quewe, meaning ‘‘curved,’’ and rukana, meaning ‘‘finger.’’ It refers the strong curvature of the movable finger of the pedipalp chela in the adult male. D IAGNOSIS : Orobothriurus quewerukana is closely related to O. curvidigitus and O. tamarugal (fig. 5), with which it shares the following characters: anterior margin of carapace with weak median projection (epistome; figs. 14C, 15C, D); pedipalp chela shape similar, with movable finger strongly curved in male (figs. 26C, 43C, 51A); pedipalp femur and patella more elongated in male than female; sternite VII with VSM carinae absent or obsolete (fig. 19A, C); metasomal segments I–IV with DL and ML carinae complete, granular, and VL carinae obsolete, smooth. Orobothriurus quewerukana may be separated from O. tamarugal by the shape of the lamina of the hemispermatophore (figs. 40E, 41D). The pedicel of the apex is narrower, and the frontal crest more developed and elongated in O. quewerukana; the ventral border of the distal portion of the apex is slightly curved in O. quewerukana, but straight in O. tamarugal; and one or two small folds, observed in the distal crest of 80%of the specimens of O. quewerukana, are absent in the distal crest of O. tamarugal. The hemispermatophore of O. quewerukana is similar to that of O. curvidigitus, except for the slightly shorter frontal crest and the absence of folds in the distal crest (figs. 28B, 40E). The two species may be distinguished by the shape and granulation of the male telson vesicle, which is slightly deeper, with a length/height ratio of 3.59–4.00 (mean 5 3.76), and granulation on the entire ventral surface (fig. 23F), in O. quewerukana, compared with O. curvidigitus, in which the length/height ratio is 3.72–4.23 (mean 5 3.96) and granulation is present only on the 70 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 anterior third of the ventral surface. The male pedipalp chela carinae are finely and densely granular in O. quewerukana and O. tamarugal (figs. 43D, E, 51B, C), but finely and sparsely granular to smooth in O. curvidigitus. These species may also be separated by the pigmentation pattern. Orobothriurus curvidigitus exhibits well-developed pigmentation on the carapace, tergites, metasoma, and pedipalps (figs. 11B, 12C, 13E), compared to O. quewerukana in which the pigmentation, especially that of the carapace and tergites, is faint or barely discernible (fig. 41) and O. tamarugal, in which it is absent (fig. 48). D ESCRIPTION : Based on holotype - and paratypes. Measurements of holotype - and paratype U recorded in table 3. Total length: - , 38.09–47.00 mm (n58, mean 541.59 mm); U , 37.20–51.50 mm (n5 8, mean 543.35). Color: General color yellowish with light brown spots on carapace, tergites, and pedipalps; metasomal segments, ventral pigmentation slightly darker, especially in specimens from Palca (fig. 41). Carapace faintly spotted, especially laterally and posteriorly; anterior margin with two spots submedially (fig. 41A, C); posterolateral surfaces faintly pigmented; median ocular tubercle and lateral ocelli dark brown to black; anterior third of anteromedian longitudinal sulcus pigmented along borders; posteromedian longitudinal sulcus faintly pigmented or unpigmented; posterolateral sulci unpigmented. Chelicerae unpigmented. Pedipalp coxa, trochanter and femur usually unpigmented or faintly pigmented along DI and DE carinae (some juveniles); patella external and, usually, internal surfaces faintly spotted; chela unpigmented. Legs, only ventral margin of femur, and dorsal and ventral margins of patella pigmented (unpigmented in some adults). Tergites I–VI each with two faint spots sublaterally, extending along anterior twothirds, becoming broader near anterior margins, delimiting broad, unpigmented median stripe (fig. 41A, C); VII unpigmented (adults) or with two faint spots sublaterally (juveniles). Sternum, genital opercula, pectines and sternites III–VII unpigmented (fig. 41B, D). Metasomal segments I–III, dorsal surfaces each with two faint subtriangular spots medially, separated by narrow unpigmented line (juveniles) or unpigmented (adults, fig. 41); lateral surfaces unpigmented, except near VL carinae; ventral surface with narrow VM stripe on III (absent on I and II), not contiguous with lateral pigmentation, and two VL stripes on I–III (occasionally faint or absent on I) becoming broader in posterior third; specimens from Mamin˜a usually less pigmented: VL stripe absent on segments I and II; VM stripe often absent on III. Segment IV, dorsal surface unpigmented; lateral and ventral surfaces as for segment III. Segment V, dorsal surface unpigmented or faintly pigmented posteriorly; lateral surfaces pigmented in distal third; ventral surface with three stripes, two VL and one VM, becoming slightly broader in posterior half but not joining with lateral pigmentation. Telson vesicle unpigmented or with lateral surfaces faintly pigmented; aculeus sclerotized, dark reddish brown. Fig. 49. Orobothriurus tamarugal, n. sp., holotype - (MACN-Ar), dextral pedipalp femur (A) and patella (B–D). A, B. Dorsal aspect. C. External aspect. D. Ventral aspect. Scale bar 51 mm. 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 71 Chelicerae: Movable finger with two subdistal teeth. Carapace: Surfaces finely granular, more coarsely so along anterior margin, anteromedian longitudinal and median ocular sulci ( - ) or smooth in anterior third, finely granular elsewhere ( U ). Anterior margin with weak median projection (epistome; fig. 14C). Anteromedian longitudinal sulcus complete, moderately developed; median ocular and posteromedian longitudinal sulci well developed; posterolateral sulci obsolete. Median ocular tubercle raised, situated anteromedially; median ocelli two ocular diameters apart. Pedipalps: Femur, DI, DE, and VI carinae complete (fig. 42A), DI carina more strongly developed than DE and VI carinae; VM carina vestigial, finely granular proximally; EM carina complete, granular ( - ) or obsolete, finely granular ( U , juveniles); dorsal surface finely granular; internal surface sparsely granular medially. Patella, DI, VI, and VE carinae complete, granular (fig. 42B–D); DE carina complete, less developed than DI carina ( - ) or obsolete ( U ); EM carina vestigial, reduced to few small granules medially ( - ) or absent ( U ); DPP carina comprising prominent granule and additional small granules proximally; VPP carina vestigial, comprising two or three small granules proximally; internal surface with prominent granule adjacent to trichobothrium inear DI carina. Chela manus slightly rounded, fingers relatively elongated (fig. 43); length/width ratio: - , 3.42–4.19, (n59, mean 53.71), U , 3.54–4.22 (n56, mean 53.87); length/height ratio: - , 2.95–3.32, (n59, mean 53.07), U , 2.87–3.34 (n56, mean 5 3.16); most carinae obsolete, finely granular (less so in U ) or absent; VM carina restricted Fig. 50. Orobothriurus tamarugal, n. sp., paratype U (MZUC), dextral pedipalp chela. A. Dorsal aspect. B. External aspect. C. Internal aspect. Scale bar 51 mm. 72 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 anteromedially; median ocelli two ocular diameters apart. Pedipalps: Femur, more elongated in - , length/width ratio: - , 4.05–4.27 (n52), U , 3.13; DI and VI carinae complete, granular; DE carina almost complete, coarsely granular in proximal three-quarters, smooth elsewhere; internal surface sparsely and coarsely granular (fig. 49A). Patella, more elongated in - , length/width ratio: - , 3.45–4.10 (n5 2), U , 2.95; DI and VI carinae almost complete, granular (fig. 49B–D); DE and VE carinae vestigial, finely and sparsely granular in - ; internal surface sparsely granular, with prominent granule adjacent to trichobothrium inear DI carina. Chela manus slightly rounded, fingers relatively elongated (figs. 49, 50); length/width ratio: - , 3.18–4.07 (n52), U , 3.83; length/height ratio: - , 2.90–3.62 (n5 2), U , 3.83; carinae obsolete, finely granular (less so in U ) or absent; VM carina restricted to proximal three-quarters of manus; DS, DMA, DI carinae complete, finely and densely granular ( - ) or smooth ( U ); D carina finely and sparsely granular; IM carina finely and sparsely granular in proximal half (fig. 51B); intercarinal surfaces finely granular ( - , fig. 51) or smooth ( U , fig. 50); internal surface with acuminate apophysis ( - ) or low bulge ( U ) near articulation of movable finger ( U ) (figs. 50A, 51B); movable finger ( - ), strongly curved, creating small gap with fixed finger when fingers closed (fig. 51A); fingers, dentate margins each with median denticle row and 5–6 pairs of internal and external accessory denticles. Trichobothria: Femur with 3 trichobothria, patella with 19, chela with 27 (figs. 49–51). Chela trichobothrium Et 3 situated in same axis as, or slightly basal to Est (figs. 50, 51). Tergites: Tergites I–VI, surfaces finely granular, more coarsely and densely so near distal margins of III–VI ( - ) or smooth ( U ). Tergite VII tetracarinate, paired DL carinae restricted to posterior two-thirds of segment, paired DSM carinae to posterior third; intercarinal surfaces granular. Legs: Femur and patella, prolateral surfaces finely granular, retrolateral surfaces smooth. Femur, ventral carinae weakly developed; other carinae absent. Patella acarinate. Telotarsi, proand retroventral rows of spiniform macrosetae with following counts on leg I, 1/1; II, 2/2; III and IV, 3/3. Pectines: Pectinal tooth count: - , 19–21 (n56; mode 520); U , 17–18 (n52). Sternites: Sternites III–VI, surfaces smooth; spiracles small, elliptical, and narrow. Sternite VII, surface smooth in anterior half, sparsely granular in posterior half; VL carinae obsolete, slightly more developed in holotype - (fig. 19F); VSM carinae absent. Metasoma: Segment I, DL carinae complete, moderately to coarsely granular; ML carinae well developed, becoming more coarsely granular in posterior two-thirds; LIM carinae moderately to coarsely granular, restricted to posterior two-thirds, oriented obliquely to antero-dorsal surface, almost joining ML carinae; one pair of macrosetae situated medially between ML and LIM carinae; VL carinae complete, obsolete, finely granular; VSM carinae weakly developed, sparsely granular; two pairs of VSM and VL macrosetae (fig. 19F). Segments II–IV, DL carinae complete, coarsely granular; one pair of DL macrosetae; ML carinae complete, coarsely granular; one pair of ML macrosetae; surfaces between ML and LIM carinae granular on segments II and III; LIM carinae reduced to few granules in posterior third of segment II, less developed on III, absent on IV; VL carinae vestigial, becoming absent toward segment IV; VSM absent or reduced to few scattered granules on segment II ( U ); three pairs of VL and VSM macrosetae. Segment V, length/width ratio, - , 1.95–2.24, U ,1.86;DL carinae reduced to few granules in anterior half; one pair of DL macrosetae; ML carinae obsolete, granular; two pairs of ML macrosetae; lateral intercarinal surfaces finely and sparsely granular; VL and VM carinae complete, granular; VL and VSM carinae fused, only separated medially (fig. 20E); ventral intercarinal surfaces granular; three pairs of VL and VSM macrosetae; two pairs of macrosetae along posterior margin. Telson: Length/height ratio: - , 2.53–3.06 (n52); U , 2.3 (n51). Vesicle elongated ( - , fig. 22H) or oval ( U , fig. 24F); dorsal surface smooth, slightly concave medially, gland not apparent ( - ); ventral surface coarsely granular. Aculeus short and curved, more so in U . 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 79 Hemispermatophore: Apex very well developed; distal crest almost straight, well developed, restricted to distal half of apex. Frontal crest weakly developed, less than half length of lamina; basal part oblique; distal part short, parallel to ventral margin of lamina, lateral projections slightly undulated with minute granulation. Basal lobe, terminal process extending almost to median part of frontal crest (fig. 47C, D). D ISTRIBUTION : All known records of this species are located in the Pampa del Tamarugal of Iquique Province, Region I (Tarapaca´), Chile (figs. 3A, 53). E COLOGY : The Pampa del Tamarugal is a desert area characterized by the near absence of rainfall (0.2–1 mm per year), high-salinity soils (mostly of alluvial origin from the Andes), depressions with salt lakes, and a forest dominated by Prosopis tamarugo and Prosopis chilensis (Mol.) Stuntz trees (fig. 3A). Prosopis tamarugo is a deciduous open-crowned tree up to 18 m tall, with a trunk up to 80 cm in diameter, a dense mat of lateral roots and a deep taproot (up to 6 m deep on trees 15 m tall; Habit et al., 1981; Serra, 1997). Temperatures in the Pampa del Tamarugal vary from 212 uC during winter nights to 36 uCon summer days, with a daily range of more than 35 uCinsummer.Thesaltlakesaredry,surface water is absent year-round, and the groundwater may be more than 60 m below the surface. The only humidity is provided by sporadic fog. Prosopis tamarugo forests occur only where groundwater is between 2–40 m below the surface (Serra, 1997). Specimens of O. tamarugal were collected at night, with UV light detection, inside the forest. The two Fig. 52. Orobothriurus Maury, 1976, locality records in central Peru. Orobothriurus ampay Ochoa and Acosta, 2003, cross; Orobothriurus huascaran, n. sp., circles; Orobothriurus parvus Maury, 1976, squares; Orobothriurus wawita Acosta and Ochoa, 2000, triangles. 80 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 males collected were active on the surface, sitting below large trees. The females and juveniles were inactive, resting under large plates of hard soil and salt (one of the forests is located near a dry salt lake). Brachistosternus donosoi Cekalovic, 1974, a larger and more active bothriurid species, was collected in sympatry. Orobothriurus wawita Acosta and Ochoa, 2000 Figures 13C, D, 16C, D, 19E, 21C, 22F, 25B, C, 27C, 52 Orobothriurus wawita Acosta and Ochoa, 2000: 137–143, figs. 1–13; 2001: 205; 2002: 18; Ochoa, 2004a: 43, 52, 55, 73, figs. 1, 2, 21, table 1; 2005: 55, 56, figs. 7, 9, table 2; Rein, 2007: 5. T YPE M ATERIAL : PERU: Cusco Department: Urubamba Province: Holotype - (MACN-Ar 9652), Pacchac, Pumahuanca, 13u139S72u069W, 3800 m, 17.vii.1998, J.A. Ochoa. Paratypes: Cusco Department: Urubamba Province: same data as holotype, 1 - , 1 U (CDA 019); same locality as holotype, 23.vii.1998, J. Flores and J.A. Ochoa, 2 - ,2 U (MHNC), 1 U (MACN-Ar 9653); Maras [13u199530S72u099220W, 3360 m], 29.x.1997, O. Mujica and J.A. Ochoa, 1 - (MHNC). Ayacucho Department: Cangallo Province: Comu´n Pampa [13u379S74u089W, 2700 m], 10.ii.1963, R. Garcia, 2 U (MUSM). N EW R ECORDS : PERU: Cusco Department: Quispicanchis Province: Lucre, Huacarpay [13u369300S71u449030W], ca. 3000 m, 14.i.2004, J.A. Ochoa, 2 juv. (AMNH [LP 3059]). Urubamba Province: Ollantaytambo [13u159180S72u159460W, 2861 m], 14.iii.1947, J.C. Palhsler, 1 U (MACN-Ar), 13.ii.1983, S. and A. Roig, 1 U (MACN-Ar); Ollantaytambo, Rı ´o Kusichaca (Inka trail) [13u149450S 72u259460W], 2750 m, 15.i.1983, S. and A. Roig, 1 U (MACN-Ar). D IAGNOSIS : Orobothriurus wawita may be distinguished from other species of the genus by the following characters. This species lacks an apophysis on the internal surface of the pedipalp chela manus of the male Fig. 53. Orobothriurus Maury, 1976, locality records in southern Peru and northern Chile. Orobothriurus atiquipa Ochoa and Acosta, 2002, square; Orobothriurus curvidigitus (Kraepelin, 1911), circles; Orobothriurus paessleri (Kraepelin, 1911), crosses; Orobothriurus quewerukana, n. sp., triangles; Orobothriurus tamarugal, n. sp., diamonds. 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 81 (fig. 25C), which is present in all other species except O. ampay, in which it is reduced to vestigial granules. The pedipalp chela manus of O. wawita is slender (fig. 23B, C) with a greater length/width ratio, 5.6–6.5 ( - ), 4.46– 5.45 ( U ), than all other species, 2.98–4.6 ( - ), 3.2–4.22 ( U ). The pigmentation on the ventral surfaces of the metasoma of O. wawita comprises irregular scattered spots, not forming a VM stripe (fig. 13D) that is present and well defined, at least on metasomal segments II–IV, in other species (figs. 11, 13A). Orobothriurus wawita is most closely related to O. parvus (fig. 5). Both species share similar reticulate pigmentation along the metasomal DL carinae (figs. 12B, 13C), macrosetal counts on the ventral surfaces of the metasoma, and hemispermatophore morphology, including an elongated lamina apex and a short frontal crest (figs. 27B, C). These two species may be distinguished by the following characters. The fixed finger of the pedipalp chela of the male is straight, such that no gap is evident when the fingers are closed, in O. wawita (fig. 25B, C), whereas it is slightly curved, creating a small gap when the fingers are closed, in O. parvus (fig. 26A). The VL and VM carinae of metasomal Fig. 54. Orobothriurus Maury, 1976, locality records in Argentina and central-northern Chile. Orobothriurus alticola (Pocock, 1899), squares; Orobothriurus calchaqui, n. sp., circle; Orobothriurus compagnuccii, n. sp., star; Orobothriurus famatina Acosta, 2001, diamonds; Orobothriurus grismadoi Ojanguren Affilastro et al., 2009, triangle; Orobothriurus ramirezi, n. sp., crosses. 82 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359 segment V are absent ( - ) or restricted to the distal third of the segment ( U )inO. wawita (figs. 21C, 22F), but complete in O. parvus (fig. 22E). The ventral margin of the apex of the hemispermatophore is straight in O. wawita (fig. 27C), but curved distally to the dorsal surface in O. parvus (fig. 27B). D ISTRIBUTION : All except one record of this species are situated in inter-Andean valleys at 2700–3800 m in the Ayacucho and Cusco departments of southern Peru (figs. 2B, C, 52). A record from Potosi (19u349S65u289W), 900 km from Cusco in Bolivia (fig. 1), is based on a single female, which, although morphologically similar to Peruvian material, possesses slight differences in pigmentation pattern and granulation of the metasomal segments (Acosta and Ochoa, 2002). This record is probably a mislabeling. E COLOGY : This species is endemic to the Queswa biogeographical region (Marı ´nMoreno, 1961; Ceballos Bendezu´, 1976; Ochoa, 2005; fig. 2B, C) and syntopic with two other bothriurids, Brachistosternus andinus and Pachakutej oscari Ochoa, 2004. ACKNOWLEDGMENTS We are grateful to Iva´n Benoit (Corporacio´n Nacional Forestal del Gobierno de Chile, CONAF) for assistance with obtaining permits to collect scorpions in Chilean National Parks, to the CONAF staff at Pampa del Tamarugal National Park, and to the Direccio´n General Forestal y de Fauna Silvestre, Ministerio de Agricultura de Peru (ex-NRENA) for permits 061-2004-INRENA-IFFS-DCB, 002-2008INRENA-IFFS-DCB, to collect scorpions in Peru. We thank John Achicahuala, Juan Carlos Chaparro, Luis Compagnucci, Roberto Gutie´rrez, Paula Korob, Juan Jose´ Martinez, Daniel Mun˜iz, Williams Paredes, Luis Piacentini, Jaime Pizarro, Elias Ponce, Aaro´nQuiroz, J.L. Velasquez, Matı ´as Vivanco, Ulrich Zanabria, and Horacio Zeballos for assisting us in the field; the following curators and collections managers for loaning and/or granting access to material from the collections in their care: Petra Sierwald and Alfred Newton (FMNH), Frantisˇek Kovar ˇı ´k(FKPC), Olintho Aguilar (MHNC), Gerardo Lamas and Diana Silva (MUSM), Alberto Chiarle (MRSN), and Hieronymus Dastych (ZMH); Luis Acosta for access to material on loan from MHNG; Wilfredo Mendoza (MUSM) for providing data on Peruvian vegetation; Ricardo Pinto da Rocha, Joyce Vitorino Villegas, Oscar Mujica, Horacio Larrain Barros, and Hugo Zamora for providing some of the photos of habitats and live habitus; Steve Thurston for assistance with preparing the plates; and two anonymous reviewers for comments on the manuscript. This research was supported by a Kalbfleisch Postdoctoral Research Fellowship from the AMNH, and a fellowship from the Fundac¸a˜o de Amparo a` Pesquisa do Estado de Sa˜o Paulo, Brazil (FAPESP 2010/00018-9) to J.A.O., grants from the Consejo Nacional de Investigaciones Cientı ´ficas y Te´cnicas, Argentina (CONICET) to A.A.O.A. and C.I.M., and a Genomics Postdoctoral Research Fellowship from the AMNH to C.I.M. Fieldwork in Chile was financially supported by the following sources: J.A.O., C.I.M. and L.P. (2003) from U.S. National Science Foundation grant EAR 0228699 to L.P.;J.A.O,C.I.M.andA.A.O.A.(2005)from the AMNH; A.A.O.A. (2006) from CONICET grant PIP 6502. Fieldwork in Peru (2008) was funded in part by a Kalbfleisch Postdoctoral Research Fellowship from the AMNH to J.A.O. Fieldwork in Argentina (2006, 2007) was financially supported by a CONICET grant PIP 6502 to A.A.O.A. REFERENCES Abalos, J.W. 1959. Scorpionida. Primeras Jornadas Entomoepidemiolo´gicas Argentinas 2: 591–593. 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Jahrbu¨cher des Nassauischen Vereins fu¨r Naturkunde 69: 79–97. 2011 OCHOA ET AL.: REVISION OF OROBOTHRIURUS MAURY, 1976 87 APPENDIX 1 C HECKLIST OF TAXA ASSIGNED TO O ROBOTHRIURUS M AURY , 1976, AT VARIOUS TIMES Bothriurus dumayi Cekalovic, 1974: transferred to Orobothriurus by Maury (1976) and returned to Bothriurus Peters, 1861, by Acosta and Ochoa (2001). Bothriurus lampei Werner, 1916: synonymized with O. curvidigitus by Maury (1973). Cercophonius brachycentrus var. bbivittatus Thorell, 1877: transferred to Orobothriurus by Acosta (2002), elevated to species (Orobothriurus bivittatus) by Acosta (2005), and synonymized with O. alticola by Ojanguren Affilastro et al. (2009). Orobothriurus alticola (Pocock, 1899): originally described as Bothriurus alticola Pocock, 1899, and transferred to Orobothriurus by Maury (1976), becoming type species of the genus. Orobothriurus ampay Ochoa and Acosta, 2003. Orobothriurus atiquipa Ochoa and Acosta, 2002. Orobothriurus calchaqui,n.sp. Orobothriurus compagnuccii, n. sp. Orobothriurus curvidigitus (Kraepelin, 1911): originally described as Bothriurus curvidigitus Kraepelin, 1911, and transferred to Orobothriurus by Maury (1976). Orobothriurus famatina Acosta, 2001. Orobothriurus grismadoi Ojanguren Affilastro et al., 2009. Orobothriurus huascaran, n. sp. Orobothriurus lourencoi Ojanguren Afillastro, 2003: although currently placed in Orobothriurus, this species will be placed in another genus (Mattoni et al., in litt.). Orobothriurus paessleri (Kraepelin, 1911): originally described as Bothriurus paessleri Kraepelin, 1911, and transferred to Orobothriurus by Maury (1976). Orobothriurus parvus Maury, 1976: previously misidentified as Bothriurus borellianus Mello-Leita˜o, 1934, Bothriurus chilensis (Molina, 1782), and Bothriurus peruvianus Mello-Leita˜o, 1948, by Bu¨- cherl, 1959b; Aguilar and Meneses, 1970; and Francke, 1974, respectively. Orobothriurus quewerukana, n. sp.: confused with Bothriurus dumayi by Maury (1976). Orobothriurus ramirezi, n. sp. Orobothriurus tamarugal, n. sp. Orobothriurus wawita Acosta and Ochoa, 2000. Pachakutej crassimanus (Maury, 1976): originally described as Orobothriurus crassimanus Maury, 1976, transferred to Pachakutej by Ochoa (2004a). Pachakutej inca (Maury, 1976): originally described as Orobothriurus inca Maury, 1976, transferred to Pachakutej by Ochoa (2004a). Pachakutej iskay (Acosta and Ochoa, 2001): originally described as Orobothriurus iskay Acosta and Ochoa, 2001, and transferred to Pachakutej by Ochoa (2004a). Pachakutej peruvianus (Mello-Leita˜o, 1948): originally described as Bothriurus peruvianus Mello-Leita˜o, 1948, transferred to Orobothriurus by Maury (1976), and to Pachakutej by Ochoa (2004a). APPENDIX 2 L IST OF 65 CHARACTERS SCORED FOR 15 SPECIES OF O ROBOTHRIURUS M AURY , 1976, AND 15 OUTGROUP TAXA , FROM M ATTONI ET AL .( IN LITT .) Character states are scored 0–3, unknown (?) and inapplicable (-). Characters from previous analyses that correspond partially or entirely to those in the present matrix are as follows: O2004 5Ochoa (2004a); OA&R2009 5Ojanguren Affilastro and Ramı ´rez (2009); P2003 5Prendini (2003). Pigmentation pattern 0. Tergites I–IV, pigmentation: entirely pigmented (0); paired spots sublaterally, unpigmented area medially (1) [O2004: 0; OA&R2009: 1]. 1. Tergite VII, pigmentation: entirely pigmented (0); paired spots sublaterally, unpigmented area medially (1); inapplicable (-). 2. Metasomal segments II and III, dorsal surfaces, pigmentation: absent, unpigmented (0); subtriangular spot medially, it may be divided by unpigmented line medially (1). 3. Metasomal segments I–III, dorsal surfaces, pigmentation along DL carinae: absent, unpigmented (0); reticulate lines (1). 4. Metasomal segments IV and V, ventral surfaces, VM stripe: contiguous with VL stripe posteriorly (0); not contiguous with VL stripe posteriorly (1); absent (2). [OA&R2009: 5]. 5. Telson vesicle, ventral and lateral surface, coloration ( -): similar to U, pigmented (0); different from U, unpigmented, with glandular, light yellow coloration (1). Carapace 6. Anterior margin, shape: sublinear or with shallow median notch (0); with weak median projection (epistome) (1). [OA&R2009: 7]. 7. Anteromedian longitudinal sulcus, length (-): complete (0); vestigial (1). [P2003: 4; O2004: 1; OA&R2009: 8]. Chelicera 8. Movable finger, subdistal teeth, number: one (0); two (1). [P2003: 9; O2004: 2; OA&R2009: 6]. Pedipalps 9. Femur, length (-): greater than three times width (0); less than three times width (1). [O2004: 3; OA&R2009: 75]. 10. Femur, dorsal surface, shape: slightly convex, DE and DI carinae situated in different axes (0); flat, DE and DI carinae situated in same axis (1). [O2004: 4]. 11. Chela, shape: similar in both sexes or narrower (length/width ratio of pedipalp chela smaller) in - than U(0); more robust (length/width ratio of 88 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 359