UROPHONIUS TRANSANDINUS SP. NOV. (BOTHRIURIDAE), A SCORPI ON FROM CENTRAL CHILE. Luis Eduardo Acosta

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1 Stud Neotrop Fauna & Environm (1998) Vol. 33: /98/ $12.00 Swets & Zeitlinger UROPHONIUS TRANSANDINUS SP. NOV. (BOTHRIURIDAE), A SCORPI ON FROM CENTRAL CHILE Luis Eduardo Acosta Faculty of Exact, Physical and Natural Sciences, National University of Cordoba, Argentina ABSTRACT Urophonius transandinus, new species belonging in the brachycentrus species-group is described. It can be separated from other species in the group by the pigmentation pattern of meso soma and ventral side of metasoma, the number of pectinal teeth, and the arrangement of the ventral submedian keels of caudal segment 1. Additionally, hemispermatophores of the new species lack a posterodistal fold (present in the rest). Records of U. transandinus sp.nov. come from the Chilean provinces Valparaiso, San Antonio, San Felipe de Aconcagua, Santiago and Cordillera. The probable zoogeographical links of Central Chile and the "peripampasic track" (Argentina - Uruguay southern Brazil), as suggested by the geonemy of the brachycentrus group, are discussed. KEYWORDS: Urophonius transandinus, Scorpiones, Bothriuridae, Chile, Neotropics. RESUMEN Se describe Urophonius transandinus, nueva especie perteneciente al grupo brachycentrus. Esta forma se distingue de otras especies en el grupo por su patr6n de pigmentos en el mesosoma y la cara ventral del metasoma, el numero de dientes pectineos, y la disposici6n de las carenas ventrales submedianas en el segmento caudal 1. Adicionalmente, la nueva especie carece de repliegue distal posterior en el hemiespermatoforo (presente en las demas). Se ha estudiado material de Urophonius transandinus sp.nov. proveniente de las pro vinci as chilenas de Valparaiso, San Antonio, San Felipe de Aconcagua, Santiago y Cordillera. Se discuten las probables vinculaciones zoogeograficas entre Chile central y el "track peripampasico" (Argentina - Uruguay - sur de Brasil), segun 10 sugiere la geonemia del grupo brachycentrus. INTRODUCTION Although the presence of the genus Urophonius Pocock in Chile was known since the end of last century, references in the literature are scarce, and concerning the species identity, some are erroneous or controversial. Cekalovic's (1983) checklist in- Address correspondence to: Luis E. Acosta, Catedra de Diversidad Animal I, Facultad de Ciencias Exactas, Fisi~ cas y Naturales, Universidad Nacional de C6rdoba. Av. Velez Sarsfield 299, 5000 C6rdoba, Argentina. lacosta@tecomnet.com.ar eludes six Urophonius species from this country, but as stated below, three of them are to be deleted from the list. The three nominate entities correctly cited for the Chilean scorpiofauna are: U. granulatus Pocock, and the obscure U. tregualemuensis Cekalovic and U. tumbensis Cekalovic. The original descriptions of U. tregualemuensis and U. tumbensis are very ambiguous, and neither the characters given nor the illustrations allow to identify them with certainty. From the publication alone (Cekalovic, 1981) it is not possible to assign them to any of the three species-groups I recognized

2 158 LUISEDUARDOACOSTA in the genus (Acosta, 1988). Only through the examination of a part of Cekalovic' s material I was able to determine that U. tregualemuensis very likely belongs in the granulatus group (two paratypes from the type locality examined). Unfortunately, holotypes of both U. tumbensis and U. tregualemuensis can no longer be found where they should be stored in the MZUC (Maury, pers. comm.), so that the identity of the former species remains unsolved. In the same paper, Cekalovic (1981) reports specimens of "u. corderoi" from several localities of Central Chile. I was able to examine most of these materials and found they are conspecific with a male from Valparaiso, cited by Kraepelin (1894) as U. brachycentrus (as U. granulatus in 1899). This form represents indeed an unnamed species of the brachycentrus group (Acosta, 1988), and is here described as Urophonius transandinus sp.nov. Aside of this new entity, in the MZUC material of Central Chile I detected some specimens of an undescribed member of the granulatus species-group, as well as few vials containing a curious "intermediate" between the groups brachycentrus and granulatus. All these concerns suggest strongly that the genus is more diverse in Chile than thought before (Mello-LeiH:io's 1945 monograph mentions no Urophonius species from Chile!), but certainly more intensive collections are needed. MATERIALS Abbreviations AND METHODS and curators of studied collections: CDA: Catedra de Diversidad Animal I, Facultad de Ciencias Exactas, Fisicas y Naturales, Universidad Nacional de Cordoba, Argentina (L.E. Acosta). MACN: Museo Argentino de Ciencias Naturales, Buenos Aires (E.A. Maury). MNHN: Museum National d'histoire Naturelle, Paris (W. Louren90). MZT: Museo di Zoologia delia Universita di Torino, Italy (0. Elter). MZUC: Museo de Zoologia, Universidad de Concepcion, Chile (T. Cekalovic). ZMH: Zoologisches Museum, Hamburg (G. Rack, H. Dastych). Material examined CHILE. 4 00, 17 ~ ~, 4 juv.: "Chili" (Gay), 1 juv. ["u. granulatus" Kraepelin det.] (MNHN.RS 581); "Chile central" (Porter), 1 ~ (MACN). Provincia Valparaiso: Valparaiso, 15 May 1893 (1. Michelsen), o holotype (ZMH); road from Casablanca to Algarrobo, 15 Jul (1. Solervicens), 1 ~ allotype (MACN 9601), 1 0 paratype (MACN 9602), 1 ~ paratype (CD A); San Jeronimo, 26 Jun (J. Solervicens), 3 juv. (MACN); Zapallar, 3 Jun (J. Solervicens), 1 ~ (MACN). Provincia San Antonio: El Tabo, Sep (B. Jackos), 1 ~ (MACN). Provincia San Felipe de Aconcagua?: "Provincia Aconcagua" (Porter), 1 0 (MZT.Sc.486). Area Metropolitana (Santiago): Farellones, 13 Sep (L. Peiia), 1 ~ (MNHN.RS 6343), 1 ~ (MZUC 601); El Arrayan, 4 Aug (Ecologia Animal coll.), 1 0 paratype (MZUC 392); Cerro Manquehue, 30 Jun (C. Sotomayor), 1 ~ (MZUC 368). Provincia Cordillera: EI Canelo, 3 Sep (G. Lopez), 1 ~ paratype (MZUC 608), 6 ~ (MZUC 469, 609, 610, 630,631,632), 1 ~ (MNHN.RS 6340); Rio Clarillo, Jul (M. Lewin), 1 ~ (MACN). Abbreviations of descriptive.terms: Keels of caudal segments: DL: Dorsal lateral LSM: Lateral supramedian LM: Lateral median LIM: Lateral inframedian VL: Ventral lateral VSM: Ventral submedian VM: Ventral median Hemispermatophore: L: Lamina c.d.: distal crest of lamina r.d.p.: posterodistal fold l.i.: internal lobe of capsule l.b.: basal lobe of capsule l.e.: external lobe of capsule RESULTS Urophonius transandinus sp. novo (Figs. 1-7,9) Urophonius brachycentrus: Kraepelin 1894 (part.):221 [example from Valparafso, ZMH, misidentified]; Cekalovic 1983 (part.):51 [reference to Kraepelin 1894]. Urophonius granulatus: Kraepelin 1899 (part.):194 [presumably the same example from Valparafso, with corrected determination label; misidentified]; Cekalovic 1983 (part.):52 [reference to Kraepelin 1899].

3 NEW CHILEAN UROPHONIUS (SCORPIONES) 159 Figs Urophonius transandinus sp.nov.: Figs. 1-4: d' holotype (ZMH). Fig. 1. right pedipalp, tibia and chela, ventromedial view (a: pro lateral apophysis); Fig. 2. caudal segment V and telson, lateral view. Figs left hemispermatophore (L: lamina, Ii: internal lobe, lb: basal lobe, Ie: external lobe; basal part was damaged during dissection), Fig. 3. internal view, Fig. 4. external view. Figs S? allotype (MACN), Fig. 5. right pedipalp, tibia and chela, ventromedial view (g: granule in the position of the apophysis of male); Fig. 6. caudal segment V and telson, lateral view; Fig. 7. Sternite V and caudal segments I-III, ventral view (VSM: Ventral submedian keels).

4 160 LUIS EDUARDO ACOSTA Urophonius corderoi: Cekalovic 1981 (part.):200; 1982:189; 1983:52 [misidentified]. Urophonius sp. [of the brachycentrus group]: Acosta 1988:25 (Fig. 4), 28. Type material Holotype 0' (ZMH): Valparaiso, 15 May 1893 (1. Michelsen), 1 0' ["U. brttdty. granulatus" (sic)]; allotype ~ (MACN 9601), 1 0' paratype (MACN 9602), 1 ~ paratype (CDA): road from Casablanca to Algarrobo, 15 July 1966 (J. Solervicens); 1 0' paratype (MZUC 392): EI Arrayan (Santiago), 4 August 1963 (Ecologia Animal); 1 ~ paratype (MZUC 608): EI Canelo (Santiago), 3 September 1967 (G. Lopez). Type locality Valparaiso, 33 03'S 7lo38'W (Province Valparaiso, V Region, Chile). Distribution Central Chile (Fig. 12): Provinces Valparaiso, San Antonio, San Felipe de Aconcagua, Area Metropolitana (Santiago) and Cordillera. Description Total length in adults: mm in males, up to 36.4 mm in females. Measurements of holotype and allotype: Table 1. brachycent rus General colour yellowish-hazel, pedipalp, chela and telson more orange or reddish; ventral surface, legs and pectines lighter; irregular pigmentation on prosoma, tergites, metasoma and appendages. Pigment of mesosoma: generalized pattern (tergite V) in Fig. 9; tergites I to III (or IV) show a median area without pigment, forming a narrow clear band. Pigment of metasoma: dark, reticulate on lateral surface of segments I-V; the ventral surface shows quite faint and irregular pigmentation, consisting primarily of an axial line and paramedian reticulate spots; axial line restricted to the proximal third of each segment, being more conspicuous in segments I-III; the paramedian parts are absent in segment I, they begin to be evident in segment III, and may extend as an open reticulation on the whole length of segment IV. Segment V: the paramedian pigment covers irregularly most of the ventral surface, the axial line is in most cases lacking. This arrangement varies among individuals, in many specimens the pigment becomes so weak that one or more of the described pigmented parts (either~axial, paramedian or both) -disappear, or leave just faint traces of the original condition. Tegument of prosoma and tergites I-VI finely granular; tergite VII with short keels in the distal half. Caudal segments I-IV: DL and LSM keels complete, of small and blunt granules. LIM keels sinuous and formed by tiny granules, only-present on the dist ransand inus 10 jheringii 11 achalensis Figs Pigmentation patterns of mesosomal tergites in the brachycentrus species-group (generalized from individuals of different sizes): Fig. 8. U. brachycentrus (Thorell). Fig 9. U. transandinus sp.nov.; Fig. 10. U. jheringii Pocock; Fig. 11. U. achalensis Abalos & Romina!.

5 NEW CHILEAN UROPHONIUS (SCORPIONES) 161 Fig. 12. Studied localities of Urophonius transandinus sp.nov. in five provinces of Central Chile. San Felipe de Ac. = San Felipe de Aconcagua; Santiago = Area Metropolitana. Right: location of the represented area. tal half of segments I-II, the distal third of segment III, and almost disappearing in segment IV. VL keels granulous and complete on segment I, with few granules on segment II, and smooth on segments III-IV. VSM keels: On segment I keels are formed by pearllike granules (more conspicuous in the female), arranged longitudinally in the posterior half and diverging obliquely in the anterior half, sometimes forming a transverse V-shaped keel (see variability below); following the keels, there are 4 pairs of ventral major setae (1 pair on the caudal border, 2 pairs along the longitudinal part of keels, the remaining 0 pair of setae near the end of the oblique branches); on segment II the keels are less defined and irregular, slightly diverging on the proximal end (4-6 pairs of setae following the keels); VSM keels of segments Table 1. Measurements (mm) of the 0 holotype (ZMH) and '2 allotype (MACN) of Urophonius transandinus sp.nov. Ho1otype <3 Allotype '2 Total length Prosoma length anterior / posterior width Mesosoma length Metasoma length Caudal segment I length / width Caudal segment II length / width Caudal segment III length / width Caudal segment IV length / width Caudal segment V length / width / height Telson length / width / height Aculeus length Pedipalp length Femur length / width Tibia length / width Chela length / width / height Movable finger length / / / / / /2.51/ /2.43 / / / /2.84/ / / / / / /2.44/ /2.29/ / / /2.01 /

6 162 LUIS EDUARDO ACOSTA III-IV represented by slight tegumentary ridges and scarce, irregularly arranged granules. Segment V: DL keel only represented by a short row of proximal granules. LM keel feeble, only present on the proximal 2/3 of the segment. VL keel complete and well defined, the proximal third smooth, the rest granulous. VM keel complete, Y -shaped (bifurcated posteriorly), only on the distal third granulous. Sternite V with four granulous keels, restricted to the posterior half. Chelicera with two subdistal teeth. Pedipalps: Chela of male much dilated, with a lobular prolateral apophysis delimiting a slight depression; on the base of the fixed finger a group of tiny granules (Fig. 1). Female chela is less inflated, bearing a blunt granule in the position of the apophysis of male (Fig. 5). Trichobothriotaxy according to the genus pattern. Number of pectinal teeth: , (holotype: 16-17, allotype: 14-13; see variability below). Tarsal spine formula: holotype TIll: 6-7 / 6-7, TIV: 7-7 /7-8; allotype TIll: 6-5 / 5-5, TIV: 6-6/ 7-5 (variability below). Hemispermatophore: L slender, slightly S-curved; c.d. well developed; r.d.p. absent; l.i. with a pair of tooth-like apophyses on the external side; in internal view the capsule is well developed, the l.b. ends in a slightly concave, spatulate structure; there is a large concavity between l.b. and l.e. Variability VSM keel of caudal segment I (n=25). In 13 specimens (including holotype and allotype) keels are separate, with a longitudinal caudal part, and diverging on the proximal third (Fig. 7). The oblique part is not connected to the longitudinal part in 12 specimens, forming a V or U shaped transversal keel with either separate (9) or joint (3) halves; the longitudinal part is less defined (sparse granules) in 6 specimens. Number of pectinal teeth. Frequencies in 00 (n=8 pectines): 15 teeth (3 pectines), 16 (4), 17 (1); 2 2 (n=34): 13 (15), 14 (14), 15 (4), 16 (1). Tarsal spine formula (in a pair, the first number refers to the retrolateral spines; Maury, 1977). Frequencies in T.III (n=46): 5-5 spines (4 tarsi), 5-6 (17), 6-5 (6), 7-5 (1), 6-6 (11), 6-7 (1), 7-6 (6). T.IV (n=45): 5-6 (1), 5-7 (1), 7-5 (2), 6-5 (1), 6-6 (8),6-7 (1), 7-6 (24), 7-7 (5), 7-8 (2). Comparisons The structure of the capsular lobes of the hemispermatophore, and the arrangement of ventral keels and setae on caudal segment I indicate clearly the inclusion of U. transandinus sp.nov. in the brachycentrus species-group (Acosta, 1988). However, VSM keel of the remaining species in the group is nearly always completely transversal (especially in U. jheringii Pocock), with few traces of the longitudinal part as seen in the new species. This feature of U. transandinus sp.nov. may reveal some kind of intermediary between a "full transverse keel" state (synapomorphic to species in the brachycentrus group) and the "longitudinal keels" condition (presumably plesiomorphic), present in the rest of the genus. The pedipalp chela of U. transandinus sp.nov. is remarkably more dimorphic -much dilated in males- than that of other Urophonius species. Pigmentation patterns of mesosomal tergites provide a good diagnostic character, as shown in Figs. 8-11: U. jheringii and U. transandinus sp.nov. are the forms with more extended pigment. A further difference concerns the ventral pigmentation pattern of caudal segments I IV: while paramedian, irregular spots are characteristic to other species in the brachycentrus group (Acosta, 1988), only U. transandinus sp.nov. shows some remains of an axial line. Finally, pectinal teeth counts serve to a separation too: U. jheringii 0 0 with 14-15, ; U. achalensis Abalos & Hominal , ; U~.~brachycentrus (Thorell) 00 with 17-20, (Maury, 1977, Acosta, 1988). The new species should be intercalated in the key provided by Acosta (1988) in couple 6, as follows: 6. VSMkeels of caudal segment I consisting of a longitudinal distal part and a oblique or transverse proximal part. VL and VM keels on segment V complete and well defined. Hemispermatophore without r.d.p U. transandinus sp.nov. 6. 'VSM keels of caudal segment I limited to the proximal parts, often coalescing in a single, transverse keel; longitudinal parts normally represented by scarce granulation. VL and VM keels limited to the distal 2/3 of segment V. Hemispermatophore with r.d.p.... remaining species of the brachycentrus group Known distribution of other Chilean Urophonius Urophonius granulatus Pocock 1898 ["exact locality doubtful, probably Chili, the specimens being contained in a tube with examples of Bothriurus

7 NEW CHILEANUROPHONIUS (SCORPIONES) 163 coriaceus, a species which has been recorded from Coquimbo, &c."] Southern Argentina (province of Santa Cruz) and Chile (province Magallanes) (Maury 1979). Urophonius tregualemuensis Cekalovic Chile: Provinces Curico and Maule. Urophonius tumbensis Cekalovic Chile: Province Concepcion. Exclusions of the Chilean checklist Urophonius brachycentrus. Mentioned from Chile in old citations, this species is known from central Argentina (Maury 1977). Pocock (1893) reports a DISCUSSION The presence of a member of the brachycentrus specimen from "Coquimbo", but according to Maury (1979), the same material was used by Pocock (1898) to describe U. granulatus. Kraepelin (1894) identified as U. brachycentrus the male here designated as holotype of U. transandinus sp.nov. Urophonius paynensis San Martin & Cekalovic 1968, recorded at the Payne Region (province Magallanes). Junior synonym of U. granulatus (cf. Maury 1979). Urophonius corderoi Mello-LeiUio: Cekalovic (1981). Junior synonym of U. jheringii (cf. Galiano & Maury 1979). Most materials examined by Cekalovic belong to the new species here described. group in central Chile poses some interesting phylogenetic and biogeographical questions. First, the fact that the group does exist in both sides of the Andes (at this latitude, an insurmountable geographic barrier for scorpions) might suggest that most synapomorphies of the group (concerning capsular lobes of the hemispermatophore, and keel/setae arrangement on caudal segment I; Acosta, 1988) arose so early as by the late Miocene or the early Pliocene, that is, before the Andes began to be an effective barrier. Ranges of the non Chilean species of the group are roughly related to old mountains, that were partially raised once again because of the Andean orogeny. These ancient remains seem to form a "peristepic" arc from the Sierras de Cordoba-San Luis (central Argentina) to southern Brazil, embracing also the low systems of southern Province of Buenos Aires (Argentina) and Uruguay. Other scorpions and a couple of harvestmen species show a similar pattern, what led to the recognition to the so called "peripampasic track" (Acosta, 1993). This track is deemed to connect the fragments of a once continuous area (Mattoni & Acosta, 1997). What concerns U. transandinus sp.nov., its relationships probably indicate very old transandean links between the peripampasic track and the biota of central Chile. Unlike most scorpions, all four species in the brachycentrus group exhibit an "invernal" surface activity pattern (Acosta, 1988, see also collection dates of U. transandinus sp.nov.); thus, evidence for affinity within the group is not restricted to morphological data. ACKNOWLEDGMENTS I am indebted to the curators of the studied collections (listed with acronyms of the Institutions) for the loan of Chilean material. The late Dr. Emilio A. Maury kindly looked for Cekalovic's types in the MZUC. REFERENCES Acosta LE (1988): Contribucion al conoclmlento taxonomico del genero Urophonius Pocock, 1893 (Scorpiones, Bothriuridae). I Arachnol16: Acosta LE (1993): Escorpiones y opiliones de la provincia de Cordoba (Argentina): diversidad y zoogeograffa. Bull Soc neuchatel Sci nat, (C.R. XU Ie ColI. europ. Arachnol.), 116: Cekalovic T (1981): Dos nuevas especies y un nuevo registro del genera Urophonius para Chile (Scorpiones, Bothriuridae). Bol Soc BioI Concepcion 52: Cekalovic K, T (1982): Estado actual de la Coleccion Arachnol6gica del Museo de Zoologfa de la Universidad de Concepcion (M.Z.U.C.), Parte Scorpiones. Mem Inst Butantdn 46: Cekalovic T (1983): CataIogo de los escorpiones de Chile. Bol Soc BioI Concepcion, 54: Galiano ME, Maury EA (1979): Lista de los ejemplares tfpicos de "Arachnida" (Araneae, Opiliones, Scorpiones y Solifugae) depositados en el Museo Argentino de Ciencias Naturales "Bernardino Rivadavia". Rev Mus Arg C Nat, Ent 5: Kraepelin K (1894): Revision der Scorpione. II. Scorpionidae und Bothriuridae. lahrb Hamb Wiss Anst 11: Kraepelin K (1899): Scorpiones und Pedipalpi. In: Bronn's, ed., Das Tierreich, 8: 1-265, Berlin. Mattoni CI, Acosta LE (1997): Scorpions of the insular Sierras in the Llanos District (Province of La Rioja, Argentina) and their zoogeographical links. Biogeographica 73:

8 164 LUIS EDUARDO ACOSTA Maury EA (1977): Comentarios sobre dos especies de escorpiones del genero Urophonius (Bothriuridae). Rev Mus Arg C Nat, Ent 5: Maury EA (1979): Escorpiofauna patagonica. II. Urophonius granulatus Pocock, 1898 (Bothriuridae). Physis Secc C 38: Mello-LeiHio C de (1945): Escorpi5es sul-americanos. Arq Mus Nac 40: Pocock RI (1893): A contribution to the study of Neotropical scorpions. Ann Mag Nat Hist, 6th ser, 12: Pocock RI (1898): Descriptions of some new scorpions from Central and South America. Ann Mag Nat Hist, 7th ser., 1: San Martin P, Cekalovic T (1968): Escorpiofauna chilena. I. Bothriuridae. Una nueva especie de Urophonius para Chile. Invest Zool Chilenas 13: Received: June 17, 1998 Accepted: November 17,1998