Taxonomy of the Genus Gabrius (Coleoptera: Staphylinidae) in Southern South America

ISSN 0373-5680 (impresa), ISSN 1851-7471 (en línea) Rev. Soc. Entomol. Argent. 70 (1-2): 01-15, 2011 1

Taxonomy of the genus Gabrius (Coleoptera: Staphylinidae) in southern South America

CHANI-POSSE, Mariana

Laboratorio de Entomología, Instituto Argentino de Investigaciones de las Zonas Áridas (IADIZA, CCT CONICET Mendoza), Casilla de Correo 507, 5500 Mendoza, Argentina; e-mail: [email protected]

Taxonomía del género Gabrius (Coleoptera: Staphylinidae) en el sur de Sudamérica

RESUMEN. Cinco especies de Gabrius Stephens, 1829 del sur de Sudamérica se redescriben e ilustran. Se conocen actualmente seis especies de Gabrius, del sur de Sudamérica: G. argentinus (Bernhauer, 1912), G. hornaditanus (Rambousek, 1925), G. jujuyensis (Bernhauer, 1921), G. nidicola (Bernhauer, 1921), G. tucumanensis (Bernhauer, 1927) y G. nigritulus (Gravenhorst, 1802). Se provee una clave para la identificación de estas especies, diagnosis e ilustraciones para G. nigritulus y un mapa de distribución.

PALABRAS CLAVE. Staphylininae. Philonthina. Gabrius. Philonthus. Taxonomía. Sudamérica.

ABSTRACT. Five species of Gabrius Stephens, 1829 from southern South America are redescribed and illustrated. Six species of Gabrius are now known from southern South America: G. argentinus (Bernhauer, 1912), G. hornaditanus (Rambousek, 1925), G. jujuyensis (Bernhauer, 1921), G. nidicola (Bernhauer, 1921), G. tucumanensis (Bernhauer, 1927) and G. nigritulus (Gravenhorst, 1802). An identification key to these species, diagnosis and illustrations for G. nigritulus and a distribution map are provided.

KEY WORDS. Staphylininae. Philonthina. Gabrius. Philonthus. Taxonomy. South America.

INTRODUCTION (Smetana, 1995; Schillhammer, 1997; Chani-Posse, 2010). The genus Gabrius is The genus Gabrius Stephens, 1829 distributed worldwide and accounts for belongs to the subtribe Philonthina in more than 300 species at present (Herman, the subfamily Staphylininae. According 2001). Most of its species are known to to Smetana (1995) Gabrius was often occur in the Nearctic, Palearctic and considered by many authors as a subgenus Oriental regions. This knowledge may of Philonthus, but nowadays is assumed change and the number of species may to be a separate genus which is closely increase as poorly known tropical faunas related to Philonthus and other genera. are studied (Schillhammer, 1997) and the These relationships have been discussed by narrower generic concepts at present applied Smetana (1995) and are still to be clarified to the Holarctic fauna (e.g. Smetana, 1995)

Recibido: 2-VIII-2010; aceptado: 12-X-2010 2 Rev. Soc. Entomol. Argent. 70 (1-2): 01-15, 2011 are applied to those. According to Herman Alsina). (2001), only three species of Gabrius are MLPA Museo de La Plata, La Plata, Argentina known to occur in America south of Mexico, (Norma B. Díaz). two of them cited from southern South NMPC National Museum (Natural History), America, G. nigritulus (Gravenhorst) and G. Prague, Czech Republic (Jiři Hajek) chiliensis Coiffait & Sáiz. The latter name NMW Naturhistorisches Museum Wien, was recently found to be a synonym of the Austria (Harald Schillhammer). former (Chani-Posse de Maus, 2009). As part MNNC Museo Nacional de Historia Natural, of a major revisionary study of the species Santiago, Chile (Mario Elgueta D.) of Philonthus Stephens, 1829 (Coleoptera: ZMHB Museum für Naturkunde der Staphylinidae) from southern South America Humboldt Universität, Berlin, (Chani-Posse de Maus, 2009; Chani-Posse, Germany (Manfred Uhlig, Bernd 2010), five more species of Gabrius are Jaeger) recognized for this area. The geographical area covered in this study, as in a previous work (Chani-Posse, 2010), is the southern Techniques for the preparation and portion of South America which comprises examination of male and female genitalia primarily the countries of Argentina, Chile, follow Smetana (1982). Descriptions were Paraguay and Uruguay. made using a Leica MZ6 dissecting scope, The objectives of this study are to and several genitalic features were examined redescribe five species of Gabrius (previously with a Leitz Wetzlar compound microscope. assigned to Philonthus) by incorporating Drawings were made with a camera lucida new characters from external morphology, attached to the compound microscope including genitalia, present a key to the or dissecting scope. Photographs were species for southern South America and taken using a digital camera attached to detail their geographic distribution. the dissecting scope. Measurements (given in millimeters) were made with an ocular MATERIAL AND METHODS micrometer. Overall body length was measured from the apex of the labrum to the The present study is based on the apex of the abdomen. Other measurements examination of about 150 adult specimens, were taken and abbreviated as follows: including type material. The material studied HW- maximum head capsule width was obtained on loan from, and are deposited HL- length of head capsule, from anterior in the following collections: margin of frontoclypeus to neck constriction AMNH American Museum of Natural (along midline) History, New York, USA (Lee H. PW- pronotum maximum width Herman). PL- pronotum length (along midline) CEUNC Colección Entomológica de la EL- eye length (seen from above) Universidad de Córdoba, Argentina TL- temple length (from the posterior (Miguel Angel Delfino). margin of the eye to the nuchal groove; seen FMNH Field Museum of Natural History, from above) Chicago, USA (Alfred F. Newton, NW- neck width Margaret K. Thayer). S1- first segment of hind tarsus length IADIZA Instituto Argentino de Investigaciones S5- last segment of hind tarsus length de las Zonas Aridas, Mendoza, EtL- elytra length at sides (from humerus Argentina (Sergio Roig Juñent). to apex; seen from above) JEBC Juan E. Barriga private collection, Etl- elytra length along suture Curicó, Chile (Juan E. Barriga). EtW- elytra width at base MACN Museo Argentino de Ciencias Terminology mainly follows Blackwelder Naturales Bernardino Rivadavia, (1936), Smetana (1995), Smetana & Davies Buenos Aires, Argentina (Arturo Roig (2000) and Schillhammer (2000) with slight Chani-Posse, M. Gabrius in southern South America 3 modifications and/ or inclusions already ridge present (Fig. 8); antennal segment 4 detailed by the author in previous studies elongate (Figs. 5, 10); maxillary palpus with (Chani-Posse, 2006). segment 2 longer than segment 3; posterior Biogeographical provinces considered in arms of gular sutures joined before neck the geographical distribution of the species (Fig. 2); submentum about 1.5 times as long follow those of Morrone (2006). All records as mentum; neck no more than 0.5 times and the general distribution given for each as wide as head at widest point; pronotum species are based strictly on specimens that slightly narrowed anteriad to parallel-sided I have examined. All locality data were (Figs. 3, 10); dorsal surface of pronotum collected from collection data. with two rows of punctures, each with five or six punctures; first segment of hind tarsus General remarks shorter than segments 2 and 3 combined, and about as long as last segment (Figs. 4, The most recent and complete description 11); abdominal terga 2 and 3 with posterior of the genus, at least for America north of basal line straight medially (Figs. 4, 11); Mexico, is available in Smetana (1995). Some male sternum 7 with apical margin straight; characters of Gabrius are also discussed in tergum 8 in both sexes with posterior margin Schillhammer (1997). truncate; styli of tergum 9 simple, moderately stout; sternum 8 with two long subapical Recognition of the currently known macrosetae at each side (Figs. 13, 19, 31); southern South American species paramere entire and reduced, completely fused to median lobe (Figs. 15, 21, 28, 33) The labial palpus with the last segment with exception of that of G. nigritulus (Fig. slender, as long as to slightly longer and 37); second gonocoxite with one strong distinctly narrower than the penultimate basal seta (Figs. 18, 25, 30, 35). segment allows recognition among the other southern South American Philonthina. The genus can also be recognized by the sternum RESULTS 9 of male genital segment with basal portion not reduced, more or less symmetrical with Key to southern South American species exception of that of G. brevipennis (Smetana, of Gabrius 1995), the East Palearctic and Oriental species of the fimetarioides group (Schillhammer, 1. Distance separating medial interocular 1997), P. argentinus, P. jujuyensis, and P. punctures on frons less than twice distance nidicola (Chani-Posse de Maus, 2009). separating medial punctures from lateral Besides these characters, Gabrius can punctures (Fig. 7) ……………………….…… be separated from Philonthus by having the ...... Gabrius jujuyensis (Bernhauer) first four segments of front tarsus in both 1’. Distance separating medial interocular sexes simple, not dilated, with ventral face punctures on frons twice or more as large as bearing only regular marginal setae, from distance separating medial punctures from Belonuchus by having the lateral puncture of lateral punctures (Figs. 1, 6, 12) …...... 2 pronotum bearing long macroseta separated from superior line of pronotal hypomeron 2. First antennal segment as long as segments by a distance about equal to diameter of 2 and 3 combined; antennal segment 8 puncture, and from Endeius by having the transverse ...... …...... last segment of maxillary palpus in general ...... Gabrius argentinus (Bernhauer) fusiform, not acutely narrowed apically (Fig. 2’. First antennal segment shorter than 12). segments 2 and 3 combined; antennal Other characters common to the southern segment 8 drate …...... ………………..……. 3 South American species of Gabrius known at present are the following: postmandibular 3. Head distinctly longer than wide (Fig. 4 Rev. Soc. Entomol. Argent. 70 (1-2): 01-15, 2011

Figs. 1-12. Gabrius argentinus: 1, head, dorsal view; 2, head, ventral view; 3, pronotum; 4, elytra and abdomen. G. hornaditanus: 5, antennal segments 1-10; 6, head, dorsal view. G. jujuyensis: 7, head, dorsal view; 8, head, lateral view; 9, elytra. G. tucumanensis: 10, head and pronotum, dorsal view; 11, elytra and abdomen. G. nigritulus: 12, head, dorsal view. Abbreviations: D= distance separating medial interocular punctures; d= distance separating medial punctures from lateral punctures; PmR= postmandibular ridge; GS= posterior arms of gular sutures; Dp= dorsal rows of punctures. Chani-Posse, M. Gabrius in southern South America 5 10); antennal segment 5 elongate (Fig. 10) 0.9); dorsal surface coarsely and moderately …...... Gabrius tucumanensis (Bernhauer) punctate, with three postocular punctures 3’. Head about as long as wide (Figs. 6, 12); and two to three epicranial punctures; antennal segment 5 quadrate (Fig. 5) ……… 4 distance separating medial interocular punctures on frons about three times as large 4. Antennal segment 3 longer than 2 (Fig. as distance separating medial punctures 5); middle tarsus shorter than middle tibia from lateral punctures (Fig. 1); infraorbital …...... Gabrius hornaditanus (Rambousek) ridge present, reaching postgenal ridge. Eyes 4’. Antennal segment 2 and 3 subequal distinctly shorter than temples (EL/TL= 0.75- in length (Fig. 12); middle tarsus as long 0.67) seen from above (Fig. 1). Antennae as to longer than middle tibia ...... … 5 with segment 1 as long as segments 2 and 3 combined, segment 2 shorter than segment 5. Antennal segment 10 transverse; front 3, segments 5-6 quadrate, segments 7-10 tarsus shorter than front tibia …...... …….. transverse. Maxillary palpus with segment 2 ...... Gabrius nigritulus (Gravenhorst) no more than twice as long as its maximum 5’. Antennal segment 10 quadrate; front width. Labial palpus with last segment twice tarsus as long as front tibia ……………...... as long as preceding segment. Neck no more ...... Gabrius nidicola (Bernhauer) than 0.5 times as wide as head at widest point (NW/HW= 0.4) (Fig. 1). Gabrius argentinus (Bernhauer, 1912) Pronotum about as long as wide (PW/ (Figs. 1-4, 13-18, 39; Figs. 1 a-b in Chani- PL= 1.07-0.94), dorsal surface of pronotum Posse de Maus, 2009) with two rows of punctures, each with five punctures (Fig. 3). Basisternum with medial Philonthus argentinus Bernhauer, 1912: 43; longitudinal carina slightly developed. Bernhauer & Schubert, 1914: 329 (cat.); Metaventrite with circum-mesocoxal ridge Bruch, 1915: 444 (cat.); Blackwelder, broadly rounded medially. Elytra at sides as 1944: 132 (list); Herman, 2001: 2751 long as to moderately longer than pronotum (cat.); Chani-Posse, 2004: 230 (list). at midline (EtL/PL=1.0-0.8), distinctly longer Gabrius argentinus; Chani-Posse de Maus, than elytra along suture (EtL/Etl= 1.7-1.5); 2009: 32, 34 (new combination) transverse distance between punctures distinctly larger than diameters of punctures Diagnosis. Gabrius argentinus may be (Fig. 4). Front and middle tarsus shorter than identified by its body shape and coloration, front and middle tibia; first segment of hind the antennal segment 1 as long as segments tarsus about as long as last segment (S1/S5= 2 and 3 combined and the apex of median 1.2) (Fig. 4). Abdominal terga moderately lobe turn downwards in its apical fourth (in punctuate basally, distance separating lateral view). It differs from G. tucumanensis punctures distinctly larger than diameters of by the antennal segment 5 quadrate and the punctures (Fig. 4). basal portion of sternum 9 asymmetrical. Male genitalia. Sternum 8 moderately emarginate medio-apically (Fig. 13). Tergum Redescription. Length of the body 10 subtruncate medio-apically (Fig. 17), 7.0-7.5 mm. Coloration. Head castaneous- with two long subapical macrosetae and piceous to black; thorax castaneous-piceous; 2-6 short apical setae. Sternum 9 with elytra and abdominal segments castaneous- basal portion asymmetrical, moderately piceous to black; antennae castaneous- emarginate apically with two to three long piceous to castaneous-brunneous; palpi and apical setae at each side (Fig. 14). Aedeagus legs castaneous-brunneous to brunneous. with median lobe gradually narrowed from Head moderately to slightly longer than apical third; apex of median lobe subacute wide (HW/HL= 0.9-0.8) (Fig. 1) and at basal (in dorsal view) and turn downwards in its third as wide as distal third in both sexes, apical fourth (in lateral view) (Figs. 15-16). about as wide as pronotum (HW/PW= 1.0- Female genitalia. Tergum 10 similar to that 6 Rev. Soc. Entomol. Argent. 70 (1-2): 01-15, 2011 of male. Second gonocoxite with one basal 2001: 2839 (cat.); Chani-Posse, 2004: strong seta and one long apical seta, without 231 (list). stylus (Fig. 18). Gabrius hornaditanus; Chani-Posse de Maus, 2009: 33, 34 (new combination). Geographical distribution. Gabrius argentinus is currently only known from Diagnosis. Gabrius hornaditanus may northwestern Argentina (Tucumán) (Fig. 39). be identified by the distance separating punctures of abdominal terga distinctly Type material. Lectotype, male, with smaller than the diameters of punctures and labels: “Rep. Argentina / PROV. TUCUMAN the deeply emarginate posterior margin of the / RICHTER / 19”, “387”, “argentinus Bernh. male sternum 8. It differs from G. jujuyensis / Typus” (yellow label), “Chicago NHMus. and G. tucumanensis by having the antennal / M. Bernhauer Collection”, “Lectotype segment 10 quadrate and the middle tarsus Philonthus argentinus Bernhauer, 1912 / shorter than middle tibia. Des. Chani-Posse de Maus 2008”. (FMNH). Paralectotypes: 2 females, “Rep. Argentina Redescription. Length of the body 5.0- / PROV. TUCUMAN / RICHTER / 19”, 5.2 mm. Coloration. Head black; thorax “395” (one specimen), “argentinus Bernh. / and abdominal segments castaneous; elytra Cotypus” (yellow label), “Chicago NHMus. testeceous-brunneus; antennae, palpi and / M. Bernhauer Collection”, “Paralectotype legs castaneous brunneous to brunneous. Philonthus argentinus Bernhauer, 1912 / Head about as long as wide (HW/HL= 1.1) Des. Chani-Posse de Maus 2008” (FMNH). and at basal third as wide as distal third in Note: Chani-Posse de Maus (2009: 32, both sexes, about as wide as to slightly wider 34) designated lectotype. than pronotum (HW/PW= 1.0-1.1); dorsal surface coarsely and moderately punctate, Additional material examined: 14 with three postocular punctures and four specimens (1 male, 4 females, 9 unsexed). epicranial punctures; distance separating ARGENTINA: Tucumán: Richter (1 CEUNC), medial interocular punctures on frons about 389 (1 MLPA), Philonthus argentinus Brh. three times as large as distance separating (1 MLPA), in green “cotypus”, Bernhauer medial punctures from lateral punctures determ. (2 females, 2 MLPA); Yerba Buena, and at basal third slightly narrower than at M. A. Jaynes, Philonthus argentinus Brh., distal third (Fig. 6); infraorbital ridge present, (1 MACN); 1938, G. M. Borg, Philonthus reaching postgenal ridge. Eyes distinctly boxi Bernh., (1 MLPA); 5 Oct 1927, in green shorter than temples (EL/TL= 0.5-0.4) seen “Typus”, 2838, in Bernahuer handwriting from above. Antennae with segment 1 Philonthus boxi Bernh. n. sp.[upside down], shorter than segments 2 and 3 combined, (1 MLPA), 2836, (1 MLPA); Villa Nougués, segment 2 shorter than segment 3, segments Dec. 11, 1971, L. Herman, 1250, 759 (1 5-10 quadrate (Fig. 5). Maxillary palpus male, 1 female AMNH); 30km, SE Tafi del with segment 2 no more than twice as long Valle, nr. El Indio, XI/20/1995, L. Herman, as its maximum width. Labial palpus with 950-1050m, 2891 (1 female AMNH). last segment 1.5 times as long as preceding segment. Neck no more than 0.5 times as Gabrius hornaditanus (Rambousek, 1925) wide as head at widest point (NW/HW= (Figs. 5, 6, 19-22, 39) 0.5). Pronotum longer than wide (PW/PL= Philonthus hornaditanus “Rambousek in 0.86-0.79), dorsal surface of pronotum litt.”; Bruch, 1915: 445 (cat.). Nomen with two rows of punctures, each with nudum. five punctures. Basisternum with medial Philonthus hornaditanus Rambousek, 1925: longitudinal carina moderately developed. 75; Scheerpeltz, 1933: 1345 (cat.); Metaventrite with circum-mesocoxal Blackwelder, 1944: 133 (list); Herman, ridge broadly rounded medially. Elytra at Chani-Posse, M. Gabrius in southern South America 7

Figs. 13-38. Gabrius argentinus: 13, sternum 8, male; 14, sternum 9; 15, aedoeagus, dorsal view; 16, aedoeagus, lateral view; 17, tergum 10, male; 18, gonocoxites of female genital segment. G. hornaditanus: 19, sternum 8, male; 20, sternum 9; 21, aedoeagus, dorsal view; 22, aedoeagus, lateral view. G. jujuyensis: 23, sternum 9; 24, aedoeagus, dorsal view; 25, gonocoxites of female genital segment. G. nidicola: 26, sternum 9; 27, tergum 10, male; 28, aedoeagus, dorsal view; 29, tergum 10, female; 30, gonocoxites of female genital segment. G. tucumanensis: 31, sternum 8; 32, tergum 10, male; 33, aedoeagus, dorsal view; 34, aedoeagus, lateral view; 35, gonocoxites of female genital segment. G. nigritulus: 36, sternum 9; 37, aedoeagus, dorsal view; 38, paramere, ventral side. 8 Rev. Soc. Entomol. Argent. 70 (1-2): 01-15, 2011 sides as long as to moderately longer than Note: Chani-Posse de Maus (2009: 32, pronotum at midline (EtL/PL=0.7), distinctly 33) designated lectotype. longer than elytra along suture (EtL/Etl= 1.9-1.7); transverse distance between Gabrius jujuyensis (Bernhauer, 1921) punctures distinctly smaller than diameters (Figs. 7-9, 23-25, 39; Figs. 2 a-b in Chani- of punctures. Front tarsus shorter than front Posse de Maus, 2009) tibia; middle tarsus shorter than middle tibia; first segment of hind tarsus as long as Philonthus jujuyensis “Bernhauer in litt.“; last segment (S1/S5= 1.0). Abdominal terga Bruch, 1915: 445 (cat.). Nomen nudum. moderately punctuate basally, distance Philonthus jujuyensis Bernhauer, 1921: separating punctures distinctly smaller than 107; Scheerpeltz, 1933: 1347 (cat.); diameters of punctures. Blackwelder, 1944: 133 (list); Herman, Male genitalia. Sternum 8 deeply 2001: 2850 (cat.); Chani-Posse, 2004: emarginate medio-apically (Fig. 19). Tergum 231 (list). 10 subtruncate medio-apically, with two Gabrius jujuyensis; Chani-Posse de Maus, long subapical macrosetae and 3-4 short 2009: 33, 34 (new combination) apical setae. Sternum 9 with basal portion more or less symmetrical, moderately Diagnosis. Gabrius jujuyensis may be emarginate apically with two long apical identified by the antennal segments 8-10 setae at each side and two subapical setae transverse, and the distance separating (Fig. 20). Aedeagus with apex subacute (in punctures on elytra distinctly larger than dorsal view) and simple in its apical fourth diameters of punctures. It differs from G. (in lateral view) (Figs. 21-22). hornaditanus and G. tucumanensis by the Female genitalia. Tergum 10 similar to antennal segments 2 and 3 subequal in that of male. Second gonocoxite similar to length, and the distance separating medial that of G. jujuyensis and G. tucumanensis, interocular punctures on frons less than with one basal strong seta and stylus with twice as large as distance separating medial two apical setae. punctures from lateral punctures.

Geographical distribution. Gabrius Redescription. Length of the body 4.5- hornaditanus is currently only known from 5.1 mm. Coloration. Head black; thorax and northwestern Argentina (Jujuy) (Fig. 39). abdomen castaneous-piceous, with first three abdominal segments castaneous brunneous; Type material. Lectotype, male, with elytra castaneous brunneous to brunneous; labels: “Argent. Jujuy / Ornadita 3600 / Xi.19 antennae, palpi and legs brunneous. Weiser” (green), “TYPUS” (red), “hornaditanus Head about as long as wide (HW/HL= Bh. Vid. Rambousek” (white), “Lectotype 1.1) (Fig. 7) and at basal third as wide as Philonthus hornaditanus Rambousek, 1925 / distal third in both sexes, about as wide as to Des. Chani-Posse de Maus 2008” (NMPC). slightly wider than pronotum (HW/PW= 1.0- Paralectotypes: five specimens (2 females) 1.1); dorsal surface coarsely and moderately with labels: “Argent. Hor- / nadita 3400 punctate, with three postocular punctures / Xi.20. Weiser” (green), “Paralectotype and four epicranial punctures; distance Philonthus hornaditanus Rambousek, 1925 / separating medial interocular punctures on Des. Chani-Posse de Maus 2008” (NMPC), frons less than twice as large as distance one of them with additional labels: “TYPUS” separating medial punctures from lateral (red), “hornaditanus Bh. Vid. Rambousek” punctures (Fig. 7); infraorbital ridge present, (white); one additional specimen, male, with reaching postgenal ridge. Eyes distinctly labels: “Argentina: Jujuy / Hornadita 3600 / shorter than temples (EL/TL= 0.5-0.4) seen Xi.19 Weiser”, “hornaditanus Rambousek” from above (Fig. 7). Antennae with segment (white label), “Chicago NHMus. / M. 1 shorter than segments 2 and 3 combined, Bernhauer Collection” (FMNH). segments 2 and 3 subequal in length, Chani-Posse, M. Gabrius in southern South America 9

Fig. 39. Distribution of Gabrius argentinus, G. hornaditanus, G. jujujuyensis, G. nidicola, G. tucumanensis and G. nigritulus in southern South America. 10 Rev. Soc. Entomol. Argent. 70 (1-2): 01-15, 2011 segments 5-7 quadrate, segments 8-10 Iturbe / 3700m”, “jujuyensis Bernh. / Typus transverse. Maxillary palpus with segment 2 ded. Bruch” (yellow label), “Chicago no more than twice as long as its maximum NHMus. / M. Bernhauer Collection”, width. Labial palpus with last segment twice “Lectotype Philonthus jujuyensis Bernhauer, as long as preceding segment. Neck no more 1921 / Des. Chani-Posse de Maus 2008” than 0.5 times as wide as head at widest (FMNH). Paralectotype: female, with labels: point (NW/HW= 0.5) (Fig. 7). “Jujuy / 10.Xi.919 / Weiser”, “Cueva Iturbe / Pronotum longer than wide (PW/PL= 0.86- 3700m”, “jujuyensis Bernh. / Cotypus ded. 0.79), dorsal surface of pronotum with two Bruch” (yellow label), “Chicago NHMus. / rows of punctures, each with five punctures. M. Bernhauer Collection”, “Paralectotype Basisternum with medial longitudinal carina Philonthus jujuyensis Bernhauer, 1921 / Des. weakly developed. Metaventrite with circum- Chani-Posse de Maus 2008” (FMNH). mesocoxal ridge broadly rounded medially. Note: Chani-Posse de Maus (2009: 33, Elytra at sides as long as to moderately longer 34) designated lectotype. than pronotum at midline (EtL/PL=0.7), distinctly longer than elytra along suture (EtL/ Additional material examined: 33 Etl= 1.9-1.7); transverse distance between specimens (5 males, 6 females, 22 unsexed) punctures distinctly larger than diameters of ARGENTINA: Catamarca: Caspinchango, punctures (Fig. 9). Front tarsus shorter than 10.III.1921 (2 MACN); La Cienaga, Walters, front tibia; middle tarsus as long as middle Philonthus jujuyensis Bernh., (1 MACN). tibia; first segment of hind tarsus as long as Jujuy: Cueva Iturbe, 10.XI.919, Weiser, last segment (S1/S5= 1.0). Abdominal terga 3700m, Philonthus jujuyensis Bernh., (1 impunctuate basally, distance separating MLPA); Hornadita, 30.11.1909, Weiser, punctures about equal to diameters of 3700m, in green “Coypus”, Philonthus punctures. jujuyensis Bernh. (5 MACN); V. 1920, C. Male genitalia. Sternum 8 slightly Bruch, Chicago NHMus Bernhauer Collection emarginate medio-apically. Tergum 10 (1 FMNH). La Rioja: Los Mistoles, XII-15- subtruncate medio-apically, with two 2000, L. Herman, 600m, 30° 48’ 04’’S, 66° long subapical macrosetae and 3-4 short 18’ 40’’W (1 male, 1 female AMNH). San apical setae. Sternum 9 with basal portion Juan: S. of Astica, Chucuma, XI/IV/1995, asymmetrical, moderately emarginate L. Herman, 885m (1 AMNH). San Luis: apically with three long apical setae at Quebrada López, 7 km S San Francisco each side and two subapical setae (Fig. del Monte de Oro, L. Herman, 32° 39’ 23). Aedeagus with median lobe gradually 37’’S, 66° 07’ 46’’W (1 AMNH). Tucuman: narrowed from base of paramere; apex of Villa Nougués, Dec. 11, 1971, L. Herman, median lobe subacute (in dorsal view) and 1350m (4 males, 4 females, 5 AMNH), simple in its apical fourth (in lateral view) (2 IADIZA); 30km, SE Tafi del Valle, nr. El (Fig. 24). Indio, XI/20/1995, L. Herman, 950-1050m Female genitalia. Tergum 10 similar to (1 female, 2 AMNH). that of male. Second gonocoxite with one basal strong seta, and stylus with two apical Gabrius nidicola (Bernhauer, 1921) setae (Fig. 25). (Figs. 26-30, 39; Fig. 4 in Chani-Posse de Maus, 2009) Geographical distribution. Gabrius jujuyensis is known from the biogeographical Philonthus nidicola “Bernhauer in litt.“; provinces of Yungas and Chaco (Neotropical Bruch, 1915: 445 (cat.). Nomen nudum. region) and from Monte and Puna in the Philonthus nidicola Bernhauer, 1921: South American transition Zone (Fig. 39). 108; Scheerpeltz, 1933: 1352 (cat.); Blackwelder, 1944: 134 (list); Herman, Type material. Lectotype, male, with 2001: 2889 (cat.); Chani-Posse, 2004: labels: “Jujuy / 10.Xi.919 / Weiser”, “Cueva 231 (list). Chani-Posse, M. Gabrius in southern South America 11 Gabrius nidicola; Chani-Posse de Maus, than elytra along suture (EtL/Etl= 2.2–1.8); 2009: 34 (new combination) transverse distance between punctures about equal to diameters of punctures. Front tarsus Diagnosis. Gabrius nidicola may as long as front tibia; middle tarsus longer be identified by the antennal segments than middle tibia; first segment of hind tarsus 8-10 quadrate, and the distance separating about as long as last segment (S1/S5= 1.2). punctures on elytra about equal to diameters Abdominal terga moderately punctuate of punctures. It differs from G. jujuyensis by basally, distance separating punctures about the distance separating medial interocular equal to diameters of punctures. punctures on frons twice as large as distance Male genitalia. Sternum 8 slightly separating medial punctures from lateral emarginate medio-apically. Tergum 10 punctures. subtruncate medio-apically (Fig. 27), with two long subapical macrosetae and 3-4 short Redescription. Length of the body 4.3- apical setae. Sternum 9 with basal portion 5.3 mm. Coloration. Head black; thorax asymmetrical, moderately emarginate castaneous-brunneous; abdomen castaneous- apically with two long apical setae at each piceous to castaneous-brunneous, with side and two subapical setae (Fig. 26). first three abdominal segments brunneous; Aedeagus with median lobe gradually elytra brunneous to testaceous-brunneous; narrowed from base of paramere; apex of antennae, palpi and legs testaceous- median lobe subacute (in dorsal view) and brunneous. simple in its apical fourth (in lateral view) Head about as long as wide (HW/HL= 1.0) (Fig. 28). and at basal third as wide as distal third in Female genitalia. Tergum 10 similar to both sexes, about as wide as to slightly wider that of male (Fig. 29). Second gonocoxite than pronotum (HW/PW= 0.86-0.96); dorsal with one basal strong seta and one long surface coarsely and moderately punctate, apical seta, without stylus (Fig. 30). with three postocular punctures and four epicranial punctures; distance separating Geographical distribution. Gabrius medial interocular punctures on frons about nidicola is known from one biogeographical twice as large as distance separating medial province of the Neotropical region (Pampa) punctures from lateral punctures; infraorbital and from Monte and Puna in the South ridge present, reaching postgenal ridge. American transition Zone (Fig. 39). Eyes distinctly shorter than temples (EL/TL= 0.60-0.56) seen from above. Antennae with Bionomics. According to the labelling segment 1 shorter than segments 2 and 3 data from type material, this species was combined, segments 2 and 3 subequal in found in association with the rodent length, segments 5-10 quadrate. Maxillary Ctenomys. palpus with segment 2 no more than twice as long as its maximum width. Labial palpus Type material. Lectotype, male, with with last segment twice as long as preceding labels: “Rep. ARGENTINA / Prov. Buenos segment. Neck no more than 0.5 times as Aires / 17.X.1919 / C. Bruch”, “Monte Veloz wide as head at widest point (NW/HW= / 17.X.1919 / ex nido Ctenomys”, “nidicola 0.5). Bernh. / Typus” (yellow label), “Chicago Pronotum longer than wide (PW/PL= NHMus. / M. Bernhauer Collection”, 0.8), dorsal surface of pronotum with two “Lectotype Philonthus nidicola Bernhauer, rows of punctures, each with five punctures. 1921 / Des. Chani-Posse de Maus 2008” Basisternum with medial longitudinal carina (FMNH). Paralectotype: female, with labels: weakly developed. Metaventrite with circum- “Rep. ARGENTINA / Prov. Buenos Aires / mesocoxal ridge broadly rounded medially. 17.X.1919 / C. Bruch”, “189”, “nidicola Bernh. Elytra at sides distinctly shorter than pronotum / Cotypus” (yellow label), “Chicago NHMus. at midline (EtL/PL=0.6-0.5), distinctly longer / M. Bernhauer Collection”, “Paralectotype 12 Rev. Soc. Entomol. Argent. 70 (1-2): 01-15, 2011 Philonthus nidicola Bernhauer, 1921 / Des. biogeographical provinces of the Andean Chani-Posse de Maus 2008” (FMNH). region (Fig. 39). Note: Chani-Posse de Maus (2009: 34) designated lectotype. Bionomics. According to Smetana (1995), Gabrius nigritulus may occur in a Additional material examined: 14 wide range of habitats with decaying organic specimens (2 males, 2 females, 10 unsexed) matter. According to collecting data, it may ARGENTINA: Buenos Aires: 17.X.1919/ be found in Nothofagus and palm forests, 30.XII.1922, C. Bruch (3, 2 MACN). Jujuy: cacti, sclerophyl woodland, dump mud and Cueva Iturbe, 10.XI.919, Weiser (1 MACN). La moss on sand. Rioja: Los Mistoles, XII-15-2000, L. Herman (1 male, 1 female, 2 AMNH). San Luis: 7km, Type material. Lectotype of Gabrius N Trapiche, XII-6-2000, L. Herman, 1060m, nigritulus, male, with labels: “6139”, 33° 06’ 03’’S, 66° 03’ 39’’W, litter near river “Lectotype Gabrius nigritulus Gravenhorst A. (1 male, 1 female, 2 AMNH). Smetana des. 1960”, “Hist.-Coll. (Coleoptera) / Nr. 6139 / Philonthus aterrimus Gyllh. / Gabrius nigritulus (Gravenhorst, 1802) Europa, Americ. sept. / Zool. Mus. Berlin” (Figs. 12, 36-38, 39 in this paper; Figs. (ZMHB). Paralectotypes: 3 males, with 1001-1007 in Smetana, 1995; Fig. 64 in labels: “Gabrius Steph. / nigritulus Grav. / det. Schillhammer, 1997) Smetana 1960”, “Hist.–Coll. (Coleoptera) / Nr. 6139 / Philonthus aterrimus Gyllh. / Staphylinus nigritulus Gravenhorst, 1802: Europa, Americ. sept. / Zool. Mus. Berlin” 41. For complete references see Herman (ZMHB). (2001). Note: The lectotype was designated by Gabrius chiliensis Coiffait & Sáiz, 1968: 348; Smetana in 1960 (see labels on type material) Sáiz, 1971: 340; Chani-Posse de Maus, and published two years later (Smetana, 2009: 40 (syn). 1962). Paratypes of Gabrius chiliensis, male, Diagnosis. This species can be with labels: “Cord. Chillan”, “Germain / distinguished from the other species of 1899”, “PARATYPE”, “Gabrius chiliensis southern South American Gabrius by having Coiff. et Saiz”, “Chile Tipo No 2238”; female, the dorsal rows on pronotum each with six with labels: “Germain / 1899”, “PARATYPE”, punctures. Additionally, the male differs by “Gabrius chiliensis Coiff. et Saiz”, “Chile the sternum 9 with the basal portion more Tipo No 2245” (MNNC). or less symmetrical (Fig. 36), the tergum 10 without apical setae and the paramere Additional material examined: 16 of aedeagus bifurcate, with sensory peg specimens (4 males, 10 females, 2 unsexed) setae (Fig. 37, 38). The female differs by the ARGENTINA: Rio Negro: Villa Regina, tergum 10 with the apex rounded and two Feb. 14, 1972, L. Herman (1 male, 1 female apical setae. AMNH). Chubut: El Hoyo, 10.IX.61 (1 female NMW). CHILE: Curicó: 20km Redescription. See Smetana (1995: E. Potrero Grande, El Relvo, 08-May-04, 619). 1100m, 36° 11.14’ S, 70° 56.1’ W, fogging Nothophagus dombeyi (2 females, 2 JEBC). Geographical distribution. Gabrius Concepción: Valle Nonguen, January nigritulus is assumed to have been introduced 15, 1978, T. Cekalovic (1 male AMNH). to various regions of the world from the Quillota: P.N. La Campana (Sector Ocoa) western portion of the Palearctic region, vic Quebrada Buitrera, 28.XII.2008, Thayer, and it is now considered a cosmopolitan Clarke, 415m, 32° 55.89’ S, 71° 05.10’ W, species (Smetana 1995). In southern South sclerophyl woodland, Jubaea chilensis America G. nigritulus is known from several palms, Trichocereus cacti; dump mud & Chani-Posse, M. Gabrius in southern South America 13 moss on sand, dry creek bed (1 male, 1 distance separating medial punctures from female FMNH). Valparaíso: Viña del Mar, lateral punctures; infraorbital ridge present, 28.12.94, Germain, Gabrius chiliensis Coiff. reaching postgenal ridge. Eyes distinctly et Saiz F. Saiz, det. 1970 (1 male, 1 female shorter than temples (EL/TL= 0.5-0.4) seen MNNC). Cachapoal: Las Cabras, 10/23. from above (Fig. 10). Antennae with segment XII.1954, Peña, 1480m, L. Peña Leg., Gabrius 1 shorter than segments 2 and 3 combined, chiliensis Coiffait & Saiz Det. Chani-Posse segment 2 shorter than segment 3, segment 2006 (1 female FMNH). Bío Bío: Recinto, 5 elongate, segments 6-8 quadrate and 9-10 I.1953, Peña, L. Peña Leg. (1 female FMNH). slightly transverse (Fig. 10). Maxillary palpus Maule: El Radal, 28/30.XI.1957, Peña, with segment 2 no more than twice as long 900m, L. Peña Leg. (1 female FMNH). as its maximum width. Labial palpus with last segment twice as long as preceding segment. Gabrius tucumanensis (Bernhauer, 1927) Neck no more than 0.5 times as wide as head (Figs. 10, 11, 31-35, 39; Figs. 3 a–b in at widest point (NW/HW= 0.5) (Fig. 10). Chani-Posse de Maus, 2009) Pronotum longer than wide (PW/PL= 0.9- 0.8) (Fig. 10), dorsal surface of pronotum Philonthus tucumanensis Bernhauer, 1927: with two rows of punctures, each with 245; Scheerpeltz, 1933: 1365 (cat.); five punctures. Basisternum with medial Blackwelder, 1944: 135 (list); Herman, longitudinal carina weakly developed. 2001: 2982 (cat.); Chani-Posse, 2004: Metaventrite with circum-mesocoxal ridge 231 (list). narrowly rounded medially. Elytra at sides as Gabrius tucumanensis; Chani-Posse de long as to moderately longer than pronotum Maus, 2009: 33, 34 (new combination) at midline (EtL/PL=0.7), distinctly longer than elytra along suture (EtL/Etl= 1.9-1.7); Diagnosis. Gabrius tucumanensis may transverse distance between punctures be identified by the antennal segments 9-10 distinctly larger than diameters of punctures transverse, the distance between punctures (Fig. 11). Front tarsus shorter than front tibia; on elytra distinctly larger than diameters of middle tarsus as long as middle tibia; first punctures and the distance separating medial segment of hind tarsus as long as last segment interocular punctures on frons twice to three (S1/S5= 1.0). Abdominal terga punctuate times as large as distance separating medial basally, distance separating punctures punctures from lateral punctures. It differs distinctly larger than diameters of punctures. from G. hornaditanus and G. jujuyensis by Male genitalia. Sternum 8 slightly the head longer than wide, and the antennal emarginate medio-apically (Fig. 31). Tergum segment 5 elongate. 10 subtruncate medio-apically, with two long subapical macrosetae and 3-4 short Redescription. Length of the body 4.5- apical setae. Sternum 9 similar to that of 5.1 mm. Coloration. Head black; thorax P. hornaditanus, with 5-6 long apical setae castaneous; abdomen castaneous-piceous; and two subapical setae. Aedeagus with elytra castaneous brunneous to brunneous; median lobe gradually narrowed from base scutellum black; antennae, palpi and legs of paramere; apex of median lobe arcuate (in brunneous. dorsal view) and simple in its apical fourth Head longer than wide (HW/HL= 0.8) (in lateral view) (Figs. 33, 34). (Fig. 10) and at basal third as wide as distal Female genitalia. Tergum 10 emarginate third in both sexes, about as wide as to medio-apically, with two long subapical slightly wider than pronotum (HW/PW= 1.0- macrosetae and 5-6 short apical setae. (Fig. 1.1); dorsal surface coarsely and moderately 32). Second gonocoxite with one basal punctate, with three postocular punctures strong seta, and stylus with two apical setae and four epicranial punctures; distance (Fig. 35). separating medial interocular punctures on frons twice to three times as large as Geographical distribution. Gabrius 14 Rev. Soc. Entomol. Argent. 70 (1-2): 01-15, 2011 tucumanensis is known from three Grant (MCZ, Harvard University), Austral biogeographical provinces of the Neotropical Staphylinidae PEET grant 0118749 to Margaret region (Chaco, Pampa and Yungas) and from K. Thayer and Alfred F. Newton, FMNH- Monte and Puna in the South American Collection Study Grant, FONCYT-PICT 01- transition Zone (Fig. 39). 11120 to Jorge Miguel Lobo and Sergio Roig, Foundation Grant BBVA as part of the project: Bionomics. According to the labelling “Design of a reserve network for protection data, this species may be found in litter near of biodiversity in Southern South America streams and rivers. using predictive distribution models with hyperdiverse taxa”, CONICET PIP 5766 to Type material. Lectotype, female, A. Marvaldi and CONICET PIP 112-200801- “San Pedro Colalao / Tucum. Weiser”, 00162 to A. E. Marvaldi and G. E. Flores. “tucumanensis Bernh. / Typus don. Bruch” (yellow label), “Chicago NHMus. / M. Bernhauer Collection”, “Lectotype Philonthus LITERATURE CITED tucumanensis Bernhauer, 1927 / Des. Chani- Posse de Maus 2008” (FMNH). 1. BERNHAUER, M. 1912. Zur Staphylinidenfauna von Südamerika (10. Beitrag). Verhandlungen der k. k. Note: Chani-Posse de Maus (2009: 33, zoologisch-Botanischen Gesellschaft in Wien 62: 26- 34) designated lectotype. 48. 2. BERNHAUER, M. 1921. Neue Staphyliniden aus Südamerika, besonders aus Argentinien. Wiener Entomologische Additional material examined: 50 Zeitung 38 : 101-108 [June 15], 169-179 [October 20]. specimens (29 males, 8 females, 13 unsexed). 3. BERNHAUER, M. 1927. Zur Staphylinidenfauna Süd- ARGENTINA: San Luis: Irideus, Quebrada Amerikas, insbesondere Argentiniens. Archiv für del Tala, 1100 m, 32°39’01’’S, 64°57’23’’W, Naturgeschichte (1925)(A) 91(12): 229-264. 4. BERNHAUER, M. & K. SCHUBERT. 1914. Staphylinidae IV. litter in moist narrow quebrada, 8.XII.2000, In: Schenkling, S. (Ed.), Coleopterorum Catalogus, L. Herman (25 males, 4 females AMNH); 5(57). W. Junk, Berlin, pp. 289-408. 5. BLACKWELDER, R. E. 1936. Morphology of the coleopterous Quebrada López, 7 km S. San Francisco del family Staphylinidae. Smithsonian Miscellaneous Monte de Oro, 3 males, 3 females, 4; 7km, Collection 94: 1-102. 6. BLACKWELDER, R. E. 1944. Checklist of the Coleopterous N Trapiche, XII-6-2000, L. Herman, 1060m, insects of Mexico, Central America, the West Indies, 33° 06’ 03’’S, 66° 03’ 39’’W, litter near river and South America. Part I. United States National (4 AMNH). Tucumán: Villa Nougués, 1350 Museum Bulletin 185: xii + 188 pp. 7. BRUCH, C. 1915. Catálogo Sistemático de los coleópteros de m, 11.XII.1971, L. Herman (1 male, 1 female la República Argentina. Pars VI. Revista del Museo de AMNH). La Rioja: Los Corrales, 15 km NW La Plata 19 (Segunda Serie)(6): 235-302. 8. CHANI POSSE, M. R. 2004. Eight Argentinean species of Famatina, Dec. 3, 1971, Lee Herman (2 dung-inhabiting Philonthus Stephens (Coleoptera: AMNH). Salta: 10km. NW Cafayate, Dec. Staphylinidae). Studies on Neotropical Fauna and Environment 39: 217-232. 17, 1971, Lee Herman (1 AMNH). Jujuy: 9. CHANI POSSE, M. 2006. Systematic revision and cladistic Cueva Iturbe, 10.XI.919, Weiser (2 NMW). analysis of the Neotropical genus Chroaptomus Sharp (Coleoptera: Staphylinidae), with descriptions of two new species. Insect Systematics & Evolution 37: 361- 383. ACKNOWLEDGEMENTS 10. CHANI-POSSE DE MAUS, M. 2009. Revision of some types of Philonthus Stephens and Gabrius Stephens from southern South America (Coleoptera: Staphylinidae), I gratefully acknowledge the curators with nomenclatural notes. Zootaxa, 2034: 31-42. 11. CHANI POSSE, M. 2010. Revision of the southern South listed in Material and methods for the American species of Philonthus Stephens (Coleoptera: loan of specimens. I am very indebted to Staphylinidae). Zootaxa 2595: 1-70. Gustavo E. Flores for valuable comments 12. GRAVENHORST, J. L. C. 1802. Coleoptera Microptera Brunsvicencia nec non exoticorum quotquot exstant that improved different aspects of this work. in collectionibus entomologorum brunvicencsium in I also thank Christian Maus for his support genera familias et species distribuit. Carolus Reichard, Brunsvigae, 206 pp. during this study. This study was supported 13. HERMAN, L. H. 2001. Catalog of the Staphylinidae (Insecta: by the Consejo Nacional de Investigaciones Coleoptera). 1758 to the end of the second millennium. Part V. Bulletin of the American Museum of Natural Científicas y Técnicas (CONICET), Argentina, History 265: 2441-3020. and the following grants: Ernst Mayr Travel Chani-Posse, M. Gabrius in southern South America 15 14. MORRONE, J. J. 2006. Biogeographic areas and transition 20. SMETANA, A. 1982. Revision of the subfamily Xantholininae zones of Latin America and the Caribbean islands of America North of Mexico (Coleoptera: based on panbiogeographic and cladistic analyses of Staphylinidae). Memories of the Entomological Society the entomofauna. Annual Review of Entomology 51: of Canada 12: 1-389. 467-494. 21. SMETANA, A. 1995. Rove beetles of the subtribe Philonthina 15. RAMBOUSEK, F. 1925. Staphylinides nouveaux de l’Amérique of America north of Mexico (Coleoptera: Staphylinidae): meridionale. (2e contribution). Časopis Československé Classification, phylogeny and taxonomic revision. Společnosti Entomologické 21: 69-76. Memoirs on Entomology, International 3: 1-946. 16. SCHEERPELTZ, O. 1933. Staphylinidae VII. In: Schenkling, 22. SMETANA, A. & A. DAVIES. 2000. Reclassification of the S. (Ed.), Coleopterorum Catalogus. Pars 129. W. Junk, north temperate taxa associated with Staphylinus Berlin, pp. 989-1500. sensu lato, including comments on relevant subtribes 17. SCHILLHAMMER, H. 1997. Taxonomic revision of the of Staphylinini (Coleoptera: Staphylinidae). American Oriental species of Gabrius Stephens. Monographs on Museum Novitates 3287: 1-88. Coleoptera 1: 1-139. 23. STEPHENS, J. F. 1829. The Nomenclature of British Insects; 18. SCHILLHAMMER, H. 2000. New species and records Being a Compendious List of Such Species as are of Oriental and East Palaearctic species of Gabrius Contained in the Systematic Cat.ue of British Insects, Stephens, 1829. Annalen des Naturhistorischen and Forming a Guide to their Classification. Baldwin & Museums 102 B: 117-142. Cradock, London, 68 pp. 19. SMETANA, A. 1962. Zwei Gabrius-Arten aus der Gravenhorstschen Sammlung (Col., Staphylinidae). Deutsche Entomologische Zeitschrift (N.F.) 9: 236- 238.