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Revista de la Sociedad Entomológica Argentina

versión impresa ISSN 0373-5680

Rev. Soc. Entomol. Argent. vol.70 no.1-2 Mendoza ene./jun. 2011

 

TRABAJOS CIENTÍFICOS

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

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

 

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: mchani@mendoza-conicet.gov.ar

Recibido: 2-VIII-2010
Aceptado: 12-X-2010

 


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.

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.


 

INTRODUCTION

The genus Gabrius Stephens, 1829 belongs to the subtribe Philonthina in the subfamily Staphylininae. According to Smetana (1995) Gabrius was often considered by many authors as a subgenus of Philonthus, but nowadays is assumed to be a separate genus which is closely related to Philonthus and other genera. These relationships have been discussed by Smetana (1995) and are still to be clarified (Smetana, 1995; Schillhammer, 1997; Chani-Posse, 2010). The genus Gabrius is distributed worldwide and accounts for more than 300 species at present (Herman, 2001). Most of its species are known to occur in the Nearctic, Palearctic and Oriental regions. This knowledge may change and the number of species may increase as poorly known tropical faunas are studied (Schillhammer, 1997) and the narrower generic concepts at present applied to the Holarctic fauna (e.g. Smetana, 1995) are applied to those. According to Herman (2001), only three species of Gabrius are known to occur in America south of Mexico, two of them cited from southern South America, G. nigritulus (Gravenhorst) and G. chiliensis Coiffait & Sáiz. The latter name was recently found to be a synonym of the former (Chani-Posse de Maus, 2009). As part of a major revisionary study of the species of Philonthus Stephens, 1829 (Coleoptera: Staphylinidae) from southern South America (Chani-Posse de Maus, 2009; Chani-Posse, 2010), five more species of Gabrius are recognized for this area. The geographical area covered in this study, as in a previous work (Chani-Posse, 2010), is the southern portion of South America which comprises primarily the countries of Argentina, Chile, Paraguay and Uruguay.
The objectives of this study are to redescribe five species of Gabrius (previously assigned to Philonthus) by incorporating new characters from external morphology, including genitalia, present a key to the species for southern South America and detail their geographic distribution.

MATERIAL AND METHODS

The present study is based on the examination of about 150 adult specimens, including type material. The material studied was obtained on loan from, and are deposited in the following collections: AMNH American Museum of Natural History, New York, USA (Lee   H. Herman). CEUNC Colección   Entomológica   de   la Universidad de Córdoba, Argentina (Miguel Angel Delfino). FMNH Field Museum of Natural History, Chicago, USA   (Alfred F. Newton, Margaret K. Thayer). IADIZA Instituto Argentino de Investigaciones de las Zonas Aridas, Mendoza, Argentina (Sergio Roig Juñent). JEBC Juan E. Barriga private collection, Curicó, Chile (Juan E. Barriga). MACN Museo Argentino de Ciencias Naturales Bernardino Rivadavia, Buenos Aires, Argentina (Arturo Roig Alsina). MLPA Museo de La Plata, La Plata, Argentina (Norma B. Díaz). NMPC  National Museum (Natural History), Prague, Czech Republic (Jiři Hajek) NMW Naturhistorisches Museum   Wien, Austria (Harald Schillhammer). MNNC Museo Nacional de Historia Natural, Santiago, Chile (Mario Elgueta D.) ZMHB  Museum für Naturkunde der Humboldt Universität, Berlin, Germany (Manfred Uhlig, Bernd Jaeger)
Techniques for the preparation and examination of male and female genitalia follow Smetana (1982). Descriptions were made using a Leica MZ6 dissecting scope, and several genitalic features were examined with a Leitz Wetzlar compound microscope. Drawings were made with a camera lucida attached to the compound microscope or dissecting scope. Photographs were taken using a digital camera attached to the dissecting scope. Measurements (given in millimeters) were made with an ocular micrometer. Overall body length was measured from the apex of the labrum to the apex of the abdomen. Other measurements were taken and abbreviated as follows: HW- maximum head capsule width HL- length of head capsule, from anterior margin of frontoclypeus to neck constriction (along midline)
PW- pronotum maximum width PL- pronotum length (along midline) EL- eye length (seen from above) TL- temple length (from the posterior margin of the eye to the nuchal groove; seen from above) NW- neck width S1- first segment of hind tarsus length S5- last segment of hind tarsus length EtL- elytra length at sides (from humerus to apex; seen from above) Etl- elytra length along suture EtW- elytra width at base Terminology mainly follows Blackwelder (1936), Smetana (1995), Smetana & Davies (2000) and Schillhammer (2000) with slight
modifications and/ or inclusions already detailed by the author in previous studies (Chani-Posse, 2006).
Biogeographical provinces considered in the geographical distribution of the species follow those of Morrone (2006). All records and the general distribution given for each species are based strictly on specimens that I have examined. All locality data were collected from collection data.

General remarks

The most recent and complete description of the genus, at least for America north of Mexico, is available in Smetana (1995). Some characters of Gabrius are also discussed in Schillhammer (1997).

Recognition of the currently known southern South American species

The labial palpus with the last segment slender, as long as to slightly longer and distinctly narrower than the penultimate segment allows recognition among the other southern South American Philonthina. The genus can also be recognized by the sternum 9 of male genital segment with basal portion not reduced, more or less symmetrical with exception of that of G. brevipennis (Smetana, 1995), the East Palearctic and Oriental species of the fimetarioides group (Schillhammer, 1997), P. argentinus, P. jujuyensis, and P . nidicola (Chani-Posse de Maus, 2009).
Besides these characters, Gabrius can be separated from Philonthus by having the first four segments of front tarsus in both sexes simple, not dilated, with ventral face bearing only regular marginal setae, from Belonuchus by having the lateral puncture of pronotum bearing long macroseta separated from superior line of pronotal hypomeron by a distance about equal to diameter of puncture, and from Endeius by having the last segment of maxillary palpus in general fusiform, not acutely narrowed apically (Fig. 12).


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.

Other characters common to the southern South American species of Gabrius known at present are the following: postmandibular ridge present (Fig. 8); antennal segment 4 elongate (Figs. 5, 10); maxillary palpus with segment 2 longer than segment 3; posterior arms of gular sutures joined before neck (Fig. 2); submentum about 1.5 times as long as mentum; neck no more than 0.5 times as wide as head at widest point; pronotum slightly narrowed anteriad to parallel-sided (Figs. 3, 10); dorsal surface of pronotum with two rows of punctures, each with five or six punctures; first segment of hind tarsus shorter than segments 2 and 3 combined, and about as long as last segment (Figs. 4, 11); abdominal terga 2 and 3 with posterior basal line straight medially (Figs. 4, 11); male sternum 7 with apical margin straight; tergum 8 in both sexes with posterior margin truncate; styli of tergum 9 simple, moderately stout; sternum 8 with two long subapical macrosetae at each side (Figs. 13, 19, 31); paramere entire and reduced, completely fused to median lobe (Figs. 15, 21, 28, 33) with exception of that of G. nigritulus (Fig. 37); second gonocoxite with one strong basal seta (Figs. 18, 25, 30, 35).


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.

RESULTS

Key to southern South American species of Gabrius

1.  Distance separating medial interocular punctures on frons less than twice distance separating medial punctures from lateral punctures (Fig. 7) ……………………….……..................  Gabrius jujuyensis (Bernhauer)
1’. Distance separating medial interocular punctures on frons twice or more as large as distance separating medial punctures from lateral punctures (Figs. 1, 6, 12) … ........  2

2. First antennal segment as long as segments 2 and 3 combined; antennal segment 8 transverse ............…................................................  Gabrius argentinus (Bernhauer)
2’. First antennal segment shorter than segments 2 and 3 combined; antennal segment 8 drate …......………………..……. 3

3.  Head distinctly longer than wide (Fig.10); antennal segment 5 elongate (Fig. 10)….........  Gabrius tucumanensis (Bernhauer)
3’. Head about as long as wide (Figs. 6, 12); antennal segment 5 quadrate (Fig. 5) ……… 4

4.  Antennal segment 3 longer than 2 (Fig. 5); middle tarsus shorter than middle tibia …......  Gabrius hornaditanus (Rambousek)
4’. Antennal segment 2 and 3 subequal in length (Fig. 12); middle tarsus as long as to longer than middle tibia .............… 5

5.  Antennal segment 10 transverse; front tarsus shorter than front tibia …........……...................  Gabrius nigritulus (Gravenhorst)
5’. Antennal segment 10 quadrate; front tarsus as long as front tibia …………….........................  Gabrius nidicola (Bernhauer)

Gabrius argentinus (Bernhauer, 1912) (Figs. 1-4, 13-18, 39; Figs. 1 a-b in Chani-Posse de Maus, 2009) Philonthus argentinus Bernhauer, 1912: 43; Bernhauer & Schubert, 1914: 329 (cat.); Bruch, 1915: 444 (cat.); Blackwelder, 1944: 132 (list); Herman, 2001: 2751 (cat.); Chani-Posse, 2004: 230 (list). Gabrius argentinus; Chani-Posse de Maus, 2009: 32, 34 (new combination)


Fig. 39. Distribution of Gabrius argentinus, G. hornaditanus, G. jujujuyensis, G. nidicola, G. tucumanensis and G. nigritulus in southern South America.

Diagnosis. Gabrius argentinus may be identified by its body shape and coloration, the antennal segment 1 as long as segments 2 and 3 combined and the apex of median lobe turn downwards in its apical fourth (in lateral view). It differs from G. tucumanensis by the antennal segment 5 quadrate and the basal portion of sternum 9 asymmetrical.

Redescription. Length of the body 7.0-7.5 mm. Coloration. Head castaneous-piceous to black; thorax castaneous-piceous; elytra and abdominal segments castaneous-piceous to black; antennae castaneous-piceous to castaneous-brunneous; palpi and legs castaneous-brunneous to brunneous.
Head moderately to slightly longer than wide (HW/HL= 0.9-0.8) (Fig. 1) and at basal third as wide as distal third in both sexes, about as wide as pronotum (HW/PW= 1.0-
0.9); dorsal surface coarsely and moderately punctate, with three postocular punctures and two to three epicranial punctures; distance separating medial interocular punctures on frons about three times as large as distance separating medial punctures from lateral punctures (Fig. 1); infraorbital ridge present, reaching postgenal ridge. Eyes distinctly shorter than temples (EL/TL= 0.75-0.67) seen from above (Fig. 1). Antennae with segment 1 as long as segments 2 and 3 combined, segment 2 shorter than segment 3, segments 5-6 quadrate, segments 7-10 transverse. Maxillary palpus with segment 2 no more than twice as long as its maximum width. Labial palpus with last segment twice as long as preceding segment. Neck no more than 0.5 times as wide as head at widest point (NW/HW= 0.4) (Fig. 1).
Pronotum about as long as wide (PW/ PL= 1.07-0.94), dorsal surface of pronotum with two rows of punctures, each with five punctures (Fig. 3). Basisternum with medial longitudinal carina slightly developed. Metaventrite with circum-mesocoxal ridge broadly rounded medially. Elytra at sides as long as to moderately longer than pronotum at midline (EtL/PL=1.0-0.8), distinctly longer than elytra along suture (EtL/Etl= 1.7-1.5); transverse distance between punctures distinctly larger than diameters of punctures (Fig. 4). Front and middle tarsus shorter than front and middle tibia; first segment of hind tarsus about as long as last segment (S1/S5= 1.2) (Fig. 4). Abdominal terga moderately punctuate basally, distance separating punctures distinctly larger than diameters of punctures (Fig. 4).
Male genitalia. Sternum 8 moderately emarginate medio-apically (Fig. 13). Tergum 10 subtruncate medio-apically (Fig. 17), with two long subapical macrosetae and 2-6 short apical setae. Sternum 9 with basal portion asymmetrical, moderately emarginate apically with two to three long apical setae at each side (Fig. 14). Aedeagus with median lobe gradually narrowed from apical third; apex of median lobe subacute (in dorsal view) and turn downwards in its apical fourth (in lateral view) (Figs. 15-16). Female genitalia. Tergum 10 similar to that
of male. Second gonocoxite with one basal strong seta and one long apical seta, without stylus (Fig. 18).

Geographical distribution. Gabrius argentinus is currently only known from northwestern Argentina (Tucumán) (Fig. 39).

Type material. Lectotype, male, with labels: “Rep. Argentina / PROV. TUCUMAN / RICHTER / 19”, “387”, “argentinus Bernh. / Typus” (yellow label), “Chicago NHMus. / M. Bernhauer Collection”, “Lectotype Philonthus argentinus Bernhauer, 1912 / Des. Chani-Posse de Maus 2008”. (FMNH). Paralectotypes: 2 females, “Rep. Argentina / PROV. TUCUMAN / RICHTER / 19”, “395” (one specimen), “argentinus Bernh. / Cotypus” (yellow label), “Chicago NHMus. / M. Bernhauer Collection”, “Paralectotype Philonthus argentinus Bernhauer, 1912 / Des. Chani-Posse de Maus 2008” (FMNH).
Note: Chani-Posse de Maus (2009: 32, 34) designated lectotype.

Additional material examined: 14 specimens (1 male, 4 females, 9 unsexed). ARGENTINA: Tucumán: Richter (1 CEUNC), 389 (1 MLPA), Philonthus argentinus Brh. (1 MLPA), in green “cotypus”, Bernhauer determ. (2 females, 2 MLPA); Yerba Buena, M. A. Jaynes, Philonthus argentinus Brh., (1 MACN); 1938, G. M. Borg, Philonthus boxi Bernh., (1 MLPA); 5 Oct 1927, in green “Typus”, 2838, in Bernahuer handwriting Philonthus boxi Bernh. n. sp.[upside down], (1 MLPA), 2836, (1 MLPA); Villa Nougués, Dec. 11, 1971, L. Herman, 1250, 759 (1 male, 1 female AMNH); 30km, SE Tafi del Valle, nr. El Indio, XI/20/1995, L. Herman, 950-1050m, 2891 (1 female AMNH).

Gabrius hornaditanus (Rambousek, 1925) (Figs. 5, 6, 19-22, 39) Philonthus hornaditanus “Rambousek   in litt.”; Bruch, 1915: 445 (cat.). Nomennudum. Philonthus hornaditanus Rambousek, 1925: 75; Scheerpeltz, 1933: 1345 (cat.); Blackwelder, 1944: 133 (list); Herman, 2001: 2839 (cat.); Chani-Posse, 2004: 231 (list). Gabrius hornaditanus; Chani-Posse de Maus, 2009: 33, 34 (new combination).

Diagnosis. Gabrius hornaditanus may be identified by the distance separating punctures of abdominal terga distinctly smaller than the diameters of punctures and the deeply emarginate posterior margin of the male sternum 8. It differs from G. jujuyensis and G. tucumanensis by having the antennal segment 10 quadrate and the middle tarsus shorter than middle tibia.

Redescription. Length of the body 5.0-5.2 mm. Coloration. Head black; thorax and abdominal segments castaneous; elytra testeceous-brunneus; antennae, palpi and legs castaneous brunneous to brunneous. Head about as long as wide (HW/HL= 1.1) and at basal third as wide as distal third in both sexes, about as wide as to slightly wider than pronotum (HW/PW= 1.0-1.1); dorsal surface coarsely and moderately punctate, with three postocular punctures and four epicranial punctures; distance separating medial interocular punctures on frons about three times as large as distance separating medial punctures from lateral punctures and at basal third slightly narrower than at distal third (Fig. 6); infraorbital ridge present, reaching postgenal ridge. Eyes distinctly shorter than temples (EL/TL= 0.5-0.4) seen from above. Antennae with segment 1 shorter than segments 2 and 3 combined, segment 2 shorter than segment 3, segments 5-10 quadrate (Fig. 5). Maxillary palpus with segment 2 no more than twice as long as its maximum width. Labial palpus with last segment 1.5 times as long as preceding segment. Neck no more than 0.5 times as wide as head at widest point (NW/HW= 0.5).
Pronotum longer than wide (PW/PL= 0.86-0.79), dorsal surface of pronotum with two rows of punctures, each with five punctures. Basisternum with medial longitudinal carina moderately developed. Metaventrite with circum-mesocoxal ridge broadly rounded medially. Elytra at
sides as long as to moderately longer than pronotum at midline (EtL/PL=0.7), distinctly longer than elytra along suture (EtL/Etl= 1.9-1.7); transverse distance between punctures distinctly smaller than diameters of punctures. Front tarsus shorter than front tibia; middle tarsus shorter than middle tibia; first segment of hind tarsus as long as last segment (S1/S5= 1.0). Abdominal terga moderately punctuate basally, distance separating punctures distinctly smaller than diameters of punctures.
Male genitalia. Sternum 8 deeply emarginate medio-apically (Fig. 19). Tergum 10 subtruncate medio-apically, with two long subapical macrosetae and 3-4 short apical setae. Sternum 9 with basal portion more or less symmetrical, moderately emarginate apically with two long apical setae at each side and two subapical setae (Fig. 20). Aedeagus with apex subacute (in dorsal view) and simple in its apical fourth (in lateral view) (Figs. 21-22).
Female genitalia. Tergum 10 similar to that of male. Second gonocoxite similar to that of G. jujuyensis and G. tucumanensis, with one basal strong seta and stylus with two apical setae.

Geographical distribution. Gabrius hornaditanus is currently only known from northwestern Argentina (Jujuy) (Fig. 39).

Type material. Lectotype, male, with labels: “Argent. Jujuy / Ornadita 3600 / XI.19 Weiser” (green), “TYPUS” (red), “hornaditanus Bh. Vid. Rambousek” (white), “Lectotype Philonthus hornaditanus Rambousek, 1925 / Des. Chani-Posse de Maus 2008” (NMPC). Paralectotypes: five specimens (2 females) with labels: “Argent. Hor- / nadita 3400 / XI.20. Weiser” (green), “Paralectotype Philonthus hornaditanus Rambousek, 1925 / Des. Chani-Posse de Maus 2008” (NMPC), one of them with additional labels: “TYPUS” (red), “hornaditanus Bh. Vid. Rambousek” (white); one additional specimen, male, with labels: “Argentina: Jujuy / Hornadita 3600 / XI.19 Weiser”, “hornaditanus Rambousek” (white label), “Chicago NHMus. / M. Bernhauer Collection” (FMNH).

Note: Chani-Posse de Maus (2009: 32, 33) designated lectotype.

Gabrius jujuyensis (Bernhauer, 1921) (Figs. 7-9, 23-25, 39; Figs. 2 a-b in Chani-Posse de Maus, 2009) Philonthus jujuyensis “Bernhauer in litt.“; Bruch, 1915: 445 (cat.). Nomen nudum. Philonthus jujuyensis Bernhauer, 1921: 107; Scheerpeltz, 1933: 1347 (cat.); Blackwelder, 1944: 133 (list); Herman, 2001: 2850 (cat.); Chani-Posse, 2004: 231 (list). Gabrius jujuyensis; Chani-Posse de Maus, 2009: 33, 34 (new combination)

Diagnosis. Gabrius jujuyensis may be identified by the antennal segments 8-10 transverse, and the distance separating punctures on elytra distinctly larger than diameters of punctures. It differs from G. hornaditanus and G. tucumanensis by the antennal segments 2 and 3 subequal in length, and the distance separating medial interocular punctures on frons less than twice as large as distance separating medial punctures from lateral punctures.

Redescription. Length of the body 4.5-5.1 mm. Coloration. Head black; thorax and abdomen castaneous-piceous, with first three abdominal segments castaneous brunneous; elytra castaneous brunneous to brunneous; antennae, palpi and legs brunneous.
Head about as long as wide (HW/HL= 1.1) (Fig. 7) and at basal third as wide as distal third in both sexes, about as wide as to slightly wider than pronotum (HW/PW= 1.0-1.1); dorsal surface coarsely and moderately punctate, with three postocular punctures and four epicranial punctures; distance separating medial interocular punctures on frons less than twice as large as distance separating medial punctures from lateral punctures (Fig. 7); infraorbital ridge present, reaching postgenal ridge. Eyes distinctly shorter than temples (EL/TL= 0.5-0.4) seen from above (Fig. 7). Antennae with segment 1 shorter than segments 2 and 3 combined, segments 2 and 3 subequal in length,
segments 5-7 quadrate, segments 8-10 transverse. Maxillary palpus with segment 2 no more than twice as long as its maximum width. Labial palpus with last segment twice as long as preceding segment. Neck no more than 0.5 times as wide as head at widest point (NW/HW= 0.5) (Fig. 7).
Pronotum longer than wide (PW/PL= 0.86-0.79), dorsal surface of pronotum with two rows of punctures, each with five punctures. Basisternum with medial longitudinal carina weakly developed. Metaventrite with circum-mesocoxal ridge broadly rounded medially. Elytra at sides as long as to moderately longer than pronotum at midline (EtL/PL=0.7), distinctly longer than elytra along suture (EtL/ Etl= 1.9-1.7); transverse distance between punctures distinctly larger than diameters of punctures (Fig. 9). Front tarsus shorter than front tibia; middle tarsus as long as middle tibia; first segment of hind tarsus as long as last segment (S1/S5= 1.0). Abdominal terga impunctuate basally, distance separating punctures about equal to diameters of punctures.
Male genitalia. Sternum 8 slightly emarginate medio-apically. Tergum 10 subtruncate medio-apically, with two long subapical macrosetae and 3-4 short apical setae. Sternum 9 with basal portion asymmetrical, moderately emarginate apically with three long apical setae at each side and two subapical setae (Fig. 23). Aedeagus with median lobe gradually narrowed from base of paramere; apex of median lobe subacute (in dorsal view) and simple in its apical fourth (in lateral view) (Fig. 24).
Female genitalia. Tergum 10 similar to that of male. Second gonocoxite with one basal strong seta, and stylus with two apical setae (Fig. 25).

Geographical distribution. Gabrius jujuyensis is known from the biogeographical provinces of Yungas and Chaco (Neotropical region) and from Monte and Puna in the South American transition Zone (Fig. 39).

Type material. Lectotype, male, with labels: “Jujuy / 10.XI.919 / Weiser”, “Cueva Iturbe / 3700m”, “jujuyensis Bernh. / Typus ded. Bruch” (yellow label), “Chicago NHMus. / M. Bernhauer Collection”, “Lectotype Philonthus jujuyensis Bernhauer, 1921 / Des. Chani-Posse de Maus 2008” (FMNH). Paralectotype: female, with labels: “Jujuy / 10.XI.919 / Weiser”, “Cueva Iturbe / 3700m”, “jujuyensis Bernh. / Cotypus ded. Bruch” (yellow label), “Chicago NHMus. / M. Bernhauer Collection”, “Paralectotype Philonthus jujuyensis Bernhauer, 1921 / Des. Chani-Posse de Maus 2008” (FMNH).

Note: Chani-Posse de Maus (2009: 33, 34) designated lectotype.

Additional material examined: 33 specimens (5 males, 6 females, 22 unsexed) ARGENTINA: Catamarca: Caspinchango, 10.III.1921 (2 MACN); La Cienaga, Walters, Philonthus jujuyensis Bernh., (1 MACN). Jujuy: Cueva Iturbe, 10.XI.919, Weiser, 3700m, Philonthus jujuyensis Bernh., (1 MLPA); Hornadita, 30.11.1909, Weiser, 3700m, in green “Coypus”, Philonthus jujuyensis Bernh. (5 MACN); V. 1920, C. Bruch, Chicago NHMus Bernhauer Collection (1 FMNH). La Rioja: Los Mistoles, XII-15-2000, L. Herman, 600m, 30° 48’ 04’’S, 66° 18’ 40’’W (1 male, 1 female AMNH). San Juan: S. of Astica, Chucuma, XI/IV/1995, L. Herman, 885m (1 AMNH). San Luis: Quebrada López, 7 km S San Francisco del Monte de Oro, L. Herman, 32° 39’ 37’’S, 66° 07’ 46’’W (1 AMNH). Tucuman: Villa Nougués, Dec. 11, 1971, L. Herman, 1350m (4 males, 4 females, 5 AMNH), (2 IADIZA); 30km, SE Tafi del Valle, nr. El Indio, XI/20/1995, L. Herman, 950-1050m (1 female, 2 AMNH).

Gabrius nidicola (Bernhauer, 1921) (Figs. 26-30, 39; Fig. 4 in Chani-Posse de Maus, 2009)
Philonthus nidicola “Bernhauer in litt.“; Bruch, 1915: 445 (cat.). Nomen nudum.
Philonthus nidicola 
Bernhauer, 1921: 108; Scheerpeltz, 1933: 1352   (cat.); Blackwelder, 1944: 134 (list); Herman, 2001: 2889 (cat.); Chani-Posse, 2004: 231 (list).
Gabrius  nidicola; Chani-Posse de Maus, 2009: 34 (new combination)

Diagnosis. Gabrius nidicola may be identified by the antennal segments 8-10 quadrate, and the distance separating punctures on elytra about equal to diameters of punctures. It differs from G. jujuyensis by the distance separating medial interocular punctures on frons twice as large as distance separating medial punctures from lateral punctures.

Redescription. Length of the body 4.3-5.3 mm. Coloration. Head black; thorax castaneous-brunneous; abdomen castaneous-piceous to castaneous-brunneous, with first three abdominal segments brunneous; elytra brunneous to testaceous-brunneous; antennae, palpi and legs testaceous-brunneous.
Head about as long as wide (HW/HL= 1.0) and at basal third as wide as distal third in both sexes, about as wide as to slightly wider than pronotum (HW/PW= 0.86-0.96); dorsal surface coarsely and moderately punctate, with three postocular punctures and four epicranial punctures; distance separating medial interocular punctures on frons about twice as large as distance separating medial punctures from lateral punctures; infraorbital ridge present, reaching postgenal ridge. Eyes distinctly shorter than temples (EL/TL= 0.60-0.56) seen from above. Antennae with segment 1 shorter than segments 2 and 3 combined, segments 2 and 3 subequal in length, segments 5-10 quadrate. Maxillary palpus with segment 2 no more than twice as long as its maximum width. Labial palpus with last segment twice as long as preceding segment. Neck no more than 0.5 times as wide as head at widest point (NW/HW= 0.5).
Pronotum longer than wide (PW/PL= 0.8), dorsal surface of pronotum with two rows of punctures, each with five punctures. Basisternum with medial longitudinal carina weakly developed. Metaventrite with circum-mesocoxal ridge broadly rounded medially. Elytra at sides distinctly shorter than pronotum at midline (EtL/PL=0.6-0.5), distinctly longer
than elytra along suture (EtL/Etl= 2.2–1.8); transverse distance between punctures about equal to diameters of punctures. Front tarsus as long as front tibia; middle tarsus longer than middle tibia; first segment of hind tarsus about as long as last segment (S1/S5= 1.2). Abdominal terga moderately punctuate basally, distance separating punctures about equal to diameters of punctures.
Male genitalia. Sternum 8 slightly emarginate medio-apically. Tergum 10 subtruncate medio-apically
Fig. 27), with two long subapical macrosetae and 3-4 short apical setae. Sternum 9 with basal portion asymmetrical, moderately emarginate apically with two long apical setae at each side and two subapical setae (Fig. 26). Aedeagus with median lobe gradually narrowed from base of paramere; apex of median lobe subacute (in dorsal view) and simple in its apical fourth (in lateral view) (Fig. 28).
Female genitalia. Tergum 10 similar to that of male
Fig. 29). Second gonocoxite with one basal strong seta and one long apical seta, without stylus (Fig. 30).

Geographical distribution. Gabrius nidicola is known from one biogeographical province of the Neotropical region (Pampa) and from Monte and Puna in the South American transition Zone (Fig. 39).

Bionomics. According to the labelling data from type material, this species was found in association with the rodent Ctenomys.

Type material. Lectotype, male, with labels: “Rep. ARGENTINA / Prov. Buenos Aires / 17.X.1919 / C. Bruch”, “Monte Veloz / 17.X.1919 / ex nido Ctenomys”, “nidicola Bernh. / Typus” (yellow label), “Chicago NHMus. / M. Bernhauer Collection”, “Lectotype Philonthus nidicola Bernhauer, 1921 / Des. Chani-Posse de Maus 2008” (FMNH). Paralectotype: female, with labels: “Rep. ARGENTINA / Prov. Buenos Aires / 17.X.1919 / C. Bruch”, “189”, “nidicola Bernh. / Cotypus” (yellow label), “Chicago NHMus. / M. Bernhauer Collection”, “Paralectotype Philonthus nidicola Bernhauer, 1921 / Des. Chani-Posse de Maus 2008” (FMNH).

Note: Chani-Posse de Maus (2009: 34) designated lectotype.

Additional material examined: 14 specimens (2 males, 2 females, 10 unsexed) ARGENTINA: Buenos Aires: 17.X.1919/ 30.XII.1922, C. Bruch (3, 2 MACN). Jujuy: Cueva Iturbe, 10.XI.919, Weiser (1 MACN). La Rioja: Los Mistoles, XII-15-2000, L. Herman (1 male, 1 female, 2 AMNH). San Luis: 7km, N Trapiche, XII-6-2000, L. Herman, 1060m, 33° 06’ 03’’S, 66° 03’ 39’’W, litter near river (1 male, 1 female, 2 AMNH).

Gabrius nigritulus (Gravenhorst, 1802) (Figs. 12, 36-38, 39 in this paper; Figs. 1001-1007 in Smetana, 1995; Fig. 64 in Schillhammer, 1997)
Staphylinus  nigritulus  Gravenhorst,  1802: 41. For complete references see Herman (2001).
Gabrius chiliensis Coiffait & Sáiz, 1968: 348; Sáiz, 1971: 340; Chani-Posse de Maus, 2009: 40 (syn).

Diagnosis. This species can be distinguished from the other species of southern South American Gabrius by having the dorsal rows on pronotum each with six punctures. Additionally, the male differs by the sternum 9 with the basal portion more or less symmetrical (Fig. 36), the tergum 10 without apical setae and the paramere of aedeagus bifurcate, with sensory peg setae (Fig. 37, 38). The female differs by the tergum 10 with the apex rounded and two apical setae.

Redescription. See Smetana (1995: 619).

Geographical distribution. Gabrius nigritulus is assumed to have been introduced to various regions of the world from the western portion of the Palearctic region, and it is now considered a cosmopolitan species (Smetana 1995). In southern South America G. nigritulus is known from several biogeographical provinces of the Andean region (Fig. 39).

Bionomics. According to Smetana (1995), Gabrius nigritulus may occur in a wide range of habitats with decaying organic matter. According to collecting data, it may be found in Nothofagus and palm forests, cacti, sclerophyl woodland, dump mud and moss on sand.

Type material. Lectotype of Gabrius nigritulus, male, with labels: “6139”, “Lectotype Gabrius nigritulus Gravenhorst A. Smetana des. 1960”, “Hist.-Coll. (Coleoptera) / Nr. 6139 / Philonthus aterrimus Gyllh. / Europa, Americ. sept. / Zool. Mus. Berlin” (ZMHB). Paralectotypes: 3 males, with labels: “Gabrius Steph. / nigritulus Grav. / det. Smetana 1960”, “Hist.–Coll. (Coleoptera) / Nr. 6139 / Philonthus aterrimus Gyllh. / Europa, Americ. sept. / Zool. Mus. Berlin” (ZMHB).

Note: The lectotype was designated by Smetana in 1960 (see labels on type material) and published two years later (Smetana, 1962).
Paratypes of Gabrius chiliensis, male, with labels: “Cord. Chillan”, “Germain / 1899”, “PARATYPE”, “Gabrius chiliensis Coiff. et Saiz”, “Chile Tipo No 2238”; female, with labels: “Germain / 1899”, “PARATYPE”, “Gabrius chiliensis Coiff. et Saiz”, “Chile Tipo No 2245” (MNNC).

Additional material examined: 16 specimens (4 males, 10 females, 2 unsexed) ARGENTINA: Rio Negro: Villa Regina, Feb. 14, 1972, L. Herman (1 male, 1 female AMNH). Chubut: El Hoyo, 10.IX.61 (1 female NMW). CHILE: Curicó: 20km E. Potrero Grande, El Relvo, 08-May-04, 1100m, 36° 11.14’ S, 70° 56.1’ W, fogging Nothophagus dombeyi (2 females, 2 JEBC). Concepción: Valle Nonguen, January 15, 1978, T. Cekalovic (1 male AMNH). Quillota: P.N. La Campana (Sector Ocoa) vic Quebrada Buitrera, 28.XII.2008, Thayer, Clarke, 415m, 32° 55.89’ S, 71° 05.10’ W, sclerophyl woodland, Jubaea chilensis palms, Trichocereus cacti; dump mud & moss on sand, dry creek bed (1 male, 1 female FMNH). Valparaíso: Viña del Mar, 28.12.94, Germain, Gabrius chiliensis Coiff. et Saiz F. Saiz, det. 1970 (1 male, 1 female MNNC). Cachapoal: Las Cabras, 10/23. XII.1954, Peña, 1480m, L. Peña Leg., Gabrius chiliensis Coiffait & Saiz Det. Chani-Posse 2006 (1 female FMNH). Bío Bío: Recinto, I.1953, Peña, L. Peña Leg. (1 female FMNH). Maule: El Radal, 28/30.XI.1957, Peña, 900m, L. Peña Leg. (1 female FMNH).

Gabrius tucumanensis (Bernhauer, 1927) (Figs. 10, 11, 31-35, 39; Figs. 3 a–b in Chani-Posse de Maus, 2009)
Philonthus tucumanensis Bernhauer, 1927: 245; Scheerpeltz, 1933: 1365 (cat.); Blackwelder, 1944: 135 (list); Herman, 2001: 2982 (cat.); Chani-Posse, 2004: 231 (list).
Gabrius tucumanensis
; Chani-Posse de Maus, 2009: 33, 34 (new combination)

Diagnosis. Gabrius tucumanensis may be identified by the antennal segments 9-10 transverse, the distance between punctures on elytra distinctly larger than diameters of punctures and the distance separating medial interocular punctures on frons twice to three times as large as distance separating medial punctures from lateral punctures. It differs from G. hornaditanus and G. jujuyensis by the head longer than wide, and the antennal segment 5 elongate.

Redescription. Length of the body 4.5-5.1 mm. Coloration. Head black; thorax castaneous; abdomen castaneous-piceous; elytra castaneous brunneous to brunneous; scutellum black; antennae, palpi and legs brunneous.
Head longer than wide (HW/HL= 0.8) (Fig. 10) and at basal third as wide as distal third in both sexes, about as wide as to slightly wider than pronotum (HW/PW= 1.0-1.1); dorsal surface coarsely and moderately punctate, with three postocular punctures and four epicranial punctures; distance separating medial interocular punctures on frons twice to three times as large as
distance separating medial punctures from lateral punctures; infraorbital ridge present, reaching postgenal ridge. Eyes distinctly shorter than temples (EL/TL= 0.5-0.4) seen from above (Fig. 10). Antennae with segment 1 shorter than segments 2 and 3 combined, segment 2 shorter than segment 3, segment 5 elongate, segments 6-8 quadrate and 9-10 slightly transverse (Fig. 10). Maxillary palpus with segment 2 no more than twice as long as its maximum width. Labial palpus with last segment twice as long as preceding segment. Neck no more than 0.5 times as wide as head at widest point (NW/HW= 0.5) (Fig. 10).
Pronotum longer than wide (PW/PL= 0.9-0.8) (Fig. 10), dorsal surface of pronotum with two rows of punctures, each with five punctures. Basisternum with medial longitudinal carina weakly developed. Metaventrite with circum-mesocoxal ridge narrowly rounded medially. Elytra at sides as long as to moderately longer than pronotum at midline (EtL/PL=0.7), distinctly longer than elytra along suture (EtL/Etl= 1.9-1.7); transverse distance between punctures distinctly larger than diameters of punctures (Fig. 11). Front tarsus shorter than front tibia; middle tarsus as long as middle tibia; first segment of hind tarsus as long as last segment (S1/S5= 1.0). Abdominal terga punctuate basally, distance separating punctures distinctly larger than diameters of punctures.
Male genitalia. Sternum 8 slightly emarginate medio-apically (Fig. 31). Tergum 10 subtruncate medio-apically, with two long subapical macrosetae and 3-4 short apical setae. Sternum 9 similar to that of P. hornaditanus, with 5-6 long apical setae and two subapical setae. Aedeagus with median lobe gradually narrowed from base of paramere; apex of median lobe arcuate (in dorsal view) and simple in its apical fourth (in lateral view) (Figs. 33, 34).
Female genitalia. Tergum 10 emarginate medio-apically, with two long subapical macrosetae and 5-6 short apical setae. (Fig. 32). Second gonocoxite with one basal strong seta, and stylus with two apical setae (Fig. 35).

Geographical distribution. Gabrius tucumanensis is known from three biogeographical provinces of the Neotropical region (Chaco, Pampa and Yungas) and from Monte and Puna in the South American transition Zone (Fig. 39).

Bionomics. According to the labelling data, this species may be found in litter near streams and rivers.

Type material. Lectotype, female, “San Pedro Colalao / Tucum. Weiser”, “tucumanensis Bernh. / Typus don. Bruch” (yellow label), “Chicago NHMus. / M. Bernhauer Collection”, “Lectotype Philonthus tucumanensis Bernhauer, 1927 / Des. Chani-Posse de Maus 2008” (FMNH).
Note: Chani-Posse de Maus (2009: 33, 34) designated lectotype.

Additional material examined: 50 specimens (29 males, 8 females, 13 unsexed). ARGENTINA: San Luis: Irideus, Quebrada del Tala, 1100 m, 32°39’01’’S, 64°57’23’’W, litter in moist narrow quebrada, 8.XII.2000, L. Herman (25 males, 4 females AMNH); Quebrada López, 7 km S. San Francisco del Monte de Oro, 3 males, 3 females, 4; 7km, N Trapiche, XII-6-2000, L. Herman, 1060m, 33° 06’ 03’’S, 66° 03’ 39’’W, litter near river (4 AMNH). Tucumán: Villa Nougués, 1350 m, 11.XII.1971, L. Herman (1 male, 1 female AMNH). La Rioja: Los Corrales, 15 km NW Famatina, Dec. 3, 1971, Lee Herman (2 AMNH). Salta: 10km. NW Cafayate, Dec. 17, 1971, Lee Herman (1 AMNH). Jujuy: Cueva Iturbe, 10.XI.919, Weiser (2 NMW).

ACKNOWLEDGEMENTS

I gratefully acknowledge the curators listed in Material and methods for the loan of specimens. I am very indebted to Gustavo E. Flores for valuable comments that improved different aspects of this work. I also thank Christian Maus for his support during this study. This study was supported by the Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Argentina, and the following grants: Ernst Mayr Travel Grant (MCZ, Harvard University), Austral Staphylinidae PEET grant 0118749 to Margaret K. Thayer and Alfred F. Newton, FMNH-Collection Study Grant, FONCYT-PICT 01-11120 to Jorge Miguel Lobo and Sergio Roig, Foundation Grant BBVA as part of the project: “Design of a reserve network for protection of biodiversity in Southern South America using predictive distribution models with hyperdiverse taxa”, CONICET PIP 5766 to A. Marvaldi and CONICET PIP 112-200801-00162 to A. E. Marvaldi and G. E. Flores.

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