Phylogenetic systematics of the genera Plochionocerus Dejean and Agrodes Nordmann (Coleoptera: Staphylinidae: Xantholinini) Author Asiain, Julieta Author Márquez, Juan Author Morrone, Juan J. text Zootaxa 2007 1584 1 53 journal article 10.5281/zenodo.178410 ba2b7965-13f3-401f-9c5a-1705938112ea 1175-5326 178410 Plochionocerus simplicicollis ( C. Waterhouse, 1879 ) ( Fig. 1 ) Sterculia simplicicollis C. Waterhouse, 1879 : 421 ; Herman, 2001 : 3746 ( Plochionocerus ). Sterculia basalis Sharp, 1885 : 469 ; Herman, 2001 : 3744 ( Plochionocerus ), syn. nov. Sterculia mandibularis Sharp, 1885 : 468 ; Herman, 2001 : 3745 ( Plochionocerus ), syn. nov. Sterculia pollens Sharp, 1885 : 469 ; Herman, 2001 : 3746 ( Plochionocerus ), syn. nov. Araeocnemus fulgens : Nordmann, 1837 : 165 ; Bernahuer & Schubert, 1914 : 315 (misidentification, non Fabricius, 1793 ). Sterculia nordmanni Fauvel, 1901 : 252 (replacement name for S. fulgens sensu Nordmann, 1837 ); Herman, 2001 : 3746 ( Plochionocerus ), syn. nov. Type material ( 9 specimens ). Lectotype of Sterculia simplicicollis , male: “Medellin 78.39 / Type / Sterculia simplicicollis ( Type ) C. Waterh. / Holotype / Lectotype Sterculia simplicicollis Asiain, Márquez & Morrone des. 2007” ( BMNH ). Lectotype of Sterculia basalis by present designation, male: “ Sterculia basalis . Type . Mexico . Coll. Guerin Meneville. (specimen on card) / Type / Mexico / Sharp Coll. 1905–313. / B. C. A. Col. I. 2. Sterculia basalis , Sharp. / Type / Lectotype Sterculia basalis Asiain, Márquez & Morrone des. 2007” ( BMNH ). Paralectotype , male: ” Sterculia basalis var. Jalapa. Mexico . Höge (specimen on card) / Jalapa Mexico . Hoege / B. C. A. Col. I. 2. Sterculia basalis Sharp / Paralectotype Sterculia basalis Asiain, Márquez & Morrone des. 2007”. ( BMNH ). Lectotype of Sterculia mandibularis by present designation, male: “ Sterculia mandibularis Type D. S. Jalapa. Mexico . Höge (specimen on card) / Type / Jalapa, Mexico . Hoege / B. C. A. Col. I. 2. Sterculia mandibularis , Sharp. / Sharp Coll. 1905–313. / Syntype / Lectotype Sterculia mandibularis Asiain, Márquez & Morrone des. 2007” ( BMNH ). Paralectotypes , males: “ Sterculia mandibularis D. S. Chontales. Nicaragua . Janson (specimen on card) / B. C. A. Col. I. 2. Sterculia mandibularis , Sharp. / Syntype / Paralectotype Sterculia mandibularis Asiain, Márquez & Morrone des. 2007” ( BMNH ); “ Sterculia mandibularis D.S. / Jalapa. Mexico . Höge (specimen on card) / B. C. A. Col. I. 2. Sterculia mandibularis , Sharp. / Syntype / Paralectotype Sterculia mandibularis Asiain, Márquez & Morrone des. 2007” ( BMNH ). Lectotype of Sterculia pollens by present designation, male: “ Sterculia pollens . Type D. S. Cache Costa Rica . Rogers (specimen on card) / Type / Cache Costa Rica . H. Rogers / B. C. A. Col. I. 2. Sterculia pollens , Sharp. / Sharp. Coll. 1905–313 / Syntype / Lectotype Sterculia pollens Asiain, Márquez & Morrone des. 2007” ( BMNH ); Paralectotypes , males: “ Sterculia pollens . Chontales. Nicaragua . Janson (specimen on card) / Chontales, Nicaragua . Janson. / B. C. A I. 2. Sterculia pollens , Sharp / Syntype / Paralectotype Sterculia pollens Asiain, Márquez & Morrone des. 2007” ( BMNH ); “Coban Vera Paz. Champion / B. C. A. Col. I. 2. Sterculia pollens var. Guatemala . Champion / Syntype / Paralectotype Sterculia pollens Asiain, Márquez & Morrone des. 2007” ( BMNH ). Additional material ( 123 specimens ). COLOMBIA : Santa Fé de Bogota, Dunckier ( FMNH , 2ɗɗ); Muzo ( FMNH , ɗ); Santa Fé de Bogota ( IRSNB , 2ɗɗ); no locality data ( IRSNB , 2ɗɗ,1Ψ); Guayaquil, Caesar Urban ded., 14.II.1898 ( IRSNB , ɗ); State of Darien, 1909, H. Godge ( BMNH , ɗ); no locality data, Amazones ( IRSNB , ɗ). COSTA RICA : Prov. Puntarenas, San Luis Monteverde, A.C. Arenal, LN 250850 449250, 1004– 1350 m , VII.1993 , Z. Fuentes ( INBC , 2ɗ); idem , except: 900 m ( INBC , ɗ); Prov. Puntarenas, San Luis, R. B. Monteverde, LN 250850 , 440850, 449250, 1040 m, 7.VIII.1992 , Z. Fuentes ( INBC , ɗ); idem , except: VIII.1992 ( INBC , Ψ); idem , except: VII.1992 ( INBC , ɗ); idem , except: 1000–1350 m , IX.1993 ( INBC , ɗ); Prov. Guanacaste, Estac. Pitilla, 9 km S Santa Cecilia, 330200, 380200, 700 m , malaise trap, 1988, GNP Blod. Sur ( INBC , Ψ); idem , except: VI.1989 , GNP Biodiversity Survey ( INBC , 3ɗɗ); idem , except: Alajuela Prov., VII.1988 , Espinoza & Chavez ( INBC , ɗ); idem , except: IX.1989 , C. Moraga & P. Ríos ( INBC , ɗ); Prov. Heredia, Sardinalito, Braulio Carrillo N. P., 257500, 531200, 360– 500 m , X.1989 , R. Aguilar & M. Zumbado ( INBC , 2ɗɗ); Prov. Limón, Manzanillo, RNFS Gandoca y Manzanillo, LS 398100, 610600, 0–100 m , 4– 12.XII.1992 , F. Quesada ( INBC , ɗ); Prov. San José, Est. Zurqui, 500 m antes de Tunel, LN 226800, 535200, 1600 m , IV.1991 , G. Maass ( INBC , ɗ); Prov. Limón, Reserva Biol. Hitoy Cerere, Estación Hitoy Cerere, Send. Espavel, LS 401558 570460, 220 m , 12–26.VI.1999 , W. Arana, Manual (red libre) ( INBC , ɗ); idem , except: Sendero Espavel, LS 570200 401500, 300 m , 24–30.V.2000 , intersección ( INBC , Ψ); idem , except: 220 m , 13–17.V.1999 , Manual (red libre) ( INBC , ɗ); idem , except: Valle La Estrella, LN 643400 184600, 100– 200 m , 4–20.IV.1994 , G. Carballo ( INBC , ɗ); Prov. Limón, Talamanca, San Miguel, Cabecur, LS 397900 576150, 325 m , 20.II.1997 , I. A. Chacon, Manual ( INBC , Ψ); Prov. Guanacaste, Est. Cacao, lado SO Vol. Cacao, P. N. Guanacaste, LS 323300 375700, 800– 1600 m , 12–17.VII.1993 , M. Reyes ( INBC , ɗ); idem , except: M. A. Zumbado ( INBC , ɗ); Prov. Limón, Sector Cerro Cocorí, Fca. de E. Rojas, LN 286000, 567500, 150 m , VI.1991 , E. Rojas ( INBC , ɗ); idem , except: V.1991 ( INBC , ɗ); idem , except: 31.I–21.II.1992 ( INBC , ɗ); Prov. Alajuela, Upala, Alb. Heliconias, Send Heliconias, LN 422600 299100, 700 m , 23.VI–2.VII.2000 , A. López, intersección ( INBC , ɗ); idem , except: P. N. Volcán Tenorio, 17–28.VI.2001 ( INBC , Ψ); Prov. Alajuela, Sector Tucanes, LN 269600 457600, 640 m , 14.X–3.XII.1998 , G. Caballo, Malaise ( INBC , ɗ); Puntarenas, Est. Biol. Las Alturas, Coto Brus, LS 322500, 591300, 1500 m , X.1991 , M. Ramírez ( INBC , ɗ); Va ra Blanca ( FMNH , ɗ); Vara Blanca, 2000 m , III.1936 (CC-UAEH, ɗ); Zapote (Mayalg), VIII.1939 ( FMNH , ɗ); San Pedro de Montes de Oca, XI.1939 ( FMNH , ɗ); San José, 11.XI.1937 , Nevermann, S. J. 2.XI.1937 ( FMNH , ɗ); Tapantí, IV.1941 , Guzmán ( FMNH , ɗ); San José, Desamparado ( FMNH , Ψ); no locality data, F. Nevermann ( FMNH , 1?); no locality data, C. H. Lankester ( BMNH , ɗ); Coronado, 1400–1500 m , T. Assmann & F. Nevermann ( BMNH , Ψ); Heredia, La Selva Res. Sta., 11–17.VI.1986 , W. Hanson & G. Bohart ( SEMC , ɗ); Heredia, Sto. Domingo de Heredia, INBIo, Cafetal, 1100 m , 25–28.VI.1997 , S. & J. Peck, ex: flight intercept trap ( SEMC , ɗ); Guanacaste, S. E. slope volcan Miravalles, nr. Rio Naranjo, Finca Agropecuaria Rio Naranjo, 28.III.1973 , cloud forest, Dr. Whitehead ( USNM , ɗ). EL SALVADOR : Depto. Ahuachapán, San Benito, El Imposible (P. Nac. El Imposible), 13°49’09”N ; 89°56’34”W , 22.VI. 1997 , 700 m, J. Davis ( MNHDS , ɗ); Depto. La Libertad, Los Chorros, 800 m , 9.II.1973 , V. Hellebuyck ( MNHDS , Ψ). GUA- TEMALA: no locality data, C. C. Hoffmann ( CNIN , ɗ); Chimaltenango, Conradt ( BMNH , ɗ); no locality data ( IRSNB , Ψ). HONDURAS : Olancho, La Unión, 14 km N, P. N. La Muralla, 1500 m , 16.VIII.1994 , wet, montane forest, flight intercept trap, S. & J. Peck ( FMNH , ɗ). MEXICO : Jalapa, Höge ( BMNH , 2ɗɗ, 3ΨΨ); Veracruz, Jalapa, Höge ( BMNH , Ψ, 3?; FMNH , 2ɗɗ); Cordova, Salle ( BMNH , ɗ); Cordova, Sallé ( BMNH , ɗ); no locality data, Flohr ( BMNH , 2ɗɗ); Durango, Villa Lerdo, Höge ( BMNH , Ψ); no locality data, Sharp Coll. ex Chevrolat ( BMNH , Ψ); Coatepec, J. Quiroz ( BMNH , ɗ); Veracruz, 7.1 km E Huatusco, 1230 m , 16.VII.1990 , R. L. Minckley, ex., Compositae ( SEMC , ɗ); Veracruz, 7 km E Huatusco, Hwy. 125, 16.VII.1990 , 1230 m, J. Ashe, K. J. Ahn & R. Leschen, ex: general collecting ( SEMC , Ψ); Veracruz, 2.3 km S Jalapa, 13.VII.1992 , 1320 m, J. S. Ashe, ex: under logs ( SEMC , 2ɗɗ); Veracruz, Coatepec, 25.X.1995 , en la banqueta, R. Arce ( IEXA , ɗ); Veracruz, Ixtacaoquitlán, Cuautlapan, Cerro Cementos, 1300 m , 14– 20.VIII.1996 , A. Hernández, M. Torres & L. Delgado ( IEXA , ɗ); Ver., Xalapa, 17.VIII.1978 , J. Peña M. ( IEXA , Ψ); Veracruz, Xalapa, 15.VIII.1996 , al vuelo, R. Arce ( IEXA , 1?); Veracruz, Banderilla, La Martinica , 1500 m , 31.VII.2004 , bosque mesófilo, volando a las 13:00 hrs., A. y L. Delgado ( IEXA , ɗ); Veracruz, Xalapa, Jardín Botánico, 7.VII.2005 , bosque mesófilo de montaña perturbado, Q. Santiago ( QJSJ , ɗ); Veracruz, Sontecomapan, Los Tuxtlas. SAP, 18°35’06”N , 95°04’30”W , ex. bajo hojarasca, 22.VIII.2000 , G. Ball ( QJSJ , Ψ); Veracruz, Coatepec, La Pitaya, selva mediana subcaducifolia perturbada, 1245 m , 19°29’28”N , 96°56’59”W , en suelo, 12.VIII.2000 , J. Asiain, Q. Santiago & J. Márquez ( MZFC , Ψ); Ver., Xalapa, P. F. Clavijero, 5.VIII.1981 , R. Ayala ( MZFC , 2ɗɗ); Veracruz, Cascadas de Xico, bosque mesófilo de montaña y cultivo de temporal, ex. suelo, 4–5.VIII. 1996 , Q. Santiago & J. Márquez ( MZFC , Ψ); Istmo, IX.1986 ( CNIN , ɗ); Ver., ca. Xalapa, 12.VIII.2004 , E. Maya, ex basurero, cerca borde arroyo ( CZUG , ɗ); Ver., Huatusco, 19.VII.1993 , bosque mesófilo de montaña, hojarasca, J. Márquez (JLN, ɗ); Orizaba ( IRSNB , 1?); Veracruz, Cayon Metlac, near Fortín, 3200 ft , 28.VII–1.VIII.1973 , walking in path, A. Newton ( FMNH , ɗ); Veracruz, 2 mi Catemaco, 15.II.1976 , F. Fisk ( FMNH , Ψ); Sallé, Cordova ( FMNH , ɗ); Cintalapa, La Pilar, 25.VIII.2001 , C. J. Morales ( CZUG , ɗ); Oaxaca ( IRSNB , Ψ); Oaxaca, Soyalapa, VI.64 ( CNIN , ɗ); Chiapas, El Sumidero Park, Mirador La Coyola, 750 m , VII.1991 , D. B. Thomas, ex: flight intercept ( SEMC , Ψ); Chiapas, Ocosingo, 1200 m , zona montañosa bosque primitivo, 6.IX.1947 , M. del Toro ( CNIN , Ψ); no locality data ( AMNH , ɗ; BMNH , ɗ; FMNH , Ψ; IRSNB , Ψ, 1?). NICARAGUA : Chontales, Janson ( FMNH , 2ɗɗ). PANAMA : Panama , Cerro Campana, nr. Capira, 790 m , 08°44’N , 79°57’W , 8.VII.1995 , A. Gillogly ( SEMC , 3ɗɗ); idem , except: 30.V.1995 , N. Upton, ex: crawling on ground ( SEMC , ɗ); idem , except: 1–5.VI.1995 , J. Ashe & R. Brooks, ex: flight intercep trap ( SEMC , Ψ); idem , except: 820 m , 8°40’N , 79°56’W , 9.VI.1984 , Stockwell ( FMNH , ɗ); Cerro Jefe (Cerro Azul), 09°19’N , 79°25’W , 600–870 m , 28.V.1995 , J. & A. Ashe ( SEMC , ɗ); Chiriquí, 5.6 km N Boquete, La Culebra Trail, 1800 m , 8°49’23”, 82°25’18”W , 15.VI.1996 , R. Anderson, ex: cloud forest litter ( SEMC , ɗ); no locality data, Cent. Am. ( AMNH , 2ɗɗ). Diagnosis. Total length 18.0– 25.2 mm ; metallic violet or green blue, or head and thorax violet-blue and abdomen green; head varied in shape (rectangular to rounded with intermediate shapes), 1.13–1.31 times as long as wide, slightly longer than pronotum (1.13–1.35 times), dorsal and ventral surface of head varied in convexity, from flat to convex, ventral surface with dense, expanded, umbilicate punctures ( 20–29 in each half of head), arranged in “v” ( Fig. 50 ); apical antennomere in males moderately oblong (length/width: 1.36–1.54) and almost as long as antennomeres 9+10 combined (0.83–1.00 times); mandibles with slightly visible, basal external channel; pronotum 1.36–1.65 times as long as wide; shorter than elytra (0.67–0.86 times its length), with two clearly visible depressed areas in posterior third ( Fig. 56 ); prosternum transverse, slightly transverse to as long as wide (length/width: 0.61–0.97); aedeagus ovate, with parameres moderately long (0.35–0.47 times as long as median lobe), apical area of median lobe 0.30–0.41 times as long as the total length of median lobe, and internal sac moderately visible ( Fig. 91 ). Variation. Specimens from Mexico , Guatemala and El Salvador are smaller than specimens from Honduras , Nicaragua , Costa Rica , Panama and Colombia . The length of the body corresponds to the length of the aedeagus. The head of larger specimens is more convex and rounded and the mandibles larger (mandible length 2.15–3.90 mm ); the head of smaller specimens is slightly convex and less rounded, and the mandibles are shorter. The body color may be violet-blue, metallic green or combinations of blue, green and violet. The mandibular channel can be slightly visible or reduced to 2–3 big punctures. Comparative notes. Based on the revision of the type specimens (except for S. nordmanni ), P. b a s a l i s ( Sharp, 1885 ), P. mandibularis ( Sharp, 1885 ) , P. p o l l e n s ( Sharp, 1885 ) and P. nordmanni ( Fauvel, 1901 ) are junior synonymies of P. simplicicollis ( Waterhouse, 1879 ) . Sharp (1885) noted that P. basalis presents a rugosity at the base of the elytra, that was slightly visible in few specimens that we analyzed. Additionally, he mentioned that the mandibles and antennae of P. mandibularis and P. basalis were similar, but different in color; now we know that color is not a character useful to distinguish species of the genus. The long mandibles of P. mandibularis are not a reliable character, because they are also long in P. pollens . In the specimens analyzed, the length of the mandibles, color, size and convexity of head were very variable, but the body, mandibles and aedeagus of the specimens from Mexico and Guatemala are smaller than specimens distributed in the south. When Fauvel (1901) described P. nordmanni , he characterized it by the shorter mandibles and smaller body length than P. fulgens , character states that are within the range of intraspecific variation of P. FIGURES 1–6 . Habitus, dorsal view: 1, Plochionocerus simplicicollis ; 2, P. s p l e n d e n s ; 3, Agrodes elegans ; 4, Renda fimetaria ; 5, Philonthus testaceipennis ; 6, Homalolinus flavipennis . Scale bar 3 mm. simplicicollis . Also, the labels of several specimens identified as P . fulgens , P . basalis , P . nordmanni or P . pollens suggest that they have been considered as variation (“var”) since their original description. Geographical distribution. El Salvador , Honduras , and Panama (first national records), Mexico , Guatemala , Nicaragua , Costa Rica , and Colombia ( Herman 2001 ).