Large ants are not easy - the taxonomy of Dinoponera Roger (Hymenoptera: Formicidae: Ponerinae) Author Dias, Amanda Martins D8E29645-E8DC-419D-ADFB-7ECA76541FCD Universidade Federal do Paraná, Departamento de Zoologia, Curitiba, Paraná, Brazil. amandamdbio@gmail.com Author Lattke, John Edwin 7AE0D94C-619B-4E77-B16F-3F18731EE390 Universidade Federal do Paraná, Departamento de Zoologia, Curitiba, Paraná, Brazil. lattke@ufpr.br text European Journal of Taxonomy 2021 2021-12-21 784 1 1 66 http://dx.doi.org/10.5852/ejt.2021.784.1603 journal article 2866 10.5852/ejt.2021.784.1603 8ddfc41f-28aa-48b4-a24d-25ba06447ef0 2118-9773 5798246 80B6E154-A9A3-49E3-AAF0-3FD2BEBF82D2 Dinoponera quadriceps Kempf, 1971 Figs 25–26 , 29B Dinoponera quadriceps Kempf, 1971: 380 , figs 5, 13–16 ( ☿ ). Dinoponera opaca Kempf, 1971: 379 ( ☿ ). Dinoponera quadriceps – Mann 1916: 409 . — Borgmeier 1937: 226 (male description). — Lenhart, Dash & Mackay 2013: 149 , figs 4b, g, l, 5a, 6a, 10a, 11a, 12b (redescription, key, male genitals). — Tozetto & Lattke 2020: 5 , figs 2–3, 6 (male genitals). Dinoponera opaca – Kempf 1975: 344 (junior synonym of D. quadriceps ). Non Dinoponera grandis mutica quadriceps – Santschi 1921: 84 (invalid name). Non Dinoponera grandis mutica opaca – Santschi 1921: 84 (invalid name). Diagnosis Female Malar area without striae or with weak longitudinal to oblique striae that do not reach anterior eye margin. Ventral head surface without strigulae. Head with short to inconspicuous pubescence. Anteroventral corner of pronotum in lateral view forms an obtuse angle. Abdominal tergite III microareolate and opaque, with sparse pubescence on dorsum. Male Antenna with long suberect to erect hairs, clearly longer than maximum scape diameter. Mesoscutum without notaulus. Gastral tergites without long hairs. In lateral view, dorsal margin of basal ring broadly concave to straight. Material examined Non-type specimens ( 200 ☿☿ , 17 ♂♂ ) BRAZIL – Bahia • 1 ☿ ; Aramari , ex Distrito de Alagoinhas ; MZSP • 1 ☿ ; Banzaê ; 10°34′37″ S , 38°36′54″ W ; Mar. 2009 ; M.D. Dantas leg.; 01; CPDC • 1 ☿ ; Barragem de Anagé [Amajés]; 19 Oct. 1996 ; Argôro Santos leg.; 5126; CPDC • 1 ☿ ; Barreiras , Serra do Mimo ; 5 Aug. 2010 ; S. Souza and B. Santos leg.; #5657; CPDC • 1 ☿ ; Bendegó , próx. Canudos ; 19–22 Nov. 1965 ; Exp. DZ Fapesp leg.; MZSP • 2 ☿☿ ; Lapa [ Bom Jesus da Lapa ]; 1 Dec. 1948 ; Gonçalves leg.; MZSP • 1 ☿ ; Bonfim ; 6 Dec. 1948 ; C.R. Gonçalves leg.; “Let OT Borgmeier set 1953 ”; MZSP • 7 ☿☿ ; Buritirama ; 22 Mar. 1958 ; E. Dente leg.; MZSP • 1 ☿ ; Caatinga do Moura ; 24 Jan. 1980 ; M.T.U. Rodrigues leg.; MZSP • 1 ☿ ; same locality as for preceding; 24–29 Jan. 1980 ; P.E. Vanzolini leg.; MZSP • 2 ☿☿ ; Caculé ; Jul. 1961 ; S. Laroca leg.; DZUP • 1 ☿ ; Caetité ; Nov. 2009 ; Jaqueline leg.; 8593; CPDC • 1 ☿ ; Camisão ; 8 Apr. 1905 ; E. Bondas leg.; n 3401 Coll Borgmeier ; MZSP • 1 ☿ ; Campo Formoso ; 20 Apr. 1974 ; R.L. Araújo leg.; n5534; MZSP • 1 ☿ ; same locality as for preceding; Jun. 1995 ; C. Moura leg.; 4940; CPDC • 1 ☿ ; Contendas do Sincorá ; 13°55′20″ S , 41°06′57″ W ; alt. 375 m ; 3 Jun. 2014 ; Santos Silva leg.; UFGD • 5 ☿☿ ; Contendas do Sincorá , F.N.C. Sincorá ; 14°00′ S , 41°10′ W ; 3 Jun. 2014 ; G. Santos Silva leg.; DZUP • 2 ☿☿ ; Cruz das Almas ; 1 Apr. 1988 ; C.A.L. Carvalho leg.; 704 188 (2/2) ME 014; MZSP • 2 ☿☿ ; Feira de Santana ; 23 Feb. 1993 ; J. Ricardo leg.; 4611; CPDC • 1 ☿ ; same locality as for preceding; 19 Jun. 2006 ; J.J. Resende leg.; 11, ninho 1 F07 ; CPDC • 1 ☿ ; Feira de Santana , UEFS ; 30 Nov. 1993 ; I.M.F. Soares leg.; 4899; CPDC • 1 ☿ ; Guanambi , EAFG ; Apr. 1997 ; A.J.S. Argolo leg.; CPDC • 3 ☿☿ ; Ibiraba ; 1 Mar. 1989 ; Pedro Rocha leg.; “ noite , ninho pé arbusto ”; MZSP • 1 ☿ ; Ibotirama ; 21 Jul. 1969 ; W. Silva leg.; MZSP • 1 ☿ ; Igaporã ; 9 Nov. 2011 ; J.A. Jesus leg.; CPDC • 1 ☿ ; Itaberaba ; 19 Aug. 2008 ; Antônio Alves leg.; CPDC • 1 ☿ ; Itaberaba , Amparo Br28 ; 16 May 1965 ; R. Grantsan leg.; MZSP • 4 ☿☿ ; Itaberaba , Faz. Riacho do Uruçu 1; 5 Dec. 1990 ; C.R.F. Brandão , J.L.M. Diniz and O.S. Oliveira leg.; 9 pm; MZSP • 3 ☿☿ ; Itaberaba , Faz. Riacho do Uruçu ; 1 Dec. 1990 ; S.T.P. Amarante leg.; “ bandeja d’água ”; MZSP • 1 ☿ ; Itatim ; 6 Jul. 1996 ; G.M. Santos leg.; 5105b; CPDC • 1 ☿ ; Itauçu , FLONA Contendas do Sincorá ; 9 Sep. 2005 ; Daniela and Péricles leg.; TR pt 14; CPDC • 1 ☿ ; Jandaíra , Costa Azul ; 11°56′58″ S , 37°36′34″ W ; 6 Oct. 2010 ; M.L.O. Travassos leg.; H1430; CPDC • 1 ☿ ; Juazeiro ; 11 Jul. 1989 ; M. Cristiano leg.; CPDC • 7 ☿☿ ; Lagoa Itaparica ; 8 Oct. 1990 ; Jorge L.M. Diniz leg.; coleção Diniz; DZUP • 1 ☿ ; Marcionílio Souza ; 24 Jul. 1993 ; N.S. Laca leg.; 4664; CPDC • 1 ♂ ; Manoel Vitorino ; 26–27 Jan. 2006 ; R.R. Hora leg.; CPDC • 1 ☿ ; Milagres ; 12°54′54.2″ S , 39°51′27.9″ W ; 22–26 Oct. 2010 ; M.A. Ulyssea , A.M. Medina and E.M. Campos leg.; T1F 3H1; MZSP • 1 ☿ ; same collection data as for preceding; T2F 2H5; MZSP • 1 ☿ ; same collection data as for preceding; T3F 4H4; MZSP • 1 ☿ ; same collection data as for preceding; 23–25 Oct. 2010 ; M.A. Ulysséia leg.; MuBio-CE Hym 507; UFGD • 1 ☿ ; Milagres , Br 116 km 561 , caatinga arbórea ; 12°52′12″ S , 39°11′31″ W ; 4 May 2009 ; J.J. Resende leg.; DZUP • 1 ☿ ; Monte Santo ; 19 Jul. 1995 ; Cardoso leg.; 4945A; CPDC • 1 ☿ ; Morro do Chapéu ; 25 Mar. 1990 ; J. Roseira leg.; #278; CPDC • 2 ☿☿ ; Morro do Cruzeiro ; 31 Jan. 1993 ; André leg.; 4610; CPDC • 1 ☿ ; Mucugê ; 6 Jul. 2009 ; Thomas Strwtzel leg.; UFSC • 1 ☿ ; Paulo Afonso , APA Serra Branca ; 19 Mar. 2009 ; Layla Marques leg.; #5596ª; DZUP • 1 ☿ ; Pilão Arcado , Barra Zacarias ; 9°59′ S , 42°48′ W ; alt. 416 m ; 5 Dec. 2005 ; P.P. Lopes and G.M.M. Santos leg.; CPDC • 1 ☿ ; same locality as for preceding; 3 Oct. 2006 ; J.J. Resende leg.; ninho 1 F30, 08; CPDC • 1 ☿ ; Poções , Faz. São João ; 9 feb. 1964 ; Eduardo leg.; CPDC • 1 ☿ ; Rio Real ; 27 Jun. 1994 ; Carmo leg.; 4859F; CPDC • 1 ☿ ; Rui Barbosa ; 16 Nov. 2012 ; Juliana Alves leg.; CPDC • 1 ☿ ; Sambaíba ; 24 Aug. 1993 ; C. Ivan leg.; 4512; CPDC • 1 ♂ ; same locality as for preceding; Mar. 2001 ; D.C. Nascimento leg.; CPDC • 1 ☿ ; S [Santa] Rosa ; 18 Jul. 1995 ; Cardoso leg.; 4944; CPDC • 1 ☿ ; Santa Maria da Vitória , Serra do Ramalho ; 13°23′52″ S , 44°11′52″ W ; Jun. 2009 ; J. Castro leg.; CPDC • 2 ☿☿ ; Sr do Bonfim [ Senhor do Bom Fim ]; 20 Jul. 1974 ; S. Laroca leg.; DZUP • 1 ☿ ; Serra da Jiboia ; 20 Jul. 2004 ; R.M. Rodrigues leg.; CPDC • 1 ☿ ; Tucano , caatinga ; 10.7798° S , 38.8928° W ; 15 Nov. 2010 ; E.M. Campos , A.M. Medina , P.L. Paixão and P.L.O. Rebouças leg.; MZSP • 2 ☿☿ ; Vila Nova ; 1 Feb. 1908 ; E. Gazbe leg.; MZSP . – Ceará • 1 ☿ ; Araripe ; 10 Nov. 1975 ; R.L. Araújo leg.; MZSP • 1 ☿ ; Chapada do Araripe ; 5 Apr. 1962 ; D. Zajuwawl leg.; MZSP • 1 ☿ ; Crateús , Croatá ; 5.1333° S , 40.9166° W ; 20–30 Apr. 2003 ; Y. Quinet leg.; MZSP • 1 ♂ ; Crateús , Serra das Almas ; 5.14170° S , 40.91620° W ; 19 May 2014 ; G. Melo and B. Rosa leg.; DZUP • 1 ♂ ; same locality as for preceding ; 19–20 May 2014 ; G. Melo and B.B. Rosa leg.; Mini-luminosa ; DZUP 549801 • 1 ♂ ; same collection data as for preceding; DZUP 548847 • 6 ☿☿ ; Fazenda Malhada Arcoverde ; Jul. 1974 ; MZSP • 1 ☿ ; Fortaleza ; 3°43′01″ S , 38°32′34″ W ; Sep. 2008 ; Y. Quinet leg.; 04; CPDC • 1 ☿ ; Maranguape , Serra de Maranguape ; 14 Nov. 2003 ; A.A. Tavares leg.; MZSP • 1 ☿ ; same locality as for preceding; 3.8809° S , 38.6958° W ; alt. 100 m ; 16. Oct. 2017 ; Y.P. Quinet leg.; manual ; DZUP • 1 ☿ ; Pacatuba ; 10 May 2016 ; G. Melo and B. Rosa leg.; DZUP • 1 ☿ ; Tianguá ; 18 Dec. 1948 ; C.R. Gonçalves leg.; MZSP . – Distrito Federal • 1 ♂ ; Brasília ; Sep. 1972 ; N. Simões leg.; MZSP . – Minas Gerais • 1 ☿ ; Poções de Sta Cruz , 10 km SE of Janaúba ; 16°53′ S , 43°16′ W ; alt. 660 m ; 19 Jan. 2005 ; Melo and Costa leg.; DZUP • 1 ☿ ; Viçosa , UFV ; 30 Oct. 1975 ; J. Freire leg.; CPDC . – Paraíba • 1 ☿ ; Areia ; 1 Jul. 1955 ; Pe. Pereira leg.; MZSP • 1 ☿ ; Barra do Rio Mamanguape , Rio Tinto ; 10 Jan. 2018 ; Marília Costa leg.; DZUP • 1 ☿ ; Brandão [ Caldas Brandão ]; 22 Sep. 1955 ; Aristóteles Silva leg.; MZSP • 1 ☿ ; Estrela Dalva , S [São] Sebastião do Umbuzeiro ; 24 Aug. 1954 ; A. Silva leg.; MZSP • 1 ☿ ; Fazenda Almas ; 20 Dec. 2006 ; J. Paulo leg.; pitfall A5 ; CPDC • 1 ☿ ; forest near Areia ; alt. 630 m ; 24 Jan. 1981 ; Robert Davidson leg.; primary forest , sweeping day ; MPEG • 1 ☿ ; João Pessoa ; 28 Nov. 2008 ; Linsmara Nunes leg.; CPDC • 1 ☿ ; same locality as for preceding; 23 Jan. 1988 ; M. Kammers and I.L. Oliveira leg.; MZSP • 1 ☿ ; same locality as for preceding; 4 Nov. 1955 ; C.A. Gonçalves leg.; MZSP • 2 ☿☿ ; same locality as for preceding; 9 Sep. 2002 ; A. Vasconcelos leg.; CPDC • 1 ♂ ; same collection data as for preceding; CPDC • 1 ☿ ; same locality as for preceding; 7°7′9.2″ S , 34°50′41.67″ W ; alt. 41 m ; 4 Sep. 2014 ; J.M. Churata S. leg.; DZUP • 1 ☿ ; João Pessoa , UFPB ; 2–5 Apr. 1995 ; M.F.S. Santos leg.; 4921; CPDC • 1 ☿ ; João Pessoa , Cabo Branco ; 18 Jul. 1993 ; N. Rabet leg.; CPDC • 1 ☿ ; João Pessoa , forest of Gargau ; alt. 45 m ; 22 Jan. 1981 ; R. Davidson leg.; primary forest , sweeping day ; MPEG • 5 ☿☿ ; Mamanguape ; 1 Jul. 1957 ; Dep. Zoologia exped .; MZSP • 1 ☿ ; Mata do campus UFPB ; 24 Apr. 1977 ; V.Py Daniel leg.; INPA • 1 ☿ ; same collection data as for preceding; Dec. 1976 ; INPA • 1 ♂ ; Matureia , Pico do Jabre ; 7.25250° S , 37.38472° W ; 19 Feb. 2018 ; A. Ferreira leg.; “ armadilha de luz ”; DZUP • 1 ♂ ; same collection data as for preceding; DZUP 5488852 • 1 ♂ ; same collection data as for preceding; DZUP 5488853 • 2 ☿☿ ; Olivedos ; 20 Aug. 1954 ; Aristóteles Silva leg.; MZSP • 1 ☿ ; Rebio Sema 3 ; 2 Jul. 2015 ; L. Brisa leg.; pitfall ; UEMS • 1 ☿ ; Rio Tinto ; 6°47′56.5″ S , 35°05′04″ W ; alt. 83 m ; 5 Jul. 2015 ; R. Silvestre leg.; pinça; UFGD • 2 ☿☿ ; São José Cordeiros , Faz. Almas ; 2 May 2003 ; A. Vasconcelos leg.; CPDC • 1 ☿ ; São José da Mata , Sítio São Miguel ; 12 Apr. 1997 ; A.D. Brescovit leg.; at night ; MZSP • 1 ☿ ; Tabuleiro , N Sta Rita ; 7 Feb. 1968 ; Dr. Dardano leg.; 9; MZSP . – Pernambuco • 3 ☿☿ ; Oct. 1929 ; Pickel leg.; MZSP • 1 ☿ ; Afrânio ; alt. 500 m ; 16 May 2007 ; J.A. Rafael and F.F. Xavier F. leg.; light ; INPA • 1 ☿ ; Araripina ; 2 Jan. 1973 ; R. Montenegro leg.; #324, 8430; MZSP • 2 ☿☿ ; Arcoverde ; 17 Jul. 1974 ; R.L. Araújo leg.; MZSP • 1 ☿ ; Brejão , Faz. Matary , R Maranhão ; 24 Apr. 1974 ; 932; MZSP • 1 ☿ ; same collection data as for preceding; coleção Diniz; DZUP • 6 ☿☿ ; Carnaubeira ; 20 May 1971 ; ABC MZUSP exped .; MZSP • 13 ☿☿ ; Floresta , Faz. Campos Bons ; 20–27 Jun. 1971 ; ABC MZUSP exped .; MZSP • 3 ☿☿ ; Garanhuns ; 13 Jan. 1967 ; F.B. Galinda leg.; DZUP • 1 ☿ ; same locality as for preceding; 29 Oct. 1938 ; A. Silva leg.; 5314; MZSP • 1 ☿ ; Itambé ; 13 Feb. 1976 ; C.J. Rosetto leg.; 931; DZUP • 1 ☿ ; Olinda ; Reichensperger leg.; 34; MZSP • 1 ☿ ; Parque Nacional do Catimbau ; 8°30′40.561″ S , 37°16′37.133″ W ; 27 Dec. 2017 ; Katherine B. Haedo leg., R27; DZUP • 1 ☿ ; same locality as for preceding; 8°32′48.38″ S , 37°15′21.74′′W ; alt. 938 m ; 27 Dec. 2017 ; Katherine B. Haedo leg.; R4; DZUP • 1 ☿ ; same locality as for preceding; 8°28′29.71′′ S , 37°21′22.36′ ′W ; alt. 987 m ; 27 Dec. 2017 ; Katherine B. Haedo leg.; R23; DZUP • 1 ☿ ; same locality as for preceding; 8°31′32.41′′ S , 37°14′57.62′′ W ; alt. 910 m ; 27 Dec. 2017 ; Katherine B. Haedo leg.; R45; DZUP • 1 ☿ ; Pesqueira ; 5 Aug. 1928 ; B. Pickel leg.; MZSP • 1 ☿ ; Petrolina ; 18 Jul. 1974 ; R.L. Araújo leg.; MZSP • 1 ☿ ; Recife ; 21 Apr. 1905 ; L. Lima Castro leg.; 2376; MZSP • 1 ☿ ; Salgueiro ; 17 Jul. 1974 ; P.L. Araújo leg.; MZUSP 5502 • 1 ☿ ; same collection data as for preceding; MZSP . – Piauí • 1 ☿ ; Bom Jesus ; 9°28′52.3″ S , 44°72′12.1″ W ; 8 Nov. 2010 ; T. Frizzo and H. Vasconcelos leg.; T2 PI 10S; DZUP • 1 ♂ ; Caracol , P.N. da Serra das Confusões ; 9.22637° S , 43.46331° W ; 3–7 Mar. 2016 ; R.M. Feitosa , G.P. Camacho and M.F.O. Martins leg.; DZUP 549804 • 1 ♂ ; Colônia do Piauí ; 27 Mar.–1 Apr. 1994 ; Boussard leg.; CPDC • 1 ☿ ; same collection data as for preceding; 4914; CPDC • 2 ☿☿ ; Curimatá , Espinhos ; 10°6′19″ S , 44°13′08″ W ; 20 Apr. 2000 ; M.P.D. Santos leg.; coleta manual ; MPEG • 1 ☿ ; Faz. Sta. Paz ; 19 Jun. 1992 ; Argolo leg.; 4635; CPDC • 1 ☿ ; Ribeirão Gonçalves , Est. Ecol. Uruçuí-Una , cerrado ; 10 Dec. 1980 ; A.I. Almeida leg.; MZSP • 1 ☿ ; São Raimundo Nonato , P.N. Serra da Capivara , Zabelê ; 12 Apr. 2011 ; A.C.A. leg.; pitfall ; MZSP . – Rio Grande do Norte • 1 ☿ ; FLONA de Nísia; Jun. 2017 ; Dina leg.; UFRN • 1 ☿ ; same locality as for preceding; 15 Sep. 2005 ; R.R. Hora leg.; 5457; CPDC • 1 ☿ ; Natal ; 24 May 2008 ; Pitfall F4 ; CPDC • 1 ☿ ; Natal , Campus Central UFRN ; Jun. 2014 – Jan. 2016 ; Dina leg.; “4 ninhos”; UFRN • 1 ☿ ; same collection data as for preceding; DZUP 549811 • 3 ♂♂ ; Campus Central UFRN , Jul. 2012 – Jan. 2016 , Dina leg., “4 ninhos”; DZUP • 1 ♂ ; same collection data as for preceding; DZUP 548850 • 1 ♂ ; same collection data as for preceding; DZUP 548851 • 1 ☿ ; Natal , Lagoa de Jenipabu ; Oct. 2010 ; R. Silvestre leg.; CPDC • 1 ☿ ; Parnamirim , Mata do Jiqui ; Oct. 2009 ; Dina leg.; UFRN . – Sergipe • 1 ☿ ; Estância ; 15 Jun. 1994 ; J. Jardim leg.; CPDC • 1 ☿ ; Itabaianinha , Mata Verde ; 15 Dec. 1991 ; B. Santos leg.; 4492; CPDC • 1 ☿ ; Pirambu , Pov. Aguilhada , restinga ; 10°41′37″ S , 36°50′37″ W ; 8 Jun. 2017 ; D.S. Mesquita and team leg.; pitfall ; MPEG • 1 ☿ ; Poço Redondo , Angico , alto sertão ; 9°41′ S , 38°31′ W ; Jun. 2008 ; F.S. Nascimento leg.; MZSP • 1 ☿ ; São Cristóvão ; 11°00′54″ S , 37°12′21″ W ; 16 Apr. 2009 ; J.O. Dantas leg.; “ ninho 2, 12 ”; CPDC • 1 ☿ ; Serra de Itabaiana ; Mar. 2008 ; T.J. leg.; CPDC . – Tocantins • 1 ☿ ; Jalapão , 19 Nov. 2012 ; H.L. Vasconcelos and T. Frizzo leg.; 405, T49-19S; UFU • 1 ☿ ; same locality as for preceding; 409, T49-17S; DZUP • 1 ☿ ; same locality as for preceding; 409, T50-11S; DZUP • 1 ☿ ; same locality as for preceding; 452, T51-2S; DZUP . Redescription Female MEASUREMENTS. Non-types (n = 32): HL 4.95–5.76; HW 4.9–5.73; MDL 3.9–4.66; SL 4.95–5.73; MSL 7.52–8.47; HFL 6.78–7.71; HBL 5.84–6.48; PL 2.15–2.59; PH 2.85–3.35; PW 1.45–1.95; ATS 7.04– 8.4; BL 25.71–29.39 (mm); CI 0.93–1.04; SI 0.96–1.09; DPI 0.64–0.81. HEAD. Malar area without striae or with weak longitudinal to oblique striae that do not reach anterior eye margin. Gena microareolate and opaque, usually without rugulae. Brownish, appressed and very short pubescence present between eye and frontal lobe, extending posteriorly to frons. Frons microareolate, opaque to silky; with very sparse brownish, flexuous, suberect to erect hairs, longer than scape width; pubescence densest laterally. Occipital corner microareolate and opaque (sometimes silky). Antennal scape microareolate and silky; long, suberect hairs usually present on antennal segments 1–3. Ventral surface of head microareolate, generally without strigulae, sometimes present only on anterolateral margin and extending to hypostomal tooth. Labrum without median longitudinal sulcus; very shallow transverse rugulae sometimes present. Mandibular dorsum weakly longitudinally strigulate on inner base, sculpture gradually fading apicad. Fig. 25. Dinoponera quadriceps Kempf, 1971 , ☿ (specimen from Natal, Rio Grande do Norte, Brazil; DZUP 549811). A . Full-face view. B . Full body in lateral view. C . Full body in dorsal view. MESOSOMA.Dorsal margin of pronotum in lateral view broadly convex, with no pronounced dorsoposterior swelling; anteroventral corner of pronotum forms obtuse angle. Pronotal dorsum weakly microareolate, opaque to silky. Metapleural-propodeal suture well-marked and usually straight. METASOMA. Petiolar node in lateral view elongate (usually DPI <0.8); anterodorsal corner usually higher than posterodorsal corner; anterior margin slightly concave to straight, forming blunt angle with dorsal margin; dorsal and posterior margins broadly convex and forming rounded angle. Node lateral face microareolate and usually opaque, rarely silky. Node anterior margin in dorsal view strongly convex, posterior margin broadly convex to straight, lateral margins broadly convex and converging anterodorsally. Abdominal tergite III microareolate and opaque; punctulae in variable density, usually denser laterally than dorsally; covered by brownish, flexuous, suberect hairs on entire surface; appressed pubescence usually sparse on dorsum, denser laterally. Male MEASUREMENTS. Non-types (n = 5): HL 2.13–2.32; HW1 2.56–2.84; MDL 0.52–0.58; SL 0.87–0.97; EL 1.5–1.58; MOD 0.53–0.58; LOD 0.51–0.57; MSL 6.75–7.16; HFL 5.5–6; PL1 1.8–1.9; PH 1.44–1.54; PW 1.17–1.27; ASL 4.35–4.75; BL1 15.56–16.53 (mm); CI1 1.13–1.24; SEI 1.59–1.77; SI1 0.33–0.36. HEAD. Frontal carina usually forming short longitudinal swelling. Lateral ocellus clearly surpassing posterior head margin in full-face view. Head punctulate, weakly microareolate and with silky sheen; with yellowish decumbent pubescence and scarce long suberect to erect hairs, longer than ocellus height in full-face view. Antenna with appressed pubescence and long suberect to erect hairs clearly longer than maximum scape diameter; third to fourth most apical antennomeres without hairs. Ventral surface of head punctulate and slightly microareolate; with subopaque to silky sheen. Fig. 26. Dinoponera quadriceps Kempf, 1971 , ♂ (specimen from Caracol, Piauí, Brazil; DZUP 549804). A . Full-face view. B . Full body in lateral view. C . Full body in dorsal view. MESOSOMA. Mesoscutum without notaulus. Mesopleural sulcus punctate or slightly scrobiculate. Scutoscutellar sulcus scrobiculate. Mesoscutellum longitudinally strigulate laterally. Metapleuralpropodeal suture with same microsculpture as rest of integument. Mesosoma mostly slightly microareolate and with silky sheen, becoming coarsely punctate on declivitous surface of propodeum; with decumbent to suberect pubescence and suberect hairs, distance between each hair usually greater than half its length. Legs densely covered by decumbent pubescence; coxae with suberect long hairs, greater than femur diameter. Protibial apex with at least one stout seta. METASOMA. Petiolar node microareolate and shining; densely covered by decumbent to suberect pubescence and sparse suberect hairs. Abdominal tergite VIII spiniform. Gaster very slightly microareolate and shining; tergites densely covered by appressed to decumbent pubescence, without long hairs. GENITALIA. Basal ring in dorsal view with slightly concave lateral margins, anteriorly slightly narrower than posteriorly; fenestra rounded to slightly transversally directed; median invagination U-shaped; dorsal margin in lateral view broadly concave to straight; anteroventral process subquadrate. Gonostylus narrow and sharp. Dorsal margin of volsella in lateral view anteriorly straight and posteriorly concave; anteroventral corner with two anterior teeth; posteroventral margin strongly concave and forming sharp triangular posterior lobe; digitus volsellaris with posterior margin straight or with a slight concavity. In lateral view, penisvalva with continuous dorsal posterior margins, ending in a subacute apex; ventral margin even and serrated, with slight concavity; anteroventral corner sharply pointed. COLOR. Body mostly chestnut brown. Remarks Females of Dinoponera quadriceps have a robust set of characters that usually permit easy separation from those of other species of the genus. However, D. quadriceps and D. gigantea , geographically close species, can sometimes cause confusion. The main characters dividing the two species are: (1) in D. quadriceps the anteroventral pronotal margin forms an obtuse angle, but in D. gigantea it is toothed or forms an acute angle; (2) the metapleural-propodeal suture in D. quadriceps is well-marked and straight, but in D. gigantea it is weaker and sinuous; (3) the cephalic venter is mostly strigulate in D. gigantea but in D. quadriceps the strigulae are absent or sometimes present just anterolaterally; (4) the dorsal region of abdominal tergite III in D. quadriceps is sparsely punctulate, but densely so in D. gigantea ; and (5) the anterodorsal corner of the petiolar node is higher in D. quadriceps than in D gigantea . Some specimens of D. quadriceps can present one or two D. gigantea characters, and viceversa, so the use of all these characters affords the most reliable identifications. It is worth noting that the difficult cases are more common among specimens from Maranhão and Tocantins , as in D. gigantea . Notwithstanding these complications, we consider the aforementioned set of diagnostic characters to provide stronger support for maintaining D. quadriceps and D gigantea as separate species than for synonymizing them. The alternative of describing a new species from Tocantins and Maranhão is not possible because the characters seem to vary randomly. Determining the degree of expression of each character is also important. Species differences are frequently a degree of difference one from another, as in the case of the anteroventral pronotal corner in D. quadriceps , which may be strongly angular but not toothed or forming an acute angle. A specimen from Itabaianinha ( Sergipe ) has a short and robust peciolar node (DPI> 0.8), characteristic of D. grandis . However, these two species differ in the absence of: (1) the pronotal anteroventral tooth; (2) striae in the malar area and (3) strigulae on the ventral head surface in D. quadriceps . The other species of Dinoponera can be easily distinguished by having abdominal tergite III smooth and shining. It was not possible to examine the holotype of D. opaca . It is kept as a junior synonym of D. quadriceps , on account of the examination of the holotype by C. Baroni Urbani in 1974 at the request of W. Kempf (1975: 344) . Dr Baroni Urbani found that the type agreed with the description of D. quadriceps . Lenhart et al. (2013) examined the type and found total coincidence with D. quadriceps . The Rio de Janeiro record for D. opaca is undoubtedly a labelling error, as there are no other records of Dinoponera for Rio de Janeiro , a heavily collected area for ants, and if there were any Dinoponera , the expected species would either be D. lucida or D. australis . Males of D. quadriceps are different from those of D. lucida , D. longipes , Dinoponera morphospecies 1 and D. grandis by having hairs on the antennae that are clearly longer than the scape diameter and the gonostylus narrow, with a sharp apex. As mentioned before, males of D. gigantea and D. quadriceps are similar in many characters, but the former is longer, more pilose and has a visible notaulus on the mesoscutum. Biology The species occurs mainly in arid or savanna areas with the temperature of the coldest month greater than 18 °C and the precipitation of the driest month less than 60 mm ( Alvares et al. 2013 ). Nests are usually built in the ground, near roots and often in direct contact with termite mounds ( Araujo et al. 1990 ; Paiva & Brandão 1995 ). However, Vasconcellos et al. (2004) found nests more than 3 m from trees in a remnant of Atlantic Forest, indicating nesting habits may vary with the environment. Each nest can have two to ten entrances surrounded by branches and some loose earth ( Araujo et al. 1990 ; Paiva & Brandão 1995 ). Nest depth varies from 10 to 120 cm and the number of chambers can reach 16 ( Araujo et al. 1990 ; Paiva & Brandão 1995 ; Vasconcellos et al. 2004 ). Assis et al. (2017) recorded D. quadriceps reusing nests of Atta sexdens (Linnaeus, 1758) , which are deep enough to ensure protection against hot and dry weather. The density of nests varies from 15 to 40 per hectare ( Vasconcellos et al. 2004 ). The size of the colony can vary from 12 to 141 workers ( Paiva & Brandão 1995 ; Monnin & Peeters 1997 , 1998 ; Vasconcellos et al. 2004 ; Nascimento et al. 2012 ; Asher et al. 2013 ). Vasconcellos et al. (2004) found males almost every month of the year, suggesting male production is not seasonal. Their diet is omnivorous, with a preference for arthropods found dead on the soil ( Araújo & Rodrigues 2006 ). Foraging is solitary and occurs during the day, avoiding the hottest hours ( Medeiros et al. 2012 ). The size of food items varies, but regardless of its weight there is no recruitment of other workers ( Araújo & Rodrigues 2006 ). Before finding food, the worker is usually slow and meanders about its usual route, moving up to 35 m from the nest. Foraging tends to be faster during the return trip and follows a relatively straight line ( Araújo & Rodrigues 2006 ; Azevedo et al. 2014 ). Araújo & Rodrigues (2006) suggested that spatial orientation during foraging may follow chemical and visual cues. Medeiros & Araújo (2014) divided the life cycle of the colony into stages. The first stage is monodomic, with the ants occupying a single nest. Subsequently the colony becomes polydomic, until gradually the ants stop visiting the parental nest and the colonies become isolated, completing the fission. The newly separated colonies go through a period of territorial conflict that results in the extinction or migration of the old colony ( Medeiros & Araújo 2014 ). Vasconcellos et al. (2004) also pointed out that intraspecific territorial competition may be one of the main factors responsible for the spatial arrangement of nests. Colonies of D. quadriceps are predominantly monogynous ( Monnin & Peeters 1997 , 1998 , 1999 ), though Araujo et al. (1990) found colonies with more than one gamergate. The dynamics that determine the hierarchy of dominance in this species is well studied, mainly by Monnin & Peeters (1999) . Hierarchical position is determined by age and agonistic interactions involving mainly the ants with high status in the colony ( Monnin & Peeters 1999 ). Newly emerged workers usually replace older ones in the highest positions of the hierarchy. Agonistic interactions are usually related to the stability of monogyny in the colony, and can be performed by the gamergate itself, inhibiting other workers ( Monnin & Peeters 1999 ). After copulation, gamergates chew away the end of the male abdomen, which remains trapped in the female genitalia, preventing further copulation ( Monnin & Peeters 1998 ). Distribution ( Fig. 29B ) Dinoponera quadriceps is endemic to northeastern Brazil , occurring predominantly in the Caatinga. The northernmost record is in Fortaleza, Ceará and the southernmost record is in Viçosa, Minas Gerais . However, Viçosa is more than 500 km away from the next southernmost record (Janaúba, Minas Gerais ), leaving doubts as to the reliability of the former record. Another unexpected record is from Brasília, which is also more than 500 km distant from any other record. These records may reflect a wider distribution in a recent past (both date from the 1970s) or just labeling errors. Lenhart et al. (2013) cited records of D. quadriceps in the cities of Santarém and Óbidos, as well as along the Tapajós River, all in the state of Pará. However, we found no records of this species within at least 1000 km of that region. This discrepancy may be due to either wrong labeling or differences in the criteria used for species identification in each study. The latter is most probably the case, given the similarity between D. quadriceps and D. gigantea .