Argentoleon irrigatus (Gerstaecker)

Girelli, Maria Eduarda, Tavares, Leon Gustavo Miranda & Machado, Renato Jose Pires, 2024, Taxonomical and biological notes on Argentoleon irrigatus (Gerstaecker) (Neuroptera: Myrmeleontidae), Zootaxa 5538 (5), pp. 401-423 : 403-419

Argentoleon irrigatus (Gerstaecker)

Formicaleo irrigatus Gerstaecker, 1894: 133 (OD) ; Penny, 1977:

40 (cit, cat); Stange, 2010: 6 (cit); Heckman, 2017: 253 (cit).

Austroleon irrigatus (Gerstaecker) , Navás, 1932: 55 (ncb); Navás, 1935: 362 (cit).

Moza irrigata (Gerstaecker) , Banks, 1943: 165 (ncb); Penny, 1977: 40 (cat, cit); Heckman, 2017: 253 (cit).

Brachynemurus irrigatus (Gerstaecker) , Penny, 1977: 40 (ncb, cat); DeMarmels & Gaiani, 2003: 407 (cat); Heckman, 2017: 253 (cit).

Myrmeleon irrigatus (Gerstaecker) , Stange, 1994: 81 (cit) Stange, 2004: 229 (cit).

Argentoleon irrigatus (Gerstaecker) , Stange, 1994: 81 (ncb); Stange, 2004: 229 (cat); Stange, 2010: 6 (rd, im);

Silva et al., 2015 (cit, ill); Petko et al., 2016: 1 (cit); Heckman, 2017: 252 (key); Machado et al., 2019: 424 (tree); Schuster & Machado, 2021: 998 (cat); Machado & Martins, 2023: 4 (cit); Tavares et al., 2023: 510, 548 (tree, cit); Machado, 2023: 140 (cat).

Brachynemurus meridionalis Banks, 1909: 2 (OD) ; Stange, 1967: 52 (cat); Penny, 1977: 41 (cat); Stange, 2004: 229 (syn, cat); Stange, 2010: 6 (cat); Heckman, 2017: 253 (cit).

Brachynemurus argentinus Banks, 1910: 146 (OD) ; Stange, 1961: 675 (cit); Stange, 1967: 52 (syn, cat); Penny, 1977: 41 (cat); Stange, 2004: 229 (cat); Stange, 2010: 6 (cit); Heckman, 2017: 253 (cit).

Austroleon argentinus (Banks) , Navás, 1917: 193 (ncb); Navás, 1919: 193 (cit); Stange, 1967: 52 (syn, cat); Penny, 1977: 41 (cat).

Clathroneuria amazonica Navás, 1914: 215 (OD) ; Esben-Petersen, 1920: 193 (syn); Stange, 1967: 53 (cat); Stange, 2004: 229 (cat); Stange, 2010: 6 (cit); Heckman, 2017: 253 (cit); Monserrat, 2021: 202 (cit).

Austroleon alienus Navás, 1915: 9 (OD) ; Navás, 1919 a: 293 (cit); Navás, 1920a: 57 (cit); Navás, 1920b: 47 (cit); Navás, 1922: 183; Navás, 1923: 771 (cit); Navás, 1926: 106 (cit); Navás, 1928a: 33 (cit); Navás, 1928b: 140 (cit); Navás, 1929: 141 (cit); Navás, 1930: 127 (cit); Navás, 1935: 362 (syn, cit); Stange, 1967: 53 (syn, cat); Penny, 1977: 41 (cat); Stange, 2004: 229 (cat); Stange, 2010: 6 (cit); Heckman, 2017: 253 (cit); Monserrat, 2021: 199 (cit).

Austroleon clavatus Navás, 1915: 126 (OD) ; Stange, 1967: 53 (syn, cat); Penny, 1977: 41 (cat); Stange, 2004: 229 (cat); Stange, 2010: 6 (cit); Heckman, 2017: 253 (cit); Monserrat, 2021: 199 (cit).

Austroleon antennatus Navás, 1919: 295 (OD) ; Stange, 1967: 53 (syn, cat); Penny, 1977: 41 (cat); Stange, 2004: 229 (cat); Stange, 2010: 6 (cit); Heckman, 2017: 253 (cit); Monserrat, 2021: 199 (cit).

Austroleon sectorius Navás, 1920a: 58 (OD) ; Stange, 1967: 53 (syn, cat); Penny, 1977: 41 (cat); Stange, 2004: 229 (cat); Stange, 2010: 6 (cit); Heckman, 2017: 253 (cit); Monserrat, 2021: 200 (cit).

Austroleon latipennis Banks, 1922: 60 (OD) ; Penny, 1977: 40 (cat); Stange, 2004: 229 (syn, cat); Stange, 2010: 6 (cit); Heckman, 2017: 253 (cit).

Brachynemurus latipennis (Banks) ; Penny, 1977: 40 (ncb).

Austroleon graphonotus Navás, 1934: 13 (OD, ill); Stange, 1967: 53 (syn, cat); Penny, 1977: 41 (cat); Monserrat, 1985: 241 (cit); Stange, 2004: 229 (cat); Stange, 2010: 6 (cit); Heckman, 2017: 253 (cit); Monserrat, 2021: 199 (cit).

Redescription. ADULT: Size AW 18.7–29.6 mm, BL 22.8–38.4 mm. Head: ( Fig. 1B ) Labrum yellow; set with a line of elongated light setae on the ventral margin. Clypeus pale; with a few elongated and a few short black setae. Frons pale with bimarginated interantennal marks dark brown to black reaching until the dorsal third, and a dark brown longitudinal medial mark, continuous or discontinuous (tree small irregular spots); set with a few short black setae. Gena pale, lighter than frons. Ocular rim setae absent. Antennae clavate; about 3 times the pronotum length; interantennal area about the same size of scape width. Vertex lightly raised set with decumbent black setae at the apex and in the central portion of medial row; mostly brown except for a pale mark between medial and posterior rows; anterior row as a transversal black line, medial row as an irregular light to dark brown line, posterior row with two submedial light brown, small, roundish marks, sometimes fused in a single mark. Antennae with nearly 30 flagellomeres; scape and pedicel light to dark brown, each with a submedial and a distal and transversal row of short black setae; basal flagellomere about twice as long as the other ones that are about as long as wide, except by the ones at the clave, much wider than long; flagellum dark brown except by the lighter flagellomeres at the base of the clave. Mandible orange to brown with internal margin black. Labium with long black setae in the submentum and scattered white short setae in the mentum and submentum. Maxillary and labial palpi mostly yellow, with the distal palpomere darker and slightly lighter at joints; with a few very short white setae in the first two palpomeres of the maxillary palpi and black short setae in the entire labial palpi. Distal labial palpomere fusiform with blunt end. Palpimacula oblong and medially located. Thorax: ( Fig. 1C ) Pronotum about as long as wide; posterior margin slightly wider than anterior; subapical furrow weakly marked; mostly pale with dark brown lateral margins, one light to dark brown submedial longitudinal band often interrupted by one or two pale medial marks; set with a pair of long white setae curved anteriorly in the lateral margins, few elongated black setae on the posterior margin, and several short black and white setae throughout the tergum. Mesonotum dark brown and pale; with two small pale spots in the prescutum, set with few erect black setae; scutum with a large pale mark encompassing a longitudinal dark brown line on each side and a small pale mark near the lateral margins, set with short black setae and erect black and white setae; scutellum with the medial area and lateral margins pale, set with short black setae and a row of long white setae in the posterior margin. Metanotum similar in color to mesonotum, except for the prescutum, mostly pale with two brown marks; with exclusively white setae, erect setae absent, metascutum with a row of setae pointing posteriorly, near metascutellum anterior margin; metascutellum with 1–2 setae in the lateroposterior margin. Pterothoracic pleura mostly dark brown, with some pale marks mostly at the sutures between the sclerites, pale areas in the metathorax generally larger; set with short white setae. Miller’s organ absent. Legs: ( Fig. 1A ) prothoracic leg femur and tarsus (without claw) about the same length and tibia slightly shorter, mesoleg femur longer than tarsus, both longer than tibia, metaleg femur as long as tibia, both longer than tarsus. Tarsomeres length: T2, T3 and T4 about the same length, T1 slightly longer than T2, T5 slightly longer than the basal four tarsomeres together; T1 slightly longer in the metathoracic leg. Pretarsal claws slightly longer than half of T5 length. Coxa and trochanter pale, except for procoxa with two brown marks in the lateral face, and one or two brown marks on the dorsal margin of the meso and metacoxa. Femora mostly pale with the dorsal surface of the profemur occasionally brown, dorsal surface of mesofemur also brown but lighter than profemur; femora with brown dots at the base of the setae. Tibia pale with brown dots at the base of the setae; protibia slightly darker than the others. Tarsomeres pale at the base and brown at the apex. All segments covered in short black setae (occasionally white in the trochanter), coxa with three elongated white bristles, femur and tibia with elongated black bristles mostly in the closing face, with a bristle dorsally and one laterally, both subapical. Pro and meso femoral sense hair longer than the femur length. Tibial spurs long, reaching T3. Wings: ( Fig. 2 ) Mesothoracic wing: elongated, broadening on the apical half, then narrowing to a pointed tip in the apical quarter. Anterior Banksian line present, posterior poorly developed. Membrane hyaline, with many brown marks at the forks and crossveins origins; veins brown and white dashed; pterostigma white to pale, but weakly marked; veins and margins with short black setae; often with a brown spot on the rhegma. RP origin at the basal quarter of the wing, CuA fork between RP origin and RP first branch; RP with 11 to 12 forks; 2 to 4 presectoral crossveins; Costal area similar in size to presectoral area with costal area veinlets mostly simple but forked at the apical third; prefork area wider than posterior area at CuA fork level. CuP origin distant from basal crossvein, near the level of the first crossvein between A1 and A2. A2 connected to A3 by crossvein. Metathoracic wing: Similar in size and shape to mesothoracic wing but narrower. Male pillula axilaris present, membrane hyaline, pterostigma white to pale, veins dashed; anterior banksian line poorly developed, posterior Banksian line absent; set with black setae. MP fork located beyond RP origin; costal area subcostal veinlets mostly simple, but forked at the apical quarter; RP with 8 to 13 forks; 2 or 3 presectoral crossveins; CuPA1 extending to the apical half of the wing.

Abdomen: ( Fig. 2 ) variable in color, but generally sternites brown to black, except for the sternite I pale with two submedial light brown to brown marks, forming a line or triangles in the center, sternite II with a pale medial line; all covered with white and/or black elongated and short setae. Tergites pale to brown, lighter than sternites; posterior tergites darker colored; covered with black setae; tergites I and II with a medial mark darker than the surrounding area that can extend as a medial darker line in the remaining tergites. Abdomen length, as long as or slightly shorter than wings in females, but longer than wings in males. Male Terminalia: ( Fig. 3E ) Ectoproct: thin; yellow to brown, lighter colored than the rest of the abdomen; set with elongated black setae; postventral lobe elongated, forcipate, about 3 times longer than the 8 th sclerites, with thicker setae at the posterior end. The 9 th tergite shorter than remaining sclerites but twice as tall as the other tergites in lateral view, lighter in color in the distal portion; set with black setae. The 9 th sternite posterior margin rounded in ventral view, projected posteriorly; set with black setae. Male genitalia ( Fig. 3A–D ) composed of three main structures: Gonarcus, mediuncus and parameres, all surrounded by the genital sac. Genital sac with three regions of short black setae, one dorsal to the medial area of the gonarcus and a pair of lobe-like sclerites located lateral to the parameres, in posterior view. Gonarcus in dorsal view arched with the lateral arms diverging at the anterior end; in lateral view with the anterior margin rounded; in posterior view as a transverse bar. Parameres fused; in ventral view with two medial rows of short black setae extending to the anterior end where they are replaced by two long and two medium black setae; anterior margin curved and narrower than the posterior region. Paramere plate short with the base enlarged anteriorly and more sclerotized, and paramere tooth long, projected dorsally with the tip acuminated and curving ventrally. In lateral view, anterior half taller than posterior half, with the anteroventral corner curved, paramere plate ventrally straight with the tip rounded. In posterior view portion of paramere anterior to hinge apically fused and sclerotized. Mediuncus in posterior view enlarged; mostly rectangular but bulging near the apex; less sclerotized than the other structures but with a medial stripe more sclerotized than the rest of the mediuncus; apex reaching the parameres at the portion of paramere anterior to hinge; in lateral view as a triangle with the dorsal margin thin and the ventral enlarged. Female terminalia: ( Fig. 4A–B ) Ectoproct with posterior margin rounded in lateral view, set with digging setae in the ventral half and elongate black setae in the dorsal half in lateral view. Lateral gonapophyses fused in one plate, set with shorter digging setae and short black setae. 7 th sternite with posterior margin straight set with elongated setae. Pregenital plate membranous, set with short black setae. Posterior gonapophyses digitiform, set with elongated black setae. Anterior gonapophyses present, lobe-like, at the base of the posterior gonapophyses, set with thick black setae. Gonapophyseal plate absent. Ventromedial plate squared, set with thick black setae in the anterior half and short black setae in the posterior half.

Description. LARVA: ( Fig. 5A–B ) Size BL 7.2–11.8 mm; ML 1.8–2.3; HL 1.7–2.5 mm; HW 1.6–1.8 mm. General coloring pale with dark brown marks, more abundant in dorsal surface. Head: ( Fig. 6A–B ) Head capsule broader than long in dorsal view, in ventral view longer than broad, dorsal side pale with dark brown marks: a pair of fusiform marks in the posterior area and a pair of rectangular marks in the lateral margins; lateral side with a large dark mark; ventral side unmarked. Dorsal side of head capsule set with black or white dolichasters of two types: the very short and sphere-like more numerous and the short cone-like dolichasters; and long black thick setae near the lateral margin, ventral surface set with short black setae that get longer near the lateral margin. Ocular tubercle prominent, with seven stemmata (six dorsal and one ventral), pale yellow in dorsal view and dark in ventral view, connecting to lateral mark on head capsule. Dorsal and ventral tentorial pits visible. Antennal tubercule with three cylindrical dolichasters. Antennae originating in a small tubercule, scape broad and black, pedicel similar to flagellomeres, 14 small flagellomeres and a longer apical one. Clypeo-labrum in dorsal view dark brown set with sphere-like black dolichasters; anterior margin set with a row of long black dolichasters and with a medial small concavity. Mandible brown with external margin and apex dark brown, about as long as ventral side of the head capsule, straight and with constant width until the last teeth then curving and acuminating towards apex. Internal margin with three teeth, proximal tooth located around the center of the straight region of the mandible, longer than the distal tooth but shorter than medial tooth, medial and distal teeth located at the distal end of the straight portion, with no interdental mandibular setae between them; interdental mandibular setae formula: (5– 7)(2–3)(0)(0). External margin set with long and short black setae, dorsal surface with sparse short setae until the second teeth, ventral surface glabrous. Maxila long and coupled into the mandibular ventral groove; cardo and stipe visible in ventral view. Prelabial lobe dark brown with sphere-like and cone-like dolichasters. Labial palpus with three palpomeres, distal palpomere bigger than basal palpomere, which is bigger than mid palpomere; base of the palpus with rowed, thick, cylindrical dolichasters at the base of each palpi. Distal palpomere enlarged and darker than remaining palpomeres. Mid palpomere with small black setae in the distal end. Basal palpomere with cone-like black dolichasters at the distal end. Thorax: General coloring: ventral surface pale speckled with brown; pronotum with a transversal gradient: brown medially, pale submedially and dark brown laterally; two small dark brown triangular marks in the posterior margin, covered with sphere-like and cone-like black dolichasters and long black setae on the anterior and lateral margins; meso and metanotum pale to light brown with scattered dark brown marks. Ventral surface with sparse long black setae. Mesothorax set with black cone-like and cylindrical dolichasters, spiracle born on brown tubercle, about twice longer than broad, with a pair of nearly bare submedial rounded light brown marks; two pairs of pedunculated setiferous processes with long black setae, anterior setiferous process darker and digitiform. Metanotum similar to mesonotum but with slightly shorter setiferous processes. Legs: general coloring pale with metaleg slightly darker, set with black setae, brown tarsal claws. Proleg femur with a dark brown irregular mark on dorsal surface; coxa set with thin long setae; trochanter with small thin setae; femur set with long setae and shorter thicker setae on the external margin; tibia similar to femur but with shorter thick setae; tarsus nearly bare except for a few short thin apical setae. Mesoleg similar to proleg. Metaleg similar to mesoleg but femur with a fringe of thin long setae tibia and tarsus fused (tibio-tarsus), set with sparse short setae. Abdomen: ( Fig. 6E ) each abdominal tergites 1–7 visually divided in three parts dorsally, the anterior and medial similar in size, and the posterior narrower; each of them pale speckled with brown, with a dark margin, set with cylindrical dolichasters; ventral surface mostly pale with a pair of submedial brown marks on each segment, with sparse short black setae. Abdominal setiferous processes reduced but set with long black bristles, located laterally, with one dorsal and the other one ventral in relation to the spiracle, on each segment. Eight spiracles not enlarged, located on short tubercles. Sternite VIII without digging-setae; odontoid process very small, pale and triangular, about as long as basal width. Sternite IX slightly broader than long, with digging setae: two rows, one anterior and one posterior, with three long digging setae on each side, and two short medial digging setae between the anterior row and the rastrum. Rastrum slightly longer than broader, each with four digging setae, the three biggest as long as the rastrum and the innermost one shorter than the other three. PUPA (6 days old): ( Fig. 7A ) Body slightly curved, mostly pale, thorax and abdomen with scattered black marks; covered in yellow setae, more abundant in the head, eyes and thorax (mostly prothorax), sparse in the abdomen and legs, and absent on the antennal flagellum. Head similar to adult head, but covered in long setae, antennae without the clave and projected ventrally. Abdomen tip of female rounded ( Fig. 7A ), male ( Fig. 7B ) projected.

Examined material. (140♀ 48♂ 2 larvae) ADULTS • Brazil • Amazonas : 1 ♀ Manicoré, P. N. Campos Amazônicos, alojamento, cerrado, [8°16’47.4”S, 61°34’28.3”W], vii.2017 • Bahia: Maracás , 23.iii.2012, Calor, Quinteiros, Duarte, Garcia (2♀, UFBA) • Mucugê, Areial , 10.i.2022, puçá/coleta manual, Tavares L. (6♀ 7♂, DZUP) • Mucugê, Córrego Medonho , 24.i.2018, Bandeja, Duarte T., Barata S., de Oliveira I. (1♀ 1♂, UFBA) • PMSV Rio Caparosa, 1.xii.2017 – 1.i.2028, malaise, Calor et al. (1♀, UFBA) • Distrito Federal: Brasília , 1.X.1974, Carrijo (1♀, INPA) • Brasília, Est. Metereológica , 10.iv.1969, I. C.B. (1♀, UnB) • Brasilia , 9.xii.1974, malaise trap, Knutson (1♀, USNM) • Ceilândia, 11.xii.1977, Cristino (1♀, UnB) • Guará, 17.xi.1975, Arlindo (1♀, INPA) • Parque Ecológico das Sucupiras, Coleta Manual (larvas), criadas em laboratório para obter o adulto, 09.ix.2022, Tavares L. (3♀ 1♂, DZUP) • Planaltina, [15°36’S, 47°44W], 960 m, 27.x.1988 (1♀, Embrapa DF) • Planaltina , [15°35’S, 47°42W], 1000 m, 13.iv.1976, V.O. Becker (2♀ 1♂, Embrapa DF) ; same 20.iv.1982, 1♀ • Goiás: Alto Paraíso , Chapada dos Veadeiros, Raizama, 700 m, 27.iii.2008, varredura, J. A. Rafael, F. F. Xavier F°., (1♂, INPA) • Alto Paraíso de Goias, Chapada veadeiros, [14°08’34’’S 47°46’00’’W], 10.iv.2018, Malaise, R. Lara & N. Perioto (1♀, DZUP) ; same, 24.iv.2018 2♀ ; same, 8.v.2018, 2♀ ; same, [14°08’34’’S, 47°46’08’’W], 10.iv.2018, 1♀ ; same, 22.v.2018, 1♀ ; same, 23.iv.2019, 1♂ ; same, [14°08’36’’S, 47°46’04’’W], 24.iv.2018, 1♀ ; same, 7.v.2019, 1♀ ; same, [14°08’36’’S, 47°46’07’’W], 24.iv.2018, 1♀ ; same, 27.iii.2018, 1♂ ; same, 8.v.2018, 1♀ ; same, [14°08’36’’S, 47°46’10’’W], 8.v.2018, 2♀ ; same, 22.v.2018, 1♀ ; same, [14°07’43’’S, 47°44’08’’W], 8.v.2018, 1♀ ; same, 12.ii.2019, 1♀ ; same, [14°07’43’’S, 47°44’06’’W], 10.iv.2018, 1♀ ; same, 24.iv.2018, 1♀ ; same, 13.iii.2018, 1♂ ; same, 12.ii.2019, 1♀ ; same, 26.iii.2019, 1♀ ; same, [14°07’44’’S, 47°44’04’’W], 13.iii.2018, 1♂ ; same, 10.iv.2018, 1♀ ; same, 6.xi.2018, 1♀ ; same, [14°07’44’’S, 47°44’00’’W], 10.iv.2018, 1♀ ; same, 24.iv.2018, 1♀ ; same, 8.v.2018, 1♀ ; same, 21.v.2019, 1♀ ; same, 27.iii.2018, 1♂ ; same, [14°07’43’’S, 47°43’97’’W], 27.iii.2018, 2♀ ; same, 22.v.2018, 1♀ 1♂ ; same, 9.iv.2019, 1♀ ; same, 7.v.2019, 3♀ • Caldas Novas, Parque Est. Serra de Caldas Novas , 700–1000 m, [17°46’13’’S, 48°39’22’’W], 22–23.iii.2008, arm. luz, J. A. Rafael F. F. Xavier F°. (2♀ 7♂, INPA) • Silvania, Faz. Umuabama , x.1991, M.V. Ferraz (1♀, MZUSP) • Maranhão: Mirador, Parque Est. Mirador, Base da Geraldina , 20–23.iv.2007, Armad. [ilha] Luminosa, F. Limeira-de-Oliveira (1♀, CZMA) • Mato Grosso: Barra Bugres , E. E. Serra Araras, 12.v.1990, Marinez Marques, (1♀, UFMT) • Comodoro, TI Nambiquara , [19°39’03’’S, 59°42’26’’W], 26.iv.2006, J. A. Rafael & F.F. Xavier F°, arm. Luz, (3♀, INPA) • Cuiabá, 10.III.1994, Luciana M. Silva (1♂, UFMT) • Cuiabá, Recanto das Seriemas , [15°35’33.7’’S, 56°01’37.94’’W], 17.vii.2017, leg. L.G.O.A. Nunes (1♀, CEMT) • Mato Grosso do Sul: Mato Grosso do Sul: Corumbá, Serra do Urucum , 30.xi.1960, K. Lenko (2♀ 2♂, MZUSP) • Minas Gerais: Abre Campo , 1–4.v.2014, Silveira, P.S.A. & Silveira, A.M. (1♀, CCT-UFMG) • Juiz de Fora, iii.1965, Angelo Machado (1♀, INPA) • Lavras, B. Centenário , [21°14’43’’S, 44°59’29.6’’W], 10.iii.2014, dentro de casa, P. Dias (1♀, CCT-UFMG) ; same, iii.2016, 2♀ • Paracatu, 17°35’S, 46°52W, 920 m, 8.xi.1993 (1♀, Embrapa DF) • Paracatu , [17°35’S, 46°52W], 920 m, 8.xi.1993 (1♀, Embrapa DF) ; same [17°13’S, 46°52’W] 1♀ • Parque Estadual do Rio Doce, 6–23.iv.1978, Maria Vulcano (10♀ 1♂, INPA) ; same, 28–31.iii.1979, 1♀ • 15 km ao sul de Ponto dos Volantes, Luz , 16°53’S, 41°29’W, 750 m, 11.ii.2010, P. Grossi, D. Parizotto & G. Melo leg. (1♀, CERPE) • Pouso Alegre, 5.iv.1964, P. Pereira (1♀, MZUSP) • Serra do Cipó, afl. Rio Capivara Froelich & Shimizu (1♀, MZUSP) • Santana do Riacho, Serra do Cipó (palácio), iii.1957, Machado col. (3♀, INPA) • São Gonçalo Rio Abaixo, Estação Ambiental Peti-Cemig , [19°53’02’’S, 43°22’21’’W], 28.iv–1.v.2012, luminosa, A. Lima, A. Kumagai & P. Dias, ( CCT-UFMG) • Serra do Salitre: RPPN Fazenda da Cachoeira , [19°09’56.7’’S, 46°34’06.7’’W], 1070 m, 20– 22.iii.2012, lençol iluminado, P Dias (9♀ 7♂, CCT-UFMG) • Paraná: Castro, Parque Est. Guartelá , [24°33’45’’S, 50°15’’32W], 1000 m, 10.xi.2007, Luz, J. A. Rafael & P. Grossi (1♂, INPA) • Curitiba, ii.1969, Mielke (1♀, DZUP) • Curitiba , III.1983, K. Zanol-leg (1♀, DZUP) • Curitiba, UFPR Politécnico , [25’26’47’’S, 49°13’55’’], 917 m, 17.v.2019, coleta ativa, Maciel, R. M. A leg. (1♀, DZUP) • Guarapuava, 19.ii.1969, O.H. Mielke leg. (1♀, DZUP) • Jaguariaíva, 3.xii.1986, Harada, Laroca leg. (1♀ 2♂, INPA) ; same, 2.xii.1986, 1♂ ; same, 5.xii.1986, 1♂ • Jaguariaíva, Parque Estadual do Cerrado , 13–16.xii.2022, Coleta Manual (larvas), criadas em laboratório para obter o adulto, Tavares, L; Silva-Junior, JO; Machado, RJP (2♀ 3♂, DZUP) • Ponta Grossa, Campo , iv.1940 (1♀, DZUP) • Ponta Grossa, Cidade , iv.1946 ( DZUP) • 1♀ Ponta Grossa, Oficinas , ii.1949, com luz ( DZUP) • 1♀ Porecatu, 20.x.1970, Becker-Hatschback ( DZUP) • 5♀ 2♂ Tibagi, Parque Estadual do Guartelá , 08–11.iii.2021, varredura, Barbosa D. N. (1♀, DZUP) • Rio Grande do Sul: Cachoeira do Sul , 18.xii.1977, N.D. Penny (1♀, INPA) • Rondônia: Vilhena , 5.xi.1979, ND Penny & JA Arias (1♀, INPA) • Santa Catarina: Nova Teutonia , 300–500 m, 27°11’S, 52°23’W, i.1977, Fritz Plauman (1♀, INPA) ; same, ii.1977, 1♀ • São Paulo: Campinas , 10.iii.1970, Reichardt (1♀, MZUSP) • Ilha da Vitória, 16–27.iii.1964, Exp [edition] Dep [artment] Zoo [logy] (1♀, MZUSP) • Itapetininga, Floresta Estadual , 27.xii.1965, Rebello (2♀, MZUSP) • Itapetininga , II.1970 (1♂, INPA) • Luiz Antonio, Est. Ecológica Jataí , [21°36’47’’S, 47°49’04’’W], mata ciliar, 5.xii.2007, arm. luminosa, R.I.R. Lara (1♀, APTA) • Piracicaba, Esalq (540 m), iii.1964, M. Menezes (1♀, ESALQ), same, 20.ii.1967, S.S. N., 1? • Piracicaba , centro, 31.iii.1965, F. Wiendl (1?, ESALQ) • Piracicaba , 12.iii.1979, V.L. Tornisielo (1?, ESALQ) • Rio Claro, iii.1969, A. Queiroz (1?, ESALQ) • Rio Claro , 9.iii.1968, Braulio Dias (1♀, UnB) • São Paulo, Santo Amaro , 24.xii.1959, F. Wiendl (1?, ESALQ) • São Paulo, Santo Amaro, Brooklin Paulista , iv.1963, L.T. F. col (1♀, MZUSP) • Tocantins: Palmas, Condomínio Beira Lago , 4.v.2015, coleta manual, Alvim BGC (4♀, CEUFT) • Porto Nacional, Fazenda São Judas Tadeu , 27–29.iii.2015, luminosa, Krolow equipe (3♀, CEUFT) • Pindorama, Rio Balsas , 340 m, [11°01’04’’S, - 47°29’08’’W], 28.iii.2023, arm. luz, J. A. Rafael F. F. Xavier F°. (1♀, INPA) • LARVAE • Brazil • Paraná: Jaguariaíva, Parque Estadual do Cerrado , 13–16.xii.2022, Coleta Manual (larvas), Tavares, L; Silva-Junior, JO; Machado, RJP (1 L, DZUP) • Distrito Federal: Brasília , Parque Ecológico das Sucupiras, 09.ix.2022, Coleta Manual (larvas), Tavares L. (1 L, DZUP) •

Additional data : ARGENTINA • Neuquén: Lácar [40.2S, 71.5W], I.1994, (USNM-GBIF) • Salta: Cafayate , Tacochuya , 6.iv.1968, Stange-Williams ( FSCA) • Mendoza: Pérez Cuesta , [32.9S, 68.8W], (MCZ-GBIF) • BRAZIL • São Paulo: Campinas , iii.1924 ( FSCA) • Villa Americana , ii.1924 ( FSCA) • Mato Grosso: Barracão Queimado , xi.1960 ( FSCA) • Cuiabá, Coxipó da Ponte , [15.5S, 55.8W], (MCZ-GBIF) • PARAGUAY • Alto Paraná: Estancia Dimas , 650’, [25°33’S, 55°13’W], 7–10.ii.2008, U. Dreschel coll. ( FSCA) • Caazapá: Colonia Neufeld , 390’, [26°28’S, 55°55’W], 24.ix–2.xi.2008, U. Dreschel coll. ( FSCA) • Canindeyú: Carapa , 230’, [24°22’S, 54°23’W], 1–10.xii.2007, U. Dreschel coll. ( FSCA) • Presidente Hayes : Teniente Martinez [25.7S, 56.9W] (MCZ-GBIF) .

Name-bearing types. Formicaleo irrigatus : ( Fig. 8A–F ) Holotype (by implicit monotypy), female, EMAU, examined. Type locality: Brazil, São Paulo. From original description: “fem. Long. corp. 25, alar. ant. et post. 21 mill.—Patria: Sao Paulo Brasiliae.” From Stange (2004; 2010): “ Holotype female, São Paulo, Brasil (EMAU)”. Condition: good; both antennae missing the flagellum, both eyes sunken in, right mesothoracic leg missing after the coxa, abdomen broken at the third somite, but glued on a tab under the specimen. Brachynemurus meridionalis : ( Fig. 9A–H ) Holotype (by implicit monotypy), female, MCZ, examined. Type locality: Paraguay, Paraguari, Sapucaí. From original description: “From Sapucay, Paraguay, 30 Jan.”. From Stange (1967): “Banks, J. N. Y. ent. Soc. 17:146. Hol. ♀, Sapucay, Paraguay, I. 30 (Cambridge!).”. From Stange (2004; 2010): “ Holotype female, Sapucay, Paraguay, 30.I. (MCZC!)”. Condition: good; right antenna missing the pedicel; both eyes slightly broken; meso and metathorax partially covered in brown substance; missing the right mesothoracic leg; left meso and metathoracic wings with a few red and yellow droplets and hole below the left mesothoracic wing.

Brachynemurus argentinus : ( Fig. 10A–H ) Lectotype (by subsequent designation by Stange (1961)), male, MCZ, examined. Type locality: Argentina, Mendoza. From original description: “From Mendoza, Argentina (Jensen-Haarup Expedition, through Dr. Petersen.)”. From Stange (1961): “ Lectotype — Mendoza, Argentina, Jensen-Haarup Expedition (♂). Lectoparatypes—same data (2 ♀ ♀). The lectotype has a red “type” label. M.C.Z. #10614.” From Stange (1967): “Banks, Proc. ent. Soc Wash. 12:146. Lect. ♀, Mendoza, Argentina, Jensen-Haarup (Cambridge!)”. From Stange (2004, 2010): “ Lectotype male, Mendoza, Argentina, Jensen-Haarup (MCZC), designated by Stange, 1961: 675”. Condition: good; covered in scales; both antennae missing the flagellum; right mesothoracic leg broken at the coxa, left mesothoracic leg seemingly missing the tarsus; missing the right mesothoracic wing. The species was described as B. argentinus , however the identification label presumably made by Banks says “ Brachynemurus argentinicus ”.

Clathroneuria amazonica : ( Figs. 11 A–B ) Holotype (by implicit monotypy), male, NHMW, examined. Type locality: Brazil, Amazon. From original description: “Patria. Brasil: «Amazon» (Mus. de Viena).”. From Esben-Petersen (1920): “The type-specimen (♂) from Amazon agrees exactly with specimens of Brachynemurus meridionalis Banks , determined by Banks himself.” From Stange (1967): “Navás, Brotéria 12:215. Hol. ♂. Amazon, Brasil (Viena!). ( Clathroneuria ).”. From Stange (2004, 2010): “ Holotype male, Amazon, Brazil (NHMW!).”. Condition: good; both antennae missing the flagellum; missing the right metathoracic wing, abdomen broken at the third somite.

Austroleon alienus : ( Figs. 12 A–B ) Holotype (by implicit monotypy), female, MLPA, examined. Type locality: Argentina, San Luis, Alto Pencoso. From original description: “Patria. República Argentina:Alto Pencoso, Provincia de San Luis, Febrero de 1914, Dr. Carlos Bruch (Mus. de La Plata).”. From Stange (1967): “Navás, Brotéria 13:9. Hol ♀, Altos Pencoso, San Luis (“ II.1914 ”) C. Bruch (La Plata!). ( Austroleon ).”. From Stange (2004; 2010):” Holotype female, Alto Pencoso, San Luis ( Argentina), II.1914, C. Bruch (MLPA!)”. Condition: good; both eyes sunken in, both antennae missing the flagellum.

Austroleon clavatus : Lectotype (by subsequent designation by Stange 1967), female, MACN, not examined, photos not available. Type locality: Argentina, Rio Negro, Tapiluque. From original description: “Patria. Rep. Argentina: Tapiluque (Río Negro), 5 de marzo de 1915; Corral Chico (Río Negro), 9 de marzo de 1915, R. Lehman Nink leg. (Mus. de La Plata).”. From Stange (1967); “Navás, Mem. R. Acad. Barcelona 12:126. Lect ♀, Tapiluque, Río Negro III.5.1915 L.N. leg. (Buenos Aires!). ( Austroleon ).” From Stange (2004; 2010): “ Lectotype female, Tapiluque, Rio Negro, Argentina, 5.III.1915, L. Nink (MACN!)”.

Austroleon antennatus : Holotype (by implicit monotypy), male, probably lost, not examined. Type locality: Argentina, Chubut, Puerto Madryn. From original description: “Patria. República Argentina: Puerto Mastryn, Chubut (Col. m.) Un ejemplar bastante deteriorado.”. From Stange (1967): “Navás. Rev. Acad. Madrid 17:295. Hol. ♂, Puerto Madryn (Mastryn!), Chubut (Zaragoza!) ( Austroleon ).”. From Stange (2004, 2010): “ Holotype male, Puerto Madryn (Mastryn sic!), Chubut (Zaragoza?destroyed)”.

Austroleon sectorius : Syntypes,male, not located and probably lost, not examined.Type locality: Argentina, Córdoba, Alta Gracia, La Granja. From original description: “Patria. República Argentina: La Granja (Alta Gracia), provincia de Córdoba, 1–8 de abril de 1920, Carlos Bruch (Col. Bruch y mía).”. From Stange (1967): “Navás, An. Soc. cient. argent. 90:58. Sintipos: La Granja, Alta Gracia. Córdoba. IV.1–8.1920, C. Bruch (¿perdido?). ( Austroleon )”. From Stange (2004, 2010): “ Syntypes, La Granja, Alta Gracia, Cordoba, 1–8.IV.1920, C. Bruch (not located).”

Austroleon latipennis : ( Fig. 13A–G ) Holotype (by implicit monotypy), male, MCZ, examined. Type locality: Brazil, Mato Grosso, Chapada dos Guimarães. From original description: “From Chapada, Brazil (H. H. Smith). Type, M. C. L., 12032”. From Stange (2004, 2010): “ Holotype, Chapada, Brazil, H.H. Smith (MCZC).”. Condition: good; except for a hole in the thorax between the left fore and hindwing and abdomen broken in the third somite.

Austroleon graphonotus : ( Fig. 14A–B ) Holotype (by implicit monotypy), female, MZBS, examined. Type locality: Argentina, Rio Negro, Arroyo Sanchez. From original description: “Patria. Argentina: Río Negro, Arroyo Sánchez. B.–D.”. From Stange (1967): “Navás, Rev. Acad. Madrid 31:13, fig. 27. Hol. ♀, Arroyo Sánchez, Río Negro (Barcelona!) ( Austroleon ).”. From Monserrat (1985): “ Austroleon graphonotus N., 1934, TIPO, Río Negro, Argentina ”. From Stange (2004, 2010): “ Holotype female, Arroyo Sanchez, Rio Negro, Argentina (MZBS!)”. Condition: good; antennae missing the flagelum; eyes broken; all legs present, but it’s not possible to determine if they are intact; right meso and metathoracic wing torn and missing after the first half; left mesothoracic wing stained with brown substance in the base.

Austroleon stictogaster Navás, 1913 ( Fig. 15A–C ): Holotype (by implicit monotypy), female, MZBS, examined. Type locality: Paraguay, Cordillera, San Bernardino. From original description: “Fundort: Paraguay, San Bernardino, 28.II K. Fiebrig leg. Type Zoolog. Staatssammlung, München.”. From Navás (1928): “El tipo es del Paraguay ”. From Monserrat (1985): “ Austroleon stictogaster N., 1913, TIPO, San Bernardino ( Paraguay) ”. From Stange (2004, 2010): “ Holotype, San Bernardino, Paraguay, 28.II., K. Fiebrig (MZBS!)”. Condition: good; both antennae broken; right meso and metathoracic wing torn in the middle, but glued in a tab under the specimen, abdomen broken at the third somite. Austroleon stictogaster was synonymized under Ar. irrigatus by Stange (2004). However, it is clearly not a synonym of Argentoleon irrigatus . The specimen lacks the characteristic dark brown patterns in the thorax, as well as the Banksian lines in the wings and the brown marks at forks and crossveins in the mesothoracic wing. Furthermore, the specimen is lighter colored with a thoracic pattern of light brown with dark brown marks in the margins and its head is proportionally smaller in relation to its body. In fact, it presents all the major characteristics of Austroleon immitis (Walker) , and by this reason, we are herein considering Austroleon stictogaster as a junior synonym of Austroleon immitis .

Life history

A total of 25 larvae were collected at the Parque Estadual do Cerrado ( Fig. 17 A–C ), they were all found in similar microhabitats and were brought to the laboratory for rearing. Among the collected larva, all but four were at the third instar, while half of the remaining larvae were at the second instar and the other half at the first. The first pupation happened 46 days after the collection event and the adult emerged after 33 days. Overall, out of the 25 larvae, nine pupated but only seven adults emerged (4 ♂ and 3 ♀). Including individuals from other collection events, the duration of the pupa stage ranged from 31 to 40 days, lasting on average 36.4 days. The two pupae that died started pupating over a year after the collection event and pupated after near 13 days of trying and failing to spin a cocoon. Additionally, five individuals died as larvae, three of them over one year after their collection day and one which was euthanized for the description. The larvae mostly buried and kept still, however, in a few occasions when it was raining or in the imminence of rain, the larvae would get agitated, and could be seen moving in the sand or pacing on top of it, even being protected from the rain at the laboratory.

Distribution

The species has records for seven different countries in South America ( Fig. 16 ): Argentina, Bolivia, Brazil, Venezuela, Paraguay and Uruguay, with records prevenient from museum specimens and the literature; the records from Colombia are based solely on observations posted at the iNaturalist and should be confirmed in the future by collected specimens. The previous record for Colombia comes from the Austroleon stictogaster type, which is now synonymized under Austroleon immitis , however, currently there are also two iNaturalist observations of Argentoleon irrigatus from Colombia. Due to their unusual and seemingly isolated position, the specimens prevenient from Venezuela were examined thorough photographs and their identification was confirmed. In Brazil, the species was recorded for the first time in the states of Bahia, Goiás, Maranhão, Mato Grosso do Sul, Rondônia, Santa Catarina and Tocantins.

Figures

FIGURE 1. Argentoleon irrigatus a) head, thorax and legs, lateral; b) thorax, dorsal; c) head, frontal. Scale bar: a = 1 mm, c = 0.5 mm.

FIGURE 2. Argentoleon irrigatus, male dorsal habitus. Scale bar = 2 mm.

FIGURE 3. male genitalia. a) posterior; b) ventro-posterior; c) lateral; d) ectoproct, lateral. Abbreviations: gon = gonarcus; par = parameres; med = mediuncus; lls = lobe-like sclerites; gs = genital sac Scale bars: a–c = 0.2 mm, d = 1mm.

FIGURE 4. female terminalia. a) lateral; b) ventral.Abbreviations: ag = anterior gonapophyses; lg = lateral gonapophyses; pg = posterior gonapophyses; ect = ectoproct; vmp = ventromedial plate. Scale bars: a= 0.2 mm, b = 0.5 mm.

FIGURE 5. Argentoleon irrigatus larvae. a) dorsal; b) ventral. Scale bar: 1 mm.

FIGURE 6. Argentoleon irrigatus and Austroleon immitis larvae a) Ar. irrigatus head, dorsal; b) Ar. irrigatus head, ventral; c) Au. immitis head, dorsal; d) Au. immitis head; ventral; e) Ar. irrigatus abdomen, ventral. Abbreviations: sld = sphere-like dolichasters; op = odontoid process. The arrows indicate the difference in mandible width. Scale bars: a–d = 0.5 mm; e = 0.2 mm.

FIGURE 7. Argentoleon irrigatus pupae. a) Female pupa, 5 days old; b) male pupa; 28 days old. The arrow points at the bigger ectoproct in the male. Scale bars: 1 mm.

FIGURE 8. Formicaleo irrigatus Gerstaecker, holotype. a) habitus; b) head and thorax, lateral; c) head, frontal; d) labels; e) thorax, dorsal; f) abdomen and terminalia, lateral. Scale bars = 1 mm.

FIGURE 9. Brachynemurus meridionalis Banks, holotype. a) habitus, lateral; b) head and pronotum, dorsal; c) habitus, dorsal; d) forewing; e) hindwing; f) head and pronotum, lateral; g) head, frontal; h) labels. Scale bars: a, c–e = 2 mm, b and f = 2 mm, g = 0.5 mm.

FIGURE 10. Brachynemurus argentinus Banks, lectotype. a) habitus, lateral; b) habitus, dorsal; c) head, frontal; d) head and pronotum, lateral; e) head and pronotum; dorsal; f) forewing; g) hindwing; h) labels. Scale bars: a, b, f and g = 2 mm; c–e = 0.5 mm.

FIGURE 11. Clathroneuria amazonica Navás, holotype. a) habitus, dorsal; b) labels. Scale bar = 0.5 cm.

FIGURE 12. Austroleon alienus Navás, holotype. a) habitus, dorsal; b) labels.

FIGURE 13. Austroleon latipennis Banks, holotype. a) habitus, dorsal; b) head and prothorax, lateral; c) head, anterior; d) habitus, lateral; e) forewing, dorsal; f) labels. Scale bars: a, d and e = 2 mm; b–c = 0.5 mm.

FIGURE 14. Austroleon graphonotus Navás, holotype. a) habitus, dorsal; b) labels.

FIGURE 15. Austroleon stictogaster Navás, holotype. a) habitus, dorsal; b) tab with wing fragments; c) labels.

FIGURE 16. Distribution of the genus Argentoleon.

FIGURE 17. Argentoleon irrigatus collection sites. a–c) Collection site in Jaguariaíva; c) Female laying eggs; d–f) Collection site in Brasília.