Ascalorphne impavida ( Walker, 1853 )

Schuster, Phillip Alves & Machado, Renato José Pires, 2022, Insights on the evolution of Ululodini (Insecta: Neuroptera: Myrmeleontidae: Ascalaphinae), focusing on the systematics of the genus Ascalorphne Banks, 1915, Revista Brasileira de Entomologia (e 20220070) 66, pp. 1-21 : 10-12

Ascalorphne impavida ( Walker, 1853)

Figs. 1B , 4-6 , 7A , 14D , 18A .

Ascalaphus impavidus Walker, 1853:443 (original description): Hagen, 1861:326 (list/ taxonomy); Hagen 1866:383 (cited); McLachlan, 1871:252 (synonym/ taxonomy); van der Weele,1908:141 (redescription/ distribution); Navás 1912b:80 (list); Penny 1977:11 (list); Penny, 1981a:637 (taxonomy).

Orphne impavida ( Walker, 1853) : McLachlan, 1871:252,401 (new combination/ taxonomy); van der Weele, 1908:141 (cited); Penny 1977:11 (list); Navás, 1912 a:223 (key); Navás, 1912b:80 (key/ list); Navás, 1913:64 (key/ redescription).

Amoea impavida ( Walker, 1853) : Oswald, 2022 (cited).

Colobopterus impavidus ( Walker, 1853) : Hagen, 1866:399 (new combination).

Ascalorphne impavida ( Walker, 1853) : Banks, 1915:350 (new combination); Navás, 1920:37 (cited); Navás, 1928:109 (cited); New, 1971:75 (cited); Penny, 1977:11 (list); Penny, 1981a:637 (redescription); Whittington, 2002:377 (distribution); Ardila-Camacho et al., 2019:3 (list); Oswald, 2022 (catalog); Machado & Martins, 2022 (catalog).

Ascalaphus intempestivus Walker, 1853:444 (original description): Hagen, 1861:326 (list/ taxonomy); Hagen 1866:383 (cited) McLachlan, 1873:252 (synonym/ taxonomy); van der Weele, 1908:141 (cited); Penny 1977:11 (list); Penny, 1981a:637 (taxonomy).

Colobopterus intempestivus ( Walker, 1853) : Hagen, 1866:399 (new combination).

Diagnosis. Tuft of long white seta on the dorsolateral margin of the clypeus. Males with a thick row of short setae on the surface of flagellomeres 8-23. Piriform antennal club, brown with yellowish tip. Thorax brown, pleural region with white pilosity subtler than in A. macrocerca , and yellow spots in the pleural region may or may not be visible, if visible, less prominent than in A. macrocerca . Coxae, trochanter and femur brown and/or yellowish with dark apex, tibia uniformly brown. Ectoproct of both sexes with circular black macula. GPC smaller than in A.macrocerca and differing from A.umbrina mainly by the wide GX 11 in lateral view and in dorsal view with the narrow GX9.

Dimensions (mm). Body: ♀ 21.08 ± 3.42/ ♂ 21.3 ± 2.32; antennae: ♀ 26.2 ± 0.88/ ♂ 25.38 ± 0.84; forewing: ♀ 24.8 ± 1.80/ ♂ 22.48 ± 1.07; hind wing: ♀ 22 ± 1.35/ ♂ 20.4 ± 1.01.

Head ( Figs. 4 , 5 C-E). As in the description of the genus, but differing in the following characteristics: tufts of long white setae on the dorsolateral margin of the clypeus; yellowish-brown labial palp; antennae brown, with black rings,, 38-39 flagellomeres, long, reaching the apex of the forewing; in males flagellomeres 8-23 have a dense row of short setae, but in females these setae are simple and elongated; antennal club pyriform, dorsally brown and ventrally brown with yellowish-brown apex.

Thorax ( Figs. 4 , 5B, C ). Pleural region brown with yellow macules and long white setae and some black; pronotum brown and medially with a light brown/yellowish macula; prescutum brown; scutum brown laterally and medially yellowish; scutellum brown with two yellowish maculae. Leg. Coxae and trochanter light brown and/or yellow; femur brown at the base and dark at the apex; tibia uniformly brown; tarsomeres brown with black apices or completely black and with short black setae.

Forewing ( Figs. 4 , 5A ). Wing membrane hyaline, veins brown, pterostigma black, with 5 crossveins; apical area after fusion of Sc+R with 3-4 cell lines; RP with 4-5 branches; MA long, without branches, extending beyond the 3rd bifurcation of RP; CuA+MP with 9-10 branches; cubital area with 3-4 cell lines. Hind wing ( Figs. 4 , 5A ). Membrane and veins as above; smaller pterostigma, black, with 4 crossveins; MA straight, ending at the 4th RP branch; CuP straight, extending to the first RP branch; cubital area with 2-3 cell lines.

Abdomen ( Figs.4 , 5B, C ). Shorter than wings, brown or dark brown. Terminalia and genitalia ( Fig. 6 ). ♂: Ectoproct oval, with a circular macula; T9 brown, triangular, in lateral view, and with longer setae on the ventral portion; GPC with GX1, in lateral view, wide and GX9, in dorsal view, narrow. ♀: Ectoproct oval, with circular macula.

Morphological variation. The eye spots mentioned by Penny (1981a), may or may not be present, as in the other species. This species can present a great color variation, from dark brown to brown/yellowish individuals and with practically absent dorsal yellow spots or with conspicuous spots.

Temporal distribution. Based on dates on the labels of the analyzed specimens, A.impavida can be collected almost throughout the entire year, with a peak of occurrence between February and May, with the month of March presenting the largest number of individuals collected; Penny (1981a) indicated that in Maranhão ( Brazil) the temporal distribution of adult’s peaks around June, however he had few available specimens.

Geographic distribution ( Fig. 12 ). Brazil: Bahia *, Espírito Santo *, Goiás *, Maranhão ( Penny, 1981a), Mato Grosso do Sul *, Mato Grosso *, Minas Gerais *, Pará ( Walker, 1853; Penny, 1981a), Pernambuco *, Piauí *, Rio de Janeiro *, Rio Grande do Norte *, Tocantins *. Bolivia: Santa Cruz *. Paraguay:Campito* ( Háva, 2019). iNaturalist. Brasil: Amazonas*, Ceará *, Pernambuco *, Sergipe *, Tocantins *. Argentina *: Missiones*.

Primary type ( Fig. 4 ). Ascalaphus impavidus , 1 ♂ NHMUK, high resolution images analyzed. Holotype (by monotypy): ♂ A.impavidus . 1 ♂ (NHMUK013803955); type locality: Santarém, Brasil; labels: 1) Brasil, Santarem, 2) impavidus , 3) impavidus Wlk , 4) NHMUK013803955; condition: excellent, abdomen slightly covered with fungus.

Primary type ( Fig. 5 ). Ascalaphus intempestivus , 1♀ NHMUK, high resolution images analyzed. Holotype (by monotypy): ♀ A.intempestivus , type locality: Santarém, Brasil; l abels: 1) Brasil,Santarem,2) intempestivus , 3) intempestivus Wlk , 4) NHMUK012803956; condition: excellent, abdomen covered with fungus.

Examined specimens. Brazil – Bahia: • Miguel Calmon, Mulungu do “chiola”; 9-16.vi.2021; 1 ♂; puçá; L. Tavares; UFBA#N610. Distrito Federal: 27.iii.1964; 1 ♀; C. E. & E. S. Ross; CASENT8472557. Piauí: • Piracuruca, Parque Nacional de Sete Cidades,Posto do ICMBIO; 04º05’57”S 41º42’34”W; 17-19.vii.2012; 2 ♀; luminosa; J.S. Pinto Junior & J.A. Rafael, cols. • idem, 1 ♂; 18-20.iv.2012; J.T.Camara & J.S. Pinto Junior cols. Mato Grosso: • Campo Novo do Parecis, BR 13; arbusto; 13.x.2017; 1 ♀; A.L. Klein.; DZUP • Cuiabá; 01.xii.1993; 1 ♀; C.T. Homes; UFMT • Chapada dos Guimarães, Rio Claro; 10.xii.1984; 1♀; Sebastião Marcolino; UFMT • Santo Antônio do Leverger, Fazendo Santa Maria; 09.iii.1992; 1 ♂; L. Moreira; UFMT • São Vicente; 19.X.1988; 1 ♀; D. Figueiredo; UFMT • Barra do Bugres, Reserva Araras; 22.ii.1987; 1 ♀; Y.O. Willis; UFMT. Mato Grosso do Sul: • “Corumbé”; 1.ii-8.iii.1966; 1 ♀; E. Golbach; FSCA00094852 • idem;?. i.1971; 1 ♀; E. Golbach; FSCA00094861 • Rio Brilhante; 21-27. i.1971; 1 ♀; V. O. Becker leg.; DZUP. Maranhão: • Balsas 525m; 8°38’S 46°43’W; 15.II.99; 2 ♀ e 4 ♂; Coleção EMBRAPA; CPACNº13.776 • Caxias, Povoado Boa Vista; 20.v.2004; 1 ♀; F. Limeira-de-Oliveira et al. cols. • idem, Reserva Ecológica Inhamum; 21.iv.2005; 1 ♂; varredura; F. Limeira-de-Oliveira. • idem, Shalon; 30-31.iv.2004; lençol e luz mista; 1♀ e 1 ♂; F.Limeira-de Oliveira, col. • Mirador, Parque Estadual Mirador, Posto Avançado do Mel; 06°43’50”S 44°58’59”W; 01-10. iii.2014; luminosa; 1 ♀ e 2 ♂; F. Limeira-de-Oliveira, L. L. M. Santos & T. T.A.Silva • idem, Base da Geraldina; 06°37’25” S 45°52’08”W; luminosa; 22.ii-01.iii.2009; 6 ♀ e 2 ♂; M.B. Aguiar-Neto & M.J.A, Holanda cols.; CZMA • idem, 29-31.i.2006; 1 ♀; F. Limeira-de-oliveira, col. • idem, 20-31.xii.2006, 1 ♂; F. Limeira-de-oliveira, col. • idem, 20-24.xii.2006; 1 ♂; R.O. Souza, J.C. Silva, et al. • idem, 17-21.ii.2007; 1 ♂; J.C Silva, M.J. Holanda. • idem, 20-23.iv.2007; 3 ♀ e 1 ♂; F. Limeira de Oliveira col. • idem, 20-23.ix.2007; 1 ♂; F. Limeira de Oliveira col. • idem, 08-13. iii.2008; 1♀; J.C.Silva & F.Limeira-de-Oliveira • idem, 07-14.v.2010; 1 ♀; J.S. Pinto Junior & M.M. Abreu. • idem, luminosa móvel; 07-15.?.2010; 1♀; L.L. Reis & L.M. Oliveira • idem, 07-14.iii.2013; 2 ♀ e 3 ♂; F. Limeira-de-Oliveira, A.A. Silva & C.F. Barros • idem, armadilha Malaise; 08-14. iii.2013; 1 ♂; F. Limeira-de-Oliveira, L.L.M. Santos & L.S. Santos • idem, Base do Mosquito; 04-08.ii.2011; 1 ♂; armadilha luminosa; F. Limeira-de-Oliveira • São Luís, APA Itapiracó; 28-31.v.2008; 1 ♂; luminosa; M.B. Aguiar-Neto, S.J.C.Silva. Minas Gerais: • Lavras, Bairro Centenário, dentro de casa 21°14’43.0”S 44°59’29.6”W; 28.ii.2014; 1 ♂; ativa; P.G.Dias leg.; UFMG INE2200019. Pernambuco: • Triunfo, Canaã Recanto da Laje, 07°52’28.5”S 38°08’15.3”W; 02.v.2018; 1♀ e 1 ♂; bandeja; Cavalcante Pereira; UFBA#356 • idem, Riacho da Laje; 07°52’28.2”S 38° 08’13.6”W; 02.v.2019; 1 ♀; bandeja; Cavalcante Pereira; UFBA#N352. Rio Grande do Norte: • Alto dos Rodrigues, “abuaser”, Arg.189; 17-18.vii.1996; B.L. equipe; 7-Margem de um declive rochoso; UFBA#N360 • Martins, Pousada Martinense; 04.v.2013; coleta manual; Ferreira, V.D.S; UFBA #N354 • Natal, Campus UFRN- CB; 13.v.2012; 1 ♂; Neto P.F.C., Dantas A.C.; UFBA#358 • idem, Capim Macio; 09.v.2012; 1 ♂; Salvino, S.C.A.; UFBA#355 • Serra de São Bento; 7.v.2006; 1♂; Freire, A.B.F.; UFBA#357 • Serra Negra do Norte, Servida, sede; iv.2005; 4 ♂ API; luz fluorescente; Varella, A.A.; UFBA#N351 • idem, ESEC Seridó, sede,?. v. 2005; 1 ♀; sob luz florescente; UFBA#N353 • idem, 25.iv.2009; 1 ♂; API (noite); Varela-Freire; UFBA#N361 • idem, Açude Campos I, margem ocidental; 05-06. vi.1997; 1♂; B.L.Varela-Freire, A.A.; UFBA#359. Tocantins: • Santa Isabel do Morro, Ilha do Bananal;?. vi.1961; FSCA00094864. Bolívia - Santa Cruz: • Buena vista, 380m; 17°27’69S,63°39’63”W; 20.ii.1999; 1 ♂; L. Stange; FSCA00094859.

Comments. The larvae are unknown and almost nothing is known about the biology of this species, including its feeding habits. Interestingly, Tjeder (1992) comments that, possibly, Ascalaphinae would not feed on Lepidoptera, due to the absence of scales in gut content. However, in two occasions during this work, collected specimens of A. impavida were observed firmly grasped and with the mandibles attached to disarticulated legs of a lepidopterans; it is possible, however, that this observation is an artifact,since the insect may have clung to another when captured in alcohol. Another specimen of A. umbrina was collected in the same situation, but clearly with the leg inside its mouth, reinforcing the hypothesis that butterflies/ moths might be an important dietary component of these owlflies. Also, the strong jaws of Ululodini indicate the ability to feed on hard-bodied prey, and by dissecting the crop of A. impavida it was observed a Coleoptera femur (Scarabaeoidea).

In the original description Walker (1853) described both sexes of A. impavida as two distinct species, which was later noticed and synonymized by McLachlan (1871). After reviewing the type material of Ascalaphus impavidus and Ascalaphus intempestivus , in addition to several specimens of both sexes of A. impavida , this synonymy is herein confirmed.

Through the work ofPenny (1981a) A.impavida was the only species of the genus to receive an updated description, in addition to being the last taxonomic work containing Ascalorphne . The author also indicated that A. impavida would be confined to the southeastern region of the Amazon basin, but based on the data presented here it is evident that this is a widely distributed species ( Fig. 12 ).

Also , it is interesting to note that the distribution of this species reaches the Bolivia – Brazil border (1 ♂, FSCA00094852 and 1 ♀, FSC 00094861, both collected in Corumbá, Mato Grosso do Sul) and a single individual was collected in Bolivia (1 ♂ FSCA 00094859, Santa Cruz), which led to the suspicion that it was, in fact, an A. umbrina specimen with a brown coloration, however the typical coloration of A. impavida and the narrow GX9 were observed in this specimen, indicating this overboard distribution .

According to van der Weele (1908) in A. impavida the veins of the wing apex are not as “dense” as in A. macrocerca and are rarely bifurcated, however, the number of vein bifurcations are interspecifically variable and cannot be used with confidence for species differentiation.

Another noteworthy comment is regarding the work by Háva (2019) which indicated a new of record of Ameropterus selysi (van der Weele, 1909) from Paraguay; however, the photo presented in the article belongs to a female Ascalorphne , probably A. impavida based on coloration.

Figures

Figure 1 Adult males of Ascalorphne macrocerca (Burmeister, 1839) (A) and Ascalorphne impavida (B) (Myrmeleontidae:Ascalaphinae: Ululodini).Photos: Lucas Rubio.

Figure 4 Ascalaphus impavidus Walker,1853 (=Ascalorphne impavida (Walker,1853)), holotype, male, NHMUK:A) Fore and hind wing; B) Body, dorsal; C) Body, lateral; D) Antennal club; E) Head, frontal; F) Labels.

Figure 5 Ascalaphus intempestivus Walker, 1853 (= Ascalorphne impavida (Walker, 1853)), holotype, female, NHMUK: A) Fore and hind wing; B) Body, dorsal; C) Body, lateral; D) Antennal club; E) Head, frontal; F) Labels.

Figure 6 Terminalia/genitalia of Ascalorphne impavida (Walker,1853). Male (A-D):A) Terminalia,lateral view;B) Idem,ventral view;C),Gonarcus-paramere complex,ventral view; D) Idem,lateral view.Female (E-F):E) Lateral view;F) Ventral view.D:distivalve, E: ectoproct, GX: gonocoxite, L: liguella, P: pelta, PU:pulvinus, S: sternite T: tergite.

Figure 7 Abdomen of gravid females of A) Ascalorphne impavida (Walker, 1853); and B) Ascalorphne macrocerca (Burmeister, 1839), showing the fertilized eggs and the repagula.E:fertilized eggs, R: repagula.

Figure 12 Distribution map of the genus Ascalorphne Banks, 1915 (Myrmeleontidae: Ascalaphinae:Ululodini).

Figure 14 Labium and labial palps of the neotropical tribes of Ascalaphinae (Neuroptera: Myrmeleontidae). A) Dimares elegans (Perty, 1833); B) Fillus amazonicus (Machado & Rafael, 2011); C) Haploglenius costatus (Burmeister, 1839); D) Albardia furcata van der Weele, 1903; E) Ascalorphne impavida (Walker, 1853).

Figure 18 Antennal pilosity of males ofAscalorphne Banks,1915 and Ameropterus dissimilis (McLachlan,1871). A)Ascalorphne impavida (Walker,1853); B)Ascalorphne umbrina (Gerstaecker, 1894); C) Ascalorphne macrocerca (Burmeister, 1839); D) Ameropterus dissimilis.