Heilus inaequalis ( Boheman, 1836 )

Lira, Aline De Oliveira, Sousa, Wesley Oliveira De, Rosado-Neto, Germano Henrique, Santos, Geane Brizzola Dos & Marques, Marinêz Isaac, 2020, A review of the South American species of Heilus Kuschel, 1955 (Curculionidae Molytinae: Molytini: Hylobiina) with emphasis on those from Brazil, Zootaxa 4861 (2), pp. 151-187 : 157-160

Heilus inaequalis ( Boheman, 1836)

( Figs. 3 A–F, 4, 18A–B, 19A, 20A, 21A, 22A, 23B)

Heilipus inaequalis Boheman, 1836:211 (description). Boheman 1843:94 (redescription); Kirsch 1889:53 (occurrence); Black-

welder 1947:818 (checklist). Hilipus inaequalis: Gemminger & Harold 1871:2427 (catalog); Dalla Torre et al. 1932:31 (catalog). Hilipus obesulus Pascoe, 1881:81 (description); Dalla Torre et al. 1932:31 (catalog); Blackwelder 1947:819 (catalog). Heilus inaequalis: Kuschel 1955:297 (new combination); Wibmer & O’Brien 1986:145 (checklist); Alonso-Zarazaga & Lyal

1999:201 (catalog).

Type Material. Heilipus inaequalis Boheman, 1836 : we examined the images of the female holotype ( Fig. 4 ) bearing the following labels: “Typus; Brasilia; Coll. v. Winth; NHRS–TOBI 000003224” and the male allotype bearing the following labels “ Allotypus; Brasilia; Coll. Westerman; NHRS–TOBI 000003225”, both deposited in the NHRS.

Redescription. Length (pronotum and elytra) of males and females, 10.6 to 13.8 and 11.0 to 13.7 mm, respectively. Integument light brown to black, opaque. Rostrum about 1.7 times longer than length of pronotum.

Thorax. Prothorax ( Figs. 4 A–B) covered by rounded granules sparsely arranged side by side on pronotal disc and sides; pronotal disc with fine, decumbent, yellowish, brown or black scales, with or without two rows of whitish and flattened scales on sides of pronotum; lateral region of metasternum covered with elongate, yellowish scales; median region covered with tiny sparse scales of the same color inserted into micro punctations, and row of larger, depressed scales on anterior margin, near joint with mesosternum. Elytra ( Figs. 4 A–B) about 1.6 times longer than width between humeri; moderately convex, wider than pronotum at base; elytral striae with strong punctures, coarse in basal half, progressively finer toward apex; striae 8 and 9 with deeper punctures; humeri covered with developed granules; transverse granules densely distributed in basal region of interstriae, sparse and smaller from median region to apical elytral calli, tiny granules present or absent at apex; interstriae 9 and 10 slightly convex; clusters of fine, yellowish scales distributed anteriorly to granules of interstriae; whitish, oblong scales, sparsely distributed among clusters of yellowish scales; fine, whitish scales in region of humeri; scutellum densely covered with whitish, short, flattened scales; macula of velvety black scales surrounded on anterior and posterior borders by whitish scales of same texture forming small, elongate macula between striae 3 and 4 of median region of elytra. First abdominal ventrite about 1.5 times wider than second; second ventrite wider than third; ventrite 5 about three times wider than long. Male genitalia. Median lobe of aedeagus ( Figs. 18 A–B) with straight apex and short sparse setae, slightly rounded on sides, parallel sides, slightly curved in lateral view; median lobe apodemes about 1.4 times longer than median lobe; internal sac sclerite ( Fig. 19A ) strongly sclerotized with two acute projections at entrance of the ejaculatory duct. Tegmen ( Fig. 20A ) with elongate, rounded dorsal lobes at apex, elongate manubrium, about 1.2 times longer than dorsal lobes. Female genitalia ( Fig. 21A ). Coxites with short setae in apical third and at insertion region of styli; cylindrical styli, with long dense setae at apex. Spermathecal duct short inserted into posterior margin of bursa copulatrix. Spermathecal capsule ( Fig. 22A ) slightly rough on margin near collum; ramus poorly developed; collum curved; cornu well-developed, with rounded apex. Spermathecal gland about 4.9 times longer than spermathecal capsule, dilated at base, narrowing toward apex.

Dimensions (mm). Males/females, respectively. TBL: 10.6–13.8/11.0–13.7; TRL: 3.1–3.7/3.6–4.7; MHx: 4.6–5.7/4.8–5.6; RAW: 0.7–0.8/0.7–0.8; RBW: 0.7–0.8/0.7–0.8; IO: 0.4/0.4–0.6; PnL: 2.6–3.4/2.7–3.6; PnMW: 3.4–4.4/3.7–4.4; PnBW: 3.3–4.3/3.4–4.1; EL: 7.4–10.0/7.8–10.0; EW: 4.7–6.3/4.8–6.3.

Non-type material examined. In total, 218 specimens were examined. BRAZIL: Pará: Colônia Rio Branco , 1♀, x.1953, F.M. Oliveira ( MNRJ) ; Mato Grosso do Sul: Dourados , 2♀, xii.1975, Lorenzoni ( DZUP) ; Minas Gerais: Mar de Hespanha , 2♀, 23.x.1909, J.F. Zikan ( FIOC) , 1♂, 30.x.1908, J.F. Zikan ( FIOC) ; Espírito Santo: Parque Soretama , 1♀, 1967 ( DZUP) , Rio Itabapoana , 1♂, 18.xi.1908, J.F. Zikan ( FIOC) , Rio Lameco , 1♀, xii.1958 ( DZUP) , Santa Teresa, 1♂, 13.xi.1966, C. Elias ( DZUP) , 1♂, xii.1964, C. Elias ( DZUP) ; Rio de Janeiro: Angra dos Reis , 1♂ ( MNRJ) , 2♂, ix.1931, L.T. ( FIOC) , 1♂, v.1932, L.T. ( FIOC) , 1♂, vii.1936, L.T. ( FIOC) , 1♂, xi.1931, L.T. ( FIOC) , Angra–Jussaral , 1♀, 29.ix.1935, Travassos & Almeida ( FIOC) , 1♂, xi. 1934, Trav. Col ( FIOC) , Corco- vado, 1♂, 1.xii.1966, Moure & Seabra ( DZUP) , 1♀, 13.xii.1988, C.A. Campos Seabra ( MNRJ) , 1♀, i.1968, Alva- renga & Seabra ( DZUP) , 1♂, iii.1952, Alvarenga & Seabra ( MNRJ) , 1♀, iii.1967, Alvarenga & Seabra ( DZUP) , 1♂, x.1938, L.T. ( MNRJ) , 1♂, xi.1967, Alvarenga & Seabra ( DZUP) , E. Dentro, 1♀, 8.v.1937, E.F.C.B ( DZUP) , Fio Cruz, 1♀, 5.x.1932, Cesar Pinto ( FIOC) , Floresta da Tijuca , 1♂, i.1951, C.A. Campos Seabra ( MNRJ) , 1♂, i.1961, C.A. Campos Seabra ( DZUP) , Guaratiba , 1, i.1956, J. H. Guimarães ( MNRJ) , 1♀, xii.1955, J.H. Guimarães ( FIOC) , Imbarie , 1♂, 15.v.1955, H. Ebert ( FIOC) , Itatiaia , 1♀, 10.i.1936, J.F. Zikan ( FIOC) , 1♀, 13.x.1929, J.F. Zikan ( FIOC) , 1♀, 16.v.1930, J.F. Zikan ( FIOC) , 1♂, 18.xii.1935, J.F. Zikan ( FIOC) , 1♂, 2.i.1927, J.F. Zikan ( FIOC) , 1♀, 24.i.1933, J.F. Zikan ( FIOC) , 1♀, 28.i.1930, J.F. Zikan ( FIOC) , 1, 30.iii.1947, J.F. Zikan ( FIOC) , 1♀, 8–13.xii.2004, Monné M.A., Monné M.L. & Mermudes ( MZSP) , Itatiaia , 1♀, xii.1983, ET R. Cunha S. Lopes ( MNRJ) , Jacarepaguá , 1♂, 20.iv.1940, Newton Santos ( MNRJ) , Jussara , 1♀, ii.1934, Dario Mendes ( MNRJ) , 1, A. Mendes ( MNRJ) , 1♂, ix.1984, Dario Mendes ( MNRJ) , 1♀, viii.1934, Dario Mendes ( MNRJ) , Km 47 Est. Rio–São Paulo , 1♂, 10.x.1958 ( MNRJ) , 1♂, 12.ii,1943, Wygodzinsky ( MNRJ) , 1♂, 18.ii.1946, Miranda L. ( MNRJ) , 1♂, 18.iii.1944, H Morit Neto ( MNRJ) , 1♂, 22.x.1942, Braga ( MNRJ) , 1♀, 1♂, 27.ii.1942, P. Vygodzinsky ( MNRJ) , 1♂, v.1954 ( MNRJ) , 1♂, vi.1955, W. Zikán ( MNRJ) , 1♀, x.1947, M. Braga ( MNRJ) , 1♂, xi.1944, Wygodzinsky ( MNRJ) , Miguel Pereira, 1♀, xii.2000, J. Carlos ( CEMT) , Nova Friburgo, 2♀, x.1997, P. Grossi ( CEMT) , Parati , 1♂, 30.xii.2011, M.L. Monné ( MNRJ) , Parque Mac. Itatiaia, 1♂, 5–25.xi.1974, H.S. e M.A. Monné ( MNRJ) , Pau da Fome , 1♂, 27.viii.1940, Newton Santos ( MNRJ) , Petrópolis , 1♀, x.1929, Mario Rosa (MNRJ) , Represa Rio Grande, 1♀, 15.xi.1967, F.M. Oliveira ( DZUP) , 1♀, 30.iv.1967 ( DZUP) , 2♀, ii.1961, F.M. Oliveira ( DZUP) , 1♀, iii.1961, F.M. Oliveira ( MNRJ) , 1♂, ix.1961, F.M. Oliveira ( MNRJ) , 1♂, xi.1961, F.M. Oliveira ( DZUP) , Rest- inga de Marambaia, 1♀, 2.xi.1936, S. Lopes ( MNRJ) , Retiro Pedro do Rio , 1♀, iii.1954, C.A. Campos Seabra ( MNRJ) , Rio de Janeiro, 1, 12.ii.1943, Wygodzinsky Leo ( MNRJ) , 1, 14.x.1943 ( MNRJ) , 1, 5.i.1943, P. Wygodzinsky ( MNRJ) , 1, 5.x.1942, Braga ( MNRJ) , 1♂, i.1959, M. Alvarenga ( MZSP) , 1, ii.1938, C.A. Campos Seabra ( MNRJ) , 1♂, ii.1959, M. Alvarenga ( MZSP) , 1, iii.1932, C.A. Campos Seabra ( MNRJ) , 1♂, iv.1999, A. Bello ( CEMT) , 1♂, iv.2002, A. Bello ( CEMT) , 1♂, ix.1958, Alvarenga & Seabra ( DZUP) , 1, ix.1962, E. Izecksohn ( MNRJ) , 1♂, x.1961, M. Alvarenga ( MZSP) , 1♂, xi.2006, A. Bello ( CEMT) , 1, xii.1948, W. Zikán ( MNRJ) , Rio Murihae , 1♂, 12.xi.1908, J.F. Zikan ( FIOC) , Santa Barbara, 1♂, 19.09.1922 ( FIOC) , Serra dos Órgãos , 2♀, xi.1940, Parko ( MNRJ) , Teresópolis , 1♀, i.1974, A. Bello ( CEMT) , 1♂, xii.1973, A. Bello ( CEMT) , Teresópolis , 1♀, 2– 5.xii.1958, Martinez, Wener, Alvarenga & Seabra ( DZUP) , Km 47 Est. Rio–São Paulo , 1♂, 27.xii.1945 ( MNRJ) ; São Paulo: Barueri , 1♂, 18.ii.1956, K. Lenko ( MNRJ) , 1♀, 1♂, 2.xi.1955, K. Lenko ( MNRJ) , Boracéa–Salesópo- lis, 1♀, 8–14.ii.1959, Travassos, Kloss & Pearson ( FIOC) , São Paulo: Embu , 1♀, x.2001, Reginaldo ( MZSP) , São Paulo: Fte. Andradas, Guarujá , 1♀ ( CEMT) , 1♀, xi.1971, A. Bello ( CEMT) , Juquei , 1♀, ii.1996, J. Carlos ( CEMT) , Santos , 1, ix.1931, Josué Deslander ( MNRJ) , São José Barreiro , 1♀, xi.1968, Alvarenga & Seabra ( DZUP) , São Paulo, 1♀, 16.i.1921, Saúde ( MNRJ) , 1♀ ( MNRJ) , Ypiranga , 1♀, 18.xii.1936 ( MZSP) , 1♀ ( MZSP) , 1♀, xi.1937, F. Lane ( MZSP) ; Paraná: Arapongas , 1♂, ii.1952, A. Maller ( MNRJ) , 1♀, 1, xii.1951, A. Maller ( MNRJ) , Arapoti , 1, x.1937, Anton Maller ( MNRJ) , Céu Azul, 1♂, 20.xii.2014, R.C. Barros & J.P. Bozina ( MZSP) , Guaraqueçaba , 1♀, 8–10.xii.1970, V. Becker ( DZUP) , Londrina , 1♀, 14.xi.1975, Pe. Moure & Rosado-Neto ( DZUP) , Monte Alegre, 1♀, 19.ix.1939, J. Leprevost ( DZUP) , Morretes , 1♀, 20.xii.1984, C.I. I.E. ( DZUP) , Ponta Grossa–Pedreira, 1♀, i.1943 ( DZUP) , 1♀, xii.1943 ( DZUP) , São José dos Pinhais , 1♀, 2♂, i.2014 – vii.2015, Alexandre C. Domahovski ( PUCRS) ; Santa Catarina: Córrego Grande , 1♂, 23.iv.1994 ( MCN) , Corupá , 1♀, A. Maller ( MNRJ) , 3, i.1938, A. Maller ( MNRJ) , 2, i.1953, A. Maller ( MNRJ) , 1, i.1954, A. Maller ( MNRJ) , 1♀, i.1956, A. Maller ( MNRJ) , Corupá , 1♀, iii.1952, A. Maller ( MNRJ) , 1♀, 2, ii.1938, A. Maller ( MNRJ) , 1, ii.1953, A. Maller ( MNRJ) , 3, iii.1938, A. Maller ( MNRJ) , 1♂, iii.1954, A. Maller ( MNRJ) , 1♀, iv.1962 ( DZUP) , 1♀, 1♂, ix.1953, A. Maller ( MNRJ) , 1♀, 2, viii.1953, A. Maller ( MNRJ) , 1, 1♂, x.1937, A. Maller ( MNRJ) , 1♀, 1, x.1953, A. Maller ( MNRJ) , 1♂, x.1954, A. Maller ( DZUP) , 1♂, x.1956, A. Maller ( MNRJ) , 1, xi.1937, A. Maller ( MNRJ) , 1, xi.1952, A. Maller ( MNRJ) , 1, xi.1953, A. Maller ( MNRJ) , Itapema , 1♀, 13–23.i.1976, L. Pereira ( DZUP) , Jaraguá do Sul, 1♀, xi.1937, A. Maller ( MNRJ) , Joinville , 1♂, Bruckner ( MNRJ) , Nova Teutônia, 1♀, 1936, B. Pohl ( MZSP) , 1♀, v.1934, B. Pohl ( MZSP) , 2♀, xi.1966, F. Plaumann ( MZSP) , Nova Teutônia , 1, 1.xii.1952, Fritz Plaumann ( MNRJ) , 1♂, 19.x.1952, F. Plau- mann ( MNRJ) , Nova Teutônia, 1♀, ii.1977, Fritz Plaumann ( DZUP) , 1♂, vi.1975, Fritz Plaumann ( DZUP) , 1♂, x.1973, Fritz Plaumann ( DZUP) , 1♀, xi.1972, Fritz Plaumann ( DZUP) , 2♀, 2♂, xi.1973, Fritz Plaumann ( DZUP) , 1♂, xii.1973, Fritz Plaumann ( DZUP) , Pinhal , 1♀, iv.1958 ( DZUP) , 1, xii.1952, A. Maller ( MNRJ) , Rancho Quei- mado , 1♀, 15–18.xi.1995, L. Moura ( MCN) , S. Bento, 1♂, i.1952, A. Maller ( MNRJ) , 2♀, 1♂, ii.1952, A. Maller ( MNRJ) , 1♀, iii.1952, A. Maller ( MNRJ) , São Bento do Sul, 1♂, 14.i.1975, D.M. Mielke ( DZUP) , 1♂, 25.i.1974, Mielke ( DZUP) , Timbó , 1♀, ii.1963, Dirings ( MZSP) ; Rio Grande do Sul: Barra do Ribeiro , 1♂, 16.v.2003, R.S. Araújo ( MCN) , Eldorado do Sul, 2♂, 05–07.i.2000, A. Franceschini, A. Bonaldo & A. Silva ( MCN) , Maquiné , 1♀, 18–20.vii.2006 ( MCN) , 1♀, 8–10.ix.2005, L. Moura ( MCN) , 1♂, 9–11.i.2006 ( MCN) , Montenegro, 2♀, 1.xii.1977, E.H. Buckup ( MCN) , 1♀, 30.xi.1977, A. Lise ( MCN) , Palmares do Sul, 1♀, 1♂, 11.xi.2003, Equipe Probio ( MCN) , Porto Alegre, 1♂, 16.ix.1999, A. Franceschini ( MCN) , 1♂, 30.x.1952 ( MCN) , S. Chico, 1♀, 17.11.1996, L. Wendel ( PUCRS) , S. Francisco de Paula , 1♀, 13–16.xii.2001, Proj. Araucária (PUCRS) , 1♀, 15–18.i.2002, Proj. Araucária (PUCRS) , 1♀, 15–18.xi.2002, Proj. Araucária (PUCRS) , 1♂, 18–21.x.2001, Proj. Araucária (PUCRS) , Triunfo, 1♂, 14.ii.2008, M. Pasolius ( MCN) , 1♀, 16–17.ix.1993, L. Moura ( MCN) , 1♂, 18.iii.2009, A. Barcellos ( MCN) , 1♀, 1♂, 18.ix.2006, A. Barcellos ( MCN) , 1♂, 20.iv.2003, T. Aguzzoli ( MCN) , 1♂, 27–28.viii.2008, R. Moraes ( MCN) , 1♀, 8.i.1997, A.B. Bonaldo ( MCN) , 1♂, 8.iii.2007, A. Barcellos ( MCN) . ARGENTINA: Missiones : 3♀, xi.1946, M.J. Viana ( MLP) .

Geographical distribution. This species occurs in Brazil and Argentina ( Wibmer & O’Brien 1986) , and in Brazil in the Atlantic Forest (Espírito Santo, Rio de Janeiro, São Paulo, Paraná, Santa Catarina, and Rio Grande do Sul states), Amazon (Pará), and Cerrado (Mato Grosso do Sul, Minas Gerais states) ( Fig. 23B ) .

Associated plants. Podkowa & Carvalho (2012) recorded adults of H. inaequalis in association with Araucaria angustifolia Bert. (Araceae) , Diaz (1997) registered it on Ilex paraguariensis A. St. Louis. Hil. (Aquifoliaceae) . Vernalha (1952), Vellozo et al. (1953), and Vernalha et al. (1966) mentioned the association of this species with pear trees ( Pirus communis L.) ( Rosaceae ) without specifying if they were adults or larvae.

Remarks. Heilus inaequalis resembles H. myops in shape and size of the body, and is distinguished by the number of granules on the pronotum and elytra. In H. inaequalis the granules are rounded and larger on the pronotum, and there are transverse granules on the elytra, the macula of velvety black scales is surrounded on the anterior and posterior borders by whitish scales of the same texture forming a small, elongate macula between striae 3 and 4 of

the median region of the elytra ( Fig. 4A ). Heilus myops has two rows of rounded dorsal granules on the pronotum, transverse and smaller granules on the elytra, and yellowish or orange scales distributed evenly, and a sub-rounded brown to black macula wrapped by a row of clear scales between striae 3 to 6 or 7, located in the center of the elytra ( Fig. 9A ).

Figures

FIGURE 3. Mouthparts of Heilus inaequalis. A. left mandible, dorsal view. B. right mandible, dorsal view. C. left maxilla, ventral view. D. right maxilla, ventral view. E. labium, ventral view. F. labium, lateral view.

FIGURE 4. Holotype Heilus inaequalis, female. A. dorsal view. B. lateral view. C. labels. The copyright of images belongs to NHRS.

FIGURE 9. Holotype Heilus myops, female. A. dorsal view. B. lateral view. C. labels. The copyright of images belongs to NHRS.

FIGURE 18. Aedeagus. A. Heilus inaequalis, dorsal view. B. Heilus inaequalis, lateral view. C. Heilus pupillatus. D. Heilus tuberculosus. E. Heilus rufescens. F. Heilus freyreissi. G. Heilus myops. H. Heilus faldermanni. I. Heilus ochrifer. J. Heilus fasciculatus. K. Heilus iniquus. L. Heilus bistigma. Legends: median lobe (ml), internal sac sclerite (sc), median lobe apodemes (mla).

FIGURE 19. Internal sac sclerite. A. Heilus inaequalis. B. Heilus pupillatus. C. Heilus tuberculosus. D. Heilus rufescens. E. Heilus freyreissi. F. Heilus myops. G. Heilus faldermanni. H. Heilus ochrifer. I. Heilus fasciculatus. J. Heilus iniquus. K. Heilus bistigma.

FIGURE 20. Tegmen. A. Heilus inaequalis. B. Heilus pupillatus. C. Heilus tuberculosus. D. Heilus rufescens. E. Heilus freyreissi. F. Heilus myops. G. Heilus faldermanni. H. Heilus ochrifer. I. Heilus fasciculatus. J. Heilus iniquus. K. Heilus bistigma. Legends: dorsal lobe (ld), manubrium (m).

FIGURE 21. Female genitalia. A. Heilus inaequalis. B. Heilus pupillatus. C. Heilus tuberculosus. D. Heilus rufescens. E. Heilus freyreissi. F. Heilus myops. G. Heilus faldermanni. H. Heilus ochrifer. I. Heilus fasciculatus. J. Heilus iniquus. K. Heilus bistigma. Legends: coxite (cx), styli (st), common oviduct (co), bursa copulatrix (bc), spermathecal duct (spd), spermathecal capsule (sp), spermathecal gland (spg).

FIGURE 22. Spermathecal capsule. A. Heilus inaequalis. B. Heilus pupillatus. C. Heilus tuberculosus. D. Heilus rufescens. E. Heilus freyreissi. F. Heilus myops. G. Heilus faldermanni. H. Heilus ochrifer. I. Heilus fasciculatus; J. Heilus iniquus; K. Heilus bistigma. Legends: cornu (cr), collum (cl), ramus (ra).

FIGURE 23. Geographical distribution of Heilus species based on labels information. A. South America Brazilian biomes and Paraguayan and Argentine ecoregions. B. Heilus inaequalis; C. Heilus pupillatus; D. Heilus tuberculosus; E. Heilus rufescens; F. Heilus freyreissi; G. Heilus myops; H. Heilus faldermanni; I Heilus ochrifer; J. Heilus fasciculatus; K. Heilus iniquus; L. Heilus bistigma.