Actinote mantiqueira Freitas, Francini, Paluch & Barbosa, 2018

Freitas, André Victor Lucci, Francini, Ronaldo Bastos, Paluch, Márlon & Barbosa, Eduardo Proença, 2018, A new species of Actinote Hübner (Nymphalidae: Heliconiinae: Acraeini) from southeast Brazil, Revista Brasileira de Entomologia (Rev. Bras. Entomol.) 62 (2), pp. 135-147 : 136-141

Actinote mantiqueira Freitas, Francini, Paluch & Barbosa , sp. nov.

Actinote alalia ; Ebert, 1969:35; Lewis, 1973:43, fig. 1; Lewis, 1975:43, fig. 1; Penz & Francini, 1996:313, fig. 4a,b,c, 317; Francini & Penz, 2006: figs. 8, 9, Plate 12; Silva-Brandão et al., 2008:519, 528, figs. 3, 4, 5; Freitas et al., 2009b:88; Freitas et al., 2010:100, 103.

Actinote alalia alalia ; Eltringham & Jordan, 1913:10.

Actinote undescribed species ; Freitas et al., 2010:103.

Diagnosis

The wing color and pattern are typical of the “orangish red mimicry complex” ( Francini, 1989), with dark brown stripes on an orangish red background on the dorsal wing surfaces ( Fig. 9 ), but the new species can be distinguished from all other previously described species in this complex by several morphological characters (see Francini and Penz, 2006). The males of Actinote mantiqueira sp.nov. can be distinguished from those of Actinote alalia by the usually broader transverse dark brown bands on the dorsal forewing ( Fig. 2A and E ). On the dorsal forewing, A. mantiqueira sp. nov. can be distinguished from A. alalia by the presence of a usually continuous dark band in the internal margin (this is mostly orange in A. alalia ). On the ventral hindwing, A. mantiqueira sp. nov. can be distinguished from A. alalia by the faintly marked, orange v-shaped transverse band (this band is conspicuously marked in A. alalia ), by the absence of a complete dark bar crossing the middle of the discal cell (this dark bar is present in A. alalia , delimiting a basal orange area), and by the absence of a distinct patch of orange scales in the humeral angle, basally to the humeral vein (this region is covered by cream scales in A. mantiqueira sp. nov.) ( Fig. 2B and F ). Females are very similar in both species ( Fig. 2C and G ), and hardly discernible; the most consistent character is the presence of a distinct patch of orange scales in the humeral angle, basally to the humeral vein, in A. alalia (cream in A. mantiqueira sp. nov.) ( Fig. 2D and H ). The male genitalia of both species are very similar, and the main difference is the profile of the saccus, which is more triangular and pointed in A. mantiqueira sp. nov., and bumped and abruptly enlarging near the base in A. alalia ( Fig. 3D and L ).

Description

Male ( Figs. 1A , 2A,B , 4A–D , 9A ): Antenna black, 11–12 mm in length (n = 11), extending to mid-costa, with 42 antennomeres, 9 of which form a well-defined club. Palpus length about 2.0 times head height. Male palpus shown in Fig. 4B . Forewing narrow and elongated, length 25–28 mm (mean = 26.8 mm, SD = 1.08 mm, n = 11); hindwing rounded, about two-thirds length of forewing, length 18–21 mm (mean = 19.7 mm, SD = 0.90 mm, n = 11). Male wing venation as shown in Fig. 4A . Male foreleg ( Fig. 4C ) with a single tarsomere. Male midleg as shown in Fig. 4D . Body dark brown, abdomen ventrally covered by cream scales. Forewing dorsal background dark orange with dark brown veins and stripes as follows: a broad dark brown margin, broad in apex and narrowing toward tornus; entire space from anal margin to 2A dark brown; three additional broad dark brown transverse stripes, first extending from costa to CuA 2, two thirds from base, second v-shaped crossing discal cell in mid portion, and third crossing space CuA 2 -2A halfway from base. Many individuals present a dark cubital spot adjacent to discal cell near wing base. Hindwing dorsal background dark orange with broad dark brown margin; all veins dark brown, and narrow dark brown stripes extending in inter-venal spaces; some individuals can present transverse patches of dark scales crossing proximally spaces M 1 -M 2, M 2 -M 3, M 3 -CuA 1 and CuA 1 -CuA 2, in some cases almost forming a v-shaped transverse band. Ventral forewing with a pattern very similar to dorsal pattern, but lacking broad dark brown margin; ventral hindwing more homogeneously cream, with dark brown veins and stripes in inter-venal spaces.

Male genitalia ( Fig. 3A–G ): Valvae elongated and curved, length about four times width of median portion; broader at base; apex rounded with a conspicuous curl toward middle line of body. In dorsal view, basal portion of uncus wide, abruptly narrowing, ending in a point. Tegumen long and broad, trapezoidal with a slight constriction at middle. Gnathos absent. In ventral view, saccus shaped as an isosceles triangle, about half length of genital capsule. Aedeagus about half length of genital capsule, ending in a sharp point, in lateral view straight. Juxta broad, about same width of base of one valva (in ventral view), tear-shaped (with an anterior pointed process); length equal to width.

Female ( Figs. 1B , 2C,D , 4E–H , 8B ): Antenna black, 12 mm in length (n = 3), extending to mid-costa, with 41 antennomeres, 10 in club. Female palpus as shown in Fig. 4F . Forewing narrow and elongated, length 29–31 mm (mean = 29.3 mm, SD = 1.53 mm, n = 3). Hindwing rounded, not translucent, length 20–22 mm (mean = 20.7 mm, SD = 1.15 mm, n = 3). Female wing venation is shown in Fig. 4E . Color pattern of female wings similar to that of males but paler. Foreleg with six segments on tarsus, with third segment bearing one short setae ( Fig. 4G ). Female midleg as shown in Fig. 4H .

Female genitalia ( Fig.3H ): Corpus bursae rounded, signa absent. Ductus bursae not sclerotized, same length as corpus bursae; sterigma trapezoidal.

Taxonomy and variation

As has been reported for most species of Actinote , A. mantiqueira sp. nov. also presents intraspecific variation, especially in female color pattern (males are more uniform). Minor variation has also been reported in wing venation and genitalia. The most variable characters are the extent of the humeral vein, which can reach or not the wing edge, and the shape of the saccus apex, which can vary from pointed to almost rounded. The populations from Serra da Bocaina (São Paulo State) are slightly different on then ventral hindwing, with the orange v-shaped transverse band more conspicuously marked.

Holotype ( Fig. 1A ). Male from Pindamonhangaba, São Paulo, Brazil (22 ◦ 46 l 3.62 llS 45 ◦ 31 l 27.08 llW). Deposited in the Museu de Zoologia da Universidade Estadual de Campinas (ZUEC), Unicamp, Campinas, São Paulo, Brazil. Labels on the holotype (five labels separated by transverse bars): /HOLOTYPUS/ Mirante do Pico do Itapeva , Pindamonhangaba, São Paulo: Brazil 28.XII.2016, 2000 m, 22 ◦ 46 l 3.62 llS 45 ◦ 31 l 27.08 llW, L.M. Magaldi, leg./Holotypus— Actinote mantiqueira Freitas, Francini, Paluch & Barbosa det. 2017/DNA voucher – BLU 936/ZUEC LEP 10004/ .

Allotype ( Fig. 1B ). Same data as Holotype. Deposited in the Museu de Zoologia da Universidade Estadual de Campinas ( ZUEC), Unicamp, Campinas, São Paulo, Brazil. Labels on the allotype (five labels separated by transverse bars): /ALLOTYPUS/ Mirante do Pico do Itapeva , Pindamonhangaba, São Paulo: Brazil 28.XII.2016, 2000 m, 22 ◦ 46 l 3.62 llS 45 ◦ 31 l 27.08 llW, L.M. Magaldi, leg./Allotypus— Actinote mantiqueira Freitas, Francini, Paluch & Barbosa det. 2017/DNA voucher – BLU 939/ZUEC LEP 10005/ .

Paratypes (All from Brazil). Minas Gerais: Itamonte, Caminho para Agulhas Negras, Parque Nacional do Itatiaia , 1800 m, 1 male, 4.XII.1991, A. V. L. Freitas leg. ( ZUEC-AVLF) . São Paulo: Piquete, Estrada para São Francisco dos Campos do Jordão , 1800 m, 1 male, 5.XII.2004, R.B.Francini leg. (R. B. Francini col. #22) ; Campos do Jordão, Alto da Boa Vista , 1800 m, 1 male, 6.XII.2007 (DNA voucher AC 110), A. V. L. Freitas leg .; Pindamonhangaba, Pico do Itapeva , 1800 m, 1 female, 31.XII.2005, A. V. L. Freitas leg.; 6 males (DNA vouchers AC111, AC112, AC113, AC114, AC115, AC116), 6.XII.2007, A.V.L.Freitas leg.( ZUEC-AVLF) ; Mirante do Pico do Itapeva , 2000 m, 3 males, 28.XII.2016, 22 ◦ 46 l 3.62 llS 45 ◦ 31 l 27.08 llW, L. M. Magaldi, leg. (DNA vouchers BLU 937, BLU 938, BLU 940) (ZUEC LEP 10006, ZUEC LEP 10007, ZUEC LEP 10008) ( ZUEC) ; Santo Antônio do Pinhal, Estrada para o Pico Agudo , 1220 m, 20.XI.2013, 22 ◦ 51 l 1 llS 45 ◦ 39 l 42 llW, T.S.Souza leg (ZUEC LEP 10003) ( ZUEC) .

Additional studied material (all from Brazil). DZUP – Minas Gerais: Camanducaia, Monteverde , 1650 m, 3 males and 6 females, 22.XII.1968, 1 male, 23.XII.1968, 4 males, 27.XII.1968, H. Ebert leg. DZ 3769, DZ 3772, DZ 3771, DZ 5848, DZ 5865, DZ 6041, DZ 5961, DZ 5937, DZ 5896, DZ 3773, DZ 3774, DZ 3474, DZ 7869, DZ 3770; Santos Dumont, Rio Novo , 850 m, 1 male, 13.XI.1953, H. Ebert leg. DZ 5849; Barbacena, Serra da Mantiqueira , 1100 m, 1 female, 4.XII.1952, H. Ebert leg. DZ 7091; Poços de Caldas , 1250 m, 1 male, 10.XII.1966, 2 males, 11.XII.1966, 1 male, 15.XII.1966, 3 males, 17.XII.1966, H. Ebert leg. DZ 3776, DZ 3786, DZ 3785, DZ 3787, DZ, 3767, DZ 3788, DZ 2977; Delfim Moreira , 15 Km SE, 1500–1700 m, 2 females, 22-23.I.2004, Mielke & Casagrande leg. DZ 9290, DZ 9250. Rio de Janeiro: P. N. Itatiaia, 1600 m, 2 females, 12.I.1973, Mielke leg. DZ 5969, DZ 5857; Imbariê , 25 m, 1 female, 29.VII.1964, H. Ebert leg. DZ 5928; Itatiaia, Oeste , 1400 m, 2 males, 22.XII.1957, H. Ebert leg. DZ 3782, DZ 3110; Teresópolis , 1000 m, 1 female, 5.I.1973, Mielke leg. DZ 7997. São Paulo: Campos do Jordão , 1700 m, 1 female, 30.I.1966, H. Ebert leg.DZ 5945, Campos do Jordão, Toriba , 2 males, 12.XI.1922, 1 male, 16.XII.1952, D Almeida & L.Travassos Filho leg. DZ 3781, DZ 3762, DZ 2974; Bananal, Bocaina , 2 males and 2 females, 2.I.1937, Travassos leg. DZ 3780, DZ 3779, DZ 5913, DZ 5904. IOC – Minas Gerais: Passa Quatro , Fazenda dos Campos, 1600 m, 1 female, 2.XII.1915, 1 female, 5.XII.1915, J.F. Zikán leg. I.O.C. no 25195, No. 25196. Rio de Janeiro: Itatiaia, Campo Belo , 2 females, 10.XII.1919, 1 female, 12.II.1920, km12, 1 female, 20.I.1925, J.F.Zikán leg. I.O.C. No. 25193, No. 25177, No. 25198, No. 25217, 1 female, 30.XI.1920, 1 female, 29.XII.1920, M. Zikán leg.I.O.C.No.25176, No. 25178. São Paulo: Bananal , 3 males and 3 females, 8.I.1937, L. Travassos Filho leg. Coll. Travassos. MNRJ – Rio de Janeiro: Nova Friburgo , 1 female, III.1934, Col. J. Oiticica Filho. São Paulo: Campos do Jordão , 3 females, 29.I.1933, Travassos & J. Oiticica Filho, Col. J. Oiticica Filho; Bananal, Bocaina , 3 females, I.1937, A. Costa leg. Col. Dr. A. Costa No. 4376, No. 4377, No. 4365. ZUEC-AVLF – Minas Gerais: Andradas , Pico do Gavião , 1600 m, 1 male, 12.XII.2017, 22 ◦ 1 l 1 llS 46 ◦ 37 l 37 llW, A.V.L. Freitas, L.M. Magaldi, J.Y.O. Carreira & A. Tacioli leg.; Poços de Caldas, Serra do Cristo , 1560 m, 2 males, 15.XII.2017, L. M. Magaldi & A. Tacioli leg. São Paulo: Silveiras, ponte sobre o Rio do Higino, Serra da Bocaina , 2 males (DNA vouchers AC176, AC180) and 2 females (DNA vouchers AC174, AC179), 17.I.2009, K. L. Silva-Brandão leg; Serra da Bocaina , 1450 m, 2 males, 19.XII.2017, 22 ◦ 47 l 54 llS 44 ◦ 42 l 48 llW, L.M. Magaldi, A. Tacioli & A.H.B. Rosa leg.

Immature stages

Egg ( Figs.5A,B , 8C ). Light yellow when first laid, changing gradually to pinkish red after 24 h; barrel-shaped with 13–16 vertical ribs and several (∼14–17) weakly marked horizontal ribs; mean height 0.683 mm (range 0.67–0.71 mm, n = 3), mean diameter 0.412 mm (range 0.37–0.44 mm, n = 9). Aeropyles localized only near base of egg, in two irregular rows (not every vertical rib bears an aeropyle).

First instar ( Figs. 6 and 7A ). Head brown, smooth, without scoli, mean width 0.33 mm (range 0.29–0.36 mm, n = 3); body pale cream, without scoli and with long pale setae arising from pinacula; legs pale brown, prolegs pale, anal plate pale brown. Prothoracic plate pale and difficult to observe under stereomicroscope. T1 presents only 2 subventral setae, similar to Actinote alalia . Head chaetotaxy and body chaetotaxy are presented in Figs. 6 and 7A , respectively. Maximum reported length 3.0 mm.

Last instar ( Figs. 7B and 8D ). Head dark brown, smooth with thin pale setae and without scoli, spines or chalazae, mean width 3.14 mm (range 3.01–3.24 mm, SD = 0.073 mm, n = 4); body dark brown dorsally, pale cream laterally and ventrally, covered with medium sized dark brown scoli bearing brown setae from T1 to A1 and from A7 to A10, and white setae from A2 to A6; legs black, prolegs pale cream; anal plate dark brown.Maximum length: 40 mm (n = 4). Scoli distribution as in Fig.7B .Prepupa changes color, becoming pale cream with a more homogeneous coloration.

Pupa ( Fig. 8E ). General profile elongated, ground color pale yellow cream with dark brown markings in wing cases and abdomen; abdominal segments mobile, with a series of five pairs of subdorsal black spines from segments A2 to A6. Total length 20–21 mm (n = 2).

Adult and immature behavior, host plants and natural history. Larvae of A. mantiqueira sp. nov. have been recorded using Eupatorium intermedium and Eupatorium sp. (a species very similar to E. intermedium ) ( Asteraceae ) ( Fig. 8A ) as host plants ( Penz and Francini, 1996; RBF and AVLF pers. obs.). Females were observed ovipositing from 12:00 to 02:00 PM on the underside of mature leaves of their host plants ( Fig. 8B ). Ovipositions are large ( Fig. 8C ), varying from 409 to 453 eggs (n = 4). Newly hatched larvae first consumed the chorion, and after 3–5 h began to feed on leaf tissue. First instars fed on the underside leaf tissue by scraping the leaf surface, while later instars were observed consuming the entire leaf. In first instars, frass pellets were glued onto the leaf by silk, not falling to the ground. Larvae were consistently gregarious in all instars, and all activities, such as feeding, resting, or moving between leaves, occurred simultaneously.

Adults were only observed on sunny days, quickly disappearing when weather conditions became cloudy. Males began to fly around 10:00 AM, usually flying 2–5 m high. Females were seldom observed, and territorial behavior courtship behavior or copulation were not perceived. All known populations of A. mantiqueira sp. nov. were recorded in well-preserved subtropical wet montane forest above 1000 m altitude, where the climate approaches temperate conditions, including cold winters with frequent frosts and temperature frequently falling below 0 ◦ C. Males were observed feeding on flowers of Croton urucurana Baill. ( Euphorbiaceae ), Chromolaena punctulata (D.C.) R. King & H. Robins ( Asteraceae ) and on several other species of forest edge plants. Adults are univoltine in all known populations, with only one flight period during the warmer months, from late November to early February. Adults of A. mantiqueira sp. nov. are sexually dimorphic, with males having a deep orange coloration ( Figs. 1A , 2A,B , 9A ), and females being much more translucent and sometimes having the subapical spot on the forewing light cream ( Figs. 1B , 2C,D ). In most studied sites individuals were never abundant, with usually 5–30 individuals observed in a typical day of field work (4–5 h of observation), most of which were males. This species is part of the “orangish red mimicry complex” of Actinote (sensu Francini, 1989) ( Fig. 9 ), and is sympatric with five other Actinote co-mimics, namely Actinote bonita Penz, 1996 ( Fig. 9B ), Actinote conspicua Jordan, 1913 ( Fig. 9C ), Actinote quadra (Schaus, 1902) ( Fig. 9D ), Actinote dalmeidai Francini, 1996 ( Fig. 9E ), and Actinote surima (Schaus, 1902) ( Fig. 9F ) and one pierid co-mimic, Dismorphia melia (Godart, [1824]) ( Pieridae : Dismorphiinae ) (see Dias et al., 2016).

Habitat. In all known localities, the species is associated with well-preserved montane ombrophilic forest, usually at altitudes above 1000 m.Males and females are easily observed in forest edges and in areas of contact between forest and high altitude natural grasslands ( Fig. 10A and B ).

Geographic distribution. The species is known from the Serra da Mantiqueira region, from the Serra da Bocaina and Serra dos Órgãos, at altitudes from 1000 to 2000 m ( Fig. 11 ).

Etymology. Mantiqueira is a word in native Tupi language meaning “rain drops” (from amana = rain and tykyra = drop) and is the name of a large mountain range in SE Brazil (Serra da Mantiqueira), where most of the known populations of A. mantiqueira sp. nov. occur.

Figures

Fig. 9. Comparative plate showing all known Brazilian species of Actinote belonging to the “orangish red mimicry complex”, dorsal left and ventral right (males only). (A) Actinote mantiqueira sp.nov.(paratype),(B) Actinote bonita, (C) Actinote conspicua, (D) Actinote quadra, (E) Actinote dalmeidai, (F) Actinote surima, (G) Actinote alalia, (H) Actinote catarina.

Fig. 2. Comparative plate showing the main differences in wing pattern between Actinote mantiqueira sp. nov.and Actinote alalia. (A) Male paratype of A. mantiqueira sp. nov. (dorsal left, ventral right); (B) detail of humeral region in male VHW; (C) female paratype of A.mantiqueira sp. nov. (dorsal left, ventral right); (D) detail of humeral region in female VHW; (E) male of A. alalia (dorsal left, ventral right); (F) detail of humeral region in male VHW; (G) female of A. alalia (dorsal left, ventral right); (H) detail of humeral region in female VHW.

Fig. 3. Male and female genital characters of Actinote mantiqueira sp. nov.(A–H) and Actinote alalia (I–P). A. mantiqueira sp. nov.:(A) lateral view of male genitalia; (B) uncus in dorsal view; (C) aedeagus dorsal (above) and lateral (below); (D) saccus, ventral view; (E) valva, ventral view; (F) juxta; G. 8th sternite; (H) female sterigma. A. alalia: (I) lateral view of male genitalia; (J) uncus in dorsal view; (K) aedeagus dorsal (above) and lateral (below); (L) saccus, ventral view; (M) valva, ventral view; (N) juxta; (O) 8th sternite; (P) female sterigma. ae, aedeagus; ju, juxta; sa, saccus; un, uncus; va, valva.

Fig. 1. Holotype male (A) and allotype female (B) of Actinote mantiqueira sp. nov. (left = dorsal; right =ventral). Scale bar =10 mm.

Fig. 4. Morphological characters of males (A–D) and females (E–H) of Actinote mantiqueira sp. nov. (A) Wing venation; (B) Palpus; (C) Foreleg; (D) Midleg; (E) Wing venation; (F) Palpus; (G) Foreleg; (H) Midleg.

Fig. 8. Natural history of A.mantiqueira sp. nov.(A–E) and Actinote alalia (F–G).(A) Shrub of Eupatorium intermedium, host plant of A.mantiqueira; (B) Female of A.mantiqueira sp. nov. ovipositing; (C) close view of an oviposition of A. mantiqueira sp. nov.; (D) Last instar of A. mantiqueira sp. nov. (lateral view); (E) Pupa of A. mantiqueira sp. nov. (latero-ventral view); (F) Last instar of A. alalia (lateral view); (G) Female of A. alalia ovipositing (photo by Lucas A. Kaminski).

Fig. 5. Eggs of Actinote mantiqueira sp. nov. (A – lateral view, B – dorsal view) and Actinote alalia (C – lateral view, D – dorsal view) in scanning electron microscopy.

Fig. 6. Chaetotaxy of first instar head capsule of Actinote mantiqueira sp. nov. (setae nomenclature on the right side and pore nomenclature on the left side).

Fig. 7. Larval body diagrams of Actinote mantiqueira sp. nov. (A) First instar body chaetotaxy; (B) last instar scoli distribution.

Fig. 10. (A and B) Habitat of Actinote mantiqueira sp. nov.; (C and D) Habitat of Actinote alalia. (A) General view of the montane forests in Campos do Jordão, São Paulo State (1400–1600m a.s.l.); (B) region of contact between grassland and high altitude forest, Pico do Itapeva, Pindamonhangaba, São Paulo State (1800m a.s.l.); (C) general view of the montane forests in São Francisco de Paula, Rio Grande do Sul State (900–1100 m a.s.l.); (D) close view of the montane forest in São Francisco de Paula, Rio Grande do Sul State (900 m a.s.l.); the characteristic Araucaria trees are visible near the trail (black arrow).

Fig. 11. Recorded distribution of Actinote mantiqueira sp. nov. (open circles) and Actinote alalia (solid circles) in Brazil. Locality data include records compiled from Paluch (2006), museum data (see text) and recent sightings (AVLF, unpublished data).