Tarema rivara Schaus, 1896

St Laurent, Ryan A., Herbin, Daniel & Mielke, Carlos G. C., 2017, Revision of the genus Tarema Schaus, 1896 (Lepidoptera, Mimallonoidea, Mimallonidae) with the description of a new species from southeastern Brazil, ZooKeys 646, pp. 119-137 : 123-127

Tarema rivara Schaus, 1896 Figs 1-6, 11, 14, 16

Tarema rivara Schaus, 1896: 55

Tarema rivara ; Schaus 1928: 670, fig. 88i ♂

Tarema rivara ; Gaede 1931

Tarema rivara ; Becker 1996

Tarema rivara ; Diniz et al. 2013: 88, figs ♂, ♀, larva, larval sack

Tarema macarina Schaus, 1928: 670, fig. 88i ♀ syn. n.

Tarema macarina ; Gaede 1931

Tarema macarina ; Becker 1996

Type material.

Tarema rivara Schaus: lectotype [here designated], ♂. BRAZIL: São Paulo: São Paulo, S.E. Brazil./ Collection, WmSchaus/ Type No. 12566 U.S.N.M/ USNM-Mimal: 1120/ Tarema rivara Type. Schaus./ LECTOTYPE male Tarema rivara designated by St Laurent, Herbin, and C. Mielke, 2017 [handwritten red label]/ St Laurent diss.: 8-22-16:3/ (USNM, examined). Type locality: Brazil: São Paulo.

Tarema macarina Schaus: lectotype [here designated], ♀. BRAZIL: São Paulo: 1 ♀, São Paulo, S.E. Brazil./ Collection, WmSchaus/ Type No. 33596 U.S.N.M/ USNM-Mimal: 1121/ Tarema macarina Type Schaus/ LECTOTYPE female Tarema macarina designated by St Laurent, Herbin, and C. Mielke, 2017/ Tarema rivara det. St Laurent 2016/ St Laurent diss.: 8-22-16:2/ (USNM, examined). Paralectotypes, 2 ♀. BRAZIL: São Paulo: 1 ♀, Brasil. m., S. Paulo 96/ Typus/ Coll. Staudinger/ Tarema macarina Schaus, co-type/ (MNHU, photograph examined). Unknown state: 1 ♀, Brasil [illegible]/ Tarema macarina co-type Schaus/ 8582/ (MNHU, photograph examined). Type locality: Brazil: São Paulo. - All paralectotypes with the following yellow label: PARALECTOTYPE ♀ Tarema macarina designated by St Laurent, Herbin, and C. Mielke, 2017.

Additional specimens examined.

(28 ♂, 14 ♀) BRAZIL: Maranhão: 1 ♂, Feira Nova do Maranhão, Retiro, 07°00'31"S, 46°26'41"W, 480 m: 16-17.II.2013, C. Mielke leg., Coll. C. Mielke 26.333 (CGCM). Bahia: 1 ♂, Barreiras, 12°9'S, 45°00'W, 700 m: 4.II.1994, Coleção EMBRAPA- CPAC No. 15.987, [Camargo leg.] (CPAC). Mato Grosso: 1 ♀, Chapada [dos] Guimarães: 25.V.1989, V.O. Becker leg., Coll. Becker 75031, USNM-Mimal: 2320 (USNM). 1 ♂, No specific locality: XII.1929, Coll. R. Spitz, Rothschild Bequest 1939-1 (NHMUK). Goiás: 1 ♂, Ponte Funda, Vianópolis: 24.X.1987, Tangerini leg., genitalia prep. D. Herbin ref H. 1010 (CDH). 1 ♀, Ipameri: 10.X.1988, V.O. Becker leg., Coll. Becker 59419, USNM-Mimal: 2323, St Laurent diss.: 8-22-16:8 (USNM). Distrito Federal: 1 ♂, 1 ♀, Estação Florestal, Cabeça do Veado, 1100 m: 17.X.1971, 23.X.1971, E.G., I. & E.A. Munroe leg., St Laurent diss.: 3-14-16:7, 3-14-16:8 (CNC). 5 ♂, 2 ♀, Planaltina, 15°35'S, 47°42'W, 1000 m: 1-10.XI.1994, Tangerini leg. (1 ♂, CDH); 25.IX.1985, 5.XI.1988, V.O. Becker leg., Coll. Becker 57771, 58871, 58872, USNM-Mimal: 2301-2304, 2321, 2322 (4 ♂, 2 ♀, USNM). 1 ♂, Planaltina: 15.X.1995 (MWM). Minas Gerais: 1 ♂, Serra do Cipó à Conceição do Mato Dentro, km 126.3, 19°14'51"S, 43°30'38"W, 1270 m: 18.XI.2012, genitalia prep. D. Herbin ref H. 1012 (CDH). 1 ♂, Malacacheta, 500 m: I.1998, H. Thöny leg., genital prep. 29.234 (MWM). 1 ♂, Sete Lagoas, 720 m: 15.III.1974, V.O. Becker leg., Coll. Becker 411, USNM-Mimal: 2300 (USNM). 1 ♂, Paracatu, 17°13'S, 46°52'W, 920 m: 5.II.1994, Coleção EMBRAPA- CPAC No. 14.664 (CPAC). 1 ♂, Iraí de Minas, 18°43'S, 47°30'W, 950 m: 9.II.1994, Coleção EMBRAPA- CPAC No. 14.145 (CPAC). 1 ♂, São Roque de Minas, São José do Barreiro, 870 m, 3.XII.2016, C. Mielke leg., Coll. C. Mielke 32.162 [point not on map] (CGCM). São Paulo: 2 ♂, Ribeirão Preto, Fazenda da Pedra, Rio Tamanduá: Travassos & Pearson leg., 12-15.X.1953 (NHMUK); HRP 643, USNM-Mimal.: 2425 (USNM). 1 ♂, Locality as for previous but 500 m: 2-5.III.1954, Pearson & Oiticica leg., Brit. Mus. 1962-112 (NHMUK). 1 ♂, Alto da Serra [Paranapiacaba]: I.1926, R. Spitz leg., Rothschild Bequest 1939-1, St Laurent diss.: 7-7-16:4 (NHMUK). 1 ♂, Miracatu, 700 m: 20.XI.1997, H. Thöny leg., genital prep. 29.233 (MWM). 2 ♂, 2 ♀, No additional locality data: E.D. Jones Coll., Brit. Mus. 1919-295 (NHMUK); Rothschild Bequest 1939-1 (NHMUK). 1 ♂, No specific locality, 750 m: E.D. Jones Coll., Brit. Mus. 1919-295, NHMUK010354581, St Laurent diss.: 8-29-16:5 (NHMUK). Paraná: 1 ♂, Campo do Tenente, 850 m: 17.X.1985, [O.] Mielke leg., DZ 15.496 (DZUP). 1 ♂, Ponta Grossa: II.1957, at light, No. 1780, Coll. F. Justus Jor (DZUP). 1 ♀, Castro, 950 m: E.D. Jones leg., E.D. Jones Coll., Brit. Mus. 1919-295 (NHMUK). 1 ♀, Tucunduva [ Sengés], 650 m: 17.II.1913, E.D. Jones leg., E.D. Jones Coll., Brit. Mus. 1919-295 (NHMUK). Santa Catarina: 1 ♂, No additional locality data: F. Hoffmann leg., USNM-Mimal: 2681, St Laurent diss.: 8-22-16:7 (USNM). No state: 1 ♀, “Bresil”, Joicey Coll. Brit. Mus. 1925-157 (NHMUK). PARAGUAY: Cordillera: 1 ♀, Pirareta, 25°29'S, 56°56'W, 200 m: 26-31.VIII.2012, [U. Drechsel] leg., genitalia prep. D. Herbin ref H. 1014 (CDH). Amambay: 1 ♀, Estancia Oliva, 22°10'S, 56°26'W, 225 m: 23-25.I.2013, [U. Drechsel leg.], (CDH). Guairá: 1 ♀, Villarica: 2.X.1925, F. Schade leg., J.J. Joicey Esq., B.M. 1929-458, St Laurent diss.: 7-7-16:5 (NHMUK). Paraguarí: 1 ♀, Sapucay [recte Sapucai]: 12.VIII.1904, W. Foster leg., Rothschild Bequest 1939-1 (NHMUK).

Diagnosis.

Tarema rivara can be differentiated from others in the genus by the pervasive orange to orange-red coloration on the wings in males, and by the faint orange hue on the wings of the females which is concentrated antemedially and/or submarginally. The more similar Tarema bruna sp. n. is more earthen brown in color than Tarema rivara , with a longer postmedial lunule that reaches the wing margin without becoming diffuse. In both sexes, Tarema rivara has a smaller wingspan than Tarema fuscosa and lacks a distinct dark brown patch of scales at the apex of the forewings. This patch of scales is distinct in Tarema fuscosa due to the contrast with the light cream color of the postmedial lunule that borders it. Genitalia are quite different between these species, in Tarema rivara the valves are reduced to small lobes connected to a flattened, spiny accessory, while the valves of Tarema fuscosa and Tarema bruna sp. n. are larger and not connected to, or lack the accessory arms. The gnathos of Tarema rivara is a rectangular plate rather than ovoid as in Tarema fuscosa , or tapered hexagonal as in Tarema bruna sp. n., the fingerlike projections at the base of the valves are smaller in Tarema rivara , and finally the phallus of Tarema rivara (and Tarema bruna sp. n.) bears a smooth dorsal projection whereas the same projection is short and spined in Tarema fuscosa . The key differences in female genitalia are the larger corpus bursae in Tarema rivara and the trilobed tergite VIII, which is a broad, singular plate in Tarema fuscosa . The female tergite can usually be examined under a microscrope after brushing off scales, without dissecting the specimen.

Description.

Male.Head: As for genus, gray with orange undertone, antenna coloration as for head. Thorax: Coloration similar to that of head, but more orange, appearing hoary due to banded gray scales interspersed amongst orange hued ones, prothorax covered almost entirely in light gray scales. Legs: As for genus but tibia mostly orange. Forewing dorsum: Forewing length: 9-13 mm, avg.: 11.5 mm, wingspan: 21.5-29.0 mm, n=16. Ground color ranging from clay-orange to almost brick red, overall generously shaded by gray scales giving the wing a hoary, layered appearance, especially medially. Postmedial line as for genus but coloration light cream and bordered externally with black scaling continuously along length. Antemedial area with salmon orange hue, medial area always lighter gray compared to orange or reddish submarginal area. Apical half of submarginal area with postmedial lunule, the latter never parallel with margin or postmedial line, either smoothly curved toward margin or angled acutely from postmedial line, becoming diffuse before reaching wing margin, basal half of submarginal area with bright orange or red patch along postmedial line. Discal spot as for genus. Fringe light gray to khaki with lighter and darker patches, including salmon colored scales. Forewing ventrum: Similar to dorsum but usually lighter due to more extensive covering of gray scales; antemedial line absent, postmedial line very faint, bulging outward toward wing margin mesally. Postmedial lunule present as on dorsum, more distinct than postmedial line. Hindwing dorsum: Coloration as for forewing dorsum, following similar patterning but antemedial line absent, postmedial line slightly concave, and submarginal area always orange to more reddish, postmedial lunule absent or just a faint suffusion, coloration usually concentrated somewhat mesally. Hindwing ventrum: Following same pattern as forewing ventrum. Abdomen: As for genus, concolorous with thorax. Genitalia: (Fig. 11) n=12. Vinculum almost circular. Uncus robust, sharp, parrot beak-like when viewed laterally, uncus dorsolaterally flattened. Gnathos a rectangular, elongated plate with slight curvature mesally. Valve short, rounded, weakly sclerotized mesally, strongly affixed to vinculum such that they do not open fully. Base of valves with pair of small, fingerlike projections, weakly sclerotized knobby area present above fingerlike projections. Valve with more heavily sclerotized, spined accessory arms attached basally to valves, arms originate from transtilla or base of valve (unclear), connected along length of valve. Accessory arms flattened and ventrally spined. Diaphragm with pair of horsetail-like seatal patches consisting of setae of variable length that extend outward over phallus for about three-quarters length of gnathos plate, setae mostly straight. Phallus broad, large, with two elongated accessory projections of variable length, projection superior to phallus smooth, straight, sharply pointed; other projection shorter, narrower, running laterally along phallus originating from within phallus, tip of second projection sharp, but variously bent, size of phallus relative to projections somewhat variable. Vesica balloon-like, slightly scobinate. Female.Head: Similar to male, but broader, antennae and labial palpi smaller. Thorax: As in male, but usually grayer or occasionally more salmon colored. Legs: As in male, though tibial spurs thicker. Forewing dorsum: Forewing length: 10-15 mm, avg.: 12.1 mm, wingspan: 22-30 mm, n=13. Sexual dimorphism strong, wing shape and markings similar to male, but wing broader, coloration usually much more subdued gray and brown, if salmon hue present, generally restricted to antemedial area, though rarely some specimens with salmon hue more pervasive, especially submarginally, otherwise submarginal area solid brown, postmedial lunule usually fainter than in male, sometimes almost absent except for small streak. Fringe with distinctly black portion apically. Forewing ventrum: Similar to forewing ventrum of male, but salmon hue generally absent, veins usually lined with contrasting yellow scales, wing grayer overall but apical region more distinctly solid brown than in male. Hindwing dorsum: Coloration and markings as for forewing dorsum, though postmedial lunule absent. Hindwing ventrum: Follows same pattern as forewing ventrum. Abdomen: As in male but slightly more robust, coloration subdued. Tergite VIII as three posteriorly directed lobes, sternite VIII as large, curved, wrinkled mass, covered in thick, branched setae. Genitalia: (Fig. 14) n=5. As for genus, two dissections with large, snake-like structure, apparently a spermatophore based on presence in one dissected male, present within corpus bursae. Papillae anales narrow.

Distribution

(Fig. 16). This species has a wide distribution in South America, and although most records come from Brazilian Cerrado in Bahia, Maranhão, Mato Grosso, Goiás, Minas Gerais, and Distrito Federal, Tarema rivara is also known from Brazilian Atlantic Forest in São Paulo, mixed ombrophilous forest in the state of Paraná, and inland forests of Paraguay. We are also aware of one specimen from Santa Catarina, though unfortunately it lacks more detailed data that would allow us to understand the habitat in which it inhabits in this state.

Natural history.

Diniz et al. (2013) report Tarema rivara larvae feeding on Vochysiaceae , including the species Qualea grandiflora , Qualea multiflora , and Qualea parviflora . The same authors describe the larval sack as being constructed from leaves, silk, and feces. These larval sacks are of the less uniformly constructed variety in Mimallonidae , and are thus more similar to those of Lacosoma Grote, 1864 and Cicinnus melsheimeri (Harris, 1841) rather than highly compacted, rigid structures as seen in Menevia Schaus, 1928, Cicinnus packardii (Grote, 1865), or Cicinnus bahamensis St Laurent & McCabe, 2016, among others (R. A. St. Laurent pers. obs.).

Remarks.

In the original description of Tarema rivara , Schaus (1896) stated: "What I believe to be the ♀ of this species has the reddish shades replaced by dark brown." Apparently Schaus was well aware of the dimorphism of Tarema rivara at the time of its original description, thus it is somewhat surprising that he described the female of Tarema rivara as a new species, Tarema macarina , over 30 years later ( Schaus 1928). We are also aware of a Tarema rivara female specimen determined as this species and not Tarema macarina at the NHMUK. However, we infer that Schaus changed this earlier determination due to the fact that he had located what he believed to be the male of Tarema macarina in the MNHU as per the following statement from Schaus (1928) in his treatment of Tarema macarina : "A male of this species is in the Berlin museum." Two syntypes of Tarema macarina are present in the MNHU with Schaus’s handwriting on the “type” labels; however, both specimens are female, as is the lectotype (here designated) in the USNM.

The complete lack of any male specimens correctly determined as Tarema macarina and the unusual disparity of female Tarema rivara led us to believe that these names are synonyms. Many records of Tarema rivara and Tarema macarina are sympatric, and thus support this hypothesis. A close analysis of Tarema rivara females reveals hints of orange coloration antemedially, a coloration abundant in the male specimens, but not in Tarema fuscosa . Given that Tarema fuscosa females can be easily determined as such due to the lack of sexual dimorphism in this species, the disparity of opposite sexes for Tarema rivara and Tarema macarina provides clear evidence that they represent a single, dimorphic species. Furthermore, a dissection of Tarema rivara females reveal the long, snake-like spermatophore seen in male Tarema rivara , unlike the smaller spermatophore of Tarema fuscosa .

Diniz et al. (2013) figure a larva and adults of both sexes of Tarema rivara , correctly figuring the female of Tarema rivara , which again, matches the type specimens of Tarema macarina .

Compared to Tarema fuscosa below, this species seems to primarily be an inhabitant of drier Cerrado but is also present in the more humid Atlantic forest in the states of São Paulo (type locality) and Paraná where it is sympatric with Tarema fuscosa , but apparently not synchronic. In these regions of sympatry, Tarema rivara flies during the summer (October through February), while Tarema fuscosa flies in the winter, though exceptions to these flight times are present in regions where both species are not found together.