Revision of the genus Tarema Schaus, 1896 (Lepidoptera, Mimallonoidea, Mimallonidae) with the description of a new species from southeastern Brazil

Abstract The genus Tarema Schaus, 1896 is revised. The species Tarema fuscosa Jones, 1908 and Tarema rivara Schaus, 1896 are redescribed, the female of the former is described and figured for the first time, and the genitalia of both sexes for each species are figured for the first time. The lectotype of Tarema macarina Schaus, 1928, syn. n. is determined to be the female of Tarema rivara. Tarema bruna sp. n. is described from São Paulo, Brazil. Lectotypes for Tarema fuscosa, Tarema rivara, and Tarema macarina are here designated.


Introduction
The genus Tarema Schaus, 1896 has been mostly overlooked in the literature since Schaus (1928), save for its mention in two published species lists of Mimallonidae (Gaede 1931, Becker 1996. Recently, however, Herbin and Mielke (2014) reported new collecting data for T. rivara Schaus, 1896 quite distant from its type locality, Figures were manipulated with Adobe Photoshop CS4 (Adobe 2008). Male genitalia are figured in natural color with CS4 "auto color" used to improve white backgrounds. Female genitalia were treated with "auto tone" in CS4 to darken characters. The map was created with SimpleMappr (Shorthouse 2010) and edited with CS4. All geographical coordinates are approximate, and are based on the localities provided on specimen labels. GPS data were acquired with Google Earth.

Results and discussion
Tarema Schaus, 1896: 55 Type species. Tarema rivara Schaus, 1896: 55, by original designation. Diagnosis. The genus Tarema is recognized among the family Mimallonidae by generous amounts of light gray scales present over the entirety of the dorsum and ventrum of the wings, as well as on the thorax and abdomen, giving the species of this genus a hoary appearance. The genitalia of Tarema are unique in the family. Male genitalia have short, ovoid valves and spike-covered projections emanating from near the base of the valves that may be associated with the transtilla and/or the gnathos. The gnathos itself is reduced to a flat, movable plate that covers the base of the uncus. Long setae emanate from above the phallus in paired, horsetail-like bunches. The phallus is thick and broad, and has two lengthwise processes terminating in a sharp tip and a curved tip respectively. Female genitalia are robust structures with a medium to large coiled ductus-corpus bursae complex. The sclerotized portion of abdominal segment VIII is broad; appearing wrinkled ventrally, and is covered in thick, branched setae. The genus Alheita Schaus, 1928 is somewhat similar to Tarema in overall small size, wing shape, and minor resemblance of male genitalia, namely the ovoid valves and odd shape of the uncus which is usually more deeply bifid in Alheita.
Description. Male. Head: Eyes large, more than two thirds area of head; antenna bipectinate to tip, though pectination reduced along distal fifth of antennal length; labial palpus reduced, three segmented, palpus usually not extending beyond frons, scales generally darker brown dorsally. Thorax: Appearing hoary due to banded gray or pale khaki scales interspersed amongst darker ones, prothorax with more heavily concentrated light gray or khaki scales. Legs: Coloration as for thorax, though lighter gray, vestiture finer, bushier. Tibial spurs narrow, sharp, mostly clothed in scales. Forewing dorsum: Forewing length: 9-16 mm, wingspan: 21.5-32.0 mm. Short, triangular, outer margin nearly straight but slightly convex mesally. Ground color ranging from brown or pale clay-orange to nearly black, overall generously shaded by cream or gray scales giving the wing a hoary, layered appearance. Antemedial line absent or nearly so, faint dark band may be present. Postmedial line nearly straight but may be somewhat inwardly or outwardly bent, line preapical such that submarginal area mostly uniform in width from tornus to apex. Apical half of submarginal area with postmedial lunule. Costa appearing lighter than most of wing due to high concentration of gray or khaki scales. Discal spot a thick streak spanning width of discal cell. Forewing ventrum: Antemedial line absent, postmedial line never straight, bulging outward toward wing margin mesally. Hindwing dorsum: Submarginal area with orange to reddish patch of scales mesally, discal mark present but smaller. Hindwing ventrum: Following same pattern as forewing ventrum. Frenulum as single bristle. Venation: Rather typical for Mimallonidae, discal cell quite broad, distal edge sharply slanted (see Schaus 1896). Abdomen: Short, barely or not extending beyond anal angle of hindwing. Genitalia: Complex; vinculum somewhat ovoid or almost circular, ventrally with reduced sac-cus. Uncus robust, sharp, parrot beak-like and dorsolaterally flattened or reduced to slightly triangular stump with slight bidentation terminally. Gnathos a flattened or curved plate concealing fingerlike sclerotization of anal tube. Valves short, rounded, weakly sclerotized mesally. Base of valves with pair of fingerlike projections, weakly sclerotized knobby area may be present above fingerlike projections. Valves with or without more heavily sclerotized, spined accessory arms that may or may not be attached basally to valves. Diaphragm with pair of horsetail-like seatal patches consisting of setae of variable length that extend outward over phallus below gnathos plate. Juxta partially fused to phallus, encircling it, lightly sclerotized, with ventral lip connecting phallus to base of vinculum (severed to excise phallus). Phallus broad, large, with two elongated accessory projections. Vesica balloon-like, slightly scobinate, separated into fairly distinct diverticula. Female. Head: Similar to male, but broader, antennae and labial palpi smaller. Thorax: As in male. Legs: As in male, though tibial spurs thicker. Forewing dorsum: Forewing length: 10.0-18.5 mm, wingspan: 22.0-34.5 mm. As in male but slightly broader, postmedial line usually more noticeably bent. Forewing ventrum: Similar to forewing ventrum of male, but veins usually lined with contrasting yellow scales. Hindwing dorsum: Coloration and markings as for forewing dorsum. Hindwing ventrum: Follows same pattern as forewing ventrum. Frenulum absent. Abdomen: As in male but slightly more robust. Tergite VIII as three posteriorly directed lobes or as single broad plate, sternite VIII as wrinkled mass consisting of one or two pieces, covered in thick, branched setae. Genitalia: Stout, robust or quite narrow overall; apophyses anteriores highly reduced, apophyses posteriores elongate, spanning length of segment IX. Lamella indistinct due to large sclerotization of sternite VIII. Corpus bursae large, bag-like, coiled, broadly connected beneath sternite VIII /ostium complex, no clear ductus bursae present, occasionally with large, snake-like spermatophore present within corpus bursae. Papillae anales typical of Mimallonidae, appearing rectangular laterally; papillae anales covered with fine setae.  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 T. bruna sp. n. is more earthen brown in color than T. rivara, with a longer postmedial lunule that reaches the wing margin without becoming diffuse. In both sexes, T. rivara has a smaller wingspan than T. fuscosa and lacks a distinct dark brown patch of scales at the apex of the forewings. This patch of scales is distinct in T. 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 T. rivara the valves are reduced to small lobes connected to a flattened, spiny accessory, while the valves of T. fuscosa and T. bruna sp. n. are larger and not connected to, or lack the accessory arms. The gnathos of T. rivara is a rectangular plate rather than ovoid as in T. fuscosa, or tapered hexagonal as in T. bruna sp. n., the fingerlike projections at the base of the valves are smaller in T. rivara, and finally the phallus of T. rivara (and T. bruna sp. n.) bears a smooth dorsal projection whereas the same projection is short and spined in T. fuscosa. The key differences in female genitalia are the larger corpus bursae in T. rivara and the trilobed tergite VIII, which is a broad, singular plate in T. fuscosa. The female tergite can usually be examined under a microscrope after brushing off scales, without dissecting the specimen.

Key to species of Tarema*
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, T. 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 T. rivara larvae feeding on Vochysiaceae, including the species Qualea grandiflora, Q. multiflora, and Q. 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, C. packardii  Remarks. In the original description of T. 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 T. rivara at the time of its original description, thus it is somewhat surprising that he described the female of T. rivara as a new species, T. macarina, over 30 years later (Schaus 1928). We are also aware of a T. rivara female specimen determined as this species and not T. 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 T. macarina in the MNHU as per the following statement from Schaus (1928) in his treatment of T. macarina: "A male of this species is in the Berlin museum." Two syntypes of T. 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 T. macarina and the unusual disparity of female T. rivara led us to believe that these names are synonyms. Many records of T. rivara and T. macarina are sympatric, and thus support this hypothesis. A close analysis of T. rivara females reveals hints of orange coloration antemedially, a coloration abundant in the male specimens, but not in T. fuscosa. Given that T. fuscosa females can be easily determined as such due to the lack of sexual dimorphism in this species, the disparity of opposite sexes for T. rivara and T. macarina provides clear evidence that they represent a single, dimorphic species. Furthermore, a dissection of T. rivara females reveal the long, snake-like spermatophore seen in male T. rivara, unlike the smaller spermatophore of T. fuscosa. Diniz et al. (2013) figure a larva and adults of both sexes of T. rivara, correctly figuring the female of T. rivara, which again, matches the type specimens of T. macarina.
Compared to T. 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 T. fuscosa, but apparently not synchronic. In these regions of sympatry, T. rivara flies during the summer (October through February), while T. fuscosa flies in the winter, though exceptions to these flight times are present in regions where both species are not found together. the apex of the forewings surrounded by pale cream markings. The male genitalia is unique in having heavily sclerotized, spiny, club-like vincular arms that are not connected lengthwise to the valves. This is also the only species in the genus with an ovoid gnathos plate. Additionally, the dorsal projection of the phallus is short and spiny, not smooth as in the other two species. The female genitalia have a smaller corpus bursae than in T. rivara and a broad singular tergite VIII, as opposed to the trilobed corresponding tergite of T. rivara.

Tarema fuscosa
Description. Male. Head: As for genus, grayish brown; antenna coloration usually as for head, though pectination darker brown than flagellum; labial palpus reduced, apparently three segmented, but third segment much reduced. Thorax: Coloration similar to that of head, though appearing hoary due to banded brown and pale khaki to cream colored scales, prothorax with more heavily concentrated khaki or cream colored scales. Legs: Coloration as for thorax, though femur and tibia darker brown, tarsus lighter, cream colored. Forewing dorsum: Forewing length: 11-16 mm, avg.: 13.8 mm, wingspan: 22-31 mm, n=17. Ground color ranging from pale reddish brown to nearly black, overall generously shaded by cream colored scales giving the wing a hoary, layered appearance. Postmedial line as for genus, but coloration light cream not bordered externally with black except for darkened region concentrated near to tornus. Ante-and median areas usually concolorous, submarginal area with reduced cream colored scales, appearing much darker red-brown, brown, to nearly black. Apical half of submarginal area with postmedial lunule, the latter either slightly curved toward wing margin, or nearly parallel with margin, especially along apical half of lunule, basal half of submarginal area with darker red-brown or black patch along postmedial line, apex with darker brown patch outlined by white lunule and cream colored patch immediately beneath darker apical patch. Costa appearing lighter than most of wing due to heavy concentration of cream or khaki colored scales. Discal spot as for genus. Fringe light gray to khaki with lighter and darker patches. Forewing ventrum: Similar to dorsum but usually lighter due to more extensive covering of cream and khaki scales, some of which appear yellowish, apical half of submarginal area darker than that of dorsum, except where interrupted by lighter band below apical patch; antemedial line always absent, postmedial line never straight, angled outward toward wing margin mesally. Hindwing dorsum: Coloration as for forewing dorsum, following similar patterning but antemedial line absent, postmedial line slightly concave, and submarginal area more uniformly dark reddish brown, dark brown, or black, always with contrasting orange patch of scales mesally. Hindwing ventrum: Following same pattern as forewing ventrum, postmedial lunule reduced to straight, faint streak. Abdomen: As for genus, concolorous with thorax. Genitalia: (Fig. 12) n=7. Vinculum somewhat ovoid, ventrally with reduced saccus. Uncus robust but reduced to slightly triangular stump with slight bidentation terminally. Gnathos an ovoid, elongated, mesally indented plate. Valves short, rounded, weakly sclerotized mesally. Base of valves with pair of long, fingerlike projections. Valves with more heavily sclerotized, spined accessory arms connected to vinculum. Accessory arms narrow and tube-like, terminating in enlarged club end with spines concentrated terminally or present along entire length of arm. Diaphragm with pair of horsetail-like setal patches consisting of very long setae that extend outward over phallus below gnathos plate, setae curled backward at end. Phallus broad, large, widened distally, with two elongated accessory projections, one projection more variable in length, superior to phallus, irregular, pointed, spined; other projection longer, narrower, running laterally along phallus originating from within phallus, tip of second projection sharp, angled backward. Vesica balloon-like, slightly scobinate, separated into fairly distinct diverticula. Female. Head: Similar to male, but broader, antennae and labial palpi smaller. Thorax: As in male, though cream colored scales may be a bit yellower. Legs: As in male. Forewing dorsum: avg.: 17.5 mm,n=3. Sexual dimorphism reduced, as in male but slightly broader, postmedial line usually more noticeably bent. Forewing ventrum: Similar to forewing ventrum of male, but veins usually lined with yellow scales. Hindwing dorsum: Coloration and markings as for forewing dorsum, orange mesal patch present in male very faint in female. Hindwing ventrum: Follows same pattern as forewing ventrum. Abdomen: As in male but slightly more robust. Tergite VIII as single broad plate, sternite VIII as wrinkled mass consisting of two pieces, covered in thick, branched setae. Genitalia: (Fig. 15) n=1. As for genus but particularly stout, robust. Corpus bursae somewhat reduced in size. Distribution (Fig. 16). Although we report a few records from central Brazil (Distrito Federal), most records of this species are restricted to the Brazilian Atlantic Forest in the states of Bahia, Espírito Santo, Rio de Janeiro, São Paulo, Paraná, and Santa Catarina. See remarks for potential issues pertaining to the Cerrado records.
Remarks. As mentioned in the remarks of T. rivara, T. fuscosa shows a trend in distribution where it is more commonly encountered in humid Atlantic Forest than elsewhere in Brazil. Although we do have some records of T. fuscosa from Distrito Federal, there is a possibility that they were mislabeled. Out of over 100 examined specimens of T. fuscosa, the only Cerrado material was from the same collector, who also collected in regions where T. fuscosa would be more expected, such as the Brazilian states of Paraná and São Paulo. We could not locate any T. fuscosa specimens from the Cerrado among the Mimallonidae specimens collected there in the USNM, NHMUK, or CPAC.
Prior to this work, the female of T. fuscosa was not reported in the literature; therefore we describe and figure it here for the first time. Diagnosis. Externally this species is most similar to T. rivara, but can be easily distinguished by the earthen brown and clay brown coloration rather than orange or red-orange in T. rivara. Additionally, in T. bruna sp. n. the postmedial lunule reaches the wing margin without becoming highly diffuse, and is parallel to the wing margin for its entire length until bending outward. The male genitalia is unique in the extreme reduction of the heavily sclerotized vincular/valve arms, present as a small extension at the base of the valves. The phallus is also unique in the thinness of the dorsal projection, which is smooth as in T. rivara, not spined as in T. fuscosa.
Description. Male. Head: As for genus, coloration earthen brown. Thorax: Coloration similar to that of head, appearing hoary due to banded gray and cream colored scales interspersed amongst brown ones, prothorax covered almost entirely in these lighter scales. Legs: As for genus. Forewing dorsum: Forewing length: 13 mm, wingspan: 26 mm, n=1. Ground color a mixture of earthen brown tones and clay-brown, overall generously shaded by cream colored scales giving the wing a hoary, layered appearance, especially medially. Antemedial line faint, brown, wavy. Postmedial line as for genus but wavier, coloration light cream, not bordered by darker scales except for a small external portion above the tornus. Antemedial area with salmon orange hue, medial area lighter brown compared to darker submarginal area. Apical half of submarginal area with postmedial lunule, the latter parallel with margin, then smoothly curved toward margin reaching wing margin without becoming diffuse, basal half of submarginal area darkest brown, apical portion external to lunule lighter brown. Discal spot as for genus. Fringe light cream with lighter and darker patches. Forewing ventrum: Similar to dorsum but lighter due to more extensive covering of gray and cream colored scales; antemedial line absent, postmedial line 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 straight and faintly outlined by black scales, postmedial lunule very faint. Hindwing ventrum: Following same pattern as forewing ventrum, though discal mark very dark, well defined as black oval. Abdomen: As for genus, concolorous with thorax. Genitalia: (Fig. 13) n=1. Vinculum somewhat ovoid, ventrally with reduced saccus. Uncus robust but reduced to slightly triangular stump. Gnathos a tapered, elongated, hexagonal plate. Valves short, rounded. Base of valves with pair of long, fingerlike projections. Slightly more strongly sclerotized, small projections emanate from base of valves. Diaphragm with pair of horsetail-like setal patches consisting of very long setae that extend outward over phallus below gnathos plate, setae curled backward at end. Phallus broad, large, widened distally, with two elongated accessory projections, one projection superior to phallus, smooth, narrow, pointed, other projection longer, narrower, running laterally along phallus originating from within phallus, tip of second projection sharp, angled forward. Vesica balloonlike. Female. Unknown. Distribution (Fig. 16). This new species is so far known only from the type locality at Paranapiacaba (previously known as Alto da Serra, a train station), São Paulo, Brazil. According to GoogleEarth, the elevation at this locality is approximately 700 m.
Etymology. This species is named for its brown (bruna Latin) coloration, which largely distinguishes it from the red or orange T. rivara and the black, gray, and creamcolored T. fuscosa.
Remarks. The discovery of a unique new species of Tarema from eastern São Paulo is surprising because this is a relatively well-surveyed region of Brazil (R. A. St. Laurent pers. obs.). Both T. rivara and T. fuscosa have been collected from the type locality of T. bruna (NHMUK), though at different times of the year. As previously mentioned in the remarks of T. rivara, that species is primarily a summer species, with records from Paranapiacaba in January, while T. fuscosa has only been collected there in the winter (June and July). More material of T. bruna will be needed to verify its voltinism.
An issue is presented by the fact that the type localities of T. rivara, T. macarina, and T. bruna are all from São Paulo, Brazil with specific type locality information from within the state only available for T. bruna. Therefore, the possibility arose that the name T. macarina could be wrongfully synonymized with T. rivara if indeed it is conspecific with the new species described herein. However, we consider the apparent rarity of T. bruna combined with the genitalia similarities between the lectotype of T. macarina and other T. rivara females dissected from São Paulo and elsewhere, including Cerrado regions, more suggestive that the name T. macarina does in fact represent the female of T. rivara, a much more commonly collected and widespread species. If future evidence were found to contradict our hypothesis, T. bruna would then be a junior and subjective synonym of T. macarina. It is therefore necessary that more material of T. bruna be found or collected, particularly in aim to locate the female of this species.