Research Article |
Corresponding author: Ryan A. St Laurent ( rstlaurent@flmnh.ufl.edu ) Academic editor: Christian Schmidt
© 2017 Ryan A. St Laurent, Daniel Herbin, Carlos G. C. Mielke.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
St Laurent RA, Herbin D, Mielke CGC (2017) Revision of the genus Tarema Schaus, 1896 (Lepidoptera, Mimallonoidea, Mimallonidae) with the description of a new species from southeastern Brazil. ZooKeys 646: 119-137. https://doi.org/10.3897/zookeys.646.10897
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The genus Tarema Schaus, 1896 is revised. The species T. fuscosa Jones, 1908 and T. 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 T. rivara. Tarema bruna sp. n. is described from São Paulo, Brazil. Lectotypes for T. fuscosa, T. rivara, and T. macarina are here designated.
Distribution, Neotropical, Paraguay, Tarema bruna sp. n., Tarema fuscosa , Tarema macarina syn. n., Tarema rivara , taxonomy
The genus Tarema Schaus, 1896 has been mostly overlooked in the literature since
Since
Dissections were performed as in
The primary types (when abdomen was present) and at least one specimen from most localities were dissected.
Specimens from the following collections were examined:
CDH Coll. Daniel Herbin, Garidech, France
CGCM Coll. Carlos G. C. Mielke, Curitiba, Paraná, Brazil
MNHU
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 (
Tarema rivara Schaus, 1896: 55, by original designation.
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.
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
1 | Male: coloration mostly clay-brown or orange to more red with abundant gray shading. Valves with heavily sclerotized, flattened, spined structures, or no accessory arms. Phallus with smooth dorsal projection. Female: mostly brown and gray, but may have clay or salmon-orange hue, especially antemedially, darker apical patch absent, tergite VIII trilobed | 2 |
- | Male: coloration mostly dark brown, to nearly black with cream colored shading; valves with heavily sclerotized, tubular, clubbed spine structures; phallus with spined dorsal projection. Female: mostly dark brown to black with gray shading, tergite VIII as singular plate | Tarema fuscosa |
2 | Ground color clay-orange to orange-red, postmedial lunule angled away from postmedial line toward wing margin, becoming diffuse before reaching wing margin. Male genitalia with heavily sclerotized, flattened, spined structures, connected lengthwise to short, rounded valve, gnathos as rectangular plate, dorsal phallus projection broad. Female: tergite VIII with trilobed plate | Tarema rivara |
– | Ground color earthen brown, postmedial lunule parallels postmedial line until bending toward wing margin, reaching it. Male genitalia lacking large, heavily sclerotized vincular/valve arms, which are instead reduced to small sclerotizations at base of valve, gnathos as tapered hexagonal plate, dorsal phallus projection thin | Tarema bruna sp. n.* |
*The female of T. bruna sp. n. is unknown.
Tarema rivara Schaus, 1896: 55
Tarema
rivara
;
Tarema
rivara
;
Tarema
rivara
;
Tarema
rivara
;
Tarema macarina Schaus, 1928: 670, fig. 88i ♀ syn. n.
Tarema
macarina
;
Tarema
macarina
;
Tarema rivara Schaus: lectotype [here designated], ♂. BRAZIL: São Paulo: São Paulo, S.E. Brazil./ Collection, WmSchaus/ Type No. 12566 U.S.N.M/
Tarema macarina Schaus: lectotype [here designated], ♀. BRAZIL: São Paulo: 1 ♀, São Paulo, S.E. Brazil./ Collection, WmSchaus/ Type No. 33596 U.S.N.M/
(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-
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.
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.
Tarema rivara adults, a dorsal b ventral. 1 ♂, Lectotype, Brazil, São Paulo (
Tarema adults, a dorsal b ventral. 7 T. bruna holotype ♂, Brazil, São Paulo, Alto da Serra [Paranapiacaba] (
Tarema male genitalia, a ventral b lateral c phallus lateral. 11 T. rivara, Brazil, Distrito Federal, Estação Florestal, Cabeça do Veado, 1100 m, St Laurent diss.: 3-14-16:7 (
Tarema female genitalia, a ventral b dorsal. 14 T. rivara, Brazil, Distrito Federal, Estação Florestal, Cabeça do Veado, 1100 m, St Laurent diss.: 3-14-16:8 (
(Fig.
Known distribution of Tarema. Numbers superimposed on the map refer to the following annotations: 1 Outlier data point for T. fuscosa in Brazil, Distrito Federal may be erroneous (see remarks for that species). 2 The point where all three symbols are found on top of each other is a single locality, that being Brazil, São Paulo, Paranapiacaba. 3 Data point for T. rivara in Brazil, Santa Catarina is placed at the center of the state because no detailed locality data is available for this species from Santa Catarina.
In the original description of T. rivara,
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.
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.
Tarema fuscosa Jones, 1908: 173–174
Tarema
fuscosa
;
Tarema
fuscosa
;
Tarema
fuscosa
;
Lectotype [here designated], ♂. BRAZIL: Paraná: Castro, Paraná, 950 m, E.D. Jones / Tarema fuscosa Type D. Jones/ E.D. Jones Coll., Brit. Mus., 1919–295/ BMNH(E) #805428/ SYN-TYPE/ NHMUK010354542/ [genitalia] VIAL NHMUK010402134/ LECTOTYPE male Tarema fuscosa designated by St Laurent, Herbin, and C. Mielke, 2017 [handwritten red label]/ (
(114 ♂, 4 ♀) BRAZIL: Distrito Federal: 1 ♂, Estação Florestal, Cabeça do Veado, 1100 m: 17.X.1971, E.G., I. & E.A. Munroe leg., St Laurent diss.: 3-14-16:10 (
Compared to the other two species in the genus, T. fuscosa is easily recognized by the very dark brown to nearly black ground color, with a dark patch at 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.
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.
(Fig.
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
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.
Holotype, ♂. BRAZIL: São Paulo: Alto de [recte da] Serra, [Paranapiacaba, Santo André], São Paulo, November, 1922. (R. Spitz [leg.])./ Rothschild Bequest BM 1939-1/ [genitalia] VIAL NHMUK010402168 / NHMUK010318286/ HOLOTYPE male Tarema bruna St Laurent, Herbin, & C. Mielke, 2017 [handwritten red label]/ (
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.
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.
(Fig.
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 cream-colored T. fuscosa.
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 (
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.
The Cornell University College of Agriculture and Life Sciences Alumni Association award and Thomas Witt (MWM) covered travel expenses of R. A. St. Laurent. Stefan Naumann (Germany) provided photographs of all Tarema specimens at MNHU. Alexey Prozorov (MWM) offered assistance in photographing specimens and performed and photographed genitalia preparations of Tarema in MWM. Alessandro Giusti (