Research Article |
Paul Z. Goldstein‡, Alberto Zilli§
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Corresponding author: Paul Z. Goldstein ( drpzgoldstein@gmail.com ) Academic editor: Donald Lafontaine
© 2019 Paul Z. Goldstein, Alberto Zilli.
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:
Goldstein PZ, Zilli A (2019) Thraumata, a new genus from South America with description of a new species from Peru (Lepidoptera, Noctuidae). ZooKeys 867: 139-160. https://doi.org/10.3897/zookeys.867.28728
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Abstract
Thraumata gen. nov. is described to accommodate three South American species, two previously placed in Phuphena Walker, 1858, namely Thraumata petrovna (Schaus, 1904), comb. nov. and Thraumata subvenata (Schaus, 1914), comb. nov.; and one, Thraumata peruviensia sp. nov., newly described from Peru. Although the larval biology is unknown, these species share several features that suggest their placement in Eriopinae and, as a consequence, a potential association with ferns (Pteridophyta) as larval host plants.
Keywords
Phuphena, Noctuidae, owlet moths, Peru, systematics
Introduction
Phuphena Walker, 1858 appears to represent a polyphyletic assemblage that includes a morphologically homogeneous, putatively monophyletic core group of 10 described species: P. cilix (Druce, 1898), P. constricta Dognin, 1912, P. diagona Hampson, 1908, P. multilinea Schaus, 1911, P. obliqua (Smith, 1900), P. parallela (Hampson, 1904), P. proselyta Schaus, 1921, P. transversa (Schaus, 1894), P. tura (Druce, 1889), P. zelotypa Schaus, 1911 and the type species P. fusipennis Walker, 1858. Outside this assemblage are Phuphena costata Schaus, 1914, which will be treated elsewhere, and two of the species treated here, P. petrovna (Schaus, 1904) and P. subvenata Schaus, 1914. Of the 15 available names for species in the genus Phuphena, William Schaus described eight in six publications between 1894 and 1921. Three of these species he originally placed in other genera, in some cases denoting the generic name with a question mark, and among these was P. petrovna (Schaus, 1904), which
Materials and methods
Genitalic preparations follow
Prior to dissection, the abdomens of several specimens underwent a 24-hour soak in a 10% proteinase K solution in the Laboratory of Analytical Biology at
Exemplars used for comparative purposes included specimens of generotypes within Phosphilini and Eriopinae, including Phuphena, Phosphila Hübner, 1818, Acherdoa Walker, 1865, and Callopistria Hübner, [1821].
Repository abbreviations
The following abbreviations refer to collections from which specimens forming the basis of this study originated:
Systematics
Phuphena
Type species
Phuphena fusipennis Walker, 1858 by monotypy.
Diagnosis
Species of Phuphena diverge with respect to wing pattern, but most share a bounded medial area. Orbicular and reniform stigmata sometimes absent or indistinct; outer forewing margin smooth in the majority of species but crenulate in cilix. Hindwing upperside coloration uniform. Male genitalia of Phuphena distinct, the valvae uniquely reduced, with no evidence of clasping architecture, narrow for most of their length, and in most species swollen apically to form a knob-like cucullus, giving them a club-shaped appearance, with a corona comprising an unorganized arrangement of setae. Vinculum V-shaped, slightly truncate in some species, and extending below the base of the valves for a distance equal to or greater than the width at its broadest point. The phallus is asymmetrically sclerotized, often with a patch of scale-like cornuti, and the vesica without well-developed cornuti or diverticula, but with a corresponding ventro-distal strip of scale-like cornuti. Basal abdominal brushes present but not visibly subdivided as in Callopistria Hübner, [1821] (type species: Noctua juventina Stoll, 1782); a smaller pair on A8, and eversible saccular hair pencils completely absent. Female genitalia simple, the corpus bursae lemon-shaped, with no appendix or signa, the ductus seminalis attached apically, to the distal (anterior) end of the corpus. This distinguishes Phuphena from all Thraumata spp. and from Callopistria juventina, in which the ductus arises from the proximal (posterior) end of the corpus, but this character varies among other Callopistria species.
Thraumata gen. nov.
Type species
Leptina ? petrovna Schaus, 1904, by present designation.
Etymology
Thraumata is the nominative neuter plural form of the Greek θραυμα, meaning shard, fragment, or potsherd, in reference to the fragmented appearance of the forewing pattern.
Dorsal habitus of Thraumata species. 1 T. subvenata Holotype ♂, Guyana, USNMENT00973242 2 T. subvenata, USNMENT01440366♀, Suriname 3 T. petrovna ♂ USNMENT01440358, Brazil 4 T. petrovna Holotype ♀ USNMENT00973124, Costa Rica 5 T. peruviensia Holotype ♂ USNMENT01440371, Peru 6 T. peruviensia ♀ USNMENT01440374, Peru 7–9 Holotype specimen labels of Thraumata species 7 Phuphena subvenata Schaus 8 Leptina petrovna Schaus 9 Thraumata peruviensia sp. nov.
Diagnosis
Species of Thraumata are readily distinguished from Phuphena and Callopistria by the fragmented appearance of forewing pattern elements, the differential coloration of the hindwing upperside, paler towards the base with more darkly shaded margins, and the configuration of male and female genitalia, as follows: Valves without clasper complex (as in Phuphena, Callopistria, and Phosphila), but tapered and not clublike, unlike Phuphena, and with no differentiated cucullus or corona. Vinculum squared, less V-shaped than in Phuphena, extending below the valves for a distance less than the width at its broadest point. Vesica with multiple basal lobes and without cornuti. Ventrally recurved basal abdominal brushes present in subvenata; saccular coremata present and, except in subvenata, less developed than in Callopistria. The female genitalia are distinct, the corpus bursae without signa, elongate and bowed, not lemon-shaped as in Phuphena, the ductus bursae attaching postero-dorsally to the corpus, and the ductus seminalis arises from the posterior end of the corpus as in Callopistria juventina, not at the anterior apex as in all known Phuphena and several Callopistria species.
Description
Head . Frons fully scaled; eye large, hairless; antenna filiform, shortly setose-ciliate in both sexes; labial palpus slightly upturned and densely scaled, third segment reduced compared to the first and second segments; proboscis well developed. Thorax. Vestiture variously brown or grayish-brown scaling intermixed with white. Wings. Forewing elongate, not broadly rounded, the apex acute and the outer margin slightly angled at the middle; distal field crossed by conspicuous dark fascia outwardly produced at middle with triangular projection. Hindwing elongate too, slightly produced in correspondence to vein M3. Sexual dimorphism most obvious in hindwing, female with slightly darker shading towards wing base than male, but this character is neither discrete nor reliable. Underside of both wings brown or grayish-brown, the most prominent marking that of the toothed forewing submarginal line; Hindwing with 1–3 elongate pale streaks. A clearly expressed M2 visible on hindwing, arising from the discoidal cell closer to M3 than in Callopistria, possibly representing an autapomorphic condition for Thraumata. Legs. Outwardly pale, silvery white in petrovna and peruviensia sp. nov.; otherwise brown or with a mixture of brown and whitish scales. Foretibial epiphysis rugose; a single pair of striped mid-tibial spurs, two pair on hind-tibia; 3 rows of tibial spines on all legs. Abdomen. Vestiture usually paler than on thorax and concolorous with hindwing surface; abdominal segments typically pale distally, more darkly banded at the anterior end of each segment, with darker scaling more uniformly diffuse below; typical trifine brush organs absent except in T. subvenata comb. nov., which bears a pair of tufts on male sternum A2, without pockets or levers; other species exhibit raised lateral flanges on sternum A2; posterior margin of 8th sternite incised; rods extending from base of sternum A8 to pleurae. Male genitalia. Tegumen raised at base of uncus; vinculum short relative to tegumen; paratergal sclerite visible but fused with tegumen. Valve tapered, not extending beyond tegumen, finely setose throughout and most heavily towards base. Uncus setose, swollen apically, terminating in a small point. Juxta tightly joined with valve and transtilla. Phallus asymmetrically sclerotized, narrow in its basal half; vesica with multiple bubble-like subbasal diverticula and a weakly to heavily sclerotized para-basal plate or lip from the distal end of the phallus; fremen of spermatophore with a well-developed nipple. Female genitalia. Ductus bursae short, membranous, without colliculum; appendix bursae present at posterior end of corpus bursae, but not well differentiated; ductus seminalis arising from appendix bursae; corpus bursae oblong, without signa. Posterior apophyses rodlike, distally modified faintly, if at all. The bowed, ventrally facing configuration of the intersegmental membrane between A8 and A9 deformed so as to orient the corpus bursa at an angle with respect to the ventrally faceing papillae anales.
Immature stages
Unknown.
Distribution
Recorded from northern South America, southeastern Brazil, southern Peru, Bolivia, and Argentina along the eastern Andean slopes (Fig.
Key to species of Thraumata based on habitus
| 1 | Ground color of FW predominantly dark brown, all markings conspicuously highlighted in silvery white | T. subvenata |
| – | FW dominated by gray, with dark brown fascia in distal field bearing outwardly produced tooth, white lining confined to outer edge of this fascia (submarginal line) | 2 |
| 2 | FW length less than 13 mm (males) and 14 mm (females), toothed projection not reaching termen | T. petrovna |
| – | FW length greater than 14 mm (males) and 15 mm (females), toothed projection extended to termen | T. peruviensia |
Thraumata petrovna , comb. nov.
Leptina
? petrovna Schaus, 1904: 152. Type locality: Brazil [Rio de Janeiro], Petropolis. Figure:
Material examined
Type material. BRAZIL: Holotype ♀ (
Ventral habitus of Thraumata species. 10 T. subvenata Holotype ♂, Peru, NHMUK010606200 11 T. subvenata, USNMENT01440366 ♀, Suriname 12 T. petrovna ♂ USNMENT01440358, Brazil 13 T. petrovna Holotype ♀ USNMENT00973124, Costa Rica 14 T. peruviensia Holotype ♂ USNMENT01440371, Peru 15 T. peruviensia ♀ USNMENT01440374, Peru.
Other material examined
BRAZIL (34♂, 39♀): Paraná (7♂, 7♀): Males: Banhado, Quatro Barras, Paraná, BRASIL – 800 m, 22.V.1971, Becker & Laroca, USNMENT01440489; Banhado, Quatro Barras, Paraná, BRASIL – 800 m 22.V.1971, Becker & Laroca; Phuphena petrovna (Schs, 1904), USNMENT01440447; Banhado, Quatro Barras, Paraná, BRASIL – 800 m, 27.II.1971, Becker & Laroca; Phuphena petrovna (Schs, 1904), USNMENT01440439; Castro. Parana.; petrovna Schs; Collection Wm Schaus, USNMENT01440466; Castro [ex coll. E.D. Jones] [
Diagnosis
This and the following species are readily differentiated from T. subvenata by the gray-brown ground coloration and less prominent white markings throughout the forewing. Thraumata petrovna is, on average, the smallest of the three species, and although its wing pattern closely tracks that of the larger peruviensia, the postmedial tooth of the forewing is shorter, not reaching the wing margin; the vesica is distinctly more sclerotized, bearing a pronounced lateral band and a conspicuous, rugose plate.
Redescription
Head. Vertex and frontal scaling predominantly white but with a mixture of light gray-brown scales throughout the vertex, frons and palpi; frontal scales elongate. Thorax. Vestiture a mix of white and grayish-brown scales. Wings. Forewing lengths: males 10.8–12.9 mm (N =13, x¯ = 12.2 mm, M = 12.5 mm); females (including holotype 12.1 mm) 11.0–13.7 mm (N =22, x¯ = 12.4 mm, M = 12.5 mm). Forewing ground color predominantly gray, with darker chocolate-brown buff in lower basal field, both below and between stigmata, and in toothed fascia of distal field; hindwing pale, brownish-gray towards margin; forewing and hindwing underside with diffuse chocolate-brown scaling, the shading on the hindwing concentrated between veins, fading towards hind margins on both forewing and hindwing; hindwing underside with a single conspicuous median pale streak. Legs. Outwardly pure white; otherwise light brown. Abdomen. Abdominal segments with cream-colored scales distally, darker brown-banded at the anterior end of each segment; brown scaling more uniformly diffuse below. Abdominal brushes lacking. Male genitalia. Tegumen rooflike, a distinct vertex and a pitch >45°. Sacculus bluntly rounded. Phallus heavily sclerotized, swollen and rugose along its outer edge; vesica with two rudimentary subbasal diverticula, and para-basal plate arising from distal end of the phallus. Female genitalia. Posterior apophyses roughly 1.3× as long as anterior apophyses; papillae anales densely setose.
Immature stages
Unknown.
Etymology
Not explained, but evidently in reference to the type locality Petropolis (“City of Peter”).
Distribution
Southeastern Brazil and along the eastern slopes of the Bolivian and Argentinian Andes.
Remarks
A partial barcode sequence (658bp – 286n = 372bp) was generated from a Brazilian male specimen USNMENT01440385 corresponding to
Thraumata peruviensia sp. nov.
Material examined
Type material. PERU (11♂, 6♀): Holotype ♂ (
Diagnosis
Thraumata peruviensia is most readily distinguished from the other two species by the combination of its larger size and distinctive vesica. In this and the previous species, the dominant ground color of forewing is gray, darker in the present species than in T. petrovna. This feature, the less pronounced white lining of pattern elements, particularly in basal and medial fields of forewing, and the comparatively reduced abdominal scent tufts in the male most readily distinguish this species pair from the more uniformly brown ground coloration of T. subvenata. In the new species the triangular tooth projecting from the dark brown distal fascia of the forewing is longer than in T. petrovna, and there are at least two distinct white streaks on the hindwing underside instead of one.
Description
Head. Vestiture as in T. petrovna. Thorax. Vestiture a mix of grayish-brown scales intermixed with white, especially within tegulae. Wings. Forewing lengths: males (including holotype 15.3 mm) 14.2–15.3 mm (N = 3, x¯ = 15.0 mm, M = 12.5 mm); females, 15.5–16.8 mm (N = 3, x¯ = 16.1 mm, M = 16.1 mm). Forewing coloration dominated by gray scaling suffused with white, lines for the most part obscured, although in some individuals antemedial and postmedial lines still discernible as faint, thin pale gray lines. Four areas of more uniform, chocolate-brown scaling: an elongate, obtuse subtriangular smudge in lower basal field from wing base to antemedial line; an oblique band in place of median shading between from reniform stigma and inner margin, dark filling between orbicular and reniform stigmata, and in fascia bearing outwardly projected median tooth of distal field, this fascia lined externally with silvery white. Forewing underside nearly uniform chocolate brown excepting paler white inner marginal area; the shading on the hindwing underside interrupted by two-three conspicuous pale streaks, fading towards wing base. Legs. White outward, otherwise as brown as in previous species. Abdomen. Abdominal segments predominantly gray, ringed with pale cream-colored scales distally; abdominal brushes lacking. Male genitalia. Sacculus squared. Sclerotized lateral band on phallus less pronounced than in petrovna, vesica with three rudimentary subbasal diverticula and reduced para-basal plate from distal end of the phallus. Female genitalia. Ductus bursae quite short relative to distended corpus bursae. Anterior and posterior apophyses roughly co-equal. Papillae anales densely setose. As in other Thraumata, the ductus seminalis arises from a bulge at the posterior end of the corpus bursae.
Immature stages
Unknown.
Etymology
The name peruviensia is a neuter pleural adjective, in reference to the country of origin of the new species, in coordination with the generic name Thraumata.
Distribution
Southeastern Peru.
Remarks
This species is very similar in appearance to T. petrovna but consistently larger and occurring at higher altitudes. An incomplete barcode sequence (658bp – 88n = 570bp) was generated from a 1905 male specimen, USNMENT01440371 corresponding to
Thraumata subvenata
Phuphena subvenata Schaus, 1914: 486. Type locality: [Guyana] British Guiana: Essequebo River, Rockstone.
Material examined
Type material. GUYANA: Holotype ♂ (
Other material examined
(3♂, 7♀). GUYANA: 1♀ Demerara; Rothschild Bequest, B.M. 1939-1 [
Diagnosis
The predominantly golden-brown coloration of the forewing upperside and the prominent white edging of its markings readily distinguished this species from T. petrovna and T. peruviensia. Valvae are more broadly articulated with the vinculum than in the other species, the costal edge <1/2 the length of the outer edge. Unlike the two predominantly gray-brown species, T. subvenata also bears basal abdominal brushes and what appear to be well-developed eversible coremata on the sacculus.
Redescription
Head
. Proboscis edged with papillae and a micro-serrated ridge. Antennae ciliate; eyes hairless; palpi extending above eyes. Frons, vertex and palpi with a mix of brown and white scales; terminal segment of the palpus whitish. Thorax. Vestiture predominantly light brown, a whitish V formed by two lines of whitish scales within the tegulae. Wings. Forewing length, males (N = 2), 11.5 (holotype)–12.0 mm; females (N = 3), 12.1 mm, 12.4 mm, 13.0 mm. Forewing coloration dominated by golden-brown scaling; the differential white shadowing, of various pattern elements, particularly the postmedial and subterminal lines, renders a fractured appearance consisting of streaks and sickle-shaped forms: two basal white streaks, 2 crescent-shaped medial markings bisected by brown inner line and two, more expansive, subterminal forms, each enclosing a brown center. These latter four simply bracket the broad, dentate subterminal band homologous to that in the previous two species. Legs. Predominantly brown, paler outwardly but not silvery white. Male fore-femur with a thick, cream-colored pencil of long scales (Figs
Immature stages
Unknown.
Distribution
Known only from northern South America, specifically Colombia, Venezuela, Guyana, Suriname and Brazil (Pará).
Remarks
Both the valvae and the legs of subvenata males are equipped with more conspicuous scent tufts than either of the two sister species. An incomplete barcode sequence (658bp – 88n = 570bp) was generated from a 1925 male specimen from Guyana USNMENT01440389 corresponding to
Discussion
The boundaries of the Eriopinae have yet to be established as several of the characters used to circumscribe the subfamily are either widespread or have been characterized vaguely, and are likely to be revisited during the course of ongoing studies. For present purposes, we suggest the possibility that placement of Thraumata within the Eriopinae on the basis of characters shared by Callopistria (= Eriopus Treitschke, 1825). These include the tapered, ear-shaped valves without a linear corona, the presence in T. subvenata of small saccular coremata on the inner face of the valves, and narrow sclerotization along the basal half of the phallus, which dissipates in the distal third (cf.
The pleural tufts on A8 of T. subvenata (Fig.
DNA barcode data, including partial sequences of the 658bp COI barcode region obtained for all three species of Thraumata, provisionally support the monophyly of both Phuphena and Thraumata and unite them with the eriopine genera Callopistria and Argyrosticta. But these results must be viewed skeptically, given their incompleteness (570bp for both T. peruviensia and T. subvenata, and 372bp for T. petrovna; see Material examined above for BOLD process ID codes of available sequences). Although the larvae of Phuphena have thus far been associated exclusively with ferns and differ in many other respects from the gaudy, semi-gregarious larvae of Phosphilini, which have been recorded only from Smilacaceae, we have as yet only been able to examine larvae indirectly, through images. Without specimens, none of the larval characters discussed by
Thraumata subvenata, holotype (male), foreleg, proboscis, and abdominal structures. 31 Right foreleg, outside lateral view, indicating femoral scale pencil (sp), epiphysis (ep), and proboscis (pr) with papillae (pap) and serrated lateral ridge (sr) 32 Right foreleg, inside lateral view, indicating femoral scale pencil (sp) and epiphysis (ep) 33 Ventral brushes on A2 34 Abdominal terminus, showing distal scale tufts.
Acknowledgements
Ben Proshek (USDA) prepared many of the dissections and figures. David Grimaldi, Courtney Richenbacher, and Suzanne Rab Green (all
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