Revision of the South American genus Gaujonia Dognin (Noctuidae, Pantheinae) with descriptions of five new genera and twenty-one new species

Abstract The endemic Neotropical genus Gaujonia Dognin is revised. Morphological characters and a phylogenetic analysis demonstrate paraphyletic relationships among the species. Four different groups are interpreted to represent four different genera. The G.arbosi group is the only remaining clade in the genus Gaujonia, and the other groups have been arranged into three new genera: Milleranagen. nov., Oculicattusgen. nov., and Cicadoformagen. nov. Additionally, two other genera Cicadomorphusgen. nov., and Gaujopteragen. nov. were found using morphological and molecular analyses based on some specimens that were misidentified as Gaujonia spp. A total of five new genera, three new combinations (Cicadoformavau-nigrum Hampson, comb. nov., Oculicattusrenifera Hampson, comb. nov., and Milleranaarbosioides Dognin, comb. nov.) and 21 new species (Cicadoformaocelotussp. nov., Cicadomorphuschicharrasp. nov., Cicadomorphuschuyasp. nov., Cicadomorphusfalkasiskasp. nov., Cicadomorphuslilianaesp. nov., Gaujoniabichusp. nov., Gaujoniachiqyaqsp. nov., Gaujoniakanakusikasp. nov., Gaujoniasourakovisp. nov., Gaujopteraamsasp. nov., Milleranaaustinisp. nov., Milleranacajassp. nov., Milleranacundinamarquensissp. nov., Milleranamatthewsaesp. nov., Milleranatigrinasp. nov., Oculicattusbolivianasp. nov., Oculicattusbrehmisp. nov., Oculicattusincasp. nov., Oculicattusraizaesp. nov., Oculicattusschmidtisp. nov., and Oculicattusuturunkusp. nov.) are established.


Materials and methods
Genitalia preparation and terminology follow the protocols of Lafontaine (1987Lafontaine ( , 2004 and Schmidt and Anweiler (2020). Genitalia were stained with 10% eosin Y and examined in 30% ethanol. Only the genitalia from type specimens were mounted on slides using Euparal; the remainder were stored in vials of pure glycerin. Pinned adults were photographed prior to dissection using a camera Canon EOS Rebel T5i with a Canon EF 100 mm f/2.8 USM Macro lens. The genitalia were photographed following mounting employing a StackShot automated focus stacking macro rail with a camera Canon EOS 6D and an Infinity long-distance microscope Model K2 DistaMax. Molecular diagnosis was performed by DNA barcoding, employing a segment from the mitochondrial cytochrome oxidase I (COI) gene for 29 species: one previously described species from Gaujonia sensu Hampson, 16 species newly described in this study, and the outgroup sequences of the rest of the jaguar moths along with a specimen from the type genus of Gaujonia (Gaujonia arbosi Dognin) and two other unrelated genera of pantheines (Meleneta Smith and Charadra Walker) sequences were taken from the BARCODE OF LIFE DATA SYSTEM v4 (http://barcodinglife.com). DNA was extracted from leg tissue removed from pinned dry specimens, and Sanger sequencing was performed by the Canadian Centre for DNA Barcoding, Guelph, Ontario (http://ccdb.ca) following the protocols of Hebert et al. (2003). The sequences were concatenated and aligned using Geneious 9.1.3 (https://www.geneious.com). Phylogenetic trees were constructed using a maximum-likelihood (ML) analysis that was performed in IQ-TREE v. 2 to determine relationships among taxa following Nguyen et al. (2015), Hoang et al. (2018), and Minh et al. (2020). Branch support was estimated by performing 1000 replicates each for both ultrafast bootstraps (UFBoot2) ('-bb' command) and SH-aLRT test (SH-aLRT) ('-alrt' command). Phylogenetic relationships with UFBoot ≥ 95 and SH-aLRT ≥ 80 are considered to have strong support. Molecular data and photographs of the voucher specimens are available at BARCODE OF LIFE DATA SYSTEM v4 (project: Life History of Pantheinae). Sequences were submitted to GenBank (https://www.ncbi.nlm.nih.gov/genbank); accession numbers are listed in Suppl. material 1: Table S1.
Specimens were obtained from the following museums and collections: spots; saccular and cucullar regions on the valva are separated; cucullar region squared and wide; aedeagus short; vesica long and wide, with two patches of thin setae; female wings hyaline with the line of scales wider than in male, with a small orbicular spot, but without a reniform spot; anal papilla wide; anterior apophysis small; appendix bursae and corpus bursae long and narrow, appendix bursae heavily sclerotized and smaller than corpus bursae. b) arbosioides group (Millerana): represented by the type Millerana arbosioides misplaced originally in Gaujonia, and five new species: M. austini sp. nov., M. cajas sp. nov., M. cundinamarquensis sp. nov., M. matthewsae sp. nov., M. tigrina sp. nov., which differ from other groups in that the male antenna is serrate, whereas that of the female is filiform; wings are not hyaline; both orbicular and reniform spots are well developed; valva is simple with most species presenting an extension of the apex; aedeagus and vesica are narrow, with a band of spines surrounding the vesica near the base. Female genitalia are unknown. c) renifera group (Oculicattus): represented by the type species Oculicattus renifera and seven new species: O. boliviana sp. nov., O. brehmi sp. nov., O. inca sp. nov., O. raizae sp. nov., O. schmidti sp. nov., and O. uturunku sp. nov; males have hyaline wings and veins covered by a narrow lines of scales; orbicular spot is barely visible; reniform spot is a long horizontal black line; valva with the saccular and cucullar regions separated; cucullar region long and narrow; aedeagus long; vesica long and narrow with one patch of thin setae near the base and two patches, one on each side of tip; female wings similar to those of male with orbicular spot large and line of scales wide; anal papillae narrow; anterior apophysis long; appendix bursae and corpus bursae long, similar in size. d) vau-nigrum group (Cicadoforma): includes the type species Cicadoforma vaunigrum and a new species C. ocelotus sp. nov. Male wings semi-hyaline (slightly translucent) with a large conspicuous orbicular spot and a small reniform spot; valva simple; aedeagus long; vesica with small patch of spines on upper side; female wings similar to those of male, but covered with scales; anal papilla wide; anterior apophysis long; appendix bursae small; corpus bursae broad.
Unfortunately, molecular data could not be obtained for seven of the species in the Gaujonia complex. However, the significant amount of morphological evidence presented in this study is sufficient to justify splitting Gaujonia sensu Dognin-Hampson into four genera, which is corroborated by the COI gene tree (Fig. 1).
Etymology. Cicadoforma refers to how people in South America confuse this group with cicadas.
Included species. The genus Cicadoforma is established to accommodate C. vaunigrum, which was previously included in Gaujonia because of wing pattern similarities; a new species is described in Cicadoforma: C. ocelotus sp. nov. However, there are remarkable differences in morphology and molecular characters, as shown here.
Diagnosis. Cicadoforma is similar to Cicadomorphus, not only externally, but internally as well; however, the phylogenetic analysis results showed enough evidence to separate Cicadoforma in a different genus from Cicadomorphus (Fig. 1). Nevertheless, both genera can be distinguished morphologically by the forewing, which is hyaline with scales only on margins and veins in Cicadoforma, whereas in Cicadomorphus has scales are more widely distributed on the forewing. Genitalia have some small indentations on outer margin of valva that are not present in Cicadomorphus; apex and lobe on valva is much wider in Cicadoforma; upper side of vesica with one patch of spines, and a narrow band of spines near the base are present in Cicadoforma. Female genitalia with square-shaped anal papillae in Cicadoforma; more rounded in Cicadomorphus. DNA barcodes show a closer relationship with Cicadomorphus (~ 5% divergent than with Gaujonia (~ 6%).
Description. Sexually dimorphic mainly in size, female slightly larger than male; forewing in male with some hyaline areas and with poorly developed pattern, whereas female forewing semi-hyaline with pattern better defined. Antenna filiform, black, or dark brown in both sexes; antenna with yellow basal line of scales; eye large, covered by long interfacetal setae; palp with black upper side and yellow underside; haustellum dark brown and reduced, but functional. Forewing with orbicular reniform spots small. Hindwing semi-hyaline with scales only on margin and veins. Male genitalia Figure 1. Maximum likelihood tree showing relationships between the Gaujonia genus group and the rest of the other jaguar moth genera based on the cytochrome c oxidase I gene (COI) marker. Nodes with black circles represent high support (UFBoot ≥ 95 and SH-aLRT ≥ 80). Nodes with gray circles represent low support (UFBoot < 95 and SH-aLRT < 80). Black branches constitute the outgroups. with simple, lightly-sclerotized valva; valva wide apically with some small indentations on outer margin; clasper absent, cucullus wide, with a broad lobe extended in front of sacculus, ear-like in shape; sacculus with a small foot-like process; uncus crooked, long and wide; aedeagus short, with a simple long vesica with a broad patch of spines on upper side and a narrow band of spines at base. Female genitalia with a large squareshaped and lightly sclerotized sterigma, and a rugose, sclerotized appendix bursae; corpus bursae unsclerotized.
Key to species of the genus Cicadoforma based on adult male morphology Etymology. The species name ocelotus is derived from the characteristic yellowish orange coloration on its body, reminiscent of the color of an ocelot.
Diagnosis. Cicadoforma ocelotus can be distinguished from its only congener, C. vaunigrum, by its remarkably brighter ground color, its larger orbicular spot, and shorter space between forewing veins R3 and R4. The valva of the male genitalia of C. ocelotus have concave outer margins, in contrast with the straight margin of C. vau-nigrum.
Description. Head. Palp short with a combination of black, yellow, and orange scales; ground color of frons orange or yellow with scattered black hair-like scales; antenna dark brown. Thorax. Yellow or orange with small spots scattered throughout dorsum. Wings. Forewing length: male 19-21 mm; female 25-27 mm; forewing yellow or orange scales covering veins and margins with semi-hyaline areas between them; subterminal line slightly visible; basal, antemedial, medial, and postmedial lines poorly developed, present only as small black dots on veins; reniform spot narrow with top and bottom surrounded by black scales and a small black dot in middle; orbicular spot small, elongated; female with antemedial, medial, postmedial, and subterminal lines slightly defined; reniform and orbicular spots similar to those of male; R4 almost entirely black; black V-shaped marking at base of CuA2; hindwing semi-hyaline with yellow or orange veins paler than forewing; fringe composed of short yellow or orange hair-like scales except on posterior margin, which has long,  pale-yellow scales. Leg. Yellow or orange with some irregular brown spots that decrease in number from prothoracic legs to metathoracic legs. Abdomen. Covered by brownish orange scales that are paler than those of thorax; brown tufts in A2-A7 with tips with same color as remainder of abdomen. Male genitalia. Cucullus wide, lobe narrow; lobe apex rounded; apex clothed with short setae that expand over entire costal margin; sacculus base narrow; saccus relatively long, V-shaped; tegumen fairly flat; juxta U-shaped on upper side; aedeagus 3 ¾ × longer than wide with opening to vesica same width as aedeagus; vesica 1 ½ × longer than aedeagus with a patch of spines on upper side with some conspicuous basal spines. Female genitalia. Anal pa-  pilla wide, petal shaped, clothed with relatively long setae; A8 membranous, relatively short; posterior apophysis almost same length as anal papilla; sterigma enlarged, lightly sclerotized, fused above ostium; anterior apophysis ⅞ × shorter than posterior apophyses; ductus bursae short, strongly sclerotized and wide; posterior ¾ of appendix bursae strongly sclerotized, remainder including corpus bursae unsclerotized; corpus bursae 1 ¼ × longer than the appendix bursae.
Immature stages. Unknown. Distribution. This species is only known from northern Colombia from moderate elevations from 2000-3000 m, especially in cloud forests (Fig. 92).

Cicadoforma vau-nigrum
Etymology. George F. Hampson likely named this species "vau-nigrum" because of the black V-shaped mark at the base of CuA2.
Diagnosis. Cicadoforma vau-nigrum is the largest species in this genus; males have a forewing length of ca. 24-26 mm, whereas that of females is ca. 30-32 mm. This species is similar to C. ocelotus, but easy to distinguish by the size, and also by the orbicular spot, which is larger in C. vau-nigrum. Another important difference is that C. vau-nigrum males have hyaline wings where only the veins and margins are covered with yellow and black scales; the abdomen has dark brown tufts in the middle of A4-A7. Cicadoforma vau-nigrum can be differentiated from other species by the male genitalia; the vesica has two patches of spines, one near to the aedeagus and the other on the upper side of the vesica. Female genitalia have heavily sclerotized ductus bursae and appendix bursae, with the corpus bursae ca. 2 × longer than the appendix bursae and 1½ × wider; the anterior apophyses are remarkably short, whereas the posterior apophyses are almost as long as the valva.

Distribution.
Cicadoforma vau-nigrum is endemic to Venezuela, known only from the states of Vargas and Aragua at moderate elevation in the northern part of the northwestern Andes (Fig. 92).
Remarks. The specimens that were examined were over 60 years old and DNA could not be obtained from them. The specimens from the CNC have rotting stains on the base of the wings, and one of the males is missing half of the abdomen. The only female has incomplete antennae.  The genus contains four new species, Cicadomorphus chicharra sp. nov., Cicadomorphus chuya sp. nov., Cicadomorphus falkasiska sp. nov., and Cicadomorphus lilianae sp. nov., which were placed at first into Cicadoforma, but the molecular analyses place them as a sister group.
Diagnosis. Cicadomorphus is closed related to Cicadoforma genetically and morphologically (see Cicadoforma diagnosis).
Description. Sexually dimorphic in size, female slightly larger than male; cells of forewing with some semi-hyaline areas in male, whereas in female forewing covered by scales; line pattern conspicuous in both sexes. Antenna in both sexes dark brown, filiform with a stripe of sulfur-yellow scales basally; haustellum functional but reduced. Forewing with orbicular spot varying in size, reniform spot narrow. Hindwing semihyaline presenting scales only on margins and veins. Male genitalia with simple valva, lightly sclerotized lacking clasper; cucullar region with apex and ear-shaped lobe extremely narrow; saccular region with a long and narrow harpoon-shaped process; uncus hooked, broad and long; aedeagus short with simple vesica with a narrow patch of spines on upper side. Female genitalia with sterigma large, rounded, lightly sclerotized; appendix bursae rugose and sclerotized; corpus bursae transparent, not sclerotized. Immature stages. Egg. Circular with soft surface with the chorion forming small square cells, which is consistent across the entire Gaujonia genus group. Larva. Passes through five to seven instars. Late instars with remarkably short secondary setae, which leave some spaces completely naked; they also present some scattered long setae especially on the prothorax, which cover the head capsule. Pupa. Covered by a dense cocoon. The three pair of legs are visible along with the antenna, but prothoracic legs are slightly concealed and micro-setae on the abdomen as in other pantheines, including in the other genera of the Gaujonia genus group.
Key to species of the genus Cicadomorphus based on adult male morphology Diagnosis. Cicadomorphus chicharra is one of the most easily identified species in the group due to its whitish yellow coloration, but also the wing pattern is thinner and paler than in related species, such as C. lilianae. The male genitalia have a wider vesica and narrower spine band on the upper side. Additionally, barcoding showed 2% divergence from C. lilianae.
Description. Head. Palp with last segment black with a small white dot; frons with greenish yellow scales; antenna brownish orange. Thorax. Greenish yellow, with small black dots dorsally; collar with margins black. Wings. Forewing length: male 23-25 mm; forewing greenish yellow, semi-hyaline areas clothed by fewer greenish yellow scales; subterminal, medial, and antemedial lines slightly visible; reniform spot poorly defined; orbicular spot long; hindwing: semi-hyaline with greenish yellow veins and fringe. Leg. Prothoracic legs black with some patches of same color as body; mesothoracic legs and metathoracic legs greenish yellow. Abdomen. Greenish yellow, paler than thorax; tufts in male on A1-A6 with characteristic small black spots on A1, A5 and A6. Male genitalia. Cucullus wide with a wide lobe; apex, outer margin, and lobe covered by long setae; apex slightly squared; sacculus wide with needle-shaped process; tegumen narrow, unsclerotized around uncus; saccus relatively wide; juxta with U-shaped concave depression on upper side; aedeagus ⅔ × as long as vesica; opening to vesica as wide as aedeagus; vesica wide with narrow transverse band of spines.
Immature stages. Unknown. Distribution. The three specimens were found in the western zone of Bolivia at moderate elevations ca. 1500 m (Fig. 92).
Remarks. The holotype is in perfect condition (Fig. 19). Initially this species was confused with Cicadomorphus lilianae, but a rigorous morphological examination and DNA barcoding showed they were separate species. Etymology. Ch'uya means pure, transparent, or glassy in the Quechua language. Diagnosis. Although Cicadomorphus chuya shares some characters with C. chicharra, it is easy to distinguish by wing pattern; C. chuya is the only species that has both the terminal and subterminal lines serrated. In addition, the genitalia have the lobe reduced, and the juxta is pentagonal-shaped. The base of the lobe opens externally diagonally from the tornus to the cucullus.

Cicadomorphus chuya
Description. Head. Third segment of palp divided with upper side black, underside white, but white areas with a few brown spots; antenna dark brown basally, distal segments paler. Thorax. Whitish yellow coated with small black dots dorsally. Wings. Pattern in both sexes well defined and visible; forewing length: male 22-24 mm; female 29-31 mm; forewing whitish yellow with pattern somewhat blurred; nearly all lines visible and slightly blurred with exception of postmedial line, which is formed by black dots on wing veins plus some dispersed scales; subterminal and terminal lines zigzag; reniform spot relatively wide, outline poorly defined; orbicular spot medium sized and elongate; hindwing hyaline with yellow scales on fringe paler than thorax; wing veins darkened. Leg. Prothoracic and mesothoracic legs whitish yellow with two patches of brown scales on femur. Abdomen. Dorsally gray, black, and whitish yellow tufts in middle segment with A1 and A3 whitish yellow; whitish yellow ventrally. Male genitalia. Cucullar region wider close to apex; lobe small, almost completely covered by long setae from middle of costal margin to lobe; tooth-like protuberance barely visible; lobe diagonal to tornus of valva; saccular region relatively short with tip of process just touching tooth-like protuberance externally; saccus thin, rhomboid shaped; juxta pentagonal shaped; tegumen narrow with a small hood on base of uncus; aedeagus short and wide; opening to vesica square shaped ventrally; vesica tapered, with a large band of spines. Female genitalia. Anal papilla square shaped with short setae; posterior apophysis almost same size as anal papilla; anterior apophysis short; sterigma V-shaped, lightly sclerotized, above ostium; ductus bursae wide posteriorly, narrow toward appendix bursae; appendix bursae ¼ × shorter than corpus bursae; corpus bursae semi-transparent. Immature stages. Unknown. Distribution. Males were found in Colombia and Peru, whereas the female was found in Bolivia, all in deciduous forest habitats at moderate to high elevations from 1800-3000 m (Fig. 92).
Remarks. The holotype is in near perfect condition ( Fig. 17) with only the right hindwing slightly bent inwards at the posterior margin. The female is missing the left antenna, otherwise the specimen is complete. The palpi in the female are broken, but still attached to the mouth parts. The female was originally thought to be a different species; however, the DNA match with the male of Cicadomorphus chuya. Etymology. Falkasiska is the combination of the names from Falk Zahlaus who collect the first female specimen and Toni Kasiske, who reared the eggs obtained from that female.
Diagnosis. Cicadomorphus falkasiska and C. chicharra share certain characters, especially in coloration and wing pattern. However, C. falkasiska is paler and the orbicular spot larger and outlined in black; also, the subterminal line is quite visible in C. chicharra, whereas in C. falkasiska it is inconspicuous. The male genitalia have a relatively wide aedeagus; the vesica is large and semi-rounded; the female genitalia have a wide appendix bursae and corpus bursae.
Description. Head. Third segment of palp in both sexes black with a pale-yellow stripe ventrally and a dot of same color dorsally; frons darker-yellow than rest of body; antenna dark brown. Thorax. Pale yellow with some black blurred spots, which are more visible in male than female. Wings. Forewing length: male 23-25 mm; female 30-32 mm; forewing pale yellow; hyaline areas in male nearly without scales, whereas forewing slightly more covered with scales in female; postmedial and subterminal lines inconspicuous in male, whereas basal, antemedial, and medial lines well defined; female with all lines weakly defined; orbicular spot large and elongate; reniform spot slightly wide, outlined with black scales and with a black dot in middle; hindwing hyaline in male, semi-hyaline in female with fringe paler yellow than forewing. Leg. Pale yellow with prothoracic legs with some brown patches. Abdomen. Whitish gray with some tufts over middle of abdomen; tufts on A1-A3 yellow with a small black dot on A2, whereas remainder tufts on other segments are black with some yellow and white scales; female with yellow tufts on A1-A4. Male genitalia. Cucullar region relatively wide with an axe-shaped lobe; apex round with a lobe and apex covered with quite long setae; saccular region wide with process of sacculus thin and curved; juxta flat on upper side but with a small V-shaped depression in middle; aedeagus relatively wide at opening to vesica; aedeagus ca. as long as vesica; vesica rounded with narrow transverse band of spines. Female genitalia. Anal papilla long and wide with posterior apophysis ca. as long as anal papilla; anterior apophysis short; sterigma large, fused above ostium; ductus bursae wide and short; and appendix bursae large, well sclerotized; corpus bursae not sclerotized 1¼ × longer than appendix bursae.
Immature stages. Egg. Pale yellow, turning dark brown close to emergence. Larva. Body black with yellowish orange verrucae and short setae; a set of long white setae on lateral verrucae, prothorax near head capsule, and on last tergite (Fig. 91A). Prepupa. Similar to last instar but with verrucae and setae dark yellow. Pupa. Dark brown.
Distribution. All specimens were found in central Peru at a high altitude of ca. 2500 m or above (Fig. 92).
Biology. Cicadomorphus falkasiska is the only species in the genus for which immature stages are known. Adults are active throughout the year, but especially so in spring and summer. Cicadomorphus falkasiska feeds on Prunus spp. especially on P. subcorymbosa Ruiz ex Koehne (JIM, pers. obs.).
Remarks. Holotype (Fig. 18) and paratypes are well preserved, but three of the specimens were destroyed and only identifiable by genitalia dissection.   for her love, charisma, and support offered during all her beautiful life.
Diagnosis. Cicadomorphus lilianae is small and the wing pattern is blurred, causing the lines to appear shapeless. The males have darker coloration than the females, and the line pattern is better defined. This species shares some features in genitalia with C. chuya, such as the tapered vesica.
Description. Head. Palp black, terminal segment admixed with brown and white scales; frons dark yellow basally with some black scales; female ground color pale yellow with gray scales; antenna brownish orange. Thorax. Ground color dark yellow with some black tufts; female same color but with gray tufts; collar with black ground color with margins yellow, female with ground color gray. Wings. Both sexes dark yellow, similar in pattern, nevertheless female is paler, and wing pattern is basally almost imperceptible; forewing length: male 25-27 mm; female 31-33 mm; forewing yellow with pattern in black, but blurrier, male darker; both sexes with semi-hyaline areas with some scattered dark yellow scales; both sexes with blurry lines; orbicular spot elongate, ca. same size in both sexes; reniform spot heavily outlined with black scales; hindwing hyaline with fringe yellow, but gray on posterior margin. Legs. Yellow, except prothoracic legs, which are brown with some black scales. Abdomen. Gray with yellow tufts in middle of abdomen and on each side; brown tufts on A3-A7; female abdomen yellow with some brown scales. Male genitalia. Cucullus wide with apex wide; costal margin without setae; outer margin sharply bent; tooth-like protuberance small; lobe with external apex large; posterior margin of lobe curved; sacculus wide with narrow process touching tooth-like protuberance; saccus thin; tegumen narrow, lightly sclerotized, barely visible around uncus; juxta wide, upper side flat; aedeagus relatively wide, V-shaped in opening; tapered vesica; band of spines with a rounded ending. Female genitalia. Anal papilla small; relatively wide posterior apophysis 1 ⅓ × longer than anal papilla; sterigma open trapezoid shaped; anterior apophysis short; ductus bursae small, well-sclerotized; appendix bursae broad, sclerotized; corpus bursae ⅓ × longer than appendix bursae. Immature stages. Unknown. Distribution. This species has been found in Ecuador and Colombia at very variable elevations from 1100-2700 m (Fig. 92).
Remarks. The tip of the left antenna from the type specimen was broken during examination (Fig. 16). The paratype female is intact; however, the left orbicular spot is incomplete (Fig. 22). Otherwise, the female forewing is more brightly colored than that of the male basally where the antemedial and basal lines are almost inconspicuous. The female paratype from FSU has a different DNA voucher label from the voucher published at http://barcodinglife.com.
Etymology. Paul Dognin probably named this genus Gaujonia after Fr. Theophile Gaujon a Catholic Lazarist missionary priest and entomologist residing in Loja, Ecuador, who helped by collecting specimens for him.
Diagnosis. Gaujonia is similar to Oculicattus morphologically, but it is most closely aligned genetically with Cicadoforma and Cicadomorphus (Fig. 1). The interfacetal setae of Gaujonia are longer than those in Cicadoforma and Cicadomorphus. Male genitalia are similar to those of Oculicattus, however, the cucullus is wider and slightly square in shape, whereas in Oculicattus the cucullus is thinner and rounded. Gaujonia has a short vesica and two patches of spines, whereas in Oculicattus the vesica is longer with three patches of spines. Female genitalia of Gaujonia are similar to that of Oculicattus, but that of Oculicattus is larger in size, however, the sterigma is smaller in that genus than in Gaujonia. DNA barcoding corroborated that Gaujonia is closer to Cicadoforma and Cicadomorphus (~ 6% divergent) than Oculicattus (~ 8%).
Description. Sexually dimorphic in size, female larger than males; forewing and hindwing hyaline with scales only on wing veins; orbicular and reniform spots in male inconspicuous or absent, whereas female has an orbicular spot and narrow reniform spot. Antenna filiform and long haustellum dark brown; eyes hairy with long interfacetal setae. Male genitalia with saccular and cucullar regions separated and clasper absent; short-beaked uncus; aedeagus wide and vesica short with two sclerotized patches of spines on each side. Female genitalia with sterigma open wide, ductus bursae and appendix bursae are heavily sclerotized, whereas corpus bursae is not.
Immature stages. Egg. Circular, slightly flattened; chorion forming large square cells making the surface slightly rugose. Larva. Undergo from five to seven instars. Late instars with dense bands of secondary setae on the abdominal tergites, which are more scattered on the thorax; verrucae have scattered long setae; two large prothoracic  Diagnosis. Gaujonia arbosi is similar to G. chiqyaq, but it can be identified by coloration, which is dark yellow in G. arbosi. Both sexes are similar with the only difference being that the female is covered with brighter yellow scales from the fold to the posterior margin of the forewing, accenting the lines. For males the forewing length ranges from 17-19 mm and for females from 23-25 mm. Palp short and black; antenna has a stripe of dark yellow scales from the base to the seventh antennomere; antennae are black, and longer than in G. chiqyaq. Male thorax dark yellow with some patches of black. Forewing with pattern similar to that of G. chiqyaq, but the lines on Sc+R1 and posterior margin are thicker; there is a small dot in the middle of the base of the cell M1, which is not present in G. chiqyaq; the V-shaped mark at the base of CuA2 smaller and thicker than in G. chiqyaq. The hindwing, base of M2 is angled diagonally forward to the base of the wing, fused to the base of CuA1; in G. chiqyaq M2 is squared and not fused with the base of CuA1; the fringe is black with few yellow scales making minute spots at the end of the veins, except the cell A2, which is completely outlined in black. The male genitalia have the cucullar region wide, slightly diagonal to the base, whereas the cucullar region in G. chiqyaq is wide, opened, almost touching the saccular region; the juxta is semicircular in shape; the saccus is short and wide. Female genitalia, the sterigma is peanut shaped, and the corpus bursae is ⅛ × larger than the appendix bursae. DNA Barcoding showed that G. arbosi and G. chiqyaq are sister species with ca. ~ 0.3% of difference, but the morphological characters mentioned above distinguish both species.
Immature stages. Egg. Bright green and turning dark brown close to emergence. Larva. Only known from the third instar, which is very similar to that of G. kanakusika, but the body of G. arbosi is white with dark brown secondary setae, whereas the body of G. kanakusika is rather whitish yellow. Pre-pupa. Unknown. Pupa. Dark brown (Guevara and Romero 2008).
Distribution. This species is endemic to coniferous and deciduous forest of Ecuador and Colombia and can be found at high elevations above 2500 m (Fig. 93).
Biology. Guevara and Romero (2008) were the first to record the larval stages in Gaujonia arbosi feeding on Alnus acuminata Kunth but they misidentified it as Gaujonia arbosioides (now Millerana arbosioides), even though the immature stages differ greatly between both genera (see Millerana immature stages). Additionally, the illustrations from Guevara and Romero (2008) lack of good quality to use for identification.
Remarks. The holotype of the species was "missing" in the P. Dognin collection (USNM), which was a female illustrated by Dognin (1894). A male was listed as the type specimen, but this is actually a different species from the illustration made by P. Dognin (Fig. 8). Presumably both specimens were used in the description of the species in his second work Dognin (1894), and is also deposited in the Dognin collection (USNM), I designated the male as the lectotype for Gaujonia arbosi Dognin under the provisions of Article 74.1.1 of the International Code of Zoological Nomenclature (1999), whereas the female is considered a different species (see Gaujonia chiqyaq remarks). Etymology. The term bichu is used to refer to an insect or small animal in the Quechua language. Since this is the smallest species of Gaujonia, this name seems appropriate.

Gaujonia bichu
Diagnosis. Gaujonia bichu is closely related to G. sourakovi, but G. bichu is the smaller. The male genitalia have the valva and the saccular region narrow, and the process of the sacculus is longer and thinner.
Description. Head. Palp black; frons yellow with some black scales. Thorax. Dorsally covered by large black patches. Wings. Male yellow with thin spots and lines on venation; forewing length: male 15-17 mm; forewing with two V-shaped marks on base of R1+R2 and R4+R5; base of cell with small black dot on M1; five black dots on Sc+R1 with a line right on apex; half of posterior margin black; hindwing with fringe black, interrupted by a yellow dot at end of each vein. Leg. Prothoracic leg black with femur and tarsi presenting yellow spots; mesothoracic legs yellow with some black spots; metathoracic legs yellow. Abdomen. Grayish yellow with yellow and black tufts in middle of abdomen dorsally on A1-A5, remainder of abdomen covered in yellow scales. Male genitalia. Valva long and narrow a little swollen from middle to apex externally; sacculus narrow; sacculus process long and narrow; juxta shield-like; tegumen wide, narrower near valva; aedeagus short and narrow; vesica elongated with two oval patches, one 2 × longer than the other. Immature stages. Unknown. Distribution. Gaujonia bichu specimens were found in Ecuador in coniferous and deciduous forests at high elevations above 2500 m (Fig. 93).
Biology. Unknown. Remarks. Three specimens are in good condition; one specimen from Loja has no black dots on the base of cell M1. Etymology. The name chiqyaq means green in the Quechua language, referencing the diagnostic green coloration of this species.
Diagnosis. Beside the features mentioned above to differentiate Gaujonia chiqyaq from G. arbosi, there are also other informative characters to identify this species. The male has a narrow reniform spot outlined by black scales along the upper and lower sides, but not in the middle region. The orbicular spot is small and greenish yellow, outlined in black. The thorax has large black tufts. The female has a larger reniform spot with the same yellow coloration. Forewing transverse lines are more sharply defined and thicker than in G. arbosi.
Description. Head. Palp with last segment black; frons with a combination of black and greenish yellow scales. Thorax. Covered with long black tufts. Wings. Both sexes greenish yellow with thin spots and lines; forewing length: male: 20-22 mm; female: 28-30 mm; forewing ground color greenish yellow with small, thin lines on venation forming pattern, female with thicker and well-defined lines; orbicular spot in male small, black, but almost imperceptible; orbicular spot larger in female and elongated; reniform spot in both sexes narrow, greenish yellow, outlined in black; both sexes with hindwing venation green, and fringe black with some greenish yellow dots. Leg. Metathoracic and metathoracic legs black with some greenish yellow patches; hind legs greenish yellow with black shading. Abdomen. Brown dorsally with green tufts on A1-A3 and some black scales in middle of the green tufts; ventrally greenish yellow; female similar to male but paler in color. Male genitalia. Cucullar region extended close to saccular region, valva relatively narrow, slightly ridged externally on apex; saccular region rounded at base and process wide; saccus wide V-shaped; juxta rectangular concave on top; tegumen wide; aedeagus short 3 × longer than wide; basal area almost same size as apical part of vesica with two patches of spines, one 2 × narrower than the other. Female genitalia. Anal papilla wide, slightly ridged; posterior apophysis ⅓ × longer than anal papilla; sterigma large oval shaped, fused above ostium; anterior apophysis short; ductus bursae heavily sclerotized and connected to appendix bursae, which is sclerotized as well; appendix bursae ½ × narrower than corpus bursae.
Immature stages. Unknown. Distribution. This species has a broad distribution from Ecuador to Peru between 2100-3200 m (Fig. 93).
Remarks. The holotype of Gaujonia chiqyaq closely resembles the type specimen of Gaujonia arbosi (Fig. 33) illustrated by Dognin (1894). However, it is a different species from the lectotype of G. arbosi, differing from it in characters of the male and female genitalia, thus it is here established as a new species. The holotype of Gaujonia arbosi and most of the paratypes are well preserved. Guevara and Romero (2008) misidentified this species as Gaujonia arbosioides, which is placed in a different genus (see Millerana arbosioides remarks). Etymology. The name is formed from the Quechua words kanaku (fire) and sika (caterpillar), based on the immature stages of this species (Fig. 91B).

Gaujonia kanakusika
Diagnosis. This species can be differentiated from other species by the sulfuryellow coloration of most of the body, including the abdomen.
Description. Head. Palp with basal and second palpal segment sulfur-yellow, last segment with white tip; frons with small black line; female with similar wing pattern, but brighter in coloration. Thorax. Sulfur-yellow, covered with small black dots. Wings. Forewing sulfur-yellow; length: male 18-20 mm; female 27-29 mm; forewing with lines visible from posterior margin to fold; basal and antemedial lines almost completely developed; no presence of a V-shaped mark; Hind wing fringe same yellow color as body. Legs. Prothoracic and mesothoracic legs black with some sulfur-yellow spots on joints; tarsal scales yellow and black. Abdomen. Sulfur-yellow with a small area of gray in middle of dorsum divided by black and yellow tufts. Male genitalia. Cucullus large, rounded apically; sacculus narrow with a long process; juxta shell-shaped with apices pointed; tegumen wide; saccus narrow; aedeagus short and wide; vesica 1 ½ x as long as aedeagus with two large spine patches ca. same size. Female genitalia. Anal papilla small, semicircular; posterior apophysis ⅓ × longer than anal papilla; sterigma narrow and large, semirectangular; anterior apophysis relatively long and heavily sclerotized; corpus bursae 1 ½ × longer than appendix bursae. Immature stages. Egg. Completely light green, turning dark brown close to emergence. Larva. In general, body pale orange with dark pink verrucae and neon-orange setae on dorsal verrucae; long pale-yellow setae on lateral verrucae; inter-tergal membrane brown with white dots (Fig. 91B). Pre-pupa. Darker than ultimate instar. Pupa. Dark brown with white setae covering the whole body.
Distribution. All specimens were found in deciduous forests from the centralnorth Colombia between 2900-3300 m (Fig. 93).
Biology. Gaujonia kanakusika is the second species in this genus known to feed on Alnus acuminata Kunth. Adults fly year-round, mainly in summer (JIM, pers. obs.).
Remarks. Holotype (Fig. 30) and paratypes in good condition. This species was the only one whose phylogenetic placement had low support in the gene tree; however, it shares apomorphic characters with other Gaujonia species, so it is maintained as a Gaujonia. Etymology. This species was found by my colleague and friend Andrei Sourakov when he was helping me locate additional specimens deposited in the MGCL, and thus I named it in his honor for his wonderful help.
Diagnosis. Gaujonia sourakovi differs from other species mainly in coloration, having the most lightly marked forewing in this group. The male genitalia have the apex of the cucullus rounded, the saccular region is wide, and two patches of long hair-like spines on the vesica, differing from the others which have wider and well-developed spines.
Description. Head. Last segment of palp black with some yellow scales; frons yellow, shaded with black scales. Thorax. Dark yellow with black dorsally. Wings. Forewing length: male 19-21 mm; forewing scales only on venation and margins; Vshaped mark at base of CuA2 widely open; dot in the middle of base of cell M1 elongated; orbicular spot small; veins lined with black; hindwing fringe black with dark yellow scales at end of each vein, whereas area on posterior margin of hindwing brown; a black line at base of vein M3. Leg. Prothoracic and mesothoracic legs dark brown with yellow scales on joints, whereas metathoracic legs dark yellow with some dark brown spots. Abdomen. Brown with scattered black scales dorsally, paler ventrally; long yellow scales laterally; black tufts with some yellow and brown scales on A1-A4. Male genitalia. Cucullar area narrow and curved with apex rounded; saccular region and process wide; juxta flat on upper side; tegumen wide; aedeagus 2 ⅔ × longer than wide; basal area of vesica 4 × longer than wide and vesica almost same width with two large patches of hair-like spines, one ovoid. Immature stages. Unknown.
Distribution. The species Gaujonia sourakovi was collected in a deciduous forest at 1800 m elevation in southeastern Peru (Fig. 93).
Biology. Unknown. Remarks. One specimen has the left forewing broken at the base and a brown patch on cell R5. Right wing with some brown patches near the outer margin between the cells R5-M2. In the other specimen the wings and left valva are broken, so it is not included as a paratype.
Type species. Gaujoptera amsa sp. nov. Etymology. Gaujoptera is derived from the similarities with the genera Gaujonia and Lichnoptera.
Included species. Gaujoptera is monotypic, the only species is Gaujoptera amsa sp. nov., which was misidentified as a Gaujonia species; however, morphological and molecular analyses resulted in recognizing a new genus that is positioned close to the genus Millerana (Fig. 1).
Diagnosis. Gaujoptera is only known from two well-preserved male specimens and it has a close relationship with Millerana, not only genetically, but also by distinct morphological characters. Nevertheless, the wing pattern is more defined in Millerana than in Gaujoptera, which has a blurry pattern that is difficult to discern. In addition, the thorax is marbled with black, gray, brown, and sulfur-yellow scales, differing from species of Gaujonia, which have the black spots and patches well defined on the thorax. Gaujoptera does not have a V-shaped mark at the base of CuA2. The most important feature to differentiate Gaujoptera from Gaujonia is the presence of small clasper on the valva, similar to that of Lichnoptera.
Description. Forewing and hindwing dark sulfur-yellow with an inconspicuous pattern, only reniform spot visible. Hindwing hyaline with scales only on margins. Antenna filiform, with a short brownish orange haustellum; eye hairy with short interfacial setae. Male genitalia simple in male with small triangular clasper; small, curved uncus; aedeagus shorter than vesica; vesica with small spines in middle. Female unknown.
Immature stages. Unknown. Biology. Unknown. Etymology. Amsa is a Quechuan word that means dark, opaque, or confused. Diagnosis. The species Gaujoptera amsa has similar morphological characters with species of Millerana, however, it is easy to identify because the thorax has no spots or patches, but is completely marbled with black, gray, brown, and sulfur-yellow scales. The forewing is dark sulfur-yellow with a blurry pattern in gray. Additionally, the genitalia have a simple valva without any lobe or protuberances, and a small clasper is present.

Gaujoptera amsa
Description. Head. Palp with last segment black with few yellow scales on tip; frons marbled. Thorax. Marbled in black, gray, brown, and sulfur-yellow. Wings. Forewing length: male 18-20 mm; forewing marbled with wing pattern in gray; pattern blurry; orbicular spot barely visible in yellow; a narrow lunate marking on reniform spot; terminal lines zigzag; hindwing hyaline with sulfur-yellow fringe. Leg. Prothoracic and mesothoracic legs sulfur-yellow with some brown spots; tarsi brown and yellow; metathoracic legs marbled in brown and yellow with tarsi sulfur-yellow. Abdomen. Sulfur-yellow with a black line of tufts in middle of abdomen on A1-A5. Male genitalia. Valva rectangular, simple; saccular region wide; apex densely covered by setae; clasper small, triangular; juxta shell-like; tegumen narrow; uncus small; aedeagus short, almost same width as vesica; vesica ovoid with a line of minute spines in middle, positioned diagonally. Immature stages. Unknown. Distribution. Gaujoptera amsa occurs from central Peru to northern Bolivia at high elevations ca. 3000 m or above (Fig. 93).
Biology. Unknown. Remarks. Most of the specimens are very damaged and could only be identified by genitalia examination, except the holotype and paratype (Figs 35, 36 Diagnosis. Millerana is the most genetically distant genus in the Gaujonia genus group and is more closely related to Gaujoptera (Fig. 1) to Cicadoforma and Cicadomorphus, but smaller in size and a slightly different forewing pattern. The genitalia valva is simple as for Cicadoforma and Cicadomorphus, but much wider, with relatively long protuberances at the apex and outer margin. The vesica has a band of spines surrounding the middle area. DNA barcoding reveals that Millerana is distant from the other genera: Gaujoptera (~ 9% divergent), Cicadoforma, and Cicadomorphus (~ 18%).
Description. Orbicular spot well developed; reniform spot with small lunate marking. Forewing pale yellow with black scales, which form the forewing pattern. Hindwing with yellow scales restricted to veins and margin, extending to fringe. Antenna dark brown or brownish orange, serrate, with a stripe of pale-yellow scales on basal % of antenna. Mouthparts reduced; eyes covered by black interfacetal setae. Male genitalia: moderately sclerotized; valva wide, without clasper; apex small with a pointed extension; uncus broad, beak-like; aedeagus short, vesica with narrow band of spines around middle part of vesica.
Immature stages. Egg. Circular and flattened with micro-square cells formed by the chorion. Larva. Like many pantheines, there are five to seven instars. Late instars resemble larvae of the genus Panthea. Secondary setae on the abdominal tergites spinelike. White lines between the spiracles similar to those of Panthea. Thorax densely covered by secondary setae, with long setae on the prothorax covering head. Pupa. Similar to those of other species in the Gaujonia genus group, but remarkably smaller (see Antenna black; forewing with a small orbicular spot (Figs 9, 37, 38, 42) .... 4 -Antenna orange; forewing with a large orbicular spot (Fig. 39)  Cucullar region presenting outer margin with protuberances (Figs 9, 72) ...5 -Cucullar region without protuberances on the outer margin (Fig. 73)  Etymology. Paul Dognin likely name this species arbosioides in reference to its similarities to Gaujonia arbosi, which was described previously.
Diagnosis. Millerana arbosioides is closely related to three species: M. austini, M. cajas, and M. cundinamarquensis; however, M. arbosioides can be recognized by the black antenna and haustellum. Forewing length in males is 16-18 mm. Millerana arbosioides shares more external characters with M. cundinamarquensis, but it can be distinguished from it by the antemedial line, which is well developed in M. arbosioides, the reniform spot is better defined, and the lunate marking is small; these features in M. cundinamarquensis are barely visible, but it presents four black tufts in the middle of 1A-4A, whereas M. arbosioides has three on A1-3A. The genitalia in M. arbosioides have a semi-squared cucullar area and some protuberances in the outer margin that are present in M. tigrina and M. cajas as well however, both species differ from M. arbosioides by their apexes that are rounded and their outer margins of the cucullar area, which are straighter.
Immature stages. Unknown. Distribution. The species is known only from coniferous forests in Ecuador at very high elevations ca. 2500-3500 m (Fig. 94).
Biology. Unknown. Remarks. One of the additional examined specimens was in very poor condition and only identifiable by genitalia examination. Etymology. The species was named after my friend Kyhl Austin, an American lepidopterist, who found this species in the dark corners of the CUIC.

Diagnosis.
Millerana austini is similar to M. cajas, but it can be differentiated by the antemedial line, which is formed by a single line, whereas M. cajas has two. The male genitalia are without protuberances on the outer margin, except for two extensions; one near the apex and the other on the lobe; they are similar to those of M. tigrina; however, the outer margin of the valve is concave in M. tigrina.
Description. Head. Palp black with a small line with pale yellow underneath; frons yellow. Thorax. Dark yellow with dots in black. Wings. Forewing dark yellow with black pattern; length 17-19 mm. All lines well developed with exception of terminal line; orbicular spot circular; reniform spot incomplete, with a small lunate marking; V-shaped mark on base of CuA2 thick, black; hind wing whitish orange, veins yellowish orange. Leg. Prothoracic legs black with some yellow spots on joints; mesothoracic legs with some black dots on tibia; metathoracic legs yellow. Abdomen. Paler yellow than thorax; small tufts in black from 1A-2A. Male genitalia. Cucullar area wide; apex and lobe with small extensions; outer margin flat; sacculus notably wide; tegumen broad; juxta inverted, triangle-shaped; aedeagus 3 ⅓ × longer than wide; vesica 1 ½ × wider than aedeagus, gradually reduced at tip; band with large spines and thick patch of spines near apex.
Immature stages. Unknown. Distribution. All specimens were found in a cloud forest at the north-central region of Ecuador at high elevations above 3500 m (Fig. 94).
Biology. Unknown. Remarks. Holotype (Fig. 38) and paratypes in perfect condition. The best way to identify M. cajas externally is to examine the medial and postmedial lines, which are fused on the forewing. Moreover, the genitalia are very distinctive. The valva is enlarged, especially in the costal region; Millerana cajas has a wide apical protuberance on the valve, similar to that of M. matthewsae; the vesica has a distal diverticulum surrounded by a spine band.
Description. Head. Palp short, black and yellow, except last segment that is black with some small yellow scales; frons yellow. Thorax. Covered with light lemon-yellow hair-like scales with some black spots. Wing. Light lemon yellow with black and gray pattern; forewing length: male 18-20 mm; forewing antemedial, medial, and subterminal lines well developed; whereas basal, postmedial, and terminal lines defined by small dots; medial line wide; orbicular spot slightly flattened; reniform spot large, incomplete, with lunate marking at base of M1 cell; V-shaped mark at base of CuA2 narrow; medial and postmedial lines fused by black scales between V-shaped mark in CuA2 and fold; hindwing whitish yellow with veins whitish orange. Leg. Prothoracic legs black with fuscous joints; mesothoracic and metathoracic legs yellow with a black spot on tibia; tarsi black. Abdomen. Paler yellow than remainder of body; tuft on middle area on A1-A5; A1-A3 black and A4-A5 yellow. Male genitalia. Cucullus wide, mainly in on costal area; apex with a wide extension followed by some protuberances on outer margin; saccular region wide; tegumen wide; juxta dentate, slightly curved on upper side; aedeagus 3 ½ × longer than wide; opening larger than rest of aedeagus; vesica long with a distal diverticulum, which is surrounded by a band of long spines; medial area with a large patch of minute spines.
Immature stages. Unknown. Distribution. This species was found in a deciduous forest at a high elevation in southwestern Ecuador ca. 3600 m (Fig. 94).
Biology. Unknown. Remarks. Holotype (Fig. 42) with the right hindwing and the left forewing slightly broken; the abdomen has a spot of dead fungi; the frons has pink marker stains accidentally applied at the place where it was previously deposited. The left wings are missing in the paratype. Etymology. The name is derived from the place (Department of Cundinamarca in Colombia) where this species was found.

Millerana cundinamarquensis
Diagnosis. Compared with other species in Millerana, M. cundinamarquensis differs in the genitalia, in which the valva is petal-like, whereas the other two species have a rectangular valva.
Description. Head. Palpi short, divided by black hair-like scales on upper side and yellow underneath; last segment coated with a mix of black, white, and yellow scales; frons with sulfur-yellow scales. Thorax. Covered by sulfur-yellow scales with some large black spots. Wing. Forewing length: male 16-18 mm; forewing sulfur-yellow with black scales defining transverse lines; antemedial, medial, and subterminal lines barely developed; basal, postmedial, and terminal lines defined by series of dots on veins; orbicular spot well-defined, oval; reniform spot blurry, with a small triangular lunate marking; CuA2 with a black V-shaped mark at base, with inferior line longer than superior line; hindwing with yellowish orange veins; fringe with pale-yellow hairlike scales; base of M2+M3 rounded. Legs. Yellow except prothoracic legs, which are black with pale yellow spots. Abdomen. Pale yellow with light-gray hair-like scales; four black tufts in the middle of A1-A4; a small pale-yellow tuft on A8 at terminus. Male genitalia. Cucullar area with rounded apex, apex extension claw-like; costa ovoid, covered with setae; saccular area wide; tegumen narrow and rounded; juxta with concave V-shaped depression on top; aedeagus 4 × longer than wide; opening to vesica % × total length of aedeagus; vesica long, 2 × wider than aedeagus, decreasing in width as it approaches to apex; a band of medium-size spines near basal area of vesica.
Immature stages. Unknown. Distribution. Only two specimens are known from a cloud forest in Cundinamarca, Colombia (Fig. 94).
Biology. Unknown. Remarks. Holotype well preserved (Fig. 39), but the other specimen, which was collected from the same location, was difficult to identify because it was practically destroyed. Thus, the identification was made by genital dissection. Etymology. In honor of my colleague and friend Dr. Deborah Matthews-Lott, who has supported me during my travels in entomology, especially on Lepidoptera trips.

Millerana matthewsae
Diagnosis. Millerana matthewsae is closely related to M. tigrina; it has brownish orange antenna and can be differentiated by the wide well-defined pattern of foreign lines, and the large orbicular and reniform spots. The most distinctive character of the male genitalia is the costal margin of the valva is strongly curved upward.
Description. Head. Palp with long black and yellow hair-like scales; frons dark yellow. Thorax. Dark yellow with some black patches on dorsal area; Wing. Forewing length: male 18-20 mm; forewing paler yellow coloration than thorax; antemedial, medial, and subterminal lines complete; postmedial line almost complete except in area between cells of CuA1 and half of CuA2; V-shaped mark at base of CuA2 with upper line longer than lower line; terminal area covered by scattered black scales except in M3 and CuA1 cells; orbicular spot very large, barely touching top and bottom of discal cell; reniform spot with large line marking inside of discal cell; hindwing white with veins and fringe pale yellow. Leg. Prothoracic and mesothoracic legs brownish orange and black with joints yellow; metathoracic legs yellow. Abdomen. Yellow, paler than remainder of body; dorsally with thin black tufts in middle of A1-A6 and with dark yellow tufts at A2-A3. Male genitalia. Cucullus wide and curved upward on costal margin; apex rounded; some protuberances in the outer margin with the upper one the largest and curved downward; sacculus wide and short; upper side of juxta has a shallow concave depression; tegumen narrow laterally; aedeagus narrow, ⅓ × longer than vesica; vesica long rounded basally; Transverse band of spines near middle of vesica.
Immature stages. Unknown. Distribution. This species has been collected only in the western zones of Peru at very high elevations almost 4000 m (Fig. 94).
Biology. Unknown. Remarks. The holotype (Fig. 40) and the paratype have both hindwings slightly broken. The other specimens were almost destroyed with only the thorax and forewings remaining. Etymology. The name refers to the tigrina, Leopardus guttulus (Hensel), in keeping with the wild feline names, as with the other genera, since they are known as "the jaguar moths."

Millerana tigrina
Diagnosis. Millerana tigrina can be distinguished from M. matthewsae by the large, well-rounded orbicular spot. Millerana tigrina also has distinctive genitalia, with the valva presenting a pronounced concavity on the outer margin, forming a moonshaped crescent.
Description. Head. Palp covered with hair-like scales with dorsal side black and ventral side whitish yellow; last segment black with small whitish yellow dot internally; frons yellow. Thorax. Covered with yellow hair-like scales with black patches. Wing. Forewing length: male 17-19 mm; forewing yellow with black scales forming wing pattern; basal line barely visible; antemedial and medial lines disrupted at discal cell; reniform spot large, with small lunate marking in middle; orbicular spot rounded, almost as high as discal cell; two V-shaped marks, one at base of R1+R2, other one at base of CuA2; hindwing whitish silver with yellow fringe; veins yellow. Leg. Prothoracic and mesothoracic legs dark brown with some patches of black and pale yellow; metathoracic legs pale yellow with dark brown from tibia to claws. Abdomen. Covered with pale-yellow, hair-like scales; three small black tufts on A1-A3; a pale-yellow terminal tuft. Male genitalia. Outer margin of cucullar area concave, with two small protuberances, one subapical and lateral to apex and other one smaller, on lower end of lobe; saccular area wide; tegumen wider dorsally and squared; juxta concave posteriorly; aedeagus 3 × longer than wide; a diagonal opening to vesica ⅓ length of aedeagus; vesica almost as long as aedeagus; transverse band of spines near base.
Immature stages. Egg. Bright green; turns dark green close to emergence. Larva. Five to seven instars; body brown with black and pale-brown secondary setae arising from pink verrucae; diagonal pinkish white lines laterally between spiracles; spiracles white; thorax covered with secondary setae; two tufts of on each side of second tergite. Pre-pupa. Similar to last instar, but darker and with secondary setae shorter. Pupa. Dark brown.
Distribution. The two specimens of Millerana tigrina were found in the highest elevations of the Andes in Ecuador (Fig. 94).
Biology. Only one species is known in this genus, Millerana tigrina, which was discussed first by Bollino and Onore (2001). Adults are active throughout the year, but mainly during summer. Larvae resemble some species of the genus Panthea, which also feed on pine trees. Larvae have been observed to feed on Podocarpus magnifolius J. Buchholz & N.E. Grayand and Quercus humboldtii Bonpl as well (O. Mahecha-Jiménez pers. comm.).
Remarks. The type specimen (Fig. 41) has a small patch of dead fungi on the ventral side of the abdomen. The specimen from Azuay is poor condition but is still identifiable. This species was misidentified as the female of Millerana arbosioides by Hampson (1913) (as Gaujonia arbosioides) and Seitz (1919Seitz ( -1944. In addition, the larvae are reported as pine tree pests in Ecuador (Gara and Onore 1989), but was misidentified as Gaujonia arbosi. Unfortunately, the pictures of the immature stages provided by O. Mahecha-Jiménez were not of good enough quality to be included in this revision.
Etymology. Oculicattus refers to the reniform spot, which is black and surrounded with yellow scales, giving it the appearance of a cat's eye.
Diagnosis. Oculicattus can be differentiated from Gaujonia externally by the presence of the large reniform spot in Oculicattus, as well as by other features (see Gaujonia diagnosis).
Description. Sexually dimorphic in size, females larger than males; forewing and hindwing hyaline with sulfur-yellow and black scales covering veins and wing margins.  Forewing with a small black or sulfur-yellow orbicular spot, sometimes barely perceptible; reniform spot elongated, outlined in black; elongated black central line surrounded by sulfur-yellow outline, except for O. raizae and O. uturunku, in which the spot is entirely black. Antenna filiform, brownish orange with a sulfur-yellow band on basal to three quarters of antenna; eyes with coppery interfacetal setae. Male genitalia slightly sclerotized; valva with saccular and cucullar regions separated, without clasper; uncus long and narrow ending in beak-like tip; vesica has spine patch; vesica wider than its base, which has two patches of spines with one patch of spines larger than other. Female genitalia medium sized; lightly sclerotized rectangular-shaped sterigma; appendix bursae elongate and rugose; corpus bursae for most species approximately half size of appendix bursae.
Key to species of the genus of Oculicattus based on adult male morphology  Napo, Cosanga, 2150m, 1 Feb. 1976. Additional examined specimens ( Etymology. The name is in honor of a great lepidopterist, Dr. Gunnar Brehm, who loaned some specimens for this research.
Diagnosis. Oculicattus brehmi is the smallest species in this genus; pattern formed by relative pale, thin markings. Orbicular spot small, black. There V-shaped mark at the base of CuA2 is small. Male genitalia have a tapered cucullar region, wider at the base; the saccus smaller and more tapered than in other species, and the saccular process is the second shortest in the genus with only O. uturunku being shorter than O. brehmi.
Description. Head. Wide, palp large, with last segment marbled in light yellow and black; frons pale yellowish gray; female similar to male, even in coloration. Thorax. Covered with dark sulfur-yellow scales and with small gray spots on dorsum. Wing. Pale yellow with pattern of thin dark gray lines; forewing length: male 16-18 mm; female: 21-23 mm; forewing with thin dots and stripes forming lines; lunate marking in reniform spot narrowly outlined by a thin yellow line; orbicular spot black, small; V-shaped mark at base of CuA2 small; hindwing with fringe yellow with long yellow lines between veins terminally. Leg. Prothoracic leg brown with joints pale yellow; mesothoracic legs marbled in brown and yellow, tarsi brown with each joint yellow; metathoracic legs yellow. Abdomen. Pale yellow with dorsal area gray; dorsal tufts sulfur-yellow along middle of abdomen, smaller on A1-A2, whereas A5-A8 are wide and combined with gray scales. Male genitalia. Cucullus tapered with small rounded apex and base wide, heavily covered with setae; sacculus wide with process long and densely clothed with setae; saccus V-shaped, and more narrowly tapered towards the end; juxta wide deeply concave posteriorly, , with the outer edges sharply pointed; tegumen wide; aedeagus 1 ¼ × length of vesica to medial cluster of spines; vesica with small patch of spines near middle; apical part of vesica bulbous, with large tapered patch of spines on each side. Female genitalia. anal papilla rounded posteriorly; posterior apophysis ⅓ × longer than anal papilla; anterior apophysis ⅔ × shorter than posterior apophysis; sterigma crescent moon shaped; corpus bursae ¼ × longer than appendix bursae. Immature stages. Unknown. Distribution. This species has been recorded only in Ecuador (Fig. 95).

Biology. Unknown.
Remarks. Holotype (Fig. 52) and paratypes are in good condition; specimens from NJD were soaked in 100% ethanol with only the wings in good condition; they were kept in -80 °C for molecular DNA analyses, so they are not included with the type series. Etymology. This is species is named after the Inca Empire, which originated in the area around Cusco.

Oculicattus inca
Diagnosis. Oculicattus inca is similar to O. renifera; however, there are some morphological features that are useful to differentiate them. O. inca is slightly smaller, darker in coloration, the orbicular spot smaller, and wing lines are thinner than in O. renifera; the lunate marking in the reniform spot is not outlined in yellow. Genitalia have a wider cucullus; the juxta is flattened posteriorly, whereas in O. renifera the posterior margin has a V-shaped invagination. The DNA barcodes differ by 5% between the two species.
Description. Head. Last segment of palp divided in black upper side and yellow under side; frons yellow with some gray scales. Thorax. Grayish yellow with some small gray patches. Wing. Forewing length: male: 19-21 mm; forewing grayish yellow with markings wide; orbicular spot small, elongated, grayish yellow, outlined with black scales and with some black scales inside orbicular spot; reniform spot with lunate marking not outlined; V-shaped mark short; hindwing with fringe black, vein ends yellow, with some black scales on vein CuA2. Leg. Prothoracic and mesothoracic legs black anteriorly, yellow posteriorly, whereas metathoracic legs all yellow. Abdomen. Dorsal area brown with some yellow scales between each tergite; a large yellow tuft on A3; last abdominal segment with long yellow scales. Male genitalia. Cucullar and saccular regions large; saccus triangular; juxta V-shaped with posterior part flattened; vesica 1 ½ × size of aedeagus; Vesica with small patch of spines near middle of vesica; bulbous apical part of vesica with large dense patch of spines posteriorly and smaller, less dense patch anteriorly. Immature stages. Unknown. Distribution. Broadly distributed in the cloud forests in Peru and Bolivia (Fig. 95). Biology. Unknown. Remarks. Holotype is in perfect condition (Fig. 47). This species was originally labeled as Oculicattus renifera, but DNA barcoding showed that the two species are genetically distinct. Behounek, grafing bei Müchen / "Gaujonia renifera ♂" (1 ♂).
Etymology. The species is dedicated to my wife, Raiza Castillo, for her love and support since the beginning of my career.
Diagnosis. Oculicattus raizae is closely related to O. uturunku; they differ from the other Oculicattus species by the darker color, and the unusual reniform spot that is completely brown or black. Oculicattus raizae can be identified by its brownish yellow coloration, and the brown forewing pattern. In the male genitalia the cucullus is longer and wider than in O. uturunku. Additionally, DNA barcodes are ca. 4% different. Description. Head. Palp reduced in size; dorsal surface covered by brown scales, ventral surface by dark sulfur-yellow scales; frons covered by dark sulfur-yellow scales, mixed with black; female similar to male externally. Thorax. Dark sulfur-yellow with some brown patches. Wing. Forewing with dark sulfur-yellow and brown scales covering venation and margins; forewing length: male: 22-24 mm; female: 27-29 mm; antemedial, postmedial, and subterminal lines not defined; orbicular spot small, poorly-defined oblong brown spot; reniform spot brown, narrow, and barely defined; base of CuA2 with Y-shaped mark large, brown; a brown line across fold on cell at CuA2; hindwing hyaline with some dark sulfur-yellow scales on venation, but more notable along margins; a Y-shape mark at base of CuA2; some small brown lines scattered through M3+CuA1+CuA2. Leg. Dark sulfur-yellow with brown patches on anterior area of prothoracic legs, which is lighter on mesothoracic and metathoracic legs. Abdomen. Clothed in grayish yellow hair-like scales with white scales at terminus and with a dark sulfur-yellow tuft on A8. Male genitalia. Cucullus wide, parallel sided with the apex rounded; costal margin swollen basally; sacculus triangular with saccular process entirely coated with setae; tegumen wide; saccus broadly V-shaped; juxta small with parallel sides and a V-shaped notch posteriorly; aedeagus 3 × longer than wide, with a wide opening to vesica ca. ¼ × total length of aedeagus; vesica with basal area ca. same width that of aedeagus, and it is ventrally curved; basal area with a large patch of spines; vesica with two patches of spines on each side, one small slightly beyond middle of dorsal wall of the vesica, and truncated anteriorly, whereas other larger apically narrow, and broader towards the aedeagus. Female genitalia. Small and truncated apically; anal papilla with posterior apophysis ¼ × shorter than anal papilla; anterior apophysis short; rectangular sterigma that is fused above ostium; ductus bursae wide and long; appendix bursae ⅔ × shorter than corpus bursae which is partially sclerotized at base. Immature stages. Unknown. Distribution. Recorded from cloud forests in Colombia, Ecuador, Peru, and Bolivia in middle and high elevations from 2000 m to above 3000 m (Fig. 95).

Remarks.
A specimen of Oculicattus raizae was misidentified as G. renifera by G. Behounek and posted on Barcode Of Life Data System v4 (www.barcodinglife.org). Holotype (Fig. 50) and paratypes are well preserved.
Diagnosis. Regarding Ocullicatus renifera, there is one species that is particularly similar to it (O. inca); however, O. renifera can be identifiable by its remarkably large yellow orbicular spot. The male is pale yellow, brighter yellow in the female. The forewing length in males is 19-21 mm and females 24-26 mm. Palp reduced, black with white tips; frons yellow and gray; antenna dark orange; male thorax dark yellow with scattered brownish black spots. Pattern on the forewing in male is created by black small lines and dots; the reniform spot has slightly curved and wide lunate marking. The hindwing has dark scales on the veins. The abdomen is brown with yellow scales dorsally, differing from O. inca in having a line of black tufts over the middle of the dorsum of the abdomen; black tufts on A1-A3 combined with yellow scales. Genitalia of male with the cucullar region narrow and the apex rounded; saccular area with the process almost same size that the cucullus; aedeagus long, almost the same size of the vesica; large patch of spines on the base of the vesica; the two spine patches large and wide similar size covering ¾ of the vesica. Genitalia in female with sterigma relatively open, with the corpus bursae almost the same size as the appendix bursae.

Biology. Unknown.
Remarks. Many of the new species are similar to this species making the process of identification difficult. The holotype is in perfect condition (Fig. 10). Diagnosis. Oculicattus schmidti has a set of remarkable characters that separate it from other species, such as the orbicular spot is rounded; the lunate marking of the reniform spot is incomplete, being similar to those of Gaujonia species; the forewing is more stylized with straighter margins. The male genitalia are similar to those of O. boliviana, but can be distinguished from them by the cucullar region, which is narrower than those of other species; the saccular is large and densely covered with setae; also the uncus is wider and the saccus longer than the other species of Oculicattus. The DNA barcode is similar to that of O. renifera; however, external and internal morphology reveal enough differences to identify the two species.

Oculicattus schmidti
Description. Head. Palp marbled in black and white with frons yellowish gray; black patch between antennae small. Thorax. Covered in yellow with some black patches on dorsum. Wing. Forewing length: male 17-19 mm; forewing, dark yellow with well-developed lines from posterior margin to fold that look similar to those of other Oculicattus species; reniform spot with lunate marking narrowed in middle of base of cell M1; orbicular spot small, outlined in black, rounded; V-shaped mark on CuA2 base small, with upper line longer, extending to lower side of reniform spot; hindwing with fringe marbled in yellow and black; posterior margin with a combination of yellow and brown hair-like scales; veins yellow with two black lines in middle of veins from M2 to CuA2; black line posterior to base of CuA2. Leg. Prothoracic and mesothoracic legs marbled in yellow and black; metathoracic legs yellow. Abdomen. Dark yellow with dorsum clothed with dark brown and black scales; yellow tufts in middle of A1-A4 with a small vertical line on each tuft. Male genitalia. Cucullus bullhorn-like, with pointed apex heavily coated with setae; sacculus narrow with its process long and wide, densely covered with setae; terminus of saccular process tapered to apex; saccus long rhomboid-shaped, tip barely rounded; juxta shieldshaped and upper side concave; juxta with an expanded upper side; wide tegumen; aedeagus ¼ × longer than basal area of vesica, remarkable curved inwards; vesica ⅔ × longer than wide; one of patches of spines on tip small slightly squared and other covering ⅓ of whole vesica. Immature stages. Unknown. Distribution. This species is distributed from Central region of Peru and to the east region of Bolivia (Fig. 95).
Biology. Unknown. Remarks. Holotype in perfect condition (Fig. 45), paratypes in good condition even though the forewings are slightly damaged, but they are still well preserved.   Etymology. The name uturunku makes reference to the jaguar Panthera onca (Linnaeus) in the Quechua language.
Diagnosis. There is only one species that has similar characters to O. uturunku, which is O. raizae. Nevertheless, they are easy to separate because the color black is predominant in O. uturunku and the yellow is secondary. The reniform spot is black and quite smaller, converging with long black lines making it look like an eyelash. Genitalia of the male are remarkably smaller than O. raizae, mainly the cucullus, which is also remarkably narrower.
Description. Head. Palp with last segment divided into three parts, base and tip yellow and middle area black; frons dark yellow, with a large black band between antennae. Thorax. Marbled in black and sulfur-yellow dorsally, and sulfur-yellow ventrally. Wing. Forewing length, male 20-22 mm; forewing black with some regions of sulfur-yellow; enormous black lines on veins define forewing pattern; black orbicular spot small and elongated; unusual reniform spot eyelash-like, black; black line through inferior region of discal cell barely touching base of CuA2; hindwing with black fringe and some sulfur-yellow scales at end of each vein paler than forewing, whereas that from posterior margin completely gray; veins black with some spots of sulfur-yellow. Leg. Prothoracic and mesothoracic legs black with some sulfur-yellow on joints, and metathoracic legs in yellow. Abdomen. Black with segments sulfur-yellow ventrally, whereas dorsally dark gray, paler on first three segments; A1-A3 with tufts in yellow and with some black scales. Male genitalia. Cucullus wider on base and apex small; costal margin curved; sacculus and process wide; saccus narrow and rhomboid-shaped; juxta square-shaped with base narrower; tegumen narrow; aedeagus 3 × longer than wide; basal area of vesica 1 ½ × longer than vesica itself; large slightly curved patch of spines close to basal area; one of patches of spines on tip small with triangular terminus and another larger covering almost ½ of vesica.
Immature stages. Unknown. Distribution. Oculicattus uturunku occurs mainly in the Western Cordillera of the Andes in Ecuador. It is found at high to very high elevations (Fig. 95).
Biology. Unknown. Remarks. Holotype (Fig. 49) and paratypes in good condition. The species Oculicattus uturunku has been confused with Gaujonia nr. renifera by Piñas et al. (2002). The DNA barcode is very similar to O. schmidti (see O. schmidti diagnosis)

Discussion
The Andean genus Gaujonia had not been properly studied since its discovery more than a century ago (Dognin 1891), and thus its life cycle and taxonomic status, were essentially unknown. Nevertheless, during this revision, I found that Gaujonia was not a monophyletic group, but included species representing five genera. It is worth mentioning that the whole generic complex showed signs of mtDNA introgression or introgressive hybridization, found by studying Cytochrome c oxidase subunit I (COI), which has been observed in other groups of noctuoids even in other member of the subfamily Pantheinae (Schmidt and Sperling 2008;Anweiler 2009;Otim et al. 2018, Schmidt andAnweiler 2020).
The above morphological characters and the DNA barcode data demonstrated that the members of the Gaujonia genus group are sister to each other, but surprisingly despite similarities between the renifera and arbosi groups, the arbosi and vaunigrum groups were more closely related; thus, all four groups were separated into different genera. The arbosi group is established now as the true Gaujonia due to significant differences with the other groups. Three new genera, Millerana gen. nov., Oculicattus gen. nov., and Cicadoforma gen. nov., are proposed for arbosioides, renifera, and vau-nigrum groups, respectively. In addition, there were other specimens that were misidentified as Gaujonia sp., but molecular and morphological analyses demonstrated that they belong to different genera (Gaujoptera gen. nov. and Cicadoforma gen. nov.). Gaujoptera gen. nov. was found to be closely related to the genus Millerana and the Gaujonia generic group, whereas Cicadomorphus gen. nov. is close to Cicadoforma gen. nov. In addition, the species Gaujonia kanakusika proved to be genetically different from the other species in the genus Gaujonia, however, the apomorphic characters observed in this species were not sufficient to separate this species into a different genus.
On the other hand, it was found that the Gaujonia genus group is represented by cryptic genera, which have similar morphology but are genetically distant, such as Gaujonia and Oculicattus, as well as Cicadoforma and Cicadomorphus. However, the same problem was observed with the molecular characters in some species groups: Gaujonia arbosi-G. chiqyaq, Oculicattus boliviana-O. uturunku, and O. schmidti-O. renifera, which showed slight differences in the COI sequences. Therefore, both morphological and molecular characters played a critical role to make an appropriate identification, not only in this genus group, but in general since there are many cryptic genera and species that are still misidentified or even undiscovered.
In addition, despite what was proposed by Schmidt and Anweiler (2020) about the closer relationship between Arctioptera Schmidt & Anweiler and the rest of the jaguar moths based on similar external morphology, the phylogenetic analysis placed Arctioptera closer to the genera Meleneta Smith and Charadra Walker. The internal morphology also shows more similar characters with the genus Colocasia than with the jaguar moths.
Another important discovery was that some endemic species (Cicadomorphus falkasiska and Gaujonia kanakusika) are specialists on at-risk plants such as Alnus acuminata Kunth and Prunus subcorymbosa Ruiz ex Koehne and thus are themselves vulnerable to extinction (IUCN 2019). Additionally, these species along with others, are restricted to small areas. Therefore, the group is an ideal target for conservation efforts.
Finally, it was found that larvae of Cicadomorphus falkasiska regurgitate a transparent highly alkaline chemical compound, which was observed to deter ants (JIM pers. obs.).