Ecuador, Intaj [possibly Intag, Imbabura province].

Druce (1885a) described H. buckleyi and H. lex. The upper- and undersides of the wings in H. buckleyi are very similar, with the colour of the hindwings generally being paler. The forewings feature a deep orange transversal band on a dark brown background, and the hindwings show metallic blue fields between the veins, with three located on the upperside between M₃ and CuA₂ and one in the cell, and eight between all veins on the underside. The pattern of the female is similar, with the blue fields extending further on the forewing, including the blotch between veins CuA₂ and A. In the female, metallic blue scales are also present at the base of the forewing at both the wing upper- and underside.

North-western Ecuador.

The largest species of the clade. The extension of the blue fields is significantly larger than in H. lex. The orange transversal band on the forewing is more saturated and more rounded than in H. catagrammina. Form and extension of the metallic blue blotches are different from those in H. catagrammina (Figs 2–3, 5–6).

Nicaragua: Chontales; Panama: Volcán de Chiriqui; Bugaba, 800–1500 ft.

Druce (1885b) described catagrammina in the same year, but separately from H. buckleyi and H. lex. The taxon was put in synonymy with buckleyi by Parsons et al. (1999). As noted by Druce, catagrammina is closely related to the other two species of the clade and particularly similar to H. buckleyi. In agreement with Druces’ original description of the three taxa, I revive the species from synonymy with H. buckleyi due to small but overall significant differences of the wing patterns. The morphological differences hint to different species, particularly given the experience from many other species complexes of Neotropical Geometridae in which often more subtle differences – ideally combined with results from genitalia morphology and barcoding – can be observed in different species.

Central America, from Nicaragua to Panama.

The extension of the blue blotches is significantly larger than in H. lex. The transversal band on the forewing is paler and straighter than in H. catagrammina. Form and extension of the metallic blue blotches are different from those in H. buckleyi (Figs 2–3).

Ecuador (east), [Pastaza], Sarayacu.

Hagnagora lex was described by Druce (1885a) together with H. buckleyi. While buckleyi was collected on the western slopes of the Andes, H. lex originates from the Amazon slopes of the Eastern Andes.

Eastern Ecuadorian Andes (Pastaza: Sarayacu).

Smaller than H. buckleyi and of similar size to H. catagrammina. The extension of the blue blotches is significantly smaller than in H. buckleyi. The form of the transversal band on the forewing is similar to that in H. buckleyi, but the band does not stretch as far towards the wing margins. H. lex is the species with the smallest extensions of metallic blue blotches on the underside, with the upperside completely devoit of these blotches.

[Colombia], Bogotá.

H. anicata was re-described with a description also of the male genitalia, by Sullivan (2013). The lectotype is illustrated in Fig. 7. A series of specimens collected in southern Ecuador (1999–2013) (Fig. 8) is indistinguishable from H. anicata and therefore regarded as conspecific. The female (Fig. 12) is larger than the male. A living specimen is shown in Fig. 41 in the typical resting position of these beautiful moths.

Figures 7–11. 

7 Hagnagora anicata (F&R), male lectotype a dorsal view b ventral view c valvae d aedeagus 8 H. anicata (F&R), male from Ecuador as reference specimen with Barcode Index Number (BIN) a dorsal view b ventral view c valvae d aedeagus 9 H. richardi sp. n., male holotype a dorsal view b ventral view c valvae d aedeagus 10 H. unnia Sullivan, male a dorsal view b ventral view 11 H. marionae Brehm & Sullivan, male a dorsal view b ventral view.

Apart from its Colombian type locality, H. anicata has recently been collected and barcoded from sites in southern Ecuador to central Bolivia at elevations ranging from 2000 to 2920 m a.s.l.

Most species of the H. anicata clade are very similar, and the most reliable current method for diagnosis is the COI barcode. H. anicata tends to be smaller than the other species occurring sympatrically, namely H. richardi and H. hedwigae: The wing length of the male (holotype) is only 17.5 mm in comparison to 19 mm in the male holotype of H. richardi. The structures of the female signum are also more complex than in H. richardi, but similar to those in H. hedwigae. The uncus of the male is smaller and shorter than in H. richardi. Aedeagi of the known males are (H. anicata and H. richardi) similar. COI barcode: The minimum observed distance to the presumbably most closely related species (H. hedwigae) is 3.1%.

Costa Rica: Alajuela Province, Poás Volcano National Park, 2500 m.

H. elianne was described and illustrated by Sullivan (2011). The species closely resembles the other species in the H. anicata clade, particularly H. unnia.

The species has recently been collected and barcoded in Honduras (Cortes Province) and in several provinces of Costa Rica at elevations ranging from 1480 to 2840 m a.s.l.

Males are on average slightly larger than males in H. unnia and can be distinguished from H. anicata by a swollen as opposed to a gently tapered distal half of the uncus and by the absence of a moderately large, upcurved spine at the end of the costa in H. elianne (Sullivan 2011). Females may be distinguished from females of H. unnia by their longer, more complex signum. COI barcode: The minimum observed distance to the presumably most closely related species (H. unnia) is 5.0%.

(holotype): Costa Rica, Cartago Province, Tapantí National Park, 1275 m.

H. unnia was recently described and illustrated by Sullivan (2011).

The species is known from several provinces in Costa Rica at elevations ranging from 587 to 2840 m a.s.l.

The species closely resembles other species of the H. anicata clade, particularly H. elianne; see there for a diagnosis. COI barcode: The minimum observed distance to the presumably most closely related species (H. elianne) is 5.0%.

Costa Rica, Heredia province, Braulio Carrillo National Park, Volcán Barva, 2730 m a.s.l.

H. marionae was described and illustrated by Brehm and Sullivan (2005).

The species has been collected only at two high mountain areas in Costa Rica at elevations > 2500 m a.s.l.

The species resembles the other species of the H. anicata clade, but is easily distinguished by large orange-yellow blotches on the forewing. Males have a spatula-shaped uncus. COI barcode: The minimum observed distance to the presumably most closely related species (H. richardi) is 6.6%.

Holotype: male (Fig. 9): Ecuador, Loja province, Parque Nacional Podocarpus, Cajanuma, 04°06.85'S, 79°10.47'W, 2916 m, 20 November 2008, G. Brehm leg. (ID 18080, genitalia preparation, barcode sequence 658 bp) (PMJ).

Paratypes: (deposited in CISEC, PMJ, RCGB, ZSM) 4 males, 2 females. 1 female: same as holotype but 04°06.86'S, 79°10.46'W, 2897 m, F. Bodner leg. (ID 15855, barcode sequence 658 bp); 1 female (Fig. 13): Ecuador, Zamora Chinchipe, Reserva Biológica San Francisco, 03°58.72'S, 79°04.44'W, 2180 m, 16 November 2008, F. Bodner leg. (ID 16285, barcode sequence 658 bp); 1 male same as previous but 28 October 1999, D. Süßenbach leg. (BC ZSM Lep 04774, barcode sequence 529 bp); 1 male: same as previous but 03°59.65'S, 79°04.10'W, 2670 m, G. Brehm leg. (ID 17328, barcode sequence 658 bp); 1 male as previous but 03°59.68'S, 79°04.10'W, 2677 m, 18 November 2008 (ID 16119, barcode sequence 658 bp); 1 male as previous but 25 November 2008 (ID 17863, barcode sequence 621 bp).

Figures 12–14. 

12 H. anicata (F&R), female from Ecuador as reference specimen with Barcode Index Number (BIN) a dorsal view b ventral view c genitalia 13 H. richardi sp. n., female, paratype a dorsal view b ventral view c genitalia 14 H. hedwigae sp. n., female, paratype a dorsal view b ventral view c genitalia.

As illustrated in Figs 9, 13. The wing length of the holotype (male) is 19 mm. The wing length of a female paratype (Fig. 13) is 21 mm.

Only known from a small region around Podocarpus National Park, provinces Zamora-Chinchipe and Loja, Ecuador, with an observed elevational range of 2180–3021 m a.s.l. Apart from the

and nearby sites, specimens were collected at elevations at ca. 3000 m at Cerro Toledo in the same National Park (04°23'S, 79°07'W). However, this record is not fully reliable because genitalia preparation or barcoding was not conducted for these specimens.

Closely resembles other species of the H. anicata clade. On average significantly larger than H. anicata, but the female has about the same size as H. hedwigae. The uncus of the male is larger and broader than in H. anicata. The signum of the bursa copulatrix is less complex than in H. anicata and H. hedwigae. Easily distinguishable from H. marionae by the cream-white colour of the blotches on the forewing. COI barcode: The minimum observed distance to the presumably most closely related species (H. marionae) is 6.6%.

Hagnagora richardi is named in honour of Richard Philipp from Jena, Germany, in recognition of his and his parents’ support for the taxonomy of Neotropical geometrid moths.

Holotype: female (Fig. 14): Ecuador, Loja province, Reserva Biológica San Francisco, 03°59.68'S, 79°04.10'W, 2677 m, 25 November 2008, G. Brehm leg. (ID 17397, genitalia preparation, barcode sequence 595 bp) (PMJ).

As illustrated in Fig. 14.

Only a single female is known from H. hedwigae collected in southern Ecuador (2677 m). The wing length of the holotype (female) is 21 mm (same size as richardi).

Resembles most closely H. anicata and H. richardi, but is larger than H. anicata, and the signum of the bursa copulatrix is more complex than in H. richardi. COI barcode: The minimum observed distance to the presumably most closely related species (H. anicata) is 3.1%.

Hagnagora hedwigae is named in memory of Hedwig Seppelt (*1919 in Baumgarten, Silesia; † 2013 in Korschenbroich, Germany). Mrs Seppelt loved nature, and she took care that birds, small animals and insects found a habitat in her garden. The name is given in recognition of support for the taxonomy of Neotropical geometrid moths provided by her daughter-in-law Irmgard and her son Winfried Seppelt.

[Ecador, (south)], Loja surroundings.

H. croceitincta was described by Dognin from southern Ecuador where it has recently been collected in montane forests (Brehm 2002). As one of the largest known Hagnagora species, it is conspicuously coloured, with orange, dark brown and white patterns. The taxon epimena (Bastelberger) remains in synonymy because the lectotype specimen (Fig. 16) does not show any particular differences to the type specimen of H. croceitincta (Fig. 15).

Figures 15–22. 

15 Hagnagora croceitincta (Dognin) female, holotype a dorsal view b ventral view 16 H. epimena (Bastelberger) male, lectotype a dorsal view b ventral view 17 H. croceitincta male from Ecuador as reference specimen with Barcode Index Number (BIN) a dorsal view b ventral view 18 H. clustimena (Druce) female, lectotype a dorsal view b ventral view 19 H. clustimena female from Costa Rica as reference specimen with Barcode Index Number (BIN) 20 H. clustimena male from Costa Rica as reference specimen with Barcode Index Number (BIN) a dorsal view b ventral view c valvae d aedeagus 21 H. mirandahenrichae Brehm sp. n. male, holotype a dorsal view b ventral view c valvae d aedeagus 22 H. mirandahenrichae female, paratype a dorsal view b ventral view c genitalia.

Recently collected and barcoded specimens were sampled from central Colombia to southeastern Peru at elevations between 1750 and 2540 m a.s.l.

On average larger than the closely related species H. clustimena and H. mirandahenrichae: Forewing length of the female holotype reaches 23 mm in comparison to about 20 mm in the other species. On the forewing, the white transversal blotch does not stretch to the costal margin as seen in the other two species, and the apical, dark-brown area reaches beyond veins 1A+2A. The species is also generally more vividly coloured than the other species in this clade, with white spots on the forewing between veins CuA₂ and 1A+2A and around M₃ on the upperside of the hindwing. COI barcode: The minimum observed distance to the presumably most closely related species (H. mirandahenrichae) is 7.1%.

Mexico: Coatepec.

H. clustimena was originally assigned by Druce to Heterusia and then transferred to Hagnagora by Parsons et al. (1999). H. clustimena and H. croceitincta appear to occur allopatrically.

Besides the type specimens described by Druce from Mexico and Panama, recently collected and barcoded specimens were sampled in Honduras and Costa Rica between 850 and 1550 m a.s.l.

On average smaller than H. croceitincta and slightly larger than H. mirandahenrichae (see H. croceitincta). The white transversal blotch on the forewing stretches to the costal margin and the apical dark-brown area reaches vein CuA₂, as also observed in mirandahenrichae. H. clustimena is slightly paler than H. mirandahenrichae. The male genitalia of both species are similar, but the valves are broader and differently shaped to mirandahenrichae. COI barcode: The minimum observed distance to the presumably most closely related species (H. mirandahenrichae) is 4.6%.

Costa Rica, Área de Conservación Guanacaste, Guanacaste province.

Holotype: male (Fig. 21): Costa Rica, Guanacaste province, Área de Conservación Guanacaste, Sector Santa Maria, Mirador Santa Maria, 10.766° N, 85.301°W, 920 m a.s.l., 20 June 2012, S. Rios & R. Franco leg. (voucher 12-SRNP-103819, genitalia preparation, barcode sequence 658 bp) (PMJ).

Paratypes: (deposited in PMJ, USNM) 5 males, 1 female. Costa Rica, Guanacaste province, Área de Conservación Guanacaste, Sector Pitilla, Estacion Pitilla, 10.989° N, 85.426° W, 675 m a.s.l.; 1 female (Fig. 22) 16 May 2007, F. Quesada & R. Franco leg. (voucher 07-SRNP-103401, genitalia preparation, barcode sequence 658 bp), 1 male same as previous but 17 May 2007 (voucher 07-SRNP-103498), 2 males, 02 Apr 2011, H. Cambronero & S. Rios leg. (vouchers 11-SRNP-102035 and 11-SRNP-102036, barcode sequences 658 bp), 1 male 12 November 2012, R. Franco & H. Cambronero leg. (voucher 12-SRNP-105462).

As illustrated in Figs 21, 22.

Only known from sectors Santa Maria and Pitilla from Área de Conservación Guanacaste, province Guanacaste, NW Costa Rica, at elevations ranging from 675–920 m a.s.l., and therefore with a lower elevational range than H. clustimena (observed: 850–1550 m a.s.l.).

Easily distinguished from H. croceitincta by its wing patterns (see diagnosis in that species). The yellow ground colour of H. mirandahenrichae is slightly more intensive than in H. clustimena. The male genitalia of both species are similar, but the valves of mirandahenrichae are narrower and have a different shape to clustimena. COI barcode: The minimum observed distance to the presumably closest relative, H. clustimena, is 4.6%.

Hagnagora mirandahenrichae is named in honour of Ms. Miranda Henrich of California in recognition of her and her mother’s critical support for understanding the taxonomy and biodiversity development of the Área de Conservación Guanacaste (ACG) in northwestern Costa Rica, where this species has been found by the ACG caterpillar inventory (Janzen et al. 2014).

Costa Rica.

H. mortipax is one of the earliest described species in the genus and among the smallest Hagnagora species. Together with H. luteoradiata it also has the largest known geographical range. The taxon flavipectus remains in synonymy because it falls within the confirmed geographical range of mortipax and shows no significant deviations from the type specimen of mortipax. In comparison to the type specimen, the extension of the large white blotch on the forewing is smaller in Ecuadorian specimens, where it does not reach the costal margin. Since the barcode sequences of Costa Rican and Ecuadorian populations are nearly identical, all respective specimens are treated as members of the same species, and slight differences in wing patterns are regarded as geographical variability.

Recently sampled and barcoded material is either from Costa Rica (Fig. 24) or Ecuador (Fig. 25), from elevations ranging from 540–2180 m a.s.l., and additional material from Ecuador falls within the same elevational range (Brehm 2002).

Figures 23–29. 

23 Hagnagora moripax (Druce) male, lectotype a dorsal view b ventral view 24 H. mortipax male from Costa Rica as reference specimen with Barcode Index Number (BIN) a dorsal view b ventral view 25 H. mortipax male from Ecuador as reference specimen with Barcode Index Number (BIN) a dorsal view b ventral view 26 H. flavipectus (Warren) male, holotype a dorsal view b ventral view 27 H. mortipax jamaicensis (Schaus) male, lectotype a dorsal view b ventral view (photo USNM) 28 H. mortipax acothysta (Schaus) female, lectotype a dorsal view b ventral view (photo USNM) 29 H. guatica (Schaus) female, lectotype a dorsal view b ventral view (photo USNM).

The upper side of the wing in H. mortipax has a dark brown base colour with a large cream-white blotch on the forewing. This blotch almost reaches the outer margin, also either reaching the costal margin (Costa Rican specimens), or scantily not (Ecuadorian specimens). The white blotch is narrower in H. jamaicensis (Fig. 27), and significantly smaller, and separated from the outer margin, in H. acothysta from Brazil. All three species are significantly larger than H. guatica.

Jamaica.

Originally described as a Heterusia species by Schaus (1901), this taxon was down-ranked as a subspecies of mortipax by Parsons et al. (1999). In my view, the significantly different wing pattern in jamaicensis justifies Schaus’ original species rank, but further evidence from barcoding is desirable in order to consolidate its species status.

Jamaica.

In contrast to the other taxa in the mortipax clade, this species displays a very narrow, cream-white transversal band on the forewings. The striation on the underside of the hindwing is reduced in comparison to mortipax and acothysta.

[Brazil], Parana, Castro.

Together with jamaicensis, Schaus (1901) originally placed this species in the genus Heterusia. It was then ranked down as a subspecies of mortipax by Parsons et al. (1999). The major characteristic of acothysta is the reduction of the white transversal band (found both in mortipax and jamaicensis) to a smaller blotch that reaches about half the area found in mortipax. As in jamaicensis, further evidence from barcoding is desirable for the consolidation of the species status.

Brazil.

Unlike mortipax and jamaicensis, this species shows no white transversal band on the forewing, but rather a reduced blotch that reaches only about 50% of the size observed in mortipax.

Guatemala, [Quetzaltenango Department], Volcán Sta. Maria.

Schaus described guatica as belonging to Scordylia Gn (a junior synonym of Heterusia). The wing pattern of guatica strongly resembles that of other members in the mortipax clade, but the species lacks the typical striation on the underside of the hindwing. Further evidence from barcoding and the study of the genitalia will help to better understand the relationships of this species with other species of the mortipax clade.

Guatemala.

By far the smallest Hagnagora species. The species lacks the typical striation on the underside of the hindwing found in all other members of the mortipax clade.

[Colombia], Bogota.

Felder & Rogenhofer described this species from Colombia. It closely resembles H. discordata and H. luteoradiata. Parsons et al. (1999) put luteoradiata in synonymy with ephestris, but freshly collected material from Costa Rica and Ecuador shows that luteoradiata consistently lacks yellow blotches on the hindwing. It appears therefore to be more likely that ephestris is a junior synonym of discordata, and an increased knowledge of COI sequences could help to solve this question. Given the current state of knowledge, it appears to be the most appropriate solution to revive luteoradiata from synonymy and to treat the other two taxa as full species.

Figures 30–35. 

30 Hagnagora ephestris (F&R) male, lectotype a dorsal view b ventral view 31 H. discordata male, lectotype a dorsal view b ventral view 32 H. discordata male (ZSM Lep 44128) from Brazil as reference specimen with Barcode Index Number (BIN) (photo ZSM) 33 H. luteoradiata (T-M) male, lectotype a dorsal view b ventral view 34 H. luteoradiata (T-M) male from Costa Rica (CR) as reference specimen with Barcode Index Number (BIN) a dorsal view b ventral view 35 H. luteoradiata (T-M) male from Ecuador as reference specimen with Barcode Index Number (BIN) a dorsal view b ventral view.

Colombia.

Both ephestris and discordata show a pronounced yellow blotch on the hindwings that is absent in luteoradiata. Different from discordata, the yellow transversal band on the forewing of H. ephestris reaches the outer margin of the wing. Moreover, the band is broader than in discordata, whereas the yellow field of the hindwing is narrower, particularly in the proximate half of the wing.

[Chile], Valparaíso [possibly incorrect locality].

The oldest described Hagnagora species, assigned by Guenée to Scordylia Gn (= Heterusia).

Given as Valparaíso, [Chile], requires confirmation. The cool-dry climate of this Chilean lowland region differs strongly from the wet montane habitats where other Hagnagora species are typically found.

Apart from the doubtful type locality in Chile, recently collected specimens were sampled in Santa Catarina, Brazil (27°S), at elevations of 1300 m a.s.l.

Both discordata and ephestris show a pronounced yellow blotch on the hindwings that is absent in luteoradiata. The yellow transversal band on the forewing is narrower than in ephestris, and it does not reach the outer margin of the wing. The yellow blotch on the hindwing is much broader than in H. discordata. COI barcode: The minimum observed distance of Brazilian H. discordata is 2.3% to H. luteoradiata from Costa Rica and 2.6% to H. luteoradiata from Ecuador. These short distances suggest a relatively young split within this species clade.

Costa Rica.

H. luteoradiata was put in synonymy with H. ephestris by Parsons et al. (1999). However, luteoradiata specimens consistently do not show any yellow blotches on the hindwing as observed in ephestris and clustimena. Barcoded specimens from Costa Rica (Fig. 34) and Ecuador (Fig. 35) are genetically very similar (distance only ca. 1.1%) and, together with the highly similar appearance, are therefore regarded as conspecific. The ephestris type specimen from Bogotá, Colombia, falls within the geographical range of the luteoradiata specimens, but shows a different wing pattern, i.e. a prominent yellow blotch on the hindwing and a different shape of the blotch of the forewing. The taxon luteoradiata is therefore revived from synonymy.

Costa Rica to Ecuador. Observed elevational range in Ecuador 1800–2890 m and 560–1480 m in Costa Rica.

The most prominent difference is the absence of any yellow blotches on the hindwing that are present both in ephestris and discordata. The transversal yellow band on the forewing is broader than in discordata, and has a different shape than in ephestris.

Peru (south east), Carabaya, Oconeque, 7000 ft.

Warren originally assigned subrosea to Cophocerotis Warren, but the genus-defining type species, C. jaspeata (Dognin), does not show the two prominent white transversal bands of the forewing present in subrosea. Parsons et al. (1999) transferred the species to Hagnagora. Barcoding and genitalia dissections of fresh specimens from this species are required, but judging from the two transversal bands, subrosea might indeed be associated with the croceitincta clade.

Peru.

H. subrosea has a unique combination of a pale brown wing colour with two white transversal bands on the forewings not found in any other species of Hagnagora.

Colombia, Bogotá, 2800–3200 m.

Chile.

Chile, Provincia de Valdivia.