Four new Neotropical Lophocampa species with a redescription of Lophocampa atriceps (Hampson) (Lepidoptera, Erebidae, Arctiinae)

Abstract Four new species of Lophocampa Harris, are described and illustrated: Lophocampa flavodorsata sp. n., Lophocampa griseidorsata sp. n., Lophocampa herbini sp. n. and Lophocampa sullivani sp. n. Lophocampa atriceps (Hampson) is redescribed, illustrated and compared to the new species.

Lepidoptera campaign (see www.lepbarcoding.org for details). DNA extraction, PCR amplification and sequencing follow the protocols described in Vaglia et al. (2008).
The taxa included in this study are listed in Table 1. Unfortunately, recently collected material of L. andensis was not available for sequencing. A set of twenty-four specimens of five species was sequenced for the 658 barcode base pairs fragment. Sequences were analyzed using maximum likelihood (ML) and maximum parsimony (MP) methods. MP analyses were carried out with MEGA 5 software (Tamura et al. 2011).

Species accounts
Lophocampa atriceps (Hampson) http://species-id.net/wiki/Lophocampa_atriceps Figs 1, 7, 13 and 19 Elysius atriceps Hampson, 1901: 113. Type material. Elysius atriceps: Male holotype [BMNH], examined. Type locality: Colombia. The holotype is labeled "sychesia atriceps type ♂ Hmpsn" (Hampson's hand); "Colombia 98-108" and "TYPE" (typeset). Redescription. Head -Antenna bipectinate, brown with yellow base and brown cilia. Frons light yellowish white, separated at base of antenna by transverse black line. Vertex yellowish white with a prominent black central spot. Palpi erect, yellow with second joint black at sides and third black and very short. Thorax -Patagia yellowish white with square black spot. Tegulae exteriorly yellowish white, medially brown, bordered with black. Thorax yellow-white with posterior medial brown spot. Legs with femur dull white spotted with brown; dorsal face of femur vivid yellow. Tibia dull white with brown spots. Tarsal segments brown with small whitish spots. Forewing -Brown irrorated with light and dark brown. One yellow spot and two small blacks dots at base. A series of bands formed by whitish spots, arranged as follows: antemedial band comprised of three spots; medial band curved, comprised of seven dark spots; postmedial band sinuous, a series of different size spots, the three adjacent to costa largest. Three spots forming triangle between median and postmedian bands, outermost on costa. Subterminal band comprised of spots of different shapes: arrowhead-shaped spot nearest tornus, second spot larger, spots above veins CuA1 and M2 very small and flattened, three near apex large, last one on apex ovate and compressed basad. Terminal line a series of white dots along margin. Ventral markings as above but paler. Hindwing -White slightly tinged with yellow marks on apex and along costa, with yellow tint along the anal border. Markings on ventral surface more contrasting. Abdomen -Tergites pale orange, almost entirely covered with long yellow hairs, and with a lateral series of black spots. Last segment pale orange, without hair. Sternites white with yellow-centered brown patches. Male genitalia -Uncus lozenge-shaped, setose, apex truncate. Tegumen and vinculum very slender. Saccus tongue-shaped, folded ventrally. Valve symmetrical, enlarged at base, slightly exceeding apex of uncus. Valvula slender, oblong and slightly twisted internally. Cucullus short, pointed and folded toward uncus. Juxta slender, with two arms fused in middle forming a stem with a elongated sclerotized patch at apex. Transtilla with two arms folded and converging, arms not fused, showing a large disconnect in the structure. Penis rectilinear, narrow with elongated caecum penis. Vesica large with two large and one small diverticulum. Five patches of cornuti: one at base, one on small diverticulum, and one on each of two large diverticuli. Female -identical to male except as follows: wingspan slightly larger. Antennae with pectinations shorter than in male; two small black points laterally on the collar, the central pair always present.
Biology and distribution. Early stages unknown. Colombia (Valle del Cauca), Ecuador (Guayas) and Costa Rica (Alajuela, Guanacaste, Puntarenas, Cartago). Hampson associated with the holotype an unspecified number of specimens from Bolivia, R[io] Tanampaya, [La Paz]. A second male specimen in the BMNH labeled atriceps is a specimen of Lophocampa griseidorsata Vincent & Laguerre, sp. n.
Remarks. The holotype in the BMNH has not been dissected. The type locality on the label states only "Colombia", so it is not possible to compare specimens from the  (Hampson, 1901), Holotype, male 2 Lophocampa hyalinipuncta (Rothschild, 1909), male syntype 3 Lophocampa flavodorsata sp. n., Holotype, male 4 Lophocampa sullivani sp. n., Holotype, male 5 Lophocampa griseidorsata sp. n., Holotype, male 6 Lophocampa herbini sp. n., Holotype, male.  (Hampson, 1901), specimen from Ecuador 8 Lophocampa hyalinipuncta (Rothschild, 1909), specimen from Ecuador 9 Lophocampa flavodorsata sp. n., Holotype 10 Lophocampa sullivani sp. n., Holotype 11 Lophocampa griseidorsata sp. n., Holotype 12 Lophocampa herbini sp. n., Holotype. same locality. However, the tips of the valves of the holotype are visible, and were compared with those of superficially identical specimens which were dissected. Specimens with valves matching those of the holotype are from the western slopes of the Andes in Colombia and Ecuador and the mountains of Costa Rica. The description for the genitalia of L. atriceps provided here is based on a male originating from the western slope of Ecuador. A female from the same locality in Ecuador with similar habitus is described and illustrated, including the genitalia. Etymology. The specific epithet, flavodorsata, refers to the yellowish hairs of the abdominal tergites.
Diagnosis. Lophocampa flavodorsata sp. n. can be distinguished from L. atriceps by the absence of black dots on the collar, the dark color of the head, collar and the base of the tegulae, as well as the different structure of both the male and female genitalia.
Description. Head -Antenna bipectinate, brown with yellowish base and brownish cilia. Frons light yellow on inferior half, deep brown on superior half. Vertex light yellow with strong brown central spot. Palpi erect, deep brown with second segment light yellow and third very short. Thorax -Patagia light yellow with a square brown spot centered with brownish orange. Tegulae exteriorly brown, medially yellow-white,  (Hampson, 1901), specimen from Ecuador 14 Lophocampa hyalinipuncta (Rothschild, 1909), specimen from Ecuador 15 Lophocampa flavodorsata sp. n., Holotype 16 Lophocampa sullivani sp. n., Holotype 17 Lophocampa griseidorsata sp. n., Holotype 18 Lophocampa herbini sp. n., Holotype. bordered with dark brown. Thorax light yellow with a median brown spot. Legs: Femur white spotted with brown, dorsal surface vivid yellow. Tibia white with brown spots. Tarsal segments brown with small white spots. Forewing -Brown irrorated with light and dark brown: one yellow spot with two small black dots at base. A series of bands formed by white spots arranged as follows: antemedial band broken; medial band slightly curved; postmedial band sinuous, originating at anal border and splitting between veins CuA2 and CuA1 into two branches reaching costa. After their separation the branches surround a strong brown reniform spot. Size of spots in this band  (Hampson, 1901), specimen from Ecuador 20 Lophocampa hyalinipuncta (Rothschild, 1909), specimen from Ecuador 21 Lophocampa flavodorsata sp. n., Allotype female 22 Lophocampa sullivani sp. n., Allotype female 23 Lophocampa griseidorsata sp. n., Allotype female 24 Lophocampa herbini sp. n., Allotype female. relatively constant. Basal branch slightly contrasting due to clear background. Subterminal band made up of different-shaped spots: spot near tornus arrowhead shaped, 2nd spot larger than first, those above veins CuA1 and M2 very small and flattened; three spots near apex larger; last one at apex ovate and compressed basally. Terminal line of white dots on margin. Ventrally as above but paler. Hindwing -Yellow-white slightly tinged with yellow-brown marks on apex and along the costa, with yellow tinge along anal border. Ventrally, marks more contrasting, deep brown centered with yellow-brown. Abdomen -Tergites pale orange and almost entirely covered with long yellow hair, with a lateral series of very faint brown spots. Last segment pale orange, without hair. Sternites white with brown patches centered with yellow. Male genitalia -Uncus rectangular, setose. Tegumen and vinculum slender. Saccus tongue-shaped, folded ventrally. Valves symmetrical, slender and long with apex slightly angled, extending beyond apex of the uncus. Valves slender, very elongate, terminating in a small tooth. Cucullus slender, elongated, folded toward uncus. Juxta very narrow, with two arms fused apically into a small V-shaped sclerotized patch. Transtilla slender, tongueshaped, interrupted in the middle. Aedoeagus. Penis rectilinear; caecum penis present. Vesica wide with four diverticuli, with four patches of cornuti, two with small spines and two with larger spines. Female -identical to male except as follows: Antennae with pectinations shorter than male. Female genitalia -Lamella antevaginalis slightly sclerotized, semicircular with a large medial U-shaped notch on the posterior margin. Apophyses posteriores straight, 0.9 mm long, apophyses anteriores slender, 0.6 mm long. Papillae anales rectangular, setose, with a pair of small pseudopapillae. Dorsal saccular pheromone glands absent. Ductus bursae asymmetrical with an extension on the right (ventral view). Corpus bursae reduced and ovate, bearing a large field of signa. Appendix bursae as large as corpus bursae. Ductus seminalis ventrally inserted at apex of corpus bursae and immediately transformed into seminal vesicle.

Lophocampa griseidorsata
Diagnosis. Lophocampa griseidorsata sp. n. is best distinguished from L. flavodorsata by the difference in the coloration of the abdominal scales, the larger whitish spots on forewings and by the structure of male and female genitalia, as given in the key. This species can also be distinguished from L. atriceps with the characters above along with the absence of black spots on the collar.
Description. Head -Antenna bipectinate, brown with basal third vivid yellow and with brownish cilia. Frons whitish with black T-shaped mark, vertex whitish anteriorly and black centered with brown posteriorly. Palpi erect, deep brown with 2nd segment whitish and 3rd segment very short. Thorax -Patagiae whitish with square brown spot centered with brownish orange. Tegulae exteriorly whitish, medially brown, bordered with dark brown. Thorax whitish with median brown line. Legs. Femur whitish, spotted with brown and with dorsum vivid yellow. Tibia whitish, spotted with brown. Tarsal segments brown with small whitish spots. Forewing -Brown irrorated with pale and deep brown. At base, one whitish spot with a yellow and two small blacks dots. Series of bands formed by whitish spots and organized as follows: broken antemedial band; slightly curved medial band, the size of the spots decreasing from anal border to  the costa; sinuous postmedial band starts on the anal border and splits between veins CuA2 and CuA1 into two branches that reach the costa. Size of spots constant on basal portion and basal branch. Apical branch constituted of larger spots. After their separation branches surround a strong brown reniform spot. Subterminal band constituted of different-shaped spots: spot near tornus forms a <-mark, 2nd spot larger, spots above veins CuA1 and M2 very small and flattened; three spots near apex large; last one at apex is large and ovate. A terminal line of white dots on the margin. Ventrally, the ornamentation is the same but paler. Hindwing -Whitish slightly tinged with grey on apex and along the costa. Ventrally, the marks are more contrasted, deep brown centered with yellowish brown. Abdomen -Tergites pale orange and almost entirely covered with long grayish brown hairs, with lateral series of black spots. Last segment pale orange without hairs. Sternites brown. Male genitalia -Uncus spatulated and setose. Tegumen and vinculum very slender. Saccus tongue shaped and folded ventrally. Valvae symmetrical, wide, reaching uncus apex. Valvae with prominent costa and acute, bifid apex, one tip short and rounded, the other very long and acute. Juxta with two curved arms fused apically to give a crescent-like sclerotized patch. Transtilla with two arms moving away at apex and which are not fused. Aedoeagus. Penis narrow and rectilinear. Caecum penis present. Vesica with a small and a large area of cornuti distally. Female -(Description based on allotype): identical to male excepted the following differences: Antennae with pectinations shorter than male. Wingspan slightly larger. Female genitalia -Lamella antevaginalis slightly sclerotized, rectangular with a medial notch on the posterior margin. Apophyses posteriores straight, enlarged at base, 1.2 mm long. Apophyses anteriores slightly curved, slender and 0.7 mm long. Papillae anales rectangular and setose, with a pair of large pseudopapillae. Dorsal saccular pheromone glands absent. Ductus bursae rectangular and striped. Corpus bursae reduced, oval, very wrinkled. Appendix bursae slightly smaller than corpus bursae, without signa. Apical insertion of the ductus seminalis.   Alfamayo, 2395m, 20-II-2009, 3 specimens will be deposited in [MUSM].
Etymology. Lophocampa herbini is named in honor of Daniel Herbin, specialist of Saturniidae and friend of the authors.
Diagnosis. Lophocampa herbini sp. n. is best distinguished from L. atriceps and L. flavodorsata sp. n. by the difference in the coloration of the abdominal pilosity, the larger whitish spots on forewings and by the conformation of male and female genitalia. This species can also be distinguished from L. griseidorsata with the same characters along with the absence of black spots on the collar. It is also the only species with a thin black line crossing the white ovate apical spot on the forewing.
Description. Head -Antenna bipectinate, brown with basal third vivid yellow and with brownish cilia. Frons whitish with a black T-shaped mark, vertex whitish anteriorly and black centered with brown posteriorly. Palpi erect, deep brown 2 nd segment whitish, 3rd segment very short. Thorax -Patagia whitish with a square brown spot centered with brownish orange. Pterygodes exteriorly whitish, medially brown, bordered with dark brown. Thorax whitish with a median brown line. Femur whitish, spotted with brown, dorsum vivid yellow. Tibia whitish, spotted with brown. Tarsal segments brown with small whitish spots. Forewing -Brown irrorated with pale or deep brown. At base, one whitish spot with one yellow and two small blacks dots. Presence of a series of bands formed by whitish spots and organized as follows: a broken antemedial band. A slightly curved medial band. The size of the spots decreasing from anal border to the costa. A sinuous postmedial band starts on the anal border and splits between veins CuA2 and CuA1 into 2 branches that reach the costa. The size of the spots is constant on the basal portion and the basal branch. The apical branch is constituted by larger spots. After their separation the branchs surround a strong brown reniform spot. Ssubterminal band constituted of different-shaped spots: spot near tornus forming a <-mark, 2nd spot larger, spots above veins CuA1 and M2 very small and flattened; three spots near apex large, last one on apex ovate. Terminal line of white dots on margin. Ventral pattern similar to dorsal, but paler.
Hindwing -Whitish, slightly tinged with grey on apex and along the costa. Ventral pattern more contrasting, deep brown centered with yellowish brown. Abdomen -Tergites pale orange, almost entirely covered with long grayish brown hairs, with lateral series of black spots. Last segment pale orange without hairs. Sternites brown. Male genitalia -Uncus setose, lozenge shaped, apex truncated and blunt. Tegumen and vinculum very slender. Saccus virtually absent. Valvae symmetrical, wide, reaching uncus apex, without prominent costa. Apex of valvae bifid, the two tips symmetrical, short and acute. Juxta with two curved arms fused apically to give a crescent-like sclerotized patch. Transtilla with two semi-rounded arms which are not fused. Aedoeagus. Penis moderately narrow and rectilinear. Caecum penis present. Vesica wide with four diverticuli and four spine patches: two with small spines and two with strong spines. Female -(description based on allotype): identical to male excepted the following differences: Antennae with pectinations shorter than male. Wingspan slightly larger. Female genitalia -lamella antevaginalis slightly sclerotized, semicircular with a large medial median U-shaped notch on the posterior margin. Apophyses posteriores straight, enlarged at base, 0.9 mm long. Apophyses anteriores narrower, slightly curved and 0.6 mm long. The left apophysis (ventral view) displays a fork at its apical extremity. Papillae anales rectangular and setose with a pair of small pseudopapillae. Dorsal saccular pheromone glands absent. Ductus bursae short, square and striate. Corpus bursae ovate, bearing a large signa. Ductus seminalis ventrally inserted at the extremity of a wide projection of corpus bursae and immediately transformed into a seminal vesicle.
Biology and distribution. Early stages unknown. Distribution: Bolivia (Santa Cruz, Cochabamba), Peru (Cuzco). Etymology. Lophocampa sullivani is named in honor of J. Bolling Sullivan who collected this species in 1988.

Lophocampa sullivani
Diagnosis. Lophocampa sullivani sp. n. can be best distinguished from L. atriceps by the absence of black dots on the collar and by the structure of the genitalia. It can be distinguished from L. griseidorsata sp. n. and L. herbini sp. n. by the difference in color of the abdominal scales. Lophocampa sullivani sp. n. is very close in appearance to L. flavodorsata sp. n. but can be separated by characters of the male genitalia, including the very prominent cucullus on the valve, the more robust genitalia and the presence of only three spine fields on the vesica. The female genitalia appear to be identical to those of L. flavodorsata.
Description. Head -pale yellowish, vertex brown with horizontal yellow bar, palpi dark fuscous with 2nd segment yellowish orange; antenna yellowish brown with brown-ish ciliae. Thorax -Patagia yellowish brown with a square brown spot centered with yellow, pterygodes striated with yellowish brown and chestnut brown, thorax yellowish with admixture of brown hairs. Legs yellowish, spotted with chestnut brown, fore coxae orange. Forewing -Yellowish irrorated with chestnut brown and a series of creamywhite spots with diffuse brown edges. At base, one orange spot with two small black dots. An antemedial, very oblique series of three creamy-white spots; a medial series angled on median vein, then very oblique, the two spots towards costa small; a spot at end of cell; an oblique postmedial series of six spots starting from anal border extending to end of cell followed by two series of three large spots, one basad and one distad, the last one just on the apex has the bottom constricted; a subterminal series of four spots, one near tornus forming a <-mark, the 2nd large, those above veins 3 and 4 smaller; a terminal line of white dots on the margin. Hindwing -Yellowish white slightly tinged with fuscous on apex and along the costa. Abdomen -entirely yellowish, with lateral series of black spots and ventral surface whitish. Male genitalia -Genitalia very robust. Uncus rectangular and setose, with slightly rounded apex. Tegumen and vinculum slender. Saccus tongue shaped and folded ventrally. Valvae symmetrical, wide and relatively short with apex notched into two points: one blunt and one acute. Cucullus prominent, slender and elongate, folded inward. Juxta very narrow, with two arms fused apically in a small V-shaped sclerotized patch. Transtilla slender, shaped like a tongue interrupted in the middle. Penis rectilinear. Caecum penis present. Vesica wide with four lobes. Presence of three areas of cornuti, two densely covered with small spines and one sparsely covered with strong spines. Female -(description based on allotype): identical to male except for antennae with pectinations shorter than male and wingspan slightly larger. Female genitalia -identical to those of Lophocampa flavodorsata.

Lophocampa andensis
Diagnosis. Lophocampa andensis is the smallest species of the group, with the length of the male forewing less than 20 mm. It is also distinguished from other species by the rounded apex of the forewings and by the spots between M2 and CuA1 on the forewing subterminal band, which are at least as large as the others.

Lophocampa hyalinipuncta
Diagnosis. Lophocampa hyalinipuncta can be distinguished from L. atriceps, L. flavodorsata sp. n. and L. sullivanni sp. n. by the different pattern of scales on the Abdomen -The spots of the subterminal band of the forewing are also significantly smaller than the spots on the postmedial band, a character that separates hyalinipuncta from the other species of the group.

Discussion
Distribution patterns. Each of the five species is restricted to only one slope of the Andean Cordillera (Table 1; Fig. 25); L. atriceps and L. sullivani are restricted to the western slope; L. flavodorsata, L. griseidorsata, L. herbini occur only on the eastern slope. Lophocampa atriceps and L. sullivani occur at different elevations, with L. sullivani at elevations of 700-1450 m, and L. atriceps from 1840 -2000 m, which will help separate the two taxa. The three taxa occurring on the eastern slope of the Andes overlap in range, and L. flavodorsata and L. griseidorsata have been collected several times at the same locality. The atriceps group occupies a very large area of the Andean Cordillera and extends north to the central mountain range in Costa-Rica at medium to high altitudes.
The Andean Cordillera appears to have created a barrier between the species studied here. Lophocampa atriceps is distributed from Central America in the north to Ecuador in the south but is restricted to the western slope in South America. A similar pattern has been observed for several plant species (Schnell 1987). In northern Peru, the presence of a very dry region on the Pacific slope prevents a more southerly range extension.
Molecular data. Twenty-four voucher specimens provided barcode sequences (Table 2). Calculations of genetic distances between species of the L. atriceps group (Table  3) support the distinctness of the five species. Intraspecific divergence ranged from 0 -0.99 % with a mean divergence of 0.21 %. Interspecific divergence ranged from 2.02 -5.46 % with a mean divergence of 3.61 %. The four new species described herein were all originally identified as L. atriceps. Differences in barcode sequences of L. atriceps and the four new species described herein range from 8.05 -9.64 %. These values are significant and confirm the significance of the morphological differences observed and the discrimination of the taxa. The greatest difference (9.64 %) is observed between L. atriceps and L. sullivani, the unique sympatric species. The geographic isolation that occurred with the species on the eastern slopes of the Andes did not result in the highest divergence values.
The lowest interspecific divergence was observed between L. griseidorsata and L. herbini with 2.02 %. Morphologically both species are in fact the closest with similar male genitalia structure. They do not appear to be sympatric because their altitudinal difference is in the range of 400 m. Similarly low interspecific divergence is observed also between L. flavodorsata and L sullivani, with 2.27 %. Again the two species are morphologically similar and show little difference in male genitalia structure, but they occur on different slopes of the Andes. Lophocampa hyalinipuncta differs from the new taxa by 3.56 -5.19 %. These values are significantly lower than for L. atriceps (9.33 %) even if the new species have hitherto been confused with L. atriceps. Specimens of L. andensis were not available for DNA sequencing.
The barcode data shows that L. flavodorsata is most closely related to L. sullivani, with each restricted to only one slope of the Andes, but with a broader geographic  distribution for L. flavodorsata. On the other hand, the pair griseidorsata + herbini is closest to hyalinipuncta and well-differentiated from flavodorsata + sullivani. Here also one species (griseidorsata) has a broad distribution whereas the second (herbini) appears to have a very restricted range in Bolivia. Lophocampa flavodorsata and L. griseidorsata which share a broad range on the Atlantic slope of the Andes display the largest difference in the COI gene sequence (4.9-5.3 %) for the whole group along with the pair sullivani + hyalinipuncta (which seems normal for such widely differentiated species).