Xylophanes cthulhu Haxaire & Vaglia

Vaglia, Thierry, Haxaire, Jean, Kitching, Ian J., Meusnier, Isabelle & Rougerie, Rodolphe, 2008, Morphology and DNA barcoding reveal three cryptic species within the Xylophanes neoptolemus and loelia species-groups (Lepidoptera: Sphingidae), Zootaxa 1923, pp. 18-36 : 31-34

Xylophanes cthulhu Haxaire & Vaglia n. sp.

( Figs. 4 a, 4b, 12a–c)

Type material: Holotype ( Figs. 4 a, 4b; deposited in coll. JH; BC-Hax4328/ SOWE 429-07): ɗ, Guatemala, Izabal department, track from Chocchoc to Cebol (Cevol), km. 2, Pueblo Cadenas, 76 m., 21.vii.2004, leg. O. Paquit & J. Haxaire. Paratypes: 2 ɗɗ, same data as holotype (BC-Hax4322/ SOWE 423-07; BC-Hax4324/ SOWE 425-07); 1 ɗ (BC-Hax4423/ SOWE 524-07), Guatemala, Succhitepequez Dept., Reserve Tarrales, 1041 m., 7.v.2007, leg. M. Lauras & D. Herbin; 1 ɗ, Guatemala, Baja Verapaz Dept., Reserve Santa Rosa, 1580 m., 23.v.2007, leg. M. Lauras & D. Herbin; 1 ɗ (BC-Hax4329/ SOWE 430-07), Guatemala, Huehuetenango Dept., Soloma, near Cruz Maltin, Aldea Crinolina, 1900 m., 16.v.2002, leg. J. Monzon Sierra; 1 ɗ (BC-Hax4323/ SOWE 424-07), Costa Rica, Guanacaste Prov., Area de Conservacion Guanacaste, Sector Pitilla, Estacion Biologica Pitilla, 800m., 4.v.2005, leg. J. Barbut & A. Lévêque; 1 ɗ (BC-Hax0765/ SOWA 772-06), genit. prep. #Hax171, Mexico, Chiapas State, road from Ococingo to Palenque, track to Salto de Agua, km. 10, 50m., 21.viii.1992, leg. J. Haxaire & D. Herbin; 1 ɗ (BC-Hax0768/ SOWA 775-06), genit. prep. #Hax169, Mexico, Chiapas State, Municipio Oxchuc, track to Pashtonticja, km. 4, 1600m., 20.viii.1992, leg. J. Haxaire & D. Herbin; 2 ɗɗ, genit. prep. #Hax168, Mexico, Veracruz State, road from Coatepec-Teocelo to Los Altos, track to Chilchotla, km. 4, 1350m., 3.viii.1992, leg. J. Haxaire & D. Herbin; 2 ɗɗ (BC-Hax0766/ SOWA 773- 06; BC-Hax0767/SOWA-774-06), Mexico, Veracruz State, road from Coatepec-Teocelo to Los Altos, track to Chilchotla, km. 4, 1350m., 24.viii.1992, leg. J. Haxaire & D. Herbin; 6 ɗɗ and 1 Ψ, Mexico, Oaxaca, Sierra Juarez, 12-17.iii.1992, leg. local collectors; 8 ɗɗ, Mexico, Veracruz, Dos Amates, 20.x.2005, leg. local collectors; 1 Ψ, Mexico, Veracruz, Catemaco, 06.viii.2004, leg. local collectors; 1 ɗ (BC-Hax0770/ SOWA 777- 06), Nicaragua, Granada, Mombacho Volcano, alt. 800 m., 02.viii.2000, leg. M. Laguerre; 3 ɗɗ, Nicaragua, Nueva Segovia, Rio Mazarite, 10-12.xi.2000, leg. J.-M. Maes; 1 ɗ, Panama, Coclé, Cerro Gaital, El Valle, 27.v.1994, leg. N. Smith & D. Mitchell; 1 ɗ (BC-Hax0769/ SOWA 776-06), Honduras, Yoro, Pijol Mountain, alt. 1500 m., 19.vii.1995, leg. T. Porion; 2 ɗɗ, Panama, Panama Prov., Cerro Jofe, 900-1000m, 9-13.v.2007, leg. J. Touroult; 1 Ψ, Panama, Chiriquí, 1980, leg. Moinier; all the above paratypes held in the collections of JH and TV, except 2 ɗɗ to be deposited in the CNIN and BMNH. In addition, 11 paratypes (6 ΨΨ, 5 ɗɗ) are designated from Costa Rica, Area de Conservacion Guanacaste: 1 ɗ, Sector Cacao, Gongora Bananal, alt. 600 m., 05.viii.2004, leg. M. Pereira; 1 Ψ, Sector Pitilla, Ingas, alt. 580 m., 29.i.2005, leg. M. Rios; 2 ɗɗ and 2 ΨΨ, Sector Pitilla, Loaiciga, alt. 445 m., 10-26.i.2005, leg. M. Rios; 2 ɗɗ and 2 ΨΨ, Sector Pitilla, Pasmonpa, alt. 440 m., 10-29.i.2005, leg. P. & M. Rios; 1 Ψ, Rincon Rainforest, Camino Rio Francia, alt. 410 m., 16.viii.2004, leg. J.Perez. All of these 11 specimens are part of the dataset used in the DNA barcoding study by Hajibabaei et al. (2006); these specimens to be deposited in the Smithsonian Institution, Washington DC, USA, and specimen data as well as sequences are available from the public BOLD project ‘ Sphingidae of the ACG1’ (code MHASA) or GenBank (accession numbers DQ276772 to DQ276782 ).

Description: This species is immediately distinguishable by its bright coloration and the acute and falcate apex of the forewings. It is widely distributed in Central America and was recorded as X. neoptolemus by Mooser (1940: 456) and Hoffmann (1943: 233) in their surveys of Mexican sphingids.

Male ( Figs. 4 a, 4b). Head and body: Dorsal part of body olive-brown; labial palpi, area above eyes and tegulae finely marked with grey. Tegula with a median longitudinal gold line. median dorsal area of thorax greyish-beige, contrasting with patagia and tegulae. Upperside of abdomen with five thin longitudinal dark beige lines. Forewing length: 35 mm. Forewing upperside: General background color olive-brown; crossed by six postmedian and two submarginal lines as in the previously described species. First and fifth postmedian lines the most heavily contrasted, delimiting between them a cream-colored band in the median area of the wing. General coloration beyond fifth postmedian line remains constant, submarginal area not strongly differentiated from this postmedian part; this region crossed by three even lines (sixth postmedian and the two submarginal). Apex acute, slightly to strongly falcate, especially in specimens from Veracruz State, Mexico. Hindwing upperside: Basal area pure black, extending toward tornus, where it turns pale grey with a white inner edge. Median band bright red, somewhat pinkish in fresher specimens; tapering progressively toward apex of wing, almost reaching it. Forewing and hindwing undersides: Contrasted but with few distinct markings; ground colour reddish yellow, of uniform aspect. Basal area of forewing grey-beige; first and fourth postmedian lines apparent, first straight and strongly marked, fourth very narrow and only visible from inner edge to vein M2, beyond which it is replaced by small black vein dots. Between these two lines, the wing is golden-yellow, contrasting with the reddish tone of the rest of the wing; orange submarginal area dentate between M2 and M 3 in most specimens. First submarginal line also usually apparent, running from the inner margin to the apex.

Female. Forewing length: 37mm. Identical to the males in terms of general wing pattern and color, both upperside and underside, differing only in the normal differences between sexes in the genus Xylophanes , i.e. larger wingspan, broader and more rounded wings, and thinner antennae.

Male genitalia ( Figs. 12 a–c). Uncus stout; setigerous lobes developed and distinctly protruding; distal projection bent ventrally, its apex is slightly spatulate and truncate. Harpe short, thick basally and tapering progressively into a thin and slightly upcurved apex. The latter, together with the internal margin of the harpe, only slightly sclerotized. Setae on the harpe few, scattered, and present mostly on the ventral side. Right lobe of apical process of the aedeagus very stout, with long and uneven teeth grouped in the apical portion of its internal margin; left lobe barely distinct, bearing about 10 minute teeth on its lateral part.

Immature stages: A large number of rearing records for this species are reported on the ACG caterpillar inventory website (http://janzen.sas.upenn.edu/); when writing this article, more than 100 pictures of the caterpillar (penultimate and ultimate instars) and pupae of Xylophanes cthulhu were displayed on this site along with detailed collecting information, including food plant identifications. X. cthulhu is reported to feed exclusively on Rubiaceae of the genus Spermacoce L. (52 records on S. exilis , 6 on S. ocymifolia and 1 on S. remota ).

Parasitoids: From the numerous rearing records of X. cthulhu in ACG, this species is reported as the host of the parasitoid wasps, Cryptophion inaequalipes ( Hymenoptera : Ichneumonidae, Campopleginae. The reported parasitoid actually represents a species complex and a provisional name, C. inaequalipes DHJ02, is currently attached to the specimens reared out of X. cthulhu caterpillars) and Charmedia chavarriai ( Ichneumonidae, Ichneumoninae ), as well as the tachinid fly, Drino incompta ( Diptera : Tachinidae ).

Distribution. This species is widely distributed in Central America, inhabiting low to medium altitude forest areas from southern Mexico (Veracruz, Chiapas and Oaxaca states) to eastern Panama ( Panama province). A possible contact zone with X. neoptolemus should be searched for in northern Colombia, where specimens of both species might be encountered.

Genetic variation: The lack of genetic variation of this species is remarkable, with a single haplotype found throughout the range from Mexico (Chiapas) to Panama ( Fig. 13 , group 3).

Figures

FIGURE 13. Strict consensus of the nine equally most-parsimonious cladograms (length = 133, CI = 0.78, RI = 0.94) resulting from the phylogenetic analysis of the complete dataset of DNA barcode sequences for 38 specimens belonging to the Xylophanes neoptolemus (yellow) and X. loelia (blue) species complexes (X. cyrene, X. aglaor and X. libya are outgroup taxa). Each specimen is identified by its SampleID code (see Table 1), and the two specimens with short sequences are highlighted in boxes. The branch lengths are proportional to the number of changes (indicated on branches, optimized under FAST optimization); Bremer support and rescaled Bremer support values are given above the branches for each node, and bootstrap support values are indicated below. Clades within the X. neoptolemus complex are named after groups 1, 2 and 3 as described in the text.

FIGURES 1 – 4. 1 a. Chaerocampa trilineata Walker, [1865], Lectotype, ♂, Venezuela; 1 b. Watercolour of the original type of Sphinx neoptolemus Cramer, 1780; 2 a and 2 b. Xylophanes balcazari n. sp., Holotype, ♂, Mexico, Guerrero; 3 a and 3 b. Sphinx neoptolemus, Neotype, ♂, Venezuela, Aragua; 4 a and 4 b. Xylophanes cthulhu n. sp., Holotype, ♂, Guatemala, Izabal. (2 a, 3 a, 4 a: dorsal view; 2 b, 3 b, 4 b: ventral view).

FIGURES 8 – 12. Drawings of diagnostic details of the male genitalia of Xylophanes loelia (Figs. 8 a – c), X. lolita n. sp. (Figs. 9 a – c), X. neoptolemus (Figs. 10 a – c), X. balcazari n. sp. (Figs. 11 a – c), and X. cthulhu n. sp. (Figs. 12 a – c); a. uncus in lateral view, b. harpe, c. posterior end of the aedeagus.