Heraclides

Figures 7–10.Heraclides rumiko type specimens: 7–8 holotype ♂ 9–10 paratype ♀ NVG-2564, data in text and Supplementary Table 1. Dorsal/ventral surfaces are in odd/even-numbered figures. Labels are shown between the images of the same specimen, exuvia and head capsules in a gelatin capsule are below, and 3-fold magnified segment of head, neck and thorax is on the left. All images are to scale (including labels), except the magnified insets.

Heraclides cf. hyppason (Hyppason Swallowtail) at Canopy Tower, Panama. Photographed on 5 April 2002www.inaturalist.org/observations/54478291

Pouso Alegre, Transpantaneira, Pocon, Mato Grosso, BRAZIL

One of a pair seen at a planter box in Picton, Ontario. They are an energetic butterfly and rarely settled for a photo.

Dorsal..

Figure 16.COI DNA barcode trees. Trees of representative sequences of Papilionini reconstructed with different methods: a Bayesian inference using MrBayes (alignment partitioned by codon position, nst=6, rates=invgamma, ratepr=variable), posterior probabilities are indicated by the nodes b Maximum likelihood method RAxML (-m GTRGAMMA), bootstrap values are indicated. Posterior probabilities are shown by the nodes (omitted within species). Names of different species are shown in different colors. Sequences obtained in this work are those with “NVG-” number (see Supplementary Table 1 for complete data), others are from GenBank (http://genbank.gov/) and are labeled by accessions (letters and numbers, no dashes). Specimens with sequences from GenBank were not examined (except where a photograph was available from the BOLD database) and their identification follows original work, locality, and DNA barcode.

Charcos, Puntarenas, Costa Rica

Figure 18.Localities of Heraclides cresphontes and Heraclides rumiko specimens with available DNA barcode information. Color of circles corresponds to species: Heraclides cresphontes - blue (based on 112 DNA COI barcode sequences, 103 obtained in this work); Heraclides rumiko - red (based on 183 barcodes, 146 obtained in this work), split red/blue circles mark localities where both Heraclides cresphontes and Heraclides rumiko were recorded. Type localities for taxa with available names are indicated with a corresponding name followed by “TL”. We treat Papilio cresphontes var. maxwelli Franck, 1919 & Papilio cresphontes pennsylvanicus F. Chermock & R. Chermock, 1945 as junior subjective synonyms of Heraclides cresphontes. Countries and states (for USA and Mexico) with records are labeled.

Local: Araatuba/SP - BrasilParque da Fazenda do EstadoBioma: Mata AtlnticaAcessrios: Trip + Trilho Foco Macro + Flash Circular Macro + Cabo Disparador Remoto

Charcos, Puntarenas, Costa Rica

Figure 24.Foodplants most commonly used by Heraclides caterpillars. a–c Zanthoxylum fagara [L.] Sarg (Colima, Lime Prickly-ash), USA: TX: Duval Co., Benavides, 19-Apr-2014, used by Heraclides rumiko A–C Zanthoxylum clava-herculis L. (Hercules’s club, Pepperwood), USA: TX: Denton Co., Grapevine Lake, Murrell Park, 26-Apr-2014, used by Heraclides cresphontes.

Figure 20.Life history: 2nd and 3rd instar caterpillars. a–r Heraclides rumiko, USA: TX: Duval Co., Benavides A–K Heraclides cresphontes, USA: TX: Denton Co., Grapevine Lake, Murrell Park a–k, A–E 2nd and k–r, F–K 3rd instar caterpillars; 0.5 cm scale shown on panels m and K refers to all images. Sexes and voucher numbers (where available) and dates: a–c paratype ♀ NVG-2564, 25-Apr-2014 d–f paratype ♀ NVG-2563, 25-Apr-2014 g–i larva #7 (died), 29-Apr-2014 j–k paratype ♂ NVG-2559, 25-Apr-2014 l–o paratype ♂ NVG-2559, 27-Apr-2014 p–r paratype ♀ NVG-2564, 29-Apr-2014 A–B, F–K ♂ NVG-2760, 22-Jun-2014 (A–B), 24-Jun-2014 (F–H), 26-Jun-2014 (I–K) C–E ♀ NVG-2741, 16-Jun-2014.

Figure 21.Life history: 4th and 5th instar caterpillars. a–n Heraclides rumiko, USA: TX: Duval Co., Benavides A–K Heraclides cresphontes, USA: TX: Denton Co., Grapevine Lake, Murrell Park a–e, A–F 4th and f–n, G–K 5th instar caterpillars; 1 cm scale shown on panels c and I refers to all images. Sexes and voucher numbers (where available) and dates: a–b, f–n is the same individual (larva #2, died, shown in Fig. 22a–c), 24-Apr-2014 (a–b), 29-Apr-2014 (f–j), 29-Apr-2014 (k–n) c–e larva #1 (died), 24-Apr-2014 A–C ♂ NVG-2740, 16-Jun-2014 D–F ♀ NVG-2741, 21-Jun-2014 G–K ♀ NVG-2741, 26-Jun-2014.

Figure 22.Life history: 5th instar caterpillars and prepupae. a–j Heraclides rumiko, USA: TX: Duval Co., Benavides A–K Heraclides cresphontes, USA: TX: Denton Co., Grapevine Lake, Murrell Park a–h, A–J 5th instar caterpillars and i–j, K prepupae; 1 cm scale shown on panels a and K refers to all images. Sexes and voucher numbers (where available) and dates: a–c larva #2, died (shown in Fig. 21a–b, f–n), 1-May-2013 d–h paratype ♀ NVG-2564, 11-May-2014 (adult Figs 9, 10) i–j paratype ♂ NVG-2559, 11-May-2014 A–E 26-Jun-2014 F–K ♂ NVG-2740, 21-Jun-2014 (F–J) & 23-Jun-2014 (K).

Figure 11.Neck pattern, male genitalia, and morphometrics. a–d Heraclides rumiko, paratype, Mexico: Baja California Sur: Buena Vista, 1-Oct-1981, leg. D. Faulkner & F. Andrews, genitalia KS017 [SDMC] A–D Heraclides cresphontes, USA: Georgia: Clark Co. July 2009, genitalia KS009 E Morphometric measurements performed on genitalia and facies and plotted in two dimensions. Horizontal axis is a weighted average of the three genitalic measures: 0.6*”U-B Angle” + 0.2* (B−A)/(C−B) + 0.2*G/H. “U-B Angle” is measured in radians. Vertical axis is a weighted average of the four facies measures: 0.15*”YL” + 0.4*F/P + 0.4*T/V + 0.05 W/D, where “YL” is equal to 0 or 1, if yellow line on the neck is separated into spots or continuous, respectively. Measured distances are indicated on the illustrations. Each of the two (genitalic and facies) linear combinations of measures completely segregates Heraclides rumiko (red points) from Heraclides cresphontes (blue points) specimens (not from central Texas) with a hiatus. Even a single measure “U-B Angle” identifies all specimens correctly, except #12, which has a brachium strongly curved dorsad. Specimen localities: Heraclides cresphontes: 1. GA: Clark Co.; 2. NY: Niagra Co., Lockport; 3. NC: Carteret Co.; 4. IN: Kosciusko Co., Silver Lake; 5. WI: Sauk Co., Sauk City; 6. LA: St. John Pa., Edgard; 7. AR: Osceola; 8. OK: Marshall Co., Lake Texoma; 9. FL: Okeechobee Co., Fort Drum; 10. OH: Montgomery Co., Dayton; 11. PA: York Co., Pinchot State Park. Heraclides rumiko: 12. AZ: Maricopa Co., North Phoenix; 13. AZ: Santa Cruz Co., Sycamore Canyon; 14. CA: Imperial Co.; 15. MX: Veracruz, Fortin de las Flores; 16. MX: Oaxaca, Yangul; 17. Costa Rica: Puntarenas, San Antonio; 18. MX: Tamaulipas, Gomez Farias; 19. MX: Colima, Colima; 20. MX: Sonora; 21. MX: Yukatan, Merida; 22. MX: Morelos, Rancho Viejo; 23. MX: BCS, Buena Vista; 24. MX: Jalisco, Ajajic; 25. CA: San Diego Co., La Jolla; 26. MX: Quintana Roo, nr. X-Can; 27. TX: Val Verde Co., Del Rio; 28. CA: San Diego Co., Pauma Valley. Central Texas specimens are from Bexar (33–38, 42–48), Williamson (39–41), Travis (31, 32), Bastrop (30), and Brazos (29) Counties. Voucher codes for these specimens are: 29. NVG-2236; 30. -2299; 31. -2300; 32. -2174; 33. -2192; 34. -2196; 35. -2205; 36. -2209; 37. -2210; 38. -2216; 39. -2301; 40. -2225; 41. -2229; 42. -2191; 43. -2197; 44. -2204; 45. -2208; 46. -2211; 47. -2215; 48. -2218. Species (color on the plot) is assigned to central Texas specimens by COI barcode. See Supplementary Table 1 for more data. Specimens 36 and 41 are apparent hybrids or the results of introgression.

Figure 12.Female genitalia. a–i Heraclides rumiko paratypes [TAMU]: a USA: TX: Cameron Co., Las Palomas WMA, Tucker Unit, 24-Oct-2001, leg. J. & F. Preston, DNA voucher NVG-2238, genitalia NVG140320-79 b, c USA: TX: Hidalgo Co., Santa Ana National Wildlife Refuge, 13-Oct-1968, leg. R. O. Kendall & C. A. Kendall, NVG-2163 | NVG140320-04 d–f USA: TX: Cameron Co., World Wildlife Management Area nr. Santa Maria, 14-Nov-1971, leg. R. O. Kendall & C. A. Kendall, NVG-2195 | NVG140320-36 g Honduras: Escuela Agrícola Panamericana, 30 km SE Tegucigalpa, 1-May-1985, leg. Vascones, NVG-2221 | NVG140320-62 h Mexico: Durango: Tlahualilo, 20-Aug-1935, leg. C. S. Rude, NVG-2230 | NVG140320-71; i Mexico: Tamaulipas: Cd. Monte, Los Arcos Ct., 8-May-1978, leg. R. O. Kendall & C. A. Kendall, NVG-2185 | NVG140320-26 A–D Heraclides cresphontes, USA: MO: A, B Phelps Co., Mark Twain National Forest, DeWitt Pond, N37.8367 W91.9385, 25-May-2006, J. C. Abbott, NVG-2293 | NVG140403-21 [TMMC] C, D Montgomery Co., NVG-2242 | NVG140320-83 [TAMU]. Ventrolateral view is shown in c, e, B, D (e is left-right inverted), others are in ventral view. All images are to scale shown under a, except f, which is half the size with scale shown to the right.

Ventral..

Figure 13.Facies differences between Heraclides rumiko (left, r) Heraclides cresphontes (right, c) indicated by red triangles and lines. These differences are as follows. 1) Dark spot on forewing: (r) almost always large; (c) variable, but often weak and sometimes absent 2) Forewing margin: (r) often straight with smaller or absent marginal spots; (c) strongly scalloped with yellow marginal spots at dips between veins 3) Forewing submarginal yellow spots: (r) smaller rarely more than three; (c) frequently larger, more than three 4) Thorax with: (r) yellow line running from head through patagia to tegulae; (c) spots instead of the line, or just few yellow scales. 5) Abdomen: (r) usually with a fainter dark band; (c) often with solid dark band 6) Inner edge of black discal band on ventral hindwing: (r) mostly straight; (c) usually curved 7) Tail: (r) mostly narrow and relatively longer; (c) typically rounder and wide, shorter. Heraclides rumiko is usually smaller than Heraclides cresphontes, despite being a southern taxon. Due to significant seasonal and individual variation, none of these characters is fully reliable and exceptions exist. The head-neck-thorax line vs. spots (Fig. 11a, A) might be the strongest single character. A combination of characters should be used for reliable identification, e.g., the one shown in Fig. 11E. Many specimens in central Texas exhibit intermediate characters, atypical character combinations, and possible hybrids can be found (Fig. 11E).

Figure 14.Variation in male genitalia. Left lateral view of genital ring (uncus, brachium, dorsolateral sclerite, tegumen, vinculum and saccus) is shown, valvae, aedeagus and last tergum with pseuduncus are removed. Heraclides cresphontes and Heraclides rumiko localities are shown in blue circles and red disks, respectively.