Heraclides ponceana (Schaus, 1911)

Zhang, Jing, Cong, Qian, Shen, Jinhui, Opler, Paul A. & Grishin, Nick V., 2020, Genomic evidence suggests further changes of butterfly names, The Taxonomic Report of the International Lepidoptera Survey 8 (7), pp. 1-41 : 6-8

Heraclides ponceana (Schaus, 1911) is a species distinct from Heraclides aristodemus (Esper, 1794)

Papilio temenes Godart, 1819 is a junior subjective synonym of Heraclides aristodemus (Esper, 1794)

The short version of the original description of Papilio temenes Godart, 1819 (page 21) is: "Dessus des ailes d'un brun-noirâtre, avec deux bandes jaunes, maculaires, disposées en sautoir sur les supérieures: les inférieures en queue: le dessous de celles-ci jaunâtre, avec une bande bleue, flexueuse, sur le milieu" ( Godart 1819). It can be translated word-for-word as: "Above the wings of a blackish-brown, with two bands yellow, macular, arranged in sautoir [=crosswise] on the forewings: the hindwings [end] in tail: the underside of these [hindwings, not tails] yellowish, with a band blue, flexuous, in the middle", and interpreted as: "Wings above blackish-brown, forewings with two yellow macular bands that cross each other, hindwings with tails, underside of hindwings yellowish with a blue flexuous band in the middle." Perhaps the most significant word here is " sautoir " defined in Dictionary.com as "A ribbon, chain, scarf, or the like, tied around the neck in such a manner that the ends cross over each other" (Dictionary.com LLC 2020), and by Merriam-Webster dictionary as "a chain, ribbon, or scarf worn about the neck with the ends forming a St. Andrew's cross in front" (Merriam-Webster 2020). It is important because this character (crosswise vs. more parallel arrangement of yellow forewing bands) is diagnostic of H. aristodemus (crosswise) vs. H. ponceana (more parallel, images in Warren et al. (2016)). Today, the name temenes is applied to a broad-banded form of H. ponceana that flies in Cuba (see below). Here we argue that this is a misidentification, and the Godart's temenes is H. aristodemus instead.

We see that, although the Godart's description of P. temenes is brief, two of its details agree better with H. aristodemus than with the broad-banded subspecies of H. ponceana from Cuba: crisscrossing bands and blue flexuous band. In the broad-banded subspecies, the outer band is more parallel to the inner band and does not give an impression of crossing it. And the blue spots on the hindwing underside look more like a row of lunules than an irregular blue band. In H. aristodemus , forewing bands indeed give an impression of crossing bands, and the blue spots on the hindwing below look like an irregular blue band. Furthermore, an extended description of P. temenes on page 63 states that the forewing bands are narrow and macular ("étroites, maculaires") ( Godart 1819), instead of being broad and continuous. Besides this additional detail, the extended description reiterates other points of the short description. We note that some H. aristodemus females may have broader and more continuous yellow bands that are more similar to those in H. ponceana ( Fig. 2e), but the P. temenes description does not mention this possibility, simplifying the application of the name.

Next, we inspected all potential syntypes of P. temenes , two specimens in Paris, France ( Fig. 2 ab) and one (only photograph inspected) in Edinburg, UK ( Bland 2019). These 3 specimens closely agree with the original description of P. temenes , carry historical labels and labels indicating their type status ("type", "co-type", or "?co-type") and therefore are likely to be true syntypes of this taxon. These specimens phenotypically are H. aristodemus and not H. ponceana , differing from the broad-banded subspecies from Cuba. Furthermore, we sequenced two syntypes in Paris (one labeled "TYPE", the other labeled "CO-TYPE") and they are H. aristodemus by genomic DNA (Fig. 1 blue), in agreement with their wing patterns. In conclusion, our analysis of the original description and the likely syntypes leaves little doubt about the identity of P. temenes as H. aristodemus , which is a species different from the broad-banded H. ponceana " temenes " found on Cuba today (Fig. 1 red).

Due to misapplication of the name temenes to a taxon different from that in the original description and for the future stability of the name, we designate the syntype in the Muséum National d'Histoire Naturelle, Paris, France (MNHP) shown in Fig. 2a and possessing all the characters stated in the original description, with the following 4 rectangular white (some faded to brownish) labels: printed red || TYPE ||; framed, lined and printed with green, first line handwritten in black || Anc. Collection | MUSÉUM DE PARIS ||; printed, with a square barcode on the right side || MNHN, Paris | EL63126 ||; printed || DNA sample ID: | NVG-18079F07 | c/o Nick V. Grishin ||, the lectotype of Papilio temenes Godart, 1819 . The red, rectangular, printed label || LECTOTYPE ♂ | Papilio temenes | Godart, 1819 | designated by Grishin || will be added to this specimen. This specimen was chosen as the lectotype because it is labeled "type" rather than "co-type", and it yielded genomic dataset of a good quality for a specimen that old.

It appears that Oberthür (1897) was the first to incorrectly apply the name P. temenes to the broad-banded subspecies of H. ponceana from Cuba, inconsistently with the original description and now with the identity of the lectotype of P. temenes . Oberthür illustrated one such specimen. While we have not investigated the reasons behind Oberthür's mistake, we note that it has been followed in subsequent literature. Interestingly, we found 3 century-old H. aristodemus (i.e. true P. temenes ) specimens labeled " Cuba ", in ZMHB and FMNH ( Fig. 2c–e). Further studies may answer the question whether both species ( H. aristodemus and H. ponceana ) co-occurred in Cuba, or the old records from Cuba were mislabeled.

In summary, we conclude that Papilio temenes Godart, 1819 , syn. n., is a junior subjective synonym of Heraclides aristodemus (Esper, 1794) . The type locality of temenes should remain as stated on page 63 ( Godart 1819): "Antilles & dans l'Amérique septentrionale", i.e., "West Indies and North America", which is not necessarily Cuba. It remains to be investigated whether H. aristodemus has been found or still occurs in Cuba. In any case, as detailed above, the broad-banded subspecies of H. ponceana from Cuba does not have a name, and it is described here as new.