This week's butterfly is another dark-winged, red-bodied swallowtail, Atrophaneura (formerly Papilio) polydorus. In the eighteenth and nineteenth century, naturalists established a tradition, no longer followed, of naming these butterflies after literary characters associated with funerals, on account of their "funereal looking" black or faded-black wings. Several royal and influential characters in ancient Greek literature were called Polydorus. One of the best known was a prince of Troy who was murdered for the gold he was carrying.
Photo from J. Michel.
These butterflies are found in Australia With their large size and high-contrast color scheme, they're hard to overlook. Australians don't see them as funereal. They see them as their Red-Bodied Swallowtails, as the Red-Bodied Swallowtails. They have posted and published a great deal of material about these butterflies. In literature from most of the world "red-bodied swallowtail" includes all the butterflies we've been studying for most of this year, but in Australian literature it means polydorus.
Males and females look pretty much alike; the red parts of their bodies and wings are much more visible on the underside. From above they seem to have black bodies with red tails (tufts of hair on the tail end of the body). The hind wings look as if these butterflies had tried to grow real "swallowtails" but not succeeded. Most writers agree that their stumpy little efforts in that direction don't count as tails, and describe the hind wings as tailless.
Photo by Martin Purvis. This one is female; females' fore wings tend to look a little less iridescent, more faded, than males', but not so consistently as in other species in the genus.
Sizes vary both among and within subspecies. The Auckland Museum's cases filled with dead specimens show individuals within a subspecies almost twice the size of others. The size range seems comparable to that of North American swallowtails, most about three inches. Getting this individual to perch on a ruler would have been more useful than getting it to perch on the man's hand:
Selfie by Fieldnotesfromthefarnorth, who mentions having snapped it in July.
Like the other red-bodied swallowtails, they can move their fore and hind wings independently, which may slightly offset the butterflies' disadvantage relative to predators in flight. While resting or eating they can keep the fore wings in a blur of motion while the hind wings might look like someone a hungry bird would prefer not to see more closely...
So much information is out there that, even though the different web sites do repeat the same basic facts, this post formed subsections. First it lists the tiny but consistent variations that define the subspecies, then it considers the life cycle of the butterfly.
Many Subspecies, Just Recognizably Different
Twenty-eight subspecies of Atrophaneura (now more often Pachliopta) polydorus are generally recognized today; more have been named. Web sites are devoted to some of these subspecies, but they seem to be either museum pages or placeholders waiting to be filled in with data from active naturalists. The subspecies godartianus, leodamas, and polydaemon were first counted as separate species. These butterflies live on several islands, and populations from different islands show different features. Naturally, some of these subspecies are better documented than others.
Although some American swallowtails eat a vine in the genus Aristolochia and have that weird, which-side-are-the-legs-on shape with which so many Atrophaneura caterpillars discourage birds, they can't live on the same vines. The Asian red-bodied swallowtails are thought to have more DNA in common with the very different looking Birdwings than they have in common with our swallowtails. Here's a detailed DNA study of the Atrophaneuras and their neighbors:
Here's the list of subspecies with the places where they are found. Where early descriptions have been published online in English they're included, although the transcription is not perfect. Nineteenth century scientists used special characters for "male," "female," "degrees," and other terms, which computer transcriptions misread in a variety of amusing ways. If you can read German, Danish, Dutch, and French well enough to correct errors produced when computers try to read text from photographs of old, sometimes badly printed or damaged, books, you'll find links at Funet.fi to some descriptions not included here.
1. Pachliopta or Atrophaneura polydorus aigmanus, Louisiade
Described in 1893, when Walter Rothschild was a mere "Hon.," the British "Honourable" that has nothing to do with a person's honor, but merely identifies a junior member of an aristocratic family:
"Comes very close to the form from the Key Islands (thessalia), and has like this a red head: but...the red hairs of the front of the head being much mixed with black ones, and in the posterior red spots of the upperside of the hindwing being more dusted over with black scales. From the Queensland form, which has the head similar in colour, it differs in the white patches in the cell of the hindwing and between veins 5 and 6 being as small as in thessalia. The forms of polydorus from New Guinea, the L)"Entrecasteaux Islands, the Bismarck Archipelago, and the Solomon Islands have black heads.
Hab. St. Aignan, Louisiade Archipelago, August to November 1897 "
Hind wings can be described as tailed, and the black color can look iridescent blue.
2. P.p. alboplagatus, Sula
Rothschild, 1924: "Allied to the typical form from the S. Moluccas.
Distinguished from the Moluccan form by the much larger white areas on both wings. Fore wing with white markings more distinct, the patches below vein 3 larger and purer white, the one in 2 just filling up the base of the cellule. Hind wing with larger discal patch, the spots closer together, and veins separating them very thinly scaled with black. Hed submarginal spots placed nearer the margin, and smaller. Sula Islands, June, July and September, W. J. C. Frost, 2 9? °."
3. P.p. aphnitis, Goodenough.
4. P.p. asinius, east New Guinea
5. P.p. auster, New Guinea
6. P.p. ceramites, Kapaur
7. P.p. dampierensis, Dampier/Karkar
8. P.p. godartianus, Artak and Onin
Walter Rothschild's remarks on godartianus reflect the confusion about it, whether it was a separate species, and whether things classified as godartianus would later be classified as additional subspecies: "Though I have about forty-five specimens of poli/dorus from (rermaii and Dutch New Guinea, Waigeu, and Salwatty, and some hundreds from the D'Entrecasteaux Islands, now before me, I cannot decide at present whether we have here one or more local races. The typical godartianus Lucas, of which leodamas Wall, is a synonym, has dark forewings, without a white patch on either side; the white adnervular streaks, though sometimes well marked below, are always narrow, and separated from each other by means of black internervular streaks which run from the outer margin of the wing to the black basal area; the whitish streak behind the lower median nervule on the underside is especially remarkably small. The white patch on the • hindwings has the cellular spot small, often smaller than that discal spot which stands before the upper median nervule; the first discal spot is small, longer than broad, often absent ; the second mark has a length of about 3 mm., and is mostly of an oblong shape, owing to the upper median and lower discoidal veins being heavilv bordered with black. Such specimens are figured by Lucas as godartianus and by Wallace as leodamas. In many individuals of godartianus the cellular and the anterior discal spots on the hindwings are much shaded with black...
Typical godartrianus I know only from the N.W. Peninsula of New Guinea (Arfak, Dorey)...and from Mysol and Salvatty.
At Humboldt Bay and all over German New Guinea there occurs a form which is remarkable for some characters: the upperside has a peculiar bluish gloss in certain lights, which is more obvious than in godartianus ; the black internervular streaks on the upperside of the forewings are not strongly pronounced, which gives the wing a peculiar soft appearance ; the white streaks in the postcellular region, though often but feebly marked above, are always large below, and form in most .specimens a large white patch, which is not interrupted by complete internervular black streaks; in my darkest specimen the white is more extended than in my...example of godartianus. On the hindwings above only the last two submarginal spots are marked or indicated ; there is no trace of the other spots, while in most specimens of godartianus, in all specimens of polydorus, queenslandicus, etc., the other submarginal spots are indicated at least by a paler brown scaling. The cellular white mark is very variable, occupying almost half the cell, or being reduced to a small spot ; the upper discal spot is large, mostly rliomboidal, often notched, seldom reduced to a small streak; the other spots are also larger than in typical godartianus, especially the posterior one; the nervules traversing the white patch are very thinly black.
From Waigeu I have six specimens which agree with the individuals from German New Guinea, except in the black internervular streaks on the upperside of the forewings being more prominent, in the submarginal spots to the hindwings being all indicated at least bv a hrown scaling, and in the first discal spot being longer and narrower. In these characters, the Waigen individuals agree better with the Halmaheran race of polydorus than with the above-described New Guinea form. The close relationship of the Waigeu polydorus with those from German New Guinea renders it impossible for me to treat the Waigeu and German New Guinea specimens as a subspecies distinct from godartianus, which occupies the interjacent country, and I think that the differences of the forms present themselves to us at present thus, as explained above, because om- knowledge of the fauna of those districts is still very insufficient. To draw, however, the attention of the explorers to the present question, I propose to call the form from German New Guinea (type from Stephansort) — ab. plagiatus...from the island of Fergusson, D'Entrecasteaux group...A. S. Meek sent a large number of specimens, which are all more or less different from either plagiatus or typical godartianus, and agree fairly well with the Waigeu polydorus, all of them having a white patch on the forewings, at least on the underside, the first discal ■white spot of the hindwings elongate (sometimes obliterated), and the submarginal red spots all indicated on the upperside, as in Waigeu specimens. As Mr. A. S. Meek will explore some more of the islands lying east of New Guinea in the course of this year, it is best to treat this Fergusson polydorus for the present as godartianus and plagatius.
Hah. Salvatty (1 ?); Mysol ; Waigeu (2 c?, 4 ?); N.W. Peninsula of New Guinea (8 J, 7 ?); Humboldt Bay and German New Guinea (11 i, 14 ?); Fergusson Island (a long series).
Note. — The Fergusson specimens are somewhat different in scaling from the New Guinea and Waigeu examples; the white scales, for example, between the lower median veins on the underside of the forewings have much longer and sharper teeth than in typical godartianus, ab. plagiatus, polydorus, etc. In all races, the upper scales are bi-, the under scales tridentate on both sides of the forewings. In some specimens from German New Guinea there appear a few red hairs behind the eyes.
(g) : p. polydorus novobritannicus sulis].. nov. [cr,?,pupa]."
9. P.p. humbldti, Humboldt Bay, New Guinea
Rothschild, 1903: "A transitional form, being intermediate between P. p. godartianus Gner. (1852) from Dutch New Guinea and plagiatus Rothsch. (1895) from German New Guinea. The greyish stripes on the upperside of the forewing are narrow and dark ; the black vein-stripes broad, extending to the cell, the black internervular streaks reaching close to it ; beneath the pale stripes are nearly white, those situated near the second median vein being widened to form patches. The white cell-spot of the hindwing smaller than the first discal spot, often very small, the first discal spot rhombiform, at least beneath.
Hab. Humboldt Bay, Dutch New Guinea, at the border of German New Guinea."
10. P.p. kajelanus, Buru
In 1864 A.R. Wallace wrote, considering leodamas as a distinct species, "Male. Above, glossy black, upper wings immaculate (the veins pale-margined in the female). Lower wings with a rounded white spot divided into six parts by fine nervures, of which the outermost and that in the cell are sometimes reduced to points ; marginal row of red spots obscured with black, and but faintly indicated.
Beneath, the white patch has a small red spot attached to the part nest the anal angle ; and the marginal row of sis red spots are clearly marked, that at the anal angle being twice the size of the rest. Wings short, much rounded, scarcely or not at all produced in the caudal region.
Expanse of wings 3 in. to 4 in.
Hab. New Guinea, Mysol (c?, ?) {Wall), Rockingham Bay (Australia), {Brit. Mus., ?)."
11. P.p. lascarus, northeastern New Guinea
12. P.p. leodamas, Misool
13. P.p. manus, Admiralty
14. P.p. meforanus, Mefor or Noemfoor, Geelvink or Geelwink Bay
Rothschild, 1903: "Forewing, above, with narrow streaks in front of the second median vein and behind it, the streaks being dark grey in c? and white in ? ; on underside these streaks widened in both sexes to form white patches, the outer light streaks along the veins also being more distinct beneath than above. The white cellspot of the hindwing about thrice as large as the first discal spot, the latter being small, the fourth discal spot anteriorly less than twice as long as it is broad distally.
Hab. Mefor, Geelvink Bay (W. Doherty)."
15. P.p. naissus, Trobriand, Papua low country, D'Entrecasteaux
16. P.p. novobritannicus, New Britain, New Ireland, Duke of York
Rothschild, 1895: "Head and thorax black, but the front of the head, and the sides of pronotum and of mesosternum, clothed with more or less obvious red hairs, besides the lilac ones. Abdomen much more extended red than in godartianus.
Forewings : the white stripes well marked...which in all the other subspecies are rather obscure. The black internervular .streaks prominent; that between the lower median veins abbreviated, the following one very short.
The submarginal spots to the hindwings are all visible above; the anterior one, standing behind the costa, is visibly red; the posterior ones are also red, but more shaded with black than in godartianus. The cellular white spot is large, reaching anteriorly at least as far as the origin of the first discoidal nervule ; the discal markings are short, but broad, the nervules not being heavily black; the first is more or less trapeziform, often broader than long, and oblique ; its length varies from 2 to 4 in.; the next two are much smaller; the fourth is considerably shorter than in the other races, and as broad as in godartianus; its average length is about 4 mm. ; the fifth is more or less rhomboidal, mostly somewhat longer than broad, shorter than in godartianus.
The specimens from New Ireland and Duke of York Island form partly a transition to the next subspecies, as their forewings are darker above than in the New Britain novobritannicus m., and the discal spots to the hindwings more rounded.
The pupa closely resembles that of P. aristolochiae Falw. ; the abdominal tubercles are rather higher than in that species.
Hab. New Britain (type; 2 J, .5 ?) ; New Ireland; Duke of York Island"
17. P.p. orinomus, Papua hill country
Rothschild: "Similar to P. p. aignanus Rothsch. (1898), the white cell-spot of the hindwing much larger than the second white discal spot, the fourth discal spot larger than the third, and the fifth submarginal spot smaller than in aignanus.
Hab. Upper Aroa River, British New Guinea (A. S. Meek)."
18. P.p. polydaemon, north Solomon Islands, Bougainville, Shortland
Rothschild found polydaemon the most distinctive of these subspecies, writing from the presupposition that species evolve in the direction of distinctiveness: "I cannot see that the specimens from various parts of the Solomon Islands belong to more than one race. The size and shape of the spots to the hindwings are rather variable in every locality ; the nervules separating the white markings are always verv broadlv black. The posterior black internervular streaks on the underside of the forewings are sometimes complete, extending from the brown border of the wing to the basal brown area.
Hah. Solomon Islands : Ugi ; Guadalcanar (4 J, 5 ¥ ) ; Alu (3 cJ, 4 ? ) ; Treaury Island. This is the most advanced form of polydorus, which I should treat as a species, if it were not for the Duke of York specimens of P. polydorus novobritannicus m. and for P. polydorus septentrionalis."
19. P.p. polydorus, Ambon, Serang, Obi
20. P.p. queenslandicus, Queensland, Torres Strait
Walter Rothschild compared this super-popular subspecies with the basic Papilio polydorus polydorus:
"Size rather smaller ; forewings more rounded. Cellular spot to the hindwings as in polydorus; the three first discal spots longer, the first also narrower; the two posterior ones as in polydorus. Below, there is an additional white mark between the upper discoidal and the subcostal veins, which is sometimes also marked above. The veins bordered with black as in polydorus. Head, thorax, and abdomen as in polydorus.
In my Thursday Island specimens the white colour on the forewings is rather restricted, and in one individual there is, below, a white transverse spot behind the costa of the hindwings at the basal side of the first submarginal red mark.
P.p. tenimberensis m. this subspecies differs especially in the much less extended white and rounder forewings, in the shorter discal spots to the hindwings, of which the first is broader, and in the presence of an additional white discal spot before the first discoidal vein.
From thessalia (Swinh.) it is distinguishable by the rounder forewings again, the much larger cellular spot to the hindwings, the larger and differently shaped first discal spot, the shorter second and fourth spots, and larger fifth one. Hab. Queensland (3 <?, 3 ?);"
Though grammatically feminine Latin nouns usually end in -a, some scientists feel that the subspecies should be queenslandica.
The forewings can look black, iridescent, drab, or black with white spots. When that special Atrophaneura, black-showing veins/nerves, look is present polydorus is likely to show paler patches in a way that really highlights the swallowtail wing structure. Three inches is a typical wingspan.
Photo by Terrymahney.
The caterpillar may be a little darker than some polydorus caterpillars, or it can be lighter.
Photo by Katina Gee. The thickened backs of swallowtail caterpillars accommodate the osmeterium, which resembles a little forked tongue and discourages birds with the suggestion that a caterpillar may be a bird-eating snake or lizard. Making the osmeterium visible is a stress reaction some caterpillars show easily and often, others rarely. This caterpillar's false eye spots, together with the placement of its osmeterium (not extended) and tubercles, suggests that it may be mimicking a lizard lying on its back the better to grab birds.
This photo, and the one of the greenish pupa, by Hugo Innes.
The pupa can look greenish, rather than the usual pale brown. (Most polydorus pupae are pale brown.)
21. P.p. schoutensis, Schouten
1924: "Intermediate between the Waigeu and Mefor races.
Male. Upperside of fore wing with more grey scaling than in either of the allied forms. Hind wing with the outer edges of the discal spots less sharply defined than in the other forms; red anal spots indistinct as in the Waigeu form.
Underside of hind wing with a small white spot in 6 placed on the edge of the red spot, and some slight white scaling on the edge of the red spot in 7; in one specimen there are only faint traces of these white marks.
Female. . Resembles the Waigeu form. Hind wing above with outer edge of discal spots less sharply defined than in the allied form, being bordered by a pale shade. Below with some white scaling on the red spots in 6 and 7. Discal spot in 5 larger than in the Mefor race and not so long as in the Waigeu form.
Habitat.—Schouten Islands, N.E. New Guinea: Biak, June, 1914..."
22. P.p. septentrionalis, Bachan, Halmahera
Rothschild's description: "Head and thorax entirely black. Larger than godartianus; upperside less deeply black, with a much feebler bluish gloss. Forewings longer, with the black internervular strijies more prominent than in typical godartianus, the posterior white stripes less marked than in polydorus...but more than in typical godartianus.
The submarginal spots to the hindwings are all visible above; the white cellular spot is about as large as in typical godartinmw, or a little larger; the discal spots are all rounded exteriorly; the first and last are triangular; the third, fourth, and fifth are shorter than in godartianus; in my single male the anterior spots are clouded with black. The nervules separating the spots are heavily black.
Halmahera (\V. Doherty, August 1892) "
23. P.p. tenimberensis, Tanimbar
Walter Rothschild's description, 1895: "Front of the head red, but the red hairs are so densely mixed with black ones that the head appears to be more black than red.
Upperside deeper black than in polydorus.
The white patch on the forewings beyond the cell purer white than in the two preceding subspecies. Hindwings a little shorter than in polydorus; the submarginal spots stand, as in thessalia, rather closer to the margin than in the typical race. The white patch resembles, in the length of the second, third, and fourth spots, that of thessalia. The cellular spot is larger than in thessalia, almost as large as in polydorus; the first discal mark is of even breadth (2 mm.) and has a length of from 4 to 7 mm.; the three following ones have an average length of 6, 7. and 8 mm. respectively ; the fifth mark is longer than in polydorus, as it is basally less obliquely cut off. The nervules are much thinner black than in polydorus and thessalia; the second and third median ones are reddish.
The posterior submarginal spots are above, as in thessalia, less suffused with black than in polydorus.
Hah. Tenimber Islands (W. Doherty, June to July 1892)"
24. P.p. thessalia, Kai.
Walter Rothschild quoted Swinhoe as observing that thessalia differed from P. polydorus polydorus "especially in the form of the white patch to the hindwings. The cellular spot is much reduced, often pointlike or even absent from the upperside; the first discal mark is, above, about four times as long (in the direction of the nervules) as broad, or reduced to a point, or absent; the second spot has a length of from 5 to 6 mm.; the third is of the length of the fourth, and reaches therefore farther on the disc than the latter; their length varies from 7 to 9 mm.; the posterior spot is smaller than in polydorus, often triangular. The second and third median nervules are mostly red within the white patch."
25. P.p. utuanensis, New Ireland, Duke of York
26. P.p. varus, Aru
27. P.p. vulcanicus, Vulcan
In 1915 Rothschild, by then signing himself Lord Rothschild, described this new subspecies: "Above. Differ in the white patches beyond cell, and the white spot in the cell being larger and less suffused with black scales.
Below. The band of red spots on hindwing is generally smaller, and the red smear below vein 2 is much reduced. On the forewing the white is more extended."
28. P.p. wangaarensis, New Guinea
No points for guessing which subspecies are the most popular in Australia. Older documents mention even more subspecies names that are no longer used.
In 1895 Walter Rothschild wrote a detailed discussion of the polydorus he'd had a chance to study. The whole article has been preserved at:
"The variation of this species according to locality is not very conspicuous, hut still important enough to justify a separation of polydorus into a number of geographical races, of which that inhabiting the Southern Moluccas is the typical one described by Linne. It has not yet been observed that several of these races have black heads and necks, while others have these parts red ; the extent of the red on the abdomen is also different in several subspecies, and so is the size and form of the white spots on the hindwings ; such differences are, however, not of specific value, as there exist intergradations of every kind. I distinguish eight subspecies, to which probably will come a ninth, from Woodlark Island "
Life Cycle of the Species
Parent butterflies "fall in love and get married" as other animals do. While mating back to back, as most butterflies do, their spots form a figure 3.
Jmason92, who donated this photo to Inaturalist, says it was taken in February--summer in polydorus's habitat.
Caterpillars are known to eat any of eight species of vines in the genus Aristolochia. Eggs are laid by ones, well enough separated that each caterpillar should find plenty of leaves to eat as it matures. The eggs have been described as "little pumpkins."
Photo shared by Don Herbison-Evans & Stella Crossley, authors of a whole book on Australian Caterpillars and Their Butterflies and Moths.
Their texture comes from beads of aristolochic acid, which discourages ants or other lifeforms that might want to eat them. Regrettably, some Australians have planted non-Australian species of Aristolochia; at least three species of Australian butterflies are known to mistake these imports for plants their offspring will be able to eat, but the caterpillars who try to eat, say, American Aristolochia don't survive. Apparently Brazilian Aristolochia contains a very different phytochemical from similar looking Australian species.
The caterpillars are also highly textured, from the day they hatch onward. Their bumps and bristles are harmless, but they are toxic to most creatures that might eat them, being full of aristolochic acid; their color comes from phytotoxins in their foodplants. Predators usually survive eating a swallowtail caterpillar but, if they have the ability to vomit, they do.
Photo shared by Herbison-Evans & Crossley.
There is a family resemblance to other caterpillars in the genera Atrophaneura, Byasa, Losaria, and Pachliopta. They all seem designed to look and smell as unappetizing to as many birds as possible. The nonverbal message is "Something already ate me, and was sorry it did." The orange tubercles on the back of the head and front of the body can suggest a face from a bird's-eye view, as well, and the fleshy osmeterium or "stink horns" can look like a snake's forked tongue.
After reaching its maximal size the caterpillar actually shrinks a bit while assuming the pupating position...
After hanging like this for a few hours it sheds its dark skin and assumes an even odder shape:
Yet another photo shared by Herbison-Evans & Crossley. Polydorus chrysalids are quite variable; they can look red, green, or dark brown in addition to the usual pale tan.
Photo by Mmpro, from Australia, so probably queenslandicus,
Photo by Questagame, also probably queenslandicus.
From this emerges the butterfly, whose remaining life expectancy is thought to be about two weeks. It is not endangered and can legally be trafficked online, but please don't. Whether some of the subspecies on smaller islands may be endangered is under investigation.
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