Papilio memnon ssp. ♂

[Leonard187]

A specimen of Papilio memnon from my friend. It is said from Bali island on the label, so Is it should be ssp. memnon? Is its phenotype conform to that of nominated ssp.? How to determine subspecies based on male phenotype？Thanks

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[adamcotton]

This should be ssp. memnon. Males of P. memnon are somewhat variable

Here is the text about males from Jordan (1909) in Seitz, Macrolepidoptera of the World vol. 9:
anceus Cr. is the race which inhabits Sumatra, Nias and Batu. In the ♂ the upper surface is very similar to that of the ♂ of agenor; beneath the cell-stripes and discal stripes of the forewing are broad and the distal-marginal area of the hindwing is blue-grey, often somewhat yellow at the anal angle, in exceptional cases the yellow tint is deeper and more extended.
memnon L. ♂: the blue streaks of the upper surface in general shorter than in the ♂ of anceus, on the hindwing commonly not reaching the cell; beneath the cell- stripes of the forewing less distinct and the discal stripes less broad basally and commonly not touching the cell; the distal area of the hindwing beneath blue-grey as in anceus, usually somewhat yellow posteriorly.
perlucidus Fruhst. ♂: the blue-grey stripes of the upper surface of both wings and of the under- side of the forewing extend close to the distal margin; those of the hindwing above are remote from the cell and form a proximally almost uniformly concave broad band; the distal-marginal area of the hindwing beneath usually blue-grey, sometimes more or less extended orange.

Adam.

[Leonard187]

This should be ssp. memnon. Males of P. memnon are somewhat variable

Here is the text about males from Jordan (1909) in Seitz, Macrolepidoptera of the World vol. 9:
anceus Cr. is the race which inhabits Sumatra, Nias and Batu. In the ♂ the upper surface is very similar to that of the ♂ of agenor; beneath the cell-stripes and discal stripes of the forewing are broad and the distal-marginal area of the hindwing is blue-grey, often somewhat yellow at the anal angle, in exceptional cases the yellow tint is deeper and more extended.
memnon L. ♂: the blue streaks of the upper surface in general shorter than in the ♂ of anceus, on the hindwing commonly not reaching the cell; beneath the cell- stripes of the forewing less distinct and the discal stripes less broad basally and commonly not touching the cell; the distal area of the hindwing beneath blue-grey as in anceus, usually somewhat yellow posteriorly.
perlucidus Fruhst. ♂: the blue-grey stripes of the upper surface of both wings and of the under- side of the forewing extend close to the distal margin; those of the hindwing above are remote from the cell and form a proximally almost uniformly concave broad band; the distal-marginal area of the hindwing beneath usually blue-grey, sometimes more or less extended orange.

Adam.

Thanks very much for your reply, and I will learn it carefully, though it seems that it is a little difficult for me to identify the males of different ssp. T_T

[adamcotton]

I posted photos of different subspecies here https://archive.insectnet.com/thread/93 ... non?page=2 and scroll down to the 12th post on the page. Of course at that time agenor was a subspecies of memnon.

Adam.

[Chuck]

I posted photos of different subspecies here https://archive.insectnet.com/thread/93 ... non?page=2 and scroll down to the 12th post on the page. Of course at that time agenor was a subspecies of memnon.

Adam.

Might you address the tailed and tailless “forms” in short? Are all the tailed one group, and tailless another? Memnon has always confused me, and I’ve disposed of most, and won’t have the time to understand the phrenology and taxonomy.

[Leonard187]

I posted photos of different subspecies here https://archive.insectnet.com/thread/93 ... non?page=2 and scroll down to the 12th post on the page. Of course at that time agenor was a subspecies of memnon.

Adam.

Thanks for your kind reply ^0^. That is really helpful for me. The phenotypes of Papilio memnon or agenor are really attractive for me

[Leonard187]

Might you address the tailed and tailless “forms” in short? Are all the tailed one group, and tailless another? Memnon has always confused me, and I’ve disposed of most, and won’t have the time to understand the phrenology and taxonomy.

I think the tailed or tailless are only 'apparent' description for female Papilio memnon or agenor. eg. f.distantianus and f. alcanor are both 'tailed' forms of female agenor, and f. agenor, f. esperi, f. rhetenorina, f. butlerrianus and some other phenotypes without tails are all 'tailless' form. So many phenotypes may be related to mimicry to other toxic species, such as Losaria doubledayi, Pachliopta aristolochiae, Atrophaneura nox erebus, or Atrophaneura varuna, etc. which are living at same region.

[adamcotton]

Might you address the tailed and tailless “forms” in short? Are all the tailed one group, and tailless another? Memnon has always confused me, and I’ve disposed of most, and won’t have the time to understand the phrenology and taxonomy.

I think the tailed or tailless are only 'apparent' description for female Papilio memnon or agenor. eg. f.distantianus and f. alcanor are both 'tailed' forms of female agenor, and f. agenor, f. esperi, f. rhetenorina, f. butlerrianus and some other phenotypes without tails are all 'tailless' form. So many phenotypes may be related to mimicry to other toxic species, such as Losaria doubledayi, Pachliopta aristolochiae, Atrophaneura nox erebus, or Atrophaneura varuna, etc. which are living at same region.

@Leonard187 is correct, but I should explain that there are two basic phenotypes in Papilio agenor and P. memnon:

one is tailless with a black abdomen and a range of different wing patterns very closely associated with these characters depending on the local distasteful Atrophaneura model species (often more than one)
and
the other is tailed with an orange/yellow sided abdomen and the wing pattern mimics the local Pachliopta or Losaria.

These characters are so closely placed on the same chromosomes that 'mixed forms' almost never (extremely rarely) occur - for instance there are only 3 recorded specimens of a tailed form distantianus with a black abdomen.
Clarke, Sheppard & Thornton (1968) investigated the genetics of the then single species and coined the term 'super-gene' for this bundle of characters.
See
https://www.jstor.org/stable/2416804
for the abstract and some further information
and
https://royalsocietypublishing.org/doi/ ... .1971.0109
for similar information about a subsequent paper by Clarke & Sheppard in 1971.

I was lucky enough to get to know Sir Cyril Clarke when I was a student (in the UK) in the late 1970s.

Adam.

[adamcotton]

I think the tailed or tailless are only 'apparent' description for female Papilio memnon or agenor. eg. f.distantianus and f. alcanor are both 'tailed' forms of female agenor, and f. agenor, f. esperi, f. rhetenorina, f. butlerrianus and some other phenotypes without tails are all 'tailless' form. So many phenotypes may be related to mimicry to other toxic species, such as Losaria doubledayi, Pachliopta aristolochiae, Atrophaneura nox erebus, or Atrophaneura varuna, etc. which are living at same region.

A couple of minor corrections to the above:
firstly 'f. butlerrianus' should be 'butlerianus', named after A. G. Butler by Lord Rothschild in 1895;

and also, nowadays the taxon varuna has become a subspecies of Atrophaneura astorion because it was recently shown that Papilio astorion Westwood, 1842 was published before Papilio varuna White, 1842.
Here is the relevant text from Hu et al. (2023) (see viewtopic.php?t=1288) :

5. Seniority of Synonyms
In many situations prevailing usage clauses were introduced in the current Code (ICZN 1999) to ensure stability
of nomenclature. There are no such clauses covering the Principle of Priority, so the older name becomes the valid
species name.
5.1 Papilio astorion Westwood, 1842: 37. Currently regarded as a valid subspecies of Papilio varuna White, 1842
(currently Atrophaneura varuna [White, 1842]) (Racheli & Cotton 2010: 29).
However, P. astorion Westwood was published on [1] March 1842 (Evenhuis 2003: 15), whereas the date of
publication of P. varuna White can only be attributed to [31] March 1842. Therefore, astorion has priority over
varuna as the valid species name.
The subspecies of this species are thus:
Atrophaneura astorion astorion (Westwood, 1842)
Atrophaneura astorion varuna (White, 1842)
Atrophaneura astorion zaleucus (Hewitson, 1865)

Note also that the various forms listed by @Leonard187 all belong to Papilio agenor. The tailed form of female P. memnon memnon with a bicoloured abdomen occurring in Java is called f. achates and mimics Losaria coon with yellow markings on the hindwings rather than red, and the tailless forms with black abdomens mimic other local Troidini. Interestingly only the local tailless mimetic forms occur in the Borneo populations of P. memnon memnon where there is no species of Losaria.

Adam.

[Chuck]

Can we sail this waaay back a notch?

Forget agenor to avoid my confusion and let’s stick with memnon.

Memnon has subspecies. So does any given ssp have both tailed an tailless forms? Does that the. Depend on location?

Thanks.

[adamcotton]

Memnon has subspecies. So does any given ssp have both tailed an tailless forms? Does that the. Depend on location?

Yes and yes. As I just posted yesterday, ssp. memnon has a tailed form in Java as well as tailless forms (the same in Bali), but in Borneo the same ssp. only has tailless forms, and they are not identical to those in Java.
Ssp. anceus in Sumatra also has both tailed and tailless forms, but the tailless ones tend to be more common, similarly ssp. eos from Kangean Island, not far from Java can have tailed females, but the various subspecies in the Lesser Sunda Islands east of Bali all only have tailless subspecies.
Subspecies lowii in Palawan is always tailed in both sexes, and interestingly males in Balabac Island (between Borneo and Palawan) range between tailed to tailless and in between.
Subspecies mayo in the Andaman Islands, previously regarded as a separate species, has a tailless male with a distinctive blue hindwing patch, but the female is always tailed, a mimic of the Andaman endemic Losaria rhodifer. There are no tailless forms there, and no tailless models.

Interestingly ssp. memnon was introduced to Ceram and Buru very many years ago (probably on Citrus plants with migrant Javanese), and both tailed and tailless forms still occur there. Luckily they didn't hybridise with the closely related local species Papilio deiphobus.

Adam.

[Chuck]

ssp. memnon has a tailed form in Java as well as tailless forms

In such a (relatively) small place, what is the driver for two different forms? Is it altitude or another environmental element? North/ south?

I presume these two different forms are genetic, thus parents with tails (for example) will produce offspring with tails. True?

I long ago quit trying to figure out memnon, back before the internet, and quit tracking developments. This now though is rather interesting.

Thanks.

[adamcotton]

Chuck,

As I said above:

one is tailless with a black abdomen and a range of different wing patterns very closely associated with these characters depending on the local distasteful Atrophaneura model species (often more than one)
and
the other is tailed with an orange/yellow sided abdomen and the wing pattern mimics the local Pachliopta or Losaria.

These characters are so closely placed on the same chromosomes that 'mixed forms' almost never (extremely rarely) occur - for instance there are only 3 recorded specimens of a tailed form distantianus with a black abdomen.
Clarke, Sheppard & Thornton (1968) investigated the genetics of the then single species and coined the term 'super-gene' for this bundle of characters.
See
https://www.jstor.org/stable/2416804
for the abstract and some further information
and
https://royalsocietypublishing.org/doi/ ... .1971.0109
for similar information about a subsequent paper by Clarke & Sheppard in 1971.

Basically, this means that the genes controlling the two basic forms are closely bundled together on the same part of the same chromosome so that the 'super-gene' bundle is almost always unchanged during gamete formation (cell division forming sperm and egg cells).

You asked:

I presume these two different forms are genetic, thus parents with tails (for example) will produce offspring with tails. True?

Actually, not necessarily! It would all depend on which 'super-genes' each parent is carrying and whether one is dominant over the other or not. Parents may carry one 'super-gene' on one side of the chromosome and one on the other or they may carry the same 'super-gene' on both sides of the chromosome, so when they mate there can be more than one offspring phenotype.

Adam.

[Leonard187]

Yes and yes. As I just posted yesterday, ssp. memnon has a tailed form in Java as well as tailless forms (the same in Bali), but in Borneo the same ssp. only has tailless forms, and they are not identical to those in Java.
Ssp. anceus in Sumatra also has both tailed and tailless forms, but the tailless ones tend to be more common, similarly ssp. eos from Kangean Island, not far from Java can have tailed females, but the various subspecies in the Lesser Sunda Islands east of Bali all only have tailless subspecies.
Subspecies lowii in Palawan is always tailed in both sexes, and interestingly males in Balabac Island (between Borneo and Palawan) range between tailed to tailless and in between.
Subspecies mayo in the Andaman Islands, previously regarded as a separate species, has a tailless male with a distinctive blue hindwing patch, but the female is always tailed, a mimic of the Andaman endemic Losaria rhodifer. There are no tailless forms there, and no tailless models.

Interestingly ssp. memnon was introduced to Ceram and Buru very many years ago (probably on Citrus plants with migrant Javanese), and both tailed and tailless forms still occur there. Luckily they didn't hybridise with the closely related local species Papilio deiphobus.

Adam.

Thanks for the corrections by Adam. I didn't pay much attention to the genus Atrophaneura and I will learn it carefully in the future. It is interested that I found that many female individuals of ssp. anceus on Sumatra have white triangles on the base of forewings instead of red ones distributed on the other islands，not only tailless ones but also tailed (Of course there are still some red ones on this island).

I have seen a specimen from N. Borneo with very strange phenotype just like a tailless one with tailed pattern on the hindwings (f anura), so what is its model? Might be Troides sp.?

Another question, do you have some specimen of ssp. eos? I've not ever seen this ssp before ^0^

Thanks

[adamcotton]

I found that many female individuals of ssp. anceus on Sumatra have white triangles on the base of forewings instead of red ones

The white triangle theoretically evolved to mimic Atrophaneura hageni.

I have seen a specimen from N. Borneo with very strange phenotype just like a tailless one with tailed pattern on the hindwings (f anura), so what is its model? Might be Troides sp.?

Yes, it is believed that some forms of ssp. memnon on Borneo mimic Troides spp.

do you have some specimen of ssp. eos?

Yes, and I posted the photo in the archive post I cited above, here it is again. The ssp. eos specimens from Kangean are the males in the middle column and two columns of females to the right.

The first two columns on the left are ssp. memnon from Bawean.

Adam.

[Leonard187]

The white triangle theoretically evolved to mimic Atrophaneura hageni.

Thanks Adam. However, it seems really different with its model Atrophaneura hageni, so does it use its white triangles to mimic the white head of A.hageni?
As picture 2 below, my friend has a tailed one also with white triangles which is really interesting, though I think it should also mimic L. coon.
Is picture 3 named as f.anura? It is not really seems like a Troides spp.

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[Chuck]

I presume these two different forms are genetic, thus parents with tails (for example) will produce offspring with tails. True?

Actually, not necessarily! It would all depend on which 'super-genes' each parent is carrying and whether one is dominant over the other or not. Parents may carry one 'super-gene' on one side of the chromosome and one on the other or they may carry the same 'super-gene' on both sides of the chromosome, so when they mate there can be more than one offspring phenotype.

Adam.

That is quite amazing. I wonder if one form will prevail, given that one is a mimic and the other (I presume) is not.

This all is amazing, study I've avoided. Adam, does anyone, including institutions, have a collection to rival yours?

[adamcotton]

That is quite amazing. I wonder if one form will prevail, given that one is a mimic and the other (I presume) is not.

No, in nature both tailed and tailless forms are mimetic, the tailed forms mimic Pachliopta and Losaria, whereas the different tailless forms mimic several Atrophaneura, so they all should survive by predators not attacking them.

On the other hand, in places where no tailless distasteful Troidini models occur it is likely that the tailless ones may be eaten and the tailed ones avoided. However, that was not the scenario that I was replying to in this case.

Adam, does anyone, including institutions, have a collection to rival yours?

Most definitely there are many much better collections than mine. I seem to remember that the Clarke/Sheppard collection of experimental rearing is in the NHM, London, where of course they also hold huge collections of wild specimens from just about everywhere.

Also there are some huge collections (both private and institutional) in Japan, and I know of one private collection at least in the US that has a better collection than mine.

Adam.

[Chuck]

That’s amazing, two forms within the same population mimicking two different inedibles. What a trip. No wonder people have studied them so intently. Thanks Adam. I’ll leave you alone now.

[adamcotton]

That’s amazing, two forms within the same population mimicking two different inedibles. What a trip. No wonder people have studied them so intently. Thanks Adam. I’ll leave you alone now.

There are usually at least 2 different tailless forms, mimicking tailless models, as well as the tailed one mimicking a tailed model.

Adam.