Chuck wrote: ↑Sun Dec 08, 2024 9:59 pm These unique populations can’t be described as subsubspecies. Until a ssp is elevated to full species.

Chuck,

The whole point here is:
a) that anything below the rank of subspecies is outside the ICZN Code, and as such any names proposed for such entities (individual forms, aberrations, sexual morphs etc) are not available names
and
b) these individuals are not recognised as comprising 'unique populations' but as variations within a population (= species or subspecies).

If subsequently a researcher finds that in fact certain individual variations found in the same place represent two different taxa THAT DO NOT INTERBREED then there is a process to decide the correct names to apply to these two:
1. Find out which of these two populations the name-bearing type belongs to
2. Name the other population according to the ICZN Code regulations as a separate NEW SPECIES (not subspecies - note that two subspecies of the same species do not normally occur in the same place, except for in relatively rare hybrid zones where the ranges of two subspecies overlap).

Adam.

repeat, the ICZN does not set ANY taxonomic 'rules', only regulations governing nomenclature, as I stated above

I knew you were going to say that. But the recognition of taxonomy is via nomenclature.


it does generate taxonomic rules by not allowing named sub sub species. These unique populations can’t be described as subsubspecies. Until a ssp is elevated to full species. THEN they can be described as a ssp. Until that point they have no nomenclature rules or protection of their informal identifier.

Thank you Adam. You truly know the best.

While reading this book, I noticed that morphologically, ssp. kooichi seems to be more similar to Taiwan's ssp. chungianus. This made me curious about the phylogenetic relationship of this species.

Ming

Yes, Cabintom's reply is exactly a very simple explanation of the situation.

Names that were previously treated as synonyms of a subspecies and USED as form names in a polymorphic species may be used as subspecies in the case that they were originally described as a species or subspecies (this includes names described as a form or variety before 1961 as long as they were originally described in a geographical sense rather than as a sexual or individual variation) but if they were described as an aberration or the original description clearly indicates the name is infrasubspecific (such as a different coloured individual given a name) then the name cannot currently be adopted with its original author/year as an available name. Thus it is important to examine the original description of each name, and for a separate geographical population the oldest available name becomes the valid subspecies name IF subspecific or specific separation is warranted.

However,

Chuck wrote: ↑Fri Dec 06, 2024 6:03 pm So what of those various regional "forms"? Suddenly when a ssp is elevated to sp, NOW it too can have ssp, so some of those forms- which are not- can be taxonomically recognized. This is not new, though it seems genetic analysis has drastically increased this situation.

Forms of the same subspecies may appear individually SLIGHTLY different at the individual gene level, in the same way as people with blue or brown eyes would, but this genetic difference would be minimal, and is just individual variation in a single population. Particularly in polymorphic mimetic butterfly species local selection pressure on individual appearance can have significant effects. For example in the P. glaucus group the presence or absence of a dark female form probably depends more on the presence or absence of the model (Battus philenor) but may also depend on other unclear factors where the model does not occur but the dark form is still present. For instance perhaps the predators move around and learn to avoid the dark distasteful butterfly elsewhere and then subsequently avoid any they meet among the P. glaucus in a different place and/or time. It is worth noting that this can also be a temporal difference ... the model may have been flying weeks or even months before the mimic, but the predators in the same place associate the colour with a nasty event and remember it.

Chuck wrote: ↑Fri Dec 06, 2024 6:03 pm How will ICZN recognize these distinct below-the-subspecies populations? Will ICZN become archaic, and ignored, while the geneticists publish these great branching trees?

Not at all. Bear in mind that the ICZN Code governs NOMENCLATURE, not taxonomy. The Code tells you how to treat names proposed by taxonomists and which names can and cannot be used. It is up to the taxonomists to apply the 'rules' in order to choose the valid name for any taxon THAT NEEDS A NAME, excluding names that do not conform to the 'rules' of the Code. Bear in mind that each branch of a tree is a 'taxon' but they don't ALL need scientific names.

Any names originally proposed below subspecies level (= infrasubspecific) are unavailable under the Code and cannot be used as valid names (see above), but if a taxonomist decides that a POPULATION is worthy of subspecies status but does not have an available name that can be used as the valid subspecies name the taxonomist will describe a new subspecies, thus providing an available name to use.

Chuck wrote: ↑Fri Dec 06, 2024 6:03 pm How do we reconcile ICZN's taxonomic rules with recent discoveries? Will there continue to be named populations that we circulate, like a secret, because the authority won't recognize it?

I repeat, the ICZN does not set ANY taxonomic 'rules', only regulations governing nomenclature, as I stated above. This is a slightly subtle issue to grasp, but there is a distinct difference between nomenclature (how animals are named) and taxonomy (how they are related to each other). The Code does not set rules about how to distinguish a species or subspecies, or indeed a genus or a family, it just governs the names that taxonomists should use once they have decided how to classify the animals they study.

Note that in the case of angelfish the different coloured ones in captive breeding are probably like comparing different breeds of dogs. A Chihuahua is genetically not different to a Great Dane and does not deserve subspecies status, so the different coloured angelfish are not actually subspecies. It is possible that different rivers contain individual colours, but ichthyologists do not separate subspecies as much as lepidopterists for various reasons which they have opinions about, and as they all interbreed they are clearly the same species. Similarly the pigeons around here can be a number of different colours (white, grey, mixed etc) but they are all the same taxon and they all interbreed. Different branches of Zoology have their own way of classifying populations, based on criteria relevant to the animals they study. Some recognise subspecies, others do not, usually with reasons based on the individual biology of the type of animal.

Adam.

Chuck wrote: ↑Fri Dec 06, 2024 3:13 pm Since we're already a bit off-topic, the dashed red "phylogenic uncertainties": does that mean the uncertainty likes only within those nodes enclosed in the red box, or the entire tree?

This is explained in the paper - different analyses produce slightly different linkages between the clades but do not affect the relationships between species in each clade and subgenus. The red dashes indicate the area where alternative topologies have been found and further investigation of the higher classification within this part of the genus is needed.

All analyses produce the same topology between the clades outside the red dashes, and the clades (3a, 3b & 3c) all definitely belong there relative to the others, but different methods can produce trees with these three subgenera in different positions relative to each other. For example in some analyses clade 3c is basal to 3a and 3b. The published tree with regard to the relative positions of these 3 clades is the best supported under the analysis in the paper.

Adam.

Wu Ｍing Hsuan wrote: ↑Sat Dec 07, 2024 5:53 am Did I misinterpret the tree in this paper, or could there be any other possible explanations for this?

No, you didn't misinterpret the tree, and yes there is an explanation.

Unfortunately the eurous group (both mullah and eurous belong here) is very poorly resolved using mitochondrial DNA alone so a tree with very few specimens (one of each) produces anomalous topography. This is why the third part of the subgenus Pazala analysis has been delayed due to the time it is taking to run full genomic analysis on the whole group.

That is why the Bozano 2nd edition did not update this species group. Bozano asked me to help with a second edition but I asked him to wait until our current analysis is finished (it has taken way longer than anticipated because genomics is needed - these species and subspecies separated relatively recently compared to the mandarinus group species which were very well delineated by mtDNA alone [genomics confirms the findings in this group by the way]) but he went ahead and published a 2nd edition without my involvement and sadly followed the 'peer-pressure' results published in Huang (2023) - many of Huang's friends didn't like the idea that there could be several species within what looked to them like a single species. A number of Huang's illustrated mandarinus group specimens were misidentified (his so-called 'hybrids' were actually 1st generation G. confucius, and several other specimens were misidentified) and he made assumptions about the Vietnamese taxa, and as a result his findings on Pazala in that paper were flawed.

Adam.

I think that if a subspecies is now recognized as a species, any "forms" that then merit being raised to subspecies level can be done so while adhering to the ICZN code. Depending on how they were described, some of those "forms" can have their status revised (pulled out of synonymy, stat. rev. or whatnot), others will have unavailable names and therefore would need to be described under a new name.

I expect Adam will have corrections, but I believe this is the essence of it.

According to the Guide to the Butterflies of the Palearctic Region(part 1 second edition, 2024), Graphium mullah is divided into three subspecies: ssp. mullah, ssp. kooichii, and ssp. chungianus. However, based on the phylogenetic tree constructed using mitochondrial genome analysis in the paper Complete Mitochondrial Genomes of Two Insular Races of Pazala Swordtails from Taiwan, China (Lepidoptera: Papilionidae: Graphium), the relationship between Graphium mullah mullah and Graphium eurous asakurae appears to be closer than that between ssp. mullah and ssp. chungianus.

Did I misinterpret the tree in this paper, or could there be any other possible explanations for this?

All the best,
Wu Ming Hsuan

Thank you!

adamcotton wrote: ↑Thu Nov 30, 2023 6:43 pm Subspecies generally have a smaller difference in COI between them than between species, but it is often best to treat subspecies as visibly distinguishable from the nominotypical population in the large majority of specimens.

With all the genetics work being done, I've seen a lot of ssp being elevated to full species status. And this brings an interesting question.

In binomial nomenclature the identity of an insect (and all else) is limited to two names: genus and species. However, ICZN recognizes subspecies with "A scientific name added as a trinomen on the end of a bionomen is taken to indicate a subspecies. However, it stops there."

So what of those various regional "forms"? Suddenly when a ssp is elevated to sp, NOW it too can have ssp, so some of those forms- which are not- can be taxonomically recognized. This is not new, though it seems genetic analysis has drastically increased this situation.

I wonder too- is the old taxonomic architecture dead? The genus-sp-ssp tiers are each equivilent, at the same level. But genetic trees show that the branches and nodes are not all at the same level. Some may dead-end at species; some appear to go beyond ssp...some of these "forms" are morphologically and genetically different than the sp/ ssp than the next branch over is to itself.

How will ICZN recognize these distinct below-the-subspecies populations? Will ICZN become archaic, and ignored, while the geneticists publish these great branching trees? It seems so in some cases; case in point the well known freshwater angelfish, Pterophyllum: there are to this day only three recognized species and zero subspecies. But the fish nuts have names for these unrecognized "populations". Why haven't the professionals bothered to describe new species and subspecies? Is it because, in their world, they all know what they're talking about, so don't need ICZN's taxonomic guidance? Is discovery moving too fast for science?

How do we reconcile ICZN's taxonomic rules with recent discoveries? Will there continue to be named populations that we circulate, like a secret, because the authority won't recognize it?

Since we're already a bit off-topic, the dashed red "phylogenic uncertainties": does that mean the uncertainty likes only within those nodes enclosed in the red box, or the entire tree?

For example, hospiton and zelicaon are on branches that meet, and are far off from the higher level branch that eventually leads to brevicauda. Does the red box mean that the close relationship of hospiton and zelicaon is in question, or that those are indeed as shown, but the ancestral tree at a much higher level is unclear? i.e., Clades 3b and 3c may be all one clade, or the extant species under them may not be at all as shown?

eurytides wrote: ↑Fri Dec 06, 2024 1:36 am Adam, in the tree you posted above, what does the asterisk beside some species indicate? For example P. brevicauda?

It indicates the species illustrated on the right of the tree.

Adam.

Adam, in the tree you posted above, what does the asterisk beside some species indicate? For example P. brevicauda?

adamcotton wrote: ↑Tue Dec 03, 2024 4:38 pm This malformation of the forewing costa was caused by incomplete closure of the forewing case on the right side of the pupa after shedding the pre-pupal skin.

If these specimens originate from southern Xishuangbanna the white inner hindwing edge is part of the mimicry of Losaria doubledayi which occurs nearby in Laos. This is true f. distantianus, whereas the darker form which mimics Pachliopta aristolochiae in China is called alcanor. Form alcanor particularly has a short white patch inside the hindwing cell, whereas distantianus has an elongate white patch in the cell.

Thanks, Adam. About f.alcanor, I've got a specimen without white patch inside the hindwing cell, recently.

adamcotton wrote: ↑Tue Dec 03, 2024 4:38 pm PS. Note that mayo is now a subspecies of Papilio memnon, rather than a separate species, and does not belong to P. agenor.

Sorry for my mistake T-T

martellat0 wrote: ↑Thu Dec 05, 2024 12:46 pm While on the topic, allow me to share some photos from an individual I've been following on Twitter. He is a collector and breeder based in Okinawa who - for the past few years, it seems - has been selectively breeding P. agenor in order to produce individuals with extreme white pigmentation. He can be found under the handle @rhetenor12.

Here is a prized specimen. The translate function provided by Twitter can sometimes be unreliable, but from what I can gather, he seems to be trying to recreate the phenotype of a certain population of P. agenor from Nagasaki in the 1970s. He has shared pictures of other specimens with varying degrees of amelanism, but in my opinion, this is one of the most striking examples.

He has also produced both male and female individuals with malformed (possibly absent) wing veins.

Here is a specimen similar to that other one. According to the caption, it is a wild-caught specimen from Taiwan.

Thanks for sharing, the white one is really amazing ^0^. Hoping for more pictures of Papilio agenor or memnon.
About the breeding pair, the round wing-shape make them seem like Ornithoptera sp.

While on the topic, allow me to share some photos from an individual I've been following on Twitter. He is a collector and breeder based in Okinawa who - for the past few years, it seems - has been selectively breeding P. agenor in order to produce individuals with extreme white pigmentation. He can be found under the handle @rhetenor12.


https://x.com/rhetenor12/status/1831644452248809632
Here is a prized specimen. The translate function provided by Twitter can sometimes be unreliable, but from what I can gather, he seems to be trying to recreate the phenotype of a certain population of P. agenor from Nagasaki in the 1970s. He has shared pictures of other specimens with varying degrees of amelanism, but in my opinion, this is one of the most striking examples.


https://x.com/rhetenor12/status/1705157856436318680
https://x.com/rhetenor12/status/1804744344093565097
https://x.com/rhetenor12/status/1762714940706828460
He has also produced both male and female individuals with malformed (possibly absent) wing veins.


https://x.com/rhetenor12/status/1198916482723266561

Leonard187 wrote: ↑Mon Dec 02, 2024 4:38 pm About the white one, I saw a wild-caught female with very white wings. It is attractive due to the white pattern on the cells on hindwings, and the red triangles seem to decreased.

Here is a specimen similar to that other one. According to the caption, it is a wild-caught specimen from Taiwan.

adamcotton wrote: ↑Tue Dec 03, 2024 7:54 pm I seem to remember this came from W Sumatra, where the yellow L. coon does not occur, but perhaps it is from S Sumatra, where L. coon is found in the far south. The pale hindwing spots (even though in the wrong position) combined with the white 'head' sends two signals.

No, form anura is rather different to this, although the hindwing markings are in a similar position, but they are cream tinged with orange (I suppose this may look bright yellow in flight), the underside is creamy with large black spots (like the Troides female) and it comes from Mt. Kinabalu, Borneo, not Sumatra.
I think this Sumatran form is actually derived from the originally tailed form which has lost the tails (note the pale sides to the abdomen), and it may well mimic a Troides female too. Basically this means the gene to include tails has been 'switched off'.

Adam.

Thanks, but pic2 is from S Sumatra based on the label.
About f.anura I only saw pictures from the paper of sir Clarke as well as you posted previously (really have yellow abdomen), and I donot know where this one (pic 3 of I posted above) collected from >-<
BTW, today on website, I've seen a specimen picture of ssp. oceani from remote Enggano island of Indonesia, and this female one seems rather similar with male. So whether we can suppose that this island might be the origin location of memnon which showed monomorphic of male/female?

Thank you for posting these photos of very important specimens.

Firstly the Batang - 'Mangkang' border specimen is from the type locality of archias (Batang, not 'Fou Lin' as stated by Fruhstorfer), and this is a first generation specimen (from May - note the broad black dorsal band on the abdomen). The type and other specimens I have seen (photos) are summer generation (July-August). Note that the data label is a little confused - Batang is in Sichuan not Tibet, and 'Mangkang' is correctly Markam County, Tibet, which is immediately west of Batang - the westernmost county in Sichuan. 'Mangkang' is the Han Chinese name for Markam. So Sichuan and Tibet are swapped on the label. The border locality is on the Jinsha River south of Batang. Byasa daemonius and Iphiclides podalirinus also occur there, but fly in June.

I have already asked the seller about the Gongga Shan locality (the specimen at the top of the thread is from the same lot). He told me that they came from a local catcher, and he is asking precisely where the locality is. These also look similar to archias, but possibly may not be that taxon, as the location is far from Batang with a very dry plateau at Litang separating the two populations and a totally different taxon 'neochinensis Sheljuzkho (an unavailable name) in Jiulong County to the west of Gongga Shan, i.e. between the two populations.

Adam.

Leonard187 wrote: ↑Tue Dec 03, 2024 12:59 pm it seems really different with its model Atrophaneura hageni, so does it use its white triangles to mimic the white head of A.hageni?

Exactly, and the silvery hindwings mimic the female of Atrophaneura nox in Sumatra. It's a case of two different warning patterns combined in the same mimic, sort of like the flashing lights on an ambulance telling the predator to keep away.

Leonard187 wrote: ↑Tue Dec 03, 2024 12:59 pm my friend has a tailed one also with white triangles which is really interesting, though I think it should also mimic L. coon.

I seem to remember this came from W Sumatra, where the yellow L. coon does not occur, but perhaps it is from S Sumatra, where L. coon is found in the far south. The pale hindwing spots (even though in the wrong position) combined with the white 'head' sends two signals.

Leonard187 wrote: ↑Tue Dec 03, 2024 12:59 pm Is picture 3 named as f.anura? It is not really seems like a Troides spp.

No, form anura is rather different to this, although the hindwing markings are in a similar position, but they are cream tinged with orange (I suppose this may look bright yellow in flight), the underside is creamy with large black spots (like the Troides female) and it comes from Mt. Kinabalu, Borneo, not Sumatra.
I think this Sumatran form is actually derived from the originally tailed form which has lost the tails (note the pale sides to the abdomen), and it may well mimic a Troides female too. Basically this means the gene to include tails has been 'switched off'.

Adam.

Chuck wrote: ↑Tue Dec 03, 2024 6:58 pm 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.