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Post by nomad on Jan 22, 2013 13:22:29 GMT -8
I thought I was making a strong argument for the separation of O. paradisea arfakensis. What else can I do but to study the specimens and information that I have in the specialist books on Ornithoptera. I believe you will find that this is what Bernard d Abrera and others authors like Oliver Schaffler did when they separated O. arfakensis from O. paradisea. I was merely pointing out the main differences that I observe when studying specimens of O. arfakensis without going into long morphological details. I also mentioned that O. arfakensis is sympatric with O. paradisea chrysanthemum in the Arfak mountains and this is another valid point. I also believe the very different larva is also another important factor. All the photographs I have seen of the other O. paradisea subspecies larva have orange-red tubercles not cream and black, yet another important fact. O. richmondia larva feed on a different hostplant. O. arfakensis also appears to feed on a different species of Pararistolochia. The O. arfakensis male also has a different genitalia to the other O. paradisea subspecies. I must mention that I am only expressing my opinion and agreeing with the specialists that have seperated O. arfakensis and who have studied Ornithoptera much more than I ever could.
Peter
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Post by nomihoudai on Jan 22, 2013 14:41:42 GMT -8
Greg Watson, which actually has seen them in the field, states that they live in different altitudes, this is not what sympatric means as they remain separated in habitat. Furthermore altitude can have a large impact on the look and genetics of a population. I can not give examples in Troidini for this (or is Ornithoptera arfakensis and Ornithoptera paradisea such an example?) but I can give two examples in Lycaenidae which I know a bit better. In Lycaenidae you have the two pairs of Lycaena hippothoe hippothoe and Lycaena hippothoe eurydame and Lycaena tityrus tityrus with Lycaena tityrus subalpinus. The latter is always a high altitude version of the former with strong differences in coloration (lack of purple or lack of orange, ventral side spot patterns sometimes reduced to over 50% of spotsize) and also observable differences in wing shape (accuter apex and anal angle). If you look at a map the regions where they occur will only be separated by a few kilometers or just a couple of hundred meters, but if you look at the altitude there is always a strong gap between both populations. Lycaena tityrus subalpinus even has a different hostplant than nominate (although within the same genus). There is hybrid specimen at regions where they can sometimes meet. The darker coloration of the high altitude populations is just a change of the present alleles within the populations not producing any reproductive isolation. We can not say A in the first story and B in the other story if the same situation is present.
I am not speaking against your opinion, I just wanted to show you that you should give more sound arguments when expressing an opinion.
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Post by wollastoni on Jan 23, 2013 1:13:15 GMT -8
nomad < about Yapen island. I have Jan's book, it is a great book with stunning pictures but I would say he tends to overexagerated how remote the places he has visited in New Guinea are... this is a common habit among Western people who have visited New Guinea. Some parts of New Guinea are nearly impossible to access but it's not the case for Biak and Yapen, if one has enough money to go there it is easy. As there is a strong species endemism on those islands, they have been very much surveyed by entomologists. Those entomologists were of course paying much attention to "famous" genera like Ornithoptera or Delias. I find strange all those entomologists missed those big Ornitho... maybe it is the case but I would not trust "dealers material". Every year those "dealers" offer me some Arfak species saying it's a new ssp. from Biak or Yapen... but they are not. On the attached picture took by a friend during our trip to Biak, you can see above the sea the shadow of Yapen island, as you can see you just have to rent a small boat to go there. (PS : yes this picture also shows that a trip to New Guinea can also have pleasant times...) Attachments:
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Post by nomad on Jan 23, 2013 2:57:07 GMT -8
Claude, Thank you for you information on the Lycaena and its changes in appearance at altitude. It is interesting that Greg Watson has seen both paradisea [arfakensis and chrysanthemum] in the Arfak Mountains at different altitudes. I originally thought that the different paradisea in the Arfak mountains might be flying at different altitudes and have mentioned that in this thread. I also mentioned that Greg is one of the few Entomologists that has observed O. paradisea in the wild. I do not know how many localities Greg saw O. paradisea chrysanthemum in the Arfak mountains, but I would say you would have to visit all of its localities in the Arfak's mountains, before you could say that it does not fly with arfakensis. Matsuka also visited the Arfak Mountains and observed O. arfakensis. In his book he shows one locality where both paradisea are found but does not mention the altitude. Abrera mentions both paradisea fly together in the Arfaks and says that this is the information he has received from collectors. I also mentioned that O. arfakensis has a different larva, hostplant and genitalia and again I would say these are all strong arguments when dealing with butterfly species. However elevation would not be a good argument for the separation of arfakensis from paradisea. If you believe as I do there are two O. paradisea species with a number of subspecies and local forms, geographical isolation would be much more important. Because if O. arfakensis is indeed a good species it would have to have occcidentalis and galatea as subspecies [ the latter may be a local form] both of which occur at a much lower altitude than arfakensis inhabits in the Arfak mountains. There are again the discoveries of the paradisea on Yapen and Supiori [If this is true?] and both of these are again very close morphologically to O. arfakensis. Whether you think there are two species or just subspecies of O. paradisea there are clearly two important groupings. I would suggest the oldest of these in the arfakensis group is in the Arfak mountains and the original dispersal came from here.
Thank you for you link to the Great O. paradisea versus O. arfakensis debate. I actually saw this thread when it was on going at the time but then I was not a member. I have since then, read the debate a number of times and like most people I believe that some of the points that were made are valid ones and some are not. I also would like to say that I would not even enter into a argument that involves comparing butterflies with mans infinite desire to do selective breeding with domestic dogs and other domestic animals for his own use. I believe with butterflies [species,subspecies] we are dealing with the hand of ' nature' not the hand of ' man'. I also believe with butterflies, it is geographical isolation that is important and barriers such as mountains and being confined to Islands. Ornithoptera are strong fliers but isolation it the key factor here. This can be seen with New Guinea's other important group of butterflies ' the Genus Delias '. In New Guinea most mountain ranges have their endemic Delias.
Peter.
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Post by nomad on Jan 23, 2013 3:04:04 GMT -8
Wollastoni Thank you for your information on Yapen. Really lovely picture of Biak, I wish I was there now,. What a place. It is snowing outside in England and its freezing. Like you I find it very strange O. paradisea has been discovered on those Islands and I await further information and specimens. Peter.
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Post by wollastoni on Jan 23, 2013 3:55:33 GMT -8
I would add than on this paradisiacal beach were flying hundreds of Graphium and Appias. In a small bush near the beach there were also huge Idea species. Another story : I made some snorkeling there for fun. 2 meters from the beach thousands of small tropical fishs, 4 meters from the beach bigger one like groupers, 10 meters even bigger like tuna... and then I came back because I did not want to meet the bigger ones that eat tuna fishs.
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Post by nomad on Jan 24, 2013 4:05:31 GMT -8
Wollastoni, sounds like a dream trip to paradise. Lots of fun and butterfly collecting. While you was on Biak did see much virgin forest and did you find any rare Delias. I know there are a number of rare Delias endemics on Biak and have they all been seen in recent years. Is there any disagreement on the naming of the species and subspecies in the Delias Genus.
nomad
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Post by nomad on Jan 24, 2013 10:31:13 GMT -8
Just to a add a bit of fuel to the fire, here are a few thoughts. I have been giving some thought to Claude's information of how the two Lycaena subspecies vary at different altitudes. Especially of note is how he reveals the two subspecies of Lycaena tityrus [ Sooty Copper] overlap from different altitudes and produce a multitude of different intermediate characters. Lycaena tityrus and Lycaena hippothoe are quite widespread and common in Europe. The butterfly Family Lycaenidae in Europe is especially known for its variation in the Adults. One example is Lysandra Bellargus [Adonis Blue] which I have observed in England over a number of years. In the Adonis Blue there is much variation in the hindwing red spotting of the female especially between the spring and summer broods. Can we really compare situation of the European Lycaenidae to the Ornithoptera paradisea of New Guinea? The O. paradisea populations in the Arfak mountains occur in isolated pockets with a much a smaller number of individual butterflies. I believe the morphological differences are important in the arfakensis and chrysanthemum adults because both are very uniform in their appearance, with very little variation occurring. It is worth pointing out that whether they fly together or not there are no known hybrids between the two. Ornithoptera are well known to have produced hybrids in the wild but this usually occurs on Indonesian breeding farms. In the Arfak mountains the high elevation O. rothschildi occasionally pairs with O. priamus which usually occurs at lower altitudes and produces the hybrid known as akakeae. Of all the known paradisea populations in New Guinea the Arfak arfakensis and the chrysanthemum are probably the oldest and most distinct. It is indeed highly unusual to have two very different paradisea populations occurring within one range of mountains and this situation is not found elsewhere in New Guinea. Why is this?. If you believe in Evolution, the answer certainly lies there. For a better understanding Haugum and Lows outstanding Birdwing Monograph gives you a good idea. They state that the Northern, western, and southern areas of New Guinea were not faunal centers because in the Pleistocene era they were repeatedly flooded. The Vogelkop [Arfak] area of West New Guinea was practically cut off during high level sea phases. Because of this, the high mountains of this part of the New Guinea are believed to have been the ancestral home of a number of the Ornithotera and is highly likely the ancestral O. paradisea came from here and produced the two very different populations that today occur in the Arfak. As sea levels fell the general dispersal of the Ornithoptera began and then they spread and evolved over the rest of New Guinea. The different genitalia of O. arfakensis is another important factor to consider. A good example is the following episode. Until quite recently in Ireland there was thought to be one species of Wood White [Leptidea sinapis]. That was until a number of Entomologists compared the genitialia of a number of different Wood White populations and although the adult butterflies were identical they found there were in fact two different butterflies species. The New butterfly was found to be the European Real's Wood White[Leptidea reali]. Both are considered good species on account of their different gentitalia. Peter.
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Post by nomihoudai on Jan 24, 2013 15:08:33 GMT -8
Hello Peter, I have read your post and I do not understand a few things. They seem contradicting to me and therefore I ask if you could further elaborate this to me. Could you please cite the source of Haugham and Lows that explains about the flooding of large parts of Papua New Guinea during Pleistocene era? As far as my knowledge goes this was what we call the last ice age (what I was already talking about before) and at that time the sea levels where on a general low. All the water was trapped in the huge ice shields on the northern hemisphere and even Australia and Papua New Guinea where connected by one land bridge. Therefore I cannot understand where the huge water masses splitting off the Vogelkopp would come from. Towards your question if we can compare Lycaenidae and Ornithoptera I would reply that we can do so very well, but we have to keep in mind that Lycaenidae are much smaller in size. This smaller size will induce more vulnerability towards environmental factors along with a different mobility. If you take these factors in considerations you can do compare them very well, as a butterfly is a butterfly and will remain a butterfly. You can even enlarge the reasoning given here to other animals when taking into account at what pace they reproduce and how mobile they are. That Ornithoptera populations are smaller in size sounds like a wild guess to me, Ornithoptera are canopy flyers as far as I know, assessing their population sizes will be rather difficult but it will be large enough to sustain them. The last paragraph is not entirely correct. The different genitalia in Leptidea is no valid argument to split these species as long as there is not proven that this difference will and has to result in reproductive isolation. Of course the spitters where already happy when hearing about these arguments and where jumping in joy, but anyone not working with pseudoscience had to wait until better arguments where shown. The reason for the split in Leptidea is that there had been several behavioural experiments where they checked if both species will mate and they did not due to selection by the females. Also the situation is more complicated than what you describe here, please refer to an older post of me for this : insectnet.proboards.com/index.cgi?action=display&board=general&thread=1192&page=1I know that discussing with an old naysayer like I am is not always funny but I would really like to hear if you could elaborate on the flooding part. Rgds Claude
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Post by nomad on Jan 24, 2013 20:32:08 GMT -8
Hi Claude. Thank you for your comments. The information I have on the flooding of New Guinea came from the Haugum and Low Birdwing monograph. Sadly the authors have both passed on and there is no way to find out where all their sources came from but here is their explanation in more detail. They state " The snow-capped peaks constitute remnants of what was an extensive glaciation in previous geological periods. The geological history of the island is complex, and it has been subjected to rather dramatic changes in the distant past. Once connected to the Australian continent in the southeast, a volcanic island chain arc existed north of the island, possibly forming a bridge from the Moluccas to the Solomons."
"Sea level fluctuations have been found to vary from a level some 70 meters, or more, lower than the present level, sometime during the Pleistocene to phases, inter- and post glacial, when the sea-level was upwards of 100m, higher than the present day level [Zeuner1942,1943b]. The latter phases may be important in that they served to 'concentrate' and isolate the fauna, whilst the former periods are of particular zoogeographic importance in that they afforded species with means of dispersal by the land bridges provided."
"During the low-level phases, New Guinea and the Aru Islands joined to Australia, whilst in the west, New Guinea was, at least parly joined with the E. Moluccas. The present day distribution of the Ornithoptera, [perplexing as it may seem at first, owing to the anomalies present], must be considered with such condition in mind. It is also evident that islands and land areas that remained relatively isolated, exhibit distinct species-groups and subspecies, whilst islands and areas that have been flooded have relatively recent forms. The Vogelkop area of West Irian Jaya is such an 'isolated area and was practically cut off from the rest of New Guinea during high-level sea phases. It is to be considered as a centre of faunal dispersal, and is particularly important with regard to the Ornithoptera. The condition also serves to illustrate the great differences that exist between the species and forms of W. Irian Jaya and Papua; the majority of the western species and subspecies are 'older' than are their eastern counterparts and representatives, some of which are relatively recent."
" The great Central Ranges and the Onin peninsular are other areas, to be considered as 'faunal centres'. The Onin area was reduced to two small islands during high-level sea phases, and the representative of O. goliath which existed here became isolated from mainland populations. Subsequent changes allowed this particular form to spread back into the mainland, where it is now represented from the neck of the Vogelkop Peninsular [O. goliath atlas range is now known to extend into the Western Vogelkop] over the Onin area to the East Weyland Mountains, as ssp atlas. Such subsequent dispersals may have occurred in any direction, and, where a return dispersal took place, the situation may well be complex. This is, apparently, what happened in the Arfak Mountain district, where the subspecies are allopatric in altitude. The occurrence of three distinct subspecies of O. priamus on the Australian continent may also be interpreted in terms of subsequent dispersal.
"The western, southwestern, and the southern lowlands of New Guinea cannot be regarded as faunal centres, as they have been repeatedly flooded during the Pleistocene period. This condition also appplies to the Northern Coastal Ranges and the Great Central Ranges [Zeuner]. Thus the ancestral Ornithoptera, spreading from the Moluccas and perhaps the Vogelkop Peninsula, in reaching mainland New Guinea, spread towards the East, following mainly the Northern Coastal and remained, at least partially, separated from the Schoenbergia that spread from the Great Central Ranges." I would like to say again that I believe the O paradisea populations in the Arfak Mountains to be old isolated relic populations. The very detailed distribution maps in Matsuka show clearly that O. arfakensis and ssp chrysanthemum are limted to a few populations in just a few localities and both are not widespread in the Arfak Mountains. Thus even if their numbers were high here of which I can assure you they are not, hence the high price of specimens. Their numbers would not in anyway compare to the widespread Lycaena that you mentioned.
This what Haugum and Low have written in full on this matter, I hope this helps. Please also see the post below
Peter.
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Post by nomad on Jan 24, 2013 23:24:27 GMT -8
I have found this article from the Australian Museum on the Pleistocene period 1.6 millon to 10,000 years ago. This article agrees with Haugum and Low.
The Pleistocene.
" Sea levels fell, creating a temporary land bridge between Mainland Australia and Tasmania in the South and Australia and New Guinea in the North. Because of the waxing and waning of the poplar icecaps, the climate in Australia continued to cycle rapidly between Icehouse phases [cold, dry conditions] and Greenhouse phases [Warmer wetter conditions]. During Icehouse phases sea level fell, creating land bridges that would be then flooded during Greenhouse phases."
This may have happened as this article states a number of times and it was during the warmer [Greenhouse phases] that the release of water flooded the then lower lying New Guinea, leaving the Fauna, Ornithoptera [Schoenbergia] etc isolated on the New Guinea Western Mountain ranges such as the Vogelkop [Doberai Peninusular] .
Peter
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Post by Adam Cotton on Jan 25, 2013 7:28:59 GMT -8
Can I just point out that the Monograph of the Birdwing Butterflies was written by Haugum and Low (not Haugham).
Jan Haugum was a great modern lepidopterist, and is sadly missed by all who knew him.
Adam.
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Post by bobw on Jan 25, 2013 8:28:14 GMT -8
Jan Haugum was indeed one of the great modern lepidopterists. Those of you who knew him may be interested in the following extract from my preface to the book: "The genus Colias Fabricius, 1807 - Jan Haugum's annotated catalogue of the Old World Colias; Grieshuber, Worthy & Lamas, 2012":
Jan Haugum (1940-2002) is best known for his “Monograph of the Birdwing Butterflies” written with Andrew Low and published in 6 parts between 1978 and 1985. According to many specialists in the group, this remains the foremost book dealing with the Birdwings and is credited with initiating a massive boom in the study of these insects. Jan also ran a group called the “Lepidoptera Group of 1968” which published the long defunct but highly regarded journal “Papilio International”, and he was generally regarded as an authority on worldwide Papilionidae. He told me many years later that the reason he gave up his studies into Birdwings was the increasing legislation and consequent difficulty of getting new material. It is less generally well-known that his main interest since boyhood had always been Colias.
My first contact with Jan had been when I corresponded with him very briefly about Ornithoptera in the 1980s, but I knew nothing of his interest in Colias. After several years with no contact Jan telephoned me out of the blue in the early 1990s, having been told by Cliff Ferris, a Colias specialist known to both of us, that I was interested in Colias. This phone call led to a friendship and close working relationship that lasted until his untimely death on 30th May 2002. I first visited him at his home in Vejle, Denmark in 1997; this then became an annual event until his death. These visits usually lasted about 5 days over the Christmas period, and consisted of fairly intensive study of various aspects of Colias taxonomy all day, followed by an evening relaxing with a bottle and discussing almost everything under the sun. Jan had a very wide range of interests and was knowledgeable on many subjects – these conversations were always fascinating and entertaining. We were often joined for part of these visits by other luminaries of the Danish entomological scene such as Harish Gaonkar, Michael Fibiger and Mary Petersen.
Whenever I visited Jan, he would produce 3 huge ring binders with page after page of closely-typed text relating to his Colias studies. This was the original manuscript for his magnum opus which he maintained and updated over many years. During my visits, many pages became heavily annotated with his hand-written notes made during our discussions, and after I left he would laboriously update pages of the manuscript on his old manual typewriter. He used to take great delight in sending me sheaves of paper with new entries for comment and editing. For the last few years of his life the pace of his work increased dramatically when he began to use a computer and I showed him how to hold all this information and update it using MSWord.
As new taxa were described and new information became available, Jan continually updated this manuscript. I often jokingly said to him that if he kept updating it he would never publish it. His reply to this was that I would have to publish it for him after his death – sadly, this proved to be prophetic. This book is based on Jan’s original manuscript.
Bob
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Post by nomad on Jan 25, 2013 9:30:54 GMT -8
Adam, thank you I have corrected the Authors name. I should get it right, I bought the monograph as it came out in parts from E. W. Classey and I often read them. Bob, that was a fascinating story, I should very much liked to have met Jan Haugum and also Andrew Low, there would be so much to ask them about the Birdwing Butterflies.
Peter.
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Post by nomihoudai on Jan 25, 2013 16:51:04 GMT -8
Hello Peter, thank you for the further informations.
It still looks to me as if the information presented in the book does not reflect the current knowledge of the formation of Papua New Guinea. First of all the cited source is from 1942, one has to be very careful about geological information from those dates. Until the mid 50's the theory about plate tectonics was not set yet, therefore the observed data got interpreted wrongly in many cases. There probably was never any flooding of the Papua mainland, I would not know in any way where this water would come from. We are currently at a very high sea level as the ice caps are melting. If all of the ice caps are completely gone the water will not rise more than 10 meters (see current global climate change papers), therefore flooding of 100 meters are not imaginable. The current theories all point towards Vogelkop being formed separately with FakFak peninsula as islands. They later joined the Papuan mainland from the west.
This was a very interesting constellation which drew my attention at it. Although there had not been as dramatic things happening as you reported it Vogelkop will have played an important role in biodiversification if you assume that Birdwings came from mainland PNG towards this new region then the landbridge would have made a selection on the allele diversity of the present specimen.
I wanted to check my assumptions and set up a computer simulation, you can see the result here. I simulated a simple model where I had up to 150000 butterflies with initially 64 different allele combinations (RGB color values, 4 values for each, makes 4*4*4 combinations) which could move up to 17km and would reproduce. In every reproduction cycle the alleles with the partner got mixed and up to 30 offsprings could survive. It is a very very simplistic population model. 500 generations have been simulated.
You can see that Vogelkop gets populated by a single type of allel combination and no other populations can spread there afterwards. The same applies in FakFak peninsula but frequently the allele combinations takes a turn and switches to a completely different set.
Mr. Haugum may have been mistaken in interpretation of some data as the background assumptions have changed but I guess his identification of Vogelkop as an important biodiversification tool are true. Such things can always, and will always happen in scientific reasoning.
It has been a very interesting discussion
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