Butterfly mimicry rings – a case of natural selection?

Since Darwin’s time mimicry is presented as one of the best example of the efficiency of natural selection. Several species should have been shaped by natural selection to resemble or mimic dangerous or poisonous species. It is supposed that protected by their shape and coloration they deceive their predators. Thus mimicry confers them survival advantage. In many cases mimicry is believed to be found among butterflies where palatable species mimic unpalatable ones (so called Batesian mimicry). In some cases two or more unpalatable species look alike. In this case they should be protected more effectively because their predators learn to avoid them only once. This is called Müllerian mimicry. And in some cases there is a whole bunch of Batesian and Müllerian mimics that look alike. This is called the mimicry ring.

One of the most famous mimicry rings are so called Heliconius mimicry rings from South America. According to M.Joron species 1-16 form “tiger” mimicry ring: heliconius1

1,Eueides isabella; 2,Heliconius pardalinus; 3,H. hecale; 4,Melinaea menophilus; 5,Tithorea harmonia; 6,Chetone histriona; 7,Napeogenes larina; 8,Mechanitis lysimnia; 9,Mec. polymnia; 10,Mec. mazaeus plagifera; 11,Ceratinia tutia; 12,Hypothyris cantobrica; 13,Dismorphia amphiona; 14,Eresiasp.; 15,Pterourus zagreus; 16,Consul fabius

According to M.Joron and J.Mallet (1998) who discussed the tiger mimicry ring :

Visual mimicry is a textbook case of natural selection because it is both intuitively understandable and has repeatedly evolved in a range of organisms: it is the ultimate example of parallel evolution. In many mimetic groups, particularly butterflies, a huge variety of colour patterns has arisen, even in closely related species.

This is a similar picture of the tiger mimicry ring by Seward, A. C. in “Darwin and modern science” (1909). See species 1-8:

heliconius2

EXPLANATION OF FIGS. 1-12.
Figs. 1-4 represent a Mimicry-ring from Eastern Brazil composed of four immune species belonging to three different sub-families and four different genera.

Fig. 1. Lycorea halia (Danainae).
Fig. 2. Heliconius narcaea (eucrate) (Heliconinae).
Fig. 3. Melinaea ethra (Ithomiinae).
Fig. 4. Mechanitis lysimnia (Ithomiinae).
Figs. 5, 6. Perrhybris pyrrha, male and female, S. American “Whites” (Pierinae). The female mimics immune Ithomiines, while the male shows only an indication of the mimetic colouring on the under surface.
Figs. 7, 8. Dismorphia astynome, male and female, also belonging to the family of S. American “whites,” and mimicking immune Ithomiines; a white patch on the posterior wing of the male and another on the corresponding surface of the under side of the upper wing, remain as traces of the original “white” coloration.
Fig. 9. Elymnias phegea, W. Africa, of the sub-family of Satyrines, mimics the succeeding species (Fig. 10).
Fig. 10. Planema epaea (gea), an immune West African species belonging to the Acraeinae.
Fig. 11. Danaida genutia, an immune Danaid from India, Burmah, etc.
Fig. 12. Elymnias undularis, female, one of the mimics of Fig. 11.

In the text we can read that “The strongest of all proofs of the theory (e.g. of origin through selection), however, is afforded by cases of true mimicry,… we can hardly hope to find more convincing proof of the actuality of the processes of selection than these cases put into our hands.

The picture might be of interest, because it was used also by entomologist Franz Heikertinger, a strong critic of Darwinistic natural selection. It can be found in his book “Das Rätsel der Mimikry und seine Lösung: eine kritische Darstellung des Werdens, des Wesens und der Widerlegung der Tiertrachthypothesen.” (1954).

Heikertinger_env

Notice that he put the word “Mimikryring” between quotation marks:

Heikertinger_tafel_IX_2

I have highlighted two species: Perrhybris pyrrha., male and female by red and Dismorphia astynome male and female by blue. Males of Perrhybris Pyrrha are white, whereas males of Dismorphia astynome have white spots on their hind wings. The question is why natural selection has not acted on them. Males of other species in the mimicry ring look like females e.g. protected. Yet we can see that some species may thrive with males not being protected at all:

pyrrha_mf
So called “female limited mimicry” – dimorphic female (top) and male (bottom) of Perrhybris pyrrha.

Franz Heikertinger strongly criticized those who saw natural selection as the cause of the similarity in the mimicry ring. He named natural selection as “die Hypothese” and the proponents of such a view as “Hypothetiker”. According to his opinion several questions must be answered before natural selection can be accepted as a plausible explanation of the phenomenon.

1) What are butterflies vision-oriented predators that select them? Do they exist?
2) Are unpalatable insects rejected by predators?
3) Are those rings maintained by natural selection or is the similarity between the species just a consequence of their relatedness?
4) Is such a coloration something that is beyond natural variation of the species?

Needless to say that Heikertinger answered to the all previous questions negatively:

Ad 1) and 2) These questions had been discussed by Heikertinger very thoroughly and repeatedly in many of his publications. He had dismissed birds as the predators of butterflies. I hope I will summarize his arguments in another place. In the case discussed I just quote J.Mallet (1994):

Heliconius(especially the erato taxonomic group) are renowned for roosting gregariously; and co-mimics roost gregariously with each other more often than with non-mimics. Gregarious roosting is therefore common between species, as well as within species.The paradoxical correlation between nocturnal roosting and visual mimicry is presumably explained by bird predation at dusk when roosts are forming, or at dawn before they have disbanded. Direct evidence of predation is lacking, but there are high rates of disturbance by birds at these times.

Ad 3)
It is important to notice that mimicry is supposed to exist only between unrelated taxa. No one will be surprised seeing two closely related species looking alike (for instance tiger and leopard, family Felidae). There are plenty of cases where butterflies of non-overlapping and distant regions look similar. Their similarity is explained by their relatedness and not by convergent evolution driven by natural selection. Even though natural selection is sometimes mentioned as well:

Hypolimnas_missipus
Hypolimnas misippus (Nymphalidae), “model” from Southeast Asia

limenitis
Limenitis albomaculata (Nymphalidae), “mimic” from West China

According to Edward Poulton the selective agent here are some unspecified migratory birds. Even Komárek (2003) named such an explanation something between a folk tale and “positive science”.

The species in the tiger mimicry ring belong predominantly to family Nymphalidae (and especially to subgroup Ithomiinae). Two exceptions have been already mentioned: Dismorphia astymone and Perrhybris Pyrrha (species with non-mimetic males). Dismorhiinae is subgroup of family Pieridae (“Whites”). Yet Heikertinger refutes such a categorization. In his view categorization based on tarsal claws and numbers of segments on butterfly forelimbs might be misleading. He noticed that we often observe reduced number of segments in butterfly males (as a consequence of Eimer’s Orthogenesis. Theodor Eimer claimed that males are more “developed” than females, which tends to retain original features of the species). Such a categorization would lead to two different groups with males and females separated. Heikertinger compared venation on wings and concluded that according this feature Dismorhiinae are more related to Nymphalidae (the base of the mimicry ring) than to Pieridae.

Heikertinger_Tafel_II
In the first row is the similarity between veins of 22.Dismorphia arsinoe and 23.Mechanitis polymnia (Ihtomiinae). The last row are veins of Perrhybris malenka (Pieridae).

In the case of Perrhybris Pyrrha Heikertinger quoted Eimer, Piepers, Dixey and Van Bemmelen that the ancestor of the species was not white (as it might be commonly believed), but varicolored. Consequently the supposed transition of P.pyrrha into varicolored mimic somehow loses its mysteriousness. On the other hand there isn’t any pattern on P. Pyrrha wings upon which natural selection could act towards the mimicry pattern. Quite the opposite – the males of P.Pyrrha have left or receded from the common pattern in the mimicry ring – despite the supposed unified pressure of natural selection. He refers to Eimer observation that females in Pieridae are often colored whereas males are not – Pieris brassicae, Harpaenia eriphia and others.

Perrhybris_pyrrha
Perrhybris. Heikertinger quotes Dixey according to whom the transition goes from males of P.locusta. P.phaloe, P.lypera, P.lorena to P.pyrrha (some of them by red arrow). Those forms make a connection but there is nothing that could resemble Ithomiinae and consequently to be of a selective value upon which selection could have acted.

Ad 4)
This is a point which Heikertinger often addresses. He recommends to readers to open an atlas of butterflies and compare related species. There is an abundance of varicolored butterflies in each taxa. Heikertinger claims that picking up just one species from a series doesn’t make case for natural selection. A resemblance can be explained by Vavilov’s homologous sequences as well. In the case of the tiger mimicry ring he recommends to open Seitz’s Butterfly atlas volume V and see that the mimicry pattern is actually recurring in many variants across many taxa.

One should compare all the fantastic coloration and patterns of Heliconius (family Nyphalidae) with tiger mimics – table 75 -78 from Seitz atlas volume V:

Heliconius_Seitz_78

Heliconius_Seitz_77

Heliconius_Seitz_76

Heliconius_Seitz_75

and see the discussed recurring type also among Actinote:

Actinote_81

Actinote_82

or the same type on the table 91, 92 the row C:

Slide2

Slide1

Table 103, the row E:

cyklogramma

Table 143, the row B:

Aricoris

or Phycoides Table 90, the row h:

Phycoides

Dismorphia, table 30:

Dismorphia

and others.

According to Heikertinger the coloration and pattern types on insects are often recurring phenomena. The resemblance repeats. Why there should be a different explanation for some arbitrary chosen resemblances than for thousands of other patterns and coloration?

—————————-
Literature:

Theodor Eimer: Orthogenesis der Schmetterlinge (1897)

Franz Heikertinger: Das Rätsel der Mimikry und seine Lösung: eine kritische Darstellung des Werdens, des Wesens und der Widerlegung der Tiertrachthypothesen. (1954)

Adalbert Seitz: Die Gross-Schmetterlinge der Erde (1906-1928)

Mathieu Moron: Mimicry (2003)

http://heliconius.zoo.cam.ac.uk/joron/joron02mim.pdf

Mathieu Joron, James L.B. Mallet: Diversity in mimicry: paradox or paradigm? (1998)

http://www2.yk.psu.edu/~mph13/diversity%20and%20evolution%20mimics.pdf

James Mallet: Why are there so many mimicry rings? Correlations between habitat, behaviour and mimicry in Heliconius butterflies.(1995)

http://www.ucl.ac.uk/taxome/jim/pap/Mallet%26Gilbert95.pdf

Michael Braby, Roger Vila, Naomi Pierce: Molecular phylogeny and systematics of the Pieridae(Lepidoptera: Papilionoidea): higher classification and biogeography. (2006)

http://oebweb-prd.oeb.harvard.edu/faculty/pierce/publications/pdfs/2006_Braby_et_al.pdf

Stanislav Komarek: Mimicry, aposematism and related phenomena. Mimetism in nature and the history of its study. (2003)

http://stanislav-komarek.cz/mimicry/

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19 responses to “Butterfly mimicry rings – a case of natural selection?

  1. MacBride also mentions the experiments by Harrison:

    “Professor Harrison proved that saw-flies after a period of years could be trained to lay their eggs on other species of plants than those to which they are accustomed, and that when these eggs are allowed to develop into saw-flies the flies emerging from them will choose the new plants to visit instead of the old ones proper to the species, even when the opportunity to visit both is provided.”

    Similar to above, it is very hard to locate many sources about these experiments. Stuff like this has been ignored sadly whilst most of the attention by the scientific community goes to the flawed and over-hyped natural selection scenario.

  2. You may be interested in the book

    http://archive.org/stream/evolutionadaptat00morg#page/n5/mode/2up

    Adaptation and Evolution by Thomas Hunt Morgan an early anti-Darwinian geneticist.

  3. In the preface to the book, Morgan admits that Darwinism is a dogma.

  4. I have just been reading a book by Brian Leith called “The Descent of Darwin: A Handbook of Doubts about Darwinism”.

    Here is a quote from it:

    “Another philosophical question regards the very definition of the word ‘selection’. One of the original formulations of selection was ‘the survival of the fittest’. If you open a standard textbook of genetics ‘fitness’ will probably be defined as ‘the ability to survive’ or something similar. But if the ‘fittest’ are defined as ‘the best survivors’ then the idea of natural selection becomes ‘the survival of those best at surviving’. So what else is new? If there is no more to Darwinism than a truism then the whole theory rests on very shaky ground.”

    He also writes:

    “The philosophers have another bone to pick with evolutionary theory, one that has haunted Darwinism for a hundred years: is the idea of natural selection a tautology … A tautology is the saying of something twice over in different words, and is therefore either a nonsense or a statement which is so self-evident as to be meaningless. The statement ‘several bachelors who were not married were at the meeting’ is nonsense because bachelors are unmarried, while the sentence implies that they are not… For a scientific statement to avoid being tautologous, therefore, it must propose some relationship in the world that is testable by experiment. The problem of tautology in Darwinism is a subtle one. It hinges on the definitions of a few crucial words: ‘the survival of the fittest.’ This is the central claim that Darwin made that only the ‘fittest’ succeed in a struggle for ‘survival’. If this basic statement does not tell us anything new about the outside world then the whole of Darwinism is in deep trouble. Unfortunately the senses in which these words are often used by biologists do turn the statement into a nonsense. If you turn to a textbook of genetics in search of a definition of ‘fitness’ you will find something like this: `The genotype with the largest survival rate is defined as the fittest … ‘ So the central statement of Darwinism, ‘the survival of the fittest’, becomes: ‘the survival of those creatures having the largest survival rate’! Immediately the problem is clear if you define fitness as ‘the ability to survive’ then the ‘survival of the fittest’ becomes a tautology, a self-evident bit of trivia. In this form the statement doesn’t tell us anything about the outside world that we didn’t know already. It doesn’t, for example, enable us to predict which members of a population will survive and reproduce, since we cannot measure survival until afterwards. In this sense the neoDarwinists must avoid a sloppy attitude to their theory or it will turn out to say nothing.”

  5. I brought an old booklet entitled “Evolution” by Ernest MacBride (1927). As far as I can see this booklet is quite rare and does not appear anywhere on the internet.

    I have typed up these paragraphs from the book that may be relevant;

    “A very popular idea is that Darwin has completely explained evolution and that the hypotheses of “natural selection” and “sexual selection” successfully account for all the varied peculiarities of structure and function which we see around us. A little critical consideration will, we think, convince our readers that, so far from this being the case, Darwin’s theory is in reality no explanation at all, but one great and striking instance of the common illusion of what one of our best contemporary philosophers calls “reification” of words – i.e., the conversion of mere general terms into imaginary things.

    That every species does in reality produce far more young than can survive is open to no possible kind of doubt. One simple instance of this will suffice. The common thrush lives on an average ten years. It begins to breed at the age of one year, and produces every summer two broods of nestlings, each consisting of four young. In the course of their lives, therefore, a single pair of thrushes bring into the world eighty young, and some of these may be breeding for nine years before the parents die. Since the whole population of thrushes remains about the same from generation to generation, it is obvious that only two of this vast army of children, grandchildren, and great grandchildren can survive their parents; all the rest come to an untimely end.

    But to put the matter in a nutshell, the fact that James is killed can make no difference to the structure of Tom.

    The implicit assumption in Darwin’s hypothesis is that continuous inheritable variation occurs constantly in all directions. But to assume this is really to beg the whole question. Variation is not a single thing but a collective term for a whole lot of different things. Continuous variation in any direction is evolution, and it is precisely this which has proved to exist, and, if possible, to be explained. Natural selection is the pruning-knife which trims the buds of the tree of life, but it does not account for the sprouting of the buds nor for the directions in which they tend to grow.”

    MacBride was an Irish marine biologist and zoologist. I look forward to checking out his other books. He was a supporter of Lamarckian evolution.

  6. Fleeming Jenkin wrote a critical review The Origin of Species in 1867.

    http://en.wikipedia.org/wiki/Fleeming_Jenkin

    Jenkin raised some serious objections to natural selection. You can read some of his objections here:

    http://www.viu.ca/liberalstudies/program%20info/outlines%20(current)/jenkin.asp

  7. “The Darwinian Theory of the Transmutation of Species” by Robert Mackenzie Beverley.

    The book was written in 1867. In the introduction of the book he writes:

    “M. Flourens has published a short answer to Mr Darwin, contenting himself chiefly with a pointing out the abuse of terms, and the verbal inaccuracies with which the Origin of Species is argued. The answer, as far as it goes, is very effective, and successfully assails the foundation of the Theory; but it is to be regretted that a writer, so well qualified for the task, should have confined himself chiefly to one view of the subject.”

    The entire book is a criticism of Darwinism. This is a rare book that seems to have been totally forgotten about.

    See his chapter IV “Natural Selection”. Here are some of his comments:

    “After all that has been said on the subject, it is to be hoped that the eyes of the reader will not be blinded with the dust of words by which this theory is made to push its way. Natural selection is, as a fact, absolutely nothing, there is no power or intellect to select anything, and nothing is selected.”

    “Natural selection, therefore, we affirm is a term to be utterly discarded. It is a verbal deception, and the only substance to be discovered, after the elimination of the metaphor, is that a certain series of events is said to have taken place, though those events are contested and denied.”

    He has much valid criticism of natural selection, and he talks about Darwin’s abuse of language. You can read it here and download it for free:

    http://www.archive.org/stream/darwiniantheory00bevegoog#page/n7/mode/2up

  8. Another forgotten critic of Darwinism:

    Emanuel Rádl

    http://en.wikipedia.org/wiki/Emanuel_R%C3%A1dl

    “We may therefore sum up the modern position in Driesch’s words: ‘For those with insight Darwinism has been dead for a long time’… Darwinism as a tyrannic doctrine, which imperiously enchains the minds of men, is dead.”

    —Rádl. The History of Biological Theories. (1930). p. 388

    His book can be found online here:

    http://archive.org/stream/historyofbiologi029197mbp#page/n5/mode/2up

  9. This website must have got stuck in 1914.

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