Final Version May 25, 2003

DISCUSSION PAPER SERIES

June 2003

No. 0302

The Recent Controversy in Japan on

Evolutionary Studies in Reference to

Explosive Speciation of the Cichlids

in Lake Victoria：Focused on Species

=Societal Theory of Evolution

Kawamiya, Nobuo *

Explosive speciation of the cichlids, elongation of giraffe's neck, and peacock's ocelli feathers are the Gordian knots for the extant theories of evolution. This paper provides a consistent solution for all these problems by the Species=Societal approach to theory of evolution introduced by K. Imanishi and developed by S. Mizuhata.

*Correspondence address: Department of Economics, Chukyo University,

101-2 Yagoto-honmachi, Showa-ku, Nagoya 466-8666 JAPAN

e-mail: [e-mail]

The Recent Controversy in Japan on Evolutionary Studies in Reference to Explosive Speciation of the Cichlids in Lake Victoria：

Focused on Species=Societal Theory of Evolution
Kawamiya, Nobuo (School of Economics, Chukyo University)

　　 Abstract

§1 Explosive speciation of the cichlids and limitations of extant theories

§2 The recent controversy in Japan on evolution theory

§3 The Neo-Imanishi theory of evolution on the Species=Society concept

§4 Giraffe's long neck explained by Species=Societal adaptation

§5 Diverse ways of mating explained by Mating Priority Rule (MPR)

§6 Peacock's ocelli feathers to imply non-adaptive evolution

§7 Mate selection: female choice or Species=Societal consensus?

§8 Evolution of the cichlids explained by the Neo-Imanishi theory

§9 Further application of the MPR and the SS partition approach

§10 Modes of evolution and the evolution of evolutionary modes

§11 Reinvestigating concepts of Species and Species=Society

§12 Methodological notes

Concluding remarks

Abstract

Evidently, (a) explosive speciation of the cichlids, (b) elongation of giraffe's neck, and (c) peacock's ocelli feathers are the Gordian knots for the extant theories of evolution. Because: (a) occurred while the species have completely retained their genetic identity or zygotic homogeneity; whereas (b) seems to have been a typical adaptive evolution there has been found no fossil showing intermediate neck length; and all efforts to attribute (c) to adaptive evolution resulted in illogical inferences. The recent discussion concerning these problems in Japan seems to reveal that the fiction of Action Controlling Genes is nothing but Alchemy of Today. In contrast a consistent solution for all these problems can be obtained by the Species=Societal approach to theory of evolution introduced by K. Imanishi and developed by S. Mizuhata.

§1 Explosive speciation of the cichlids and limitations of extant theories

Observation on the haplochromine cichlids in Lake Victoria (LV) shows the following facts and the interpretations thereto:

1) The cichlids in LV seem to have speciated into more than five hundred species in less than 12500 years [Goldschmidt 1996, 244a]. On an average, a new species appeared continually in every 2-3 generations. During this period, effective gene mutation of the cichlids leading to speciation must have been negligible because the replacement of amino acid in molecular evolution is confirmed to be very slow and stable [Kimura 1988, 202]. Some reports suggest that the cichlids’ evolution might have started earlier than 12500 years [Verheyehn et al. 2003]. Yet they do not refute that the major diversification occurred in the present LV.

2) Morphological divergence of the cichlids has been enormous, although　 the structure of their genoms seems to have remained unchanged. Interspecific hybrids are fertile and fully viable [Seehausen 1997a, b; 2000]. According to Mayr, “A completely consistent scenario can be inferred without any reference to the genic basis” [Mayr 2001]. In fact massive hybridization occurred when the lake water turned muddy and mate discrimination by body colors became difficult.

3) W. Dominey and other researchers attributed the cichlids' explosive speciation to disruptive selection by females on male characteristic variants (especially in body colors) [Dominey 1984]/ [Goldschmidt 1996]. Their observation and interpretation cannot be refutable. However, the importance of female choice does not necessarily imply that the hypothesis of “female choosy gene” (FCG) is tenable.

4) Consequently, in the cichlids' speciation, populations having identical genoms started mate choices oriented toward different directions, resulting in reproductive isolation. And then each population began to prefer a different niche or feeding habit [Seehausen et al. 1999]. This process must have occurred frequently.

5) No extant theory can explain this remarkable divergence without any inconsistency: both the runaway and the indicator hypotheses are based on individual gene mutation and the change in mutant gene frequency in the population. Even if, however, the “choosy gene” should really exist, it cannot mutate and diverge so rapidly in only 125 centuries.

§2 The recent controversy in Japan on evolution theory

The said cichlid problems ask for a fundamental reinvestigation of extant theories of evolution. What is a species? What is speciation? And what is evolution? There have occurred interesting discussions in Japan concerning these topics including the cichlid case, which deserve international recognition.

Firstly the cichlid problems concern the species concept. Paterson's definition based on conspecific recognition will be appropriate for this case because one has to discriminate genetically identical populations [Paterson 1985].

To Define species by conspecies recognition is not a tautology because one can verify the extent of “conspecific recognition” by means of observation. This definition, however, can apply only to animals having the brain: though this limitation is not defective but advantageous as discussed later on.

K. Imanishi had proposed a more advanced concept on conspecies recognition. According to him [Imanishi 1941, Ikeda & Sibatani 1994]:

1) Individuals of a species not only share mutual conspecific recognition but also cooperatively obey “the law of species” to live the identical life type.

2) Thereby they form a Species=Society (in short, SS). Imanishi proposed a term “specia” for “Shu(Species)＋Shakai(Society)”. The term is, however, not appropriate because the SS should encompass both species and society.

3) All the species=societies form the total biotic community in concert.

4) A new species population would evolve while maintaining their unity as a species=society and interacting with the total biotic community [Nature 1985-87].

5) Speciation occurs as a splitting or partition of species=society, or “Sumiwake” as called by Imanishi [Imanishi 1980, 1986]. @1
@1: “Sumiwake” has been translated as “habitat segregation”. This translation is no good because it sounds similar to geographical isolation. Ikeda and Sibatani renamed it as “lifestyle partition” [Ikeda & Sibatani 1995]: In this paper, we further rename it as “species=societal partition” after Imanishi's original idea.
His theory had a clear advantage to other evolutionary studies because the concept of species=societal partition, for describing speciation, enables to explain both the continuity of a population as a reproductive community and the discontinuity by reproductive isolation in species bifurcation. No genic theory has ever given a consistent explanation of these continuity and discontinuity during speciation.

He compared gradual (intraspecific) evolution and intermittent partition (speciation) of a species=society to a history of a language family [Imanishi 1980], where a continuous diachronic development causes a discontinuous synchronic diversity. It can be a reasonable analogy because both language and conspecific recognition are kinds of cerebral program.
A population going to form a new species would come to acquire a new life type while retaining their mutual conspecific recognition intact. They renovate their life type without hurting parental/filial and mating relation. So this partition is, essentially, an intentional and cooperative process and not an unintentional (incidental) and individual process. Imanishi gave a clear account for discontinuity of speciation and continuity in reproduction [Imanishi 1980; 1986], which had been an insolvable Gordian knot in the genic theory of speciation.

Meanwhile, Imanishi's original theory of species=society comprised a serious fault: He estimated the homogeneity of individuals within a species so highly that he came to deny that competition among morphological variants could be vitally important in evolution. This fault prevented him from further consideration of evolutionary mechanisms based on his species=society concept. Due to this difficulty, most of Imanishi's numerous disciples deserted him without trying to revise and improve his theory. And instead they sealed it as an embarrassing and untouchable “taboo”.

Recently S. Mizuhata published “SHIN-IMANISHI SHINKARON” (The Neo-Imanishi Theory of Evolution) [Mizuhata 2002], where he proposed an improved theory on species=societal aspects of evolution. He successfully eliminated the said theoretical fault of Imanishi's, while retaining and developing the species=societal theory of evolution. However, the Neo-Imanishi theory of evolution had to suffer an almost unanimous refusal by Japanese evolutionists and biologists because it proposed a re-evaluation of Imanishi's Theory, disclosing the once buried taboo. In fact they seemed as if detesting any discussion to remind Imanishi.

Such a situation, however, is now being mitigated owing to a fair treatment to the new theory by several well-known biologists, such as T. Hidaka, Y. Kishi, T. Yahara, etc. Especially T. Hidaka, once a crucial critique of Imanishi's theory, proposed re-evaluation of Imanishi's evolutionary thought in response to Mizuhata's new theory, which startled Japanese biologists [Hidaka 2003].

Improvement of Imanishi's theory by Mizuhata lies on the following:

Imanishi did not recognize the vital importance of competition among morphological variants because he respected equality of individuals in a species very highly and, besides, he favored orthogenetic view of evolution. Mizuhata found that mate selection is vitally important in evolution and that the mate choice system is essentially of species=societal nature, distinct from the traditional concept of sexual selection.

This paper tries to introduce this new theory and thereto relevant discussions in anticipation that world biologists and evolutionists would treat these studies without prejudice, although Imanishi's reputation may not be so good also in the West except among some anthropologists and ape researchers. Anyway the author considers that the species=societal approach to evolutionary study will soon make an indispensable methodology. Moreover the lifework of Imanishi “SEIBUTU NO SEKAI” (the World of Living Things) was published in English as a result of assiduous translation by P. J. Asquith et al. [Imanishi 2002].

§3 The Neo-Imanishi theory of evolution on the Species=Society concept

Improvement of Imanishi's theory by Mizuhata lies on the following:

Imanishi did not recognize the vital importance of competition among morphological variants because he respected equality of individuals in a species extremely highly and, besides, he favored orthogenetic view of evolution. In contrast Mizuhata found that mate choice is vitally important in evolution and that the system of mate choice is essentially of species=societal nature, distinct from the traditional concept of sexual selection.

Main points of Mizuhata's theory are:

1) “Sumiwake” (Imanishi) or “species=societal partition” is a key concept to solve the speciation problem: this is a reconfirmation of the original Imanishi theory. @2

2) The origin of species=society (and its partition) traces back to the Cambrian Great Explosion and the appearance of cerebral animals. Species does not have any origin but only has bifurcation (partition) with sibling species. The true origin of species (not of species=society) traces back to the first appearance of life.

3) Mizuhata redefined the species=society as populations that share the identical brain program, or “the species=societal software”, having the dual objectives: individual survival (viability) and achievement of mating (reproduction). It is this cerebral software that governs ethological behaviors of animals. Also beware that this software acts as a self-organizing information processor. @3 This concept is not so unorthodox because it corresponds to: what Darwin called “Mental Powers” of animals; what Lorenz called “Releasing Mechanisms”; or, what Imanishi called “Subjectivity/ Autonomy” of animals [Mizuhata 2002, 146].
@2: Imanishi supposed that all species (including non-animals) should have their “species=society”. His supposition was, however, inappropriate.

@3: Self-organization of the brain program was assigned firstly to learning (ontogenetic) alone [M 2002] and then also to “down-loading” [M 2003a], of which the latter implies formation of standing memories in the brain besides learning.
4) Splitting of a species=society occurs when a population of the original (parent) species comes to change (modify) its species=societal software. If the change occurs in the way of mating, it will directly lead to reproductive isolation and species=societal partition (speciation), where mating occurs in accordance with a proper-to-species rule: male's “priority” for mating firmly concerns a species-proper character of males, e.g., long necks of giraffes, ocelli feathers of peacocks, antlers of deer, nuptial colors of fish etc. This mating rule is named “Mating Priority Rule (MPR)” [Mizuhata 2002, 111].

5) Besides adaptation, there is another motive force for evolution: struggle to reinforce one's own species=societal unity and identification (or differentiation) against close kin species. This motive force is named species=societal coherence (SSC) [Mizuhata 2003]. The SSC explains the true cause of “character displacement”, which implies that a rapid evolution occurs in differentiation of close kin species.

This character displacement has been interpreted in terms of adaptive evolution: where one has attributed the accelerated evolution to enhanced “struggle for life” among close kin species. In fact the rapid cichlids' speciation seems to have been driven not by struggle for life but by struggle for species identification and differentiation. When the cichlids' struggle for life became extremely hard due to Nile perch and lake water deterioration, it resulted in disruption of the specific diversity instead of accelerating speciation.
The evolutionary mechanism of (4) and (5) are the newly found aspects in the species=societal theory of evolution, which explain a mechanism (or dynamics) governing evolution. This implies a fundamental improvement on the former Imanishi's theory that lacked any mechanism of evolution.

Thus the Neo-Imanishi theory of evolution attributes the cause for species=societal partition to the following:

1) Evolution for species=societal adaptation by which a population in the precedent species comes to seek for a new niche or life type. Beware that this is an intentional and cooperative activity, distinct from the Darwinian process of adaptation due to natural (stochastic) selection, where “Intentional” means “through brain activity”.

2) Evolution for species=societal coherence by which a population in the precedent species comes to set up a new species=societal symbol, making it a new criterion for mate selection (or MPR). In the cichlids, newly appeared male variants in body colors and patterns frequently got adopted as a new species=societal symbol, resulting in species=societal partitions (i.e. speciation).

Also beware that these factors imply not the results but the cause of evolution. The two moments, SS adaptation and SS coherence, are mutually interactive. Which to prevail depends on occasion. In cichlids, it is confirmed that the second proceeded in advance [Seehausen et al. 1999]. Combination of the both factors is possible as seen in the seeking for long necks in the species=society of giraffe (see below).

§4 Giraffe's long neck explained by Species=Societal adaptation

A long neck of the giraffe is “adaptive” for eating acacia leaves, their most favorite food. Darwin firmly believed that this must be the result of natural selection [Darwin 1996]. However, there is no fossil evidence for intermediate neck length to show a gradual elongation of giraffe necks. Transition from the short-necked proto-giraffe to the present type must have been too fast to leave any trace of “intermediate” giraffe necks.

Simmons and Scheepers attributed neck elongation of the giraffe to sexual selection [Simmons & Scheepers 1996]: however, they denied an adaptive aspect of the long neck without sufficient reasoning. Mizuhata explained the rapid and adaptive elongation of giraffe necks as an adaptive evolution driven by the MPR, i.e. species=societal motive force [Mizuhata 2002, 41].

　 Progenitors of the giraffe (a kind of Paleotraguinae) were a ruminant like the okapi: they were short-necked and lived in thick forests. A population of this proto-giraffe moved out of forests into savannas, seeking for a new life type: this implied a start of species=societal partition from the proto-giraffe that had remained in the forests. Ancient giraffes having moved to savannas had to change and adjust their life type toward the new environment. They became especially fond of acacia leaves, delicious and eutrophic, where long necks were effective to reach wide range of acacia trees. Thus they began to want having a longer neck and this propensity began to control mate choice (till this time this population might have remained conspecific with the former species) [Mizuhata 2002].

Meanwhile, ardent seeking for long necks resulted in the species habit of “mating contest” among males to compete for their neck length and thus for their priority to mate with females. When this “Mating Priority Rule” came to be shared by all the populations living in savannas, their reproductive isolation from the former species must have been completed. This is a case of speciation through SS partition, as first proposed by Imanishi. Further the mating contest became increasingly fiercer and giraffe males began “neck combats” using their neck and skull like a hammer. The impact of neck blows became ever serious, even lethal, as their neck got longer [Simmons & Scheepers 1996].

It is easy to understand that the MPR, once established, would greatly accelerate evolution. In fact, superior males monopolize the mating priority and females reject courtship of shorter necked males. In contrast to the former theory of Imanishi, the MPR theory is consistent with population genetics: the MPR will reduce the effective population size, raise the inbreeding coefficient, and halt the Hardy-Weinberg law: all, in concert, will have effects to accelerate evolution, as is well known in the ordinary theory of population genetics.

§5 Diverse ways of mating explained by Mating Priority Rule (MPR)

　 All the animals have a species-proper way of mating, identifying the conspecific mate of the species=society. Most insects, birds, fish etc. have a proper procedure for mating courtship and consent, whether selective or monogamic. Rather, one may say that the existence of species and species=society lies on having a species-proper way of mating [Hidaka 1966].

There is a wide variety of mating processes in species=societies, from monogamy to holding a large harem. For example, In leks of the Sage Grouse (Centrocercus urophasianus) or harems of Elephant Seal (Mirounga), a superior male mates with more than one hundred females, excluding most fellow males from mating. A species=society of this type seems to be in an “Enhancement Mode” of MPR. When the morphological evolution turns into a saturation, the species=society will enter into a “Relaxation Mode” of MPR, and then probably into an “Equilibrium Mode” of MPR, such as monogamy.

A species=society now having an enhanced MPR may change its mode, resulting in a relaxed MPR. In turn, another SS now having a stationary MPR might have had an enhanced MPR. There seems to be a general trend that characteristic differences between males and females may be large in the enhancement mode, diminishing in the relaxation mode, and disappearing in the stationary mode. For example, the present MPR of giraffe's SS is considered to be in the relaxation mode and therefore its sexual dimorphism has been sufficiently reduced [Mizuhata 2003a] though still existent [Simmons & Scheepers 1996].

What causes the diversity in ways of mating? According to Mizuhata, this diversity can be explained in terms of MPR and its diachronic modal transition: each species=society itinerates diachronically through the said several modes of the MPR. If one takes a synchronic view (cross section) of the distribution on various MPR modes, it will show a wide variety [Mizuhata 2003a].

The relation between synchronic variety and diachronic shifts of mating modes are as follows:

First a population in a SS begins to strongly cherish a special male character such as long necks, splendid tail feathers, etc. and makes it their species sex-attracting symbol. This implies a beginning of SS partition from the original (parental) SS. Then the population will enter into an enhanced mode of MPR and perform a fierce mating contest, resulting in accelerated character evolution. However, this evolution will in time reach saturation and the MPR will enter into a relaxation mode, where the mating contest may continue with reducing intensity. Finally the MPR will turn into a next equilibrium mode [Mizuhata 2003a].

The modal transition will occur in proper order. However, it may depend on occasion in which mode each species=society now resides. Therefore a synchronic observation will correspond to an incidental distribution of MPRs into various modes: equilibrium, enhancement, relaxation, or transitional. Thus one can confirm that the synchronic diversity of MPRs represents a projection of diachronic transition through various MPR modes. This gives a solution for the riddle why ways of mating is so diverse, which is an interesting result achieved by the Neo-Imanishi theory [Mizuhata 2003a].

§6 Peacock's ocelli feathers to imply non-adaptive evolution

T. Hidaka confirmed that peacock's ocelli feathers couldn’t be explained as a result of adaptive evolution [Hidaka-Mizuhata 2003]. His confirmation is grave because numerous evolutionists have been trying to explain the ocelli feathers of peacocks as indicating viability [Cronin 1991]. Hidaka assertion implies that the handicap hypothesis or its revision, the indicator hypothesis, is not an achievement in the evolution theory. Rather, these theories might be a product of psychological obsession of evolutionists to attribute every evolution to adaptation. The popularity of these hypotheses can be attributed to the fact that so many biologists share the same obsession.

Compare their obsession with Darwin's good sense: he wonders, if females seek to know male's “general vigour”, why they focus their attention to a special trait confined to male, enormously developed due to sexual selection, instead of seeing a more common indicator of “general vigour” [Darwin 1998, chapt.4].

Also if Darwin heard Hidaka's testimony, he might say why Hidaka repeats what he already said in 1860s: animals have a proper “Sense of Beauty” and “mental powers” enough to make careful mate selection [Darwin 1999, 95]. However, Darwin should have said that animals have a “Sense of Species Identification and Individual Discrimination”. Then no one could refute him. Anyway there has been no necessity to attribute mate choice to adaptation or to the “choosy gene” fiction.

As far as one thinks in line with Darwin, Hidaka, and Mizuhata, the sociobiological approach to evolution becomes untenable because it attempts to see evolution in a closed cycle of genes--fitness--genes.

How far is it possible to treat genes as an autonomous self-duplicator?

1) If it is genes alone that determine both survival and reproduction, then evolution can be explained in terms of genes and fitness alone. @4

2) If factors other than genes should cause a selective effect on gene reproduction, it is no more possible to attribute evolution to a closed loop of gene-fitness-gene. In such a case, what select genes of next generation is not genes but mating behaviors (e.g. MPR) of non-genic origin.
@4： The term “fitness” in sociobiology sounds inconsistent because fitness is a synonym of viability which is inseparable with adaptation, so the word is inappropriate for representing fertility. Moreover sociobiologists suppose that maturity brings forth reproduction, thus reducing reproduction to viability. This is not correct for highly selective mating. In the Japanese language, this inconsistency is far more serious because the word Tekioudo (fitness) is synthesized from Tekiou (adaptation) and Do (rate), literally meaning ”adaptation rate”. A more neutral word like “genic copy rate” or “genic reproduction rate is required.
Then what is a motive force for evolution besides adaptation? It is to seek for “species=societal coherence”. Species-proper characters such as ocelli patterns of peacock, long necks of giraffe, nuptial colors of fish, antlers of deer etc. are considered to be a species sex-attracting symbol determined by each SS, and not by the choosy gene. Such species-proper male characters are “the colors of each species=society” to symbolize its own unity and identity against other SSs (especially of close kin) [Mizuhata 2003].

Ethological behaviors due to MPRs and the relevant mating procedures are not the “result of natural selection” but, on the contrary, the “cause (motive force) for SS evolution”. This assertion will clearly distinguish the MPR theory from the runaway hypothesis that supposes the “runaway” of a species trait to be a result of choosy gene increase owing to its higher fitness. This theory admits the occurrence of non-adaptive evolution, however, but does not explain why it occurs with so enormous diversity in so short a period.

A species=society subjectively adopts a specific character as their sex-attracting symbol in order to establish and enhance the identity of the species=society. Wide variety of the species=societally unique characters is caused by requirement to represents interspecific differences and contrasts. Therefore evolution for species=societal coherence can occur toward adaptive, neutral, or non-adaptive directions.

Recently studies on the “culture of animals” are becoming popular. This approach of “gene-culture coevolution” is similar to this theory in that both deal with a non-genic gene selection mechanism [Aoki 2001]. However, this theory is fundamentally based on gene/meme analogy, where the 'meme frequency' changes just like genes: due to natural selection, sexual selection and analog of mutation etc. Therefore, the “culture” in this theory lacks the concept of species=society.