An extension of group selection theory.
See this paper. Abstract, emphasis added:
Two or more independent species lineages can fuse through an evolutionary transition to form a single lineage, such as in the case of eukaryotic cells, lichens, and coral. The fusion of two or more independent lineages requires intermediary steps of increasing selective interdependence between these lineages. We argue a precursory selective regime of such a transition can be Multilevel Selection 1 (MLS1). We propose that intraspecies MLS1 can be extended to ecological multispecies arrangements. We develop a trait group selection (MLS1) model applicable to multispecies mutualistic interactions. We then explore conditions under which such a model could apply to mutualistic relationships between pollinators and plants. We propose that MLS1 could drive transitions towards higher interdependency between mutualists and stabilise obligate mutualisms in the face of invasion by cheater variants. This represents a radical extension of multilevel selection theory, applying it to the evolution of multispecies populations, and indicating new avenues for researching ecological community evolution.
There are two points to be made before getting to the main discussion. First, I am not going to analyze this paper in fine detail; the reader can do so if they wish. Instead, I am going to briefly summarize its main points and then apply that to areas of human biopolitical interest. Second, since this is a paper focused on evolution and selection, there is discussion on evolutionary selection for altruism, which my readers know is not an area of particular interest to me, other than the requirement to refute critics of Salter who wrongly cite “the evolution of altruism” in their pathetic “arguments.” I would therefore like to make the following points:
1. The “evolution of altruism” is not, as I have comprehensively previously argued, anything required for the rational and intentional pursuit of ethnic genetic interests, a pursuit that is adaptive.
2. Gibbering of anti-Salterites about ‘free riding” has already been disproven via papers on ethnocentrism, which I have also previously discussed at length
3. It stands to reason that altruistic behavior of at least some sort will be involved in certain aspects of Multilevel Selection (MLS) and this general background behavior, which has been shown can be evolutionarily stable, does not require endless commentary.
4. The authors in a footnote distinguish strong altruism, in which a non-altruist converting to altruism would suffer a loss of (individual) fitness; and weak altruism, where the loss of fitness would not occur. Putting aide breathless obsession of anti-Salterians about a single sentence in On Genetic Interests that mentions a theoretical example of strong altruism, the fact remains that most of what Salter prescribes for the pursuit of ethnic genetic interests is weak altruism (or at least a form of strong altruism that would impose minimal fitness costs on an individual level). But as altruism can be evolutionarily stable, net positive adaption would be enhanced for the individual via inclusive fitness mechanisms.
5. Even if you are concerned with “genes for altruism” then it can be argued that you if you are an altruist, then you are on average more likely to find these “genes for altruism” in co-ethnics who are more closely related to you as opposed to alien ethnies who evolved under different conditions (potentially not selecting for altruism).
Keep in mind as well the definition of mutualism – an interaction of two (or more, typically two) species in which both (all) each have a net benefit.
That out of the way, we can briefly contrast MLS Type 1 (MLS1) from MLS Type 2 (MLS2). MLS2 is what most people think about when they hear group selection; simply put, one group directly outcompetes another. To adopt and adapt an example given by the authors, consider a group containing types A, B, and C. This ABC group can outcompete other groups, such as a DEF group, and thus we will observe more ABC groups. Thus, the non-ABC groups are culled over time, and the representation of the A,B, and C types would increase, even if group membership imposed some costs on each type on an individual basis. What about MLS1? Imagine a situation in which the different groups exist. In a given environment, the ABC grouping is more fit than alternatives; thus there are more A, B, and C types in the ABC group than there, are, say, D,E, and F types. The individual types are then released into a common pool. Because of the advantages of the ABC group, the A, B, and C types are heavily represented in the common pool. When groups randomly reform in the next generation, there will be more ABC groups than others, simply because there are more A, B, and C types in the preceding common pool. If the same environmental conditions obtain, then the ABC group will again be more fit, its constituent types will increase more than others, and so the next generation will see even more ABC groups forming. Over time, we will observe a dominance of ABC group similar to that of the MLS2 mechanism in which stable groups over time directly compete on a pure group basis. Note that, according to the authors, it is not necessary for the MLS1 mechanism to have, at the onset, any evolved mechanism to detect and punish free riding “cheaters” as it is expected that the groups that contain free riders would not manifest as many individuals as those composed of “honest” individuals; the “honest” would overwhelm the “cheaters” through “force of numbers” (and it this goes on long enough, it would provide MLS sufficient stability over time to allow a more directed anti-”cheating’ mechanism to evolve). In the MLS2 form, direct mechanisms against “cheating” would by necessity would have evolved, see below.
In the MLS1 mechanism, the fitness of an individual is a composite of their particular genotype-phenotype “character state” plus a component derived from the average “character state” of members of their group. In MLS2, the entire group is assigned fitness as a group (based on the “character state” of the integrated group as a whole).
All of this is straightforward when considering altruistic cooperation within a single species. What if multiple species are considered as composing the groups and if intra-group altruism is replaced by inter-species mutualism? The MLS2 mechanism for multiple species is one that definitely has occurred, for example, leading to the formation of eukaryotic cells. Thus, groups of cells in which a mutualistic association between the cell and an endosymbiont (symbiotic organisms that lives within another) exist would directly outcompete other groups of cells (and free-living versions of the endosymbionts) and thus a stable new form would emerge; e.g., a mammalian cell containing mitochondria.
However, what about the situation in which the species remain separate? The authors claim that in most cases there is not vertical transmission (parent-to-offspring) of the mutualistic interaction; instead there is horizontal transmission; the “partnerships” need to be re-established each generation. Imagine a species of bird that lives on the back of some large mammal, eating that mammal’s skin parasites. Both benefit. However, when the mammal reproduces, a bird is not carried along to the offspring (and vice versa). A new mammal-bird pairing occurs with a new generation of the mammal and a new generation of the bird; the ‘partner” is acquired horizontally through the environment, not vertically through heredity. Thus, the authors argue that MLS involving distinct species, analogous to this example, must be of the MLS1 type. It is not a stable integrated group over time (which would be MLS2), since every generation requires re-establishment of the group; it is instead a MLS1 mechanism, in which individuals join a common pool and the groups reform. In the example given, mammals that engage in this mutualism would outnumber those that do not; the same with the birds. Thus, there will be more mammal-bird groups in the next generation than lone mammals and lone birds. Of course, a MLS1 mechanism an evolve into a MLS2 form, although it is not clear how that would or could) occur in the mammal-bird example, but certainly could (and has) occur with an endosymbiont. Indeed, it may be that MLS2 interactions must pass through a MLS1 phase; in any case, the latter would be expected to be found in nature more often than the former. If MLS2 evolves from MSL1 then there would be enough time for the MSL2 mechanism to have developed direct anti-“cheating” approaches to control free riding.
I will now extend this discussion to the human situation. MacDonald’s “group evolutionary strategies” fit with the MLS2 mechanism – there are stable groups over time, group fitness as an integrated unit, and sufficient time to develop strategies against free-riding (and the papers on ethnocentrism show that ethnocentric group strategies outcompete universalist, free-riding, and traitorous ones). Given the paradigm that MLS2 evolves from MSL1, we can consider that MLS2 human groups can have evolved from individuals and then small tribes via MLS2 mechanisms; those that cooperated produced more members than those that did not and reforming the groups each generation resulting in more cooperating groups. Ethnocentrism would develop by having these groups form with individuals who are more closely related genetically (and phenotypically) and thus fitness is enhanced through inclusive fitness. Today’s homogeneous ethnies were no doubt formed by MLS1 mechanisms of tribal integration over time, forming integrated groups (that can then practice ethnocentrism). But how are such ethnocentric groups stably maintained? Aren’t the human groups reforming each generation and thus always MLS1?
In humans, there are also group identity mechanisms, including culture, national/ethnic affiliation, and other markers of kinship that can be used, in addition to biological phenotype. All of these mechanisms can be used to keep the group intact over time so that even if on a purely objective level there is a “common pool” from which individuals form groups, subjectively, IF cultural group mechanisms are maintained there it is a de facto MLS2 mechanism (including proofing against free riders). If the group is lucky enough to form a homogeneous unit in an exclusive territory then the reforming of the group from a common pool has no other outcome than stable maintenance, as there would be no other types present. Thus, MLS2 group maintenance is easier in such “ethnostate” circumstances, although Diaspora groups have successfully used cultural mechanisms to maintain group stability in more trying circumstances.
What about “purer” MLS1 mechanisms for human today? There would be two major types, one unstable from the standpoint of ethnic genetic interests and one stable. The unstable mechanism would be of group forming between individuals who are genetically very distant from each other, such as from the major racial groups. Formation of such groups is difficult (as multicultural societies have learned), given biological and cultural differences that impede cohesion as well as pre-existing MSL2 group mechanisms that promote competition vs. cooperation. These issues explain not only the problems of multicultural societies but also why attempted civic nationalist solutions often need to be coercive. But even if the coercion is “successful,” this approach is unstable because of the genetic distance of the groups and the gulf of genetic interests. This is compounded if MLS1 leads to MLS2; any integration (such as intermixture) under these circumstances, would lead to a major decrease in group genetic interests. Certainly, highly admixed human groups exist today, but their formation was typically manifested with strife, they represent a loss of genetic interest for the original groups compared to what would have obtained otherwise (e.g., Spaniards of today [assuming we agree that genetic interests should always be forward-looking and not dwell on the interests of past peoples] would have greater ethnic genetic interests if Latin America was populated exclusively by pure Spaniards rather than mostly by hybrids), and the societies are unstable and hence there are adaptive costs due to the proximate phenotypic effects of admixture. Again, this is an unstable strategy.
On the other hand, MLS1 mechanisms between closely related ethnies would avoid many of the practical difficulties and a MLS2 outcome would not be as damaging to ethnic genetic interests given the relative genetic similarity of the groups. Indeed, the loss of genetic interests may be small enough so that they would be outweighed by the gain of genetic interests from the adaptive utility of the group – a net gain of genetic interests. Indeed, as mentioned above, extant successful ethnies and successful MLS2 group evolutionary strategies most likely originated from MLS1 mechanisms, in which smaller human groups played the role of “multiple species” in the scenario outlined by the authors. In one sense, the worldwide European race is already an early stage MLS2; in another sense, focusing more on the still-existing divisions between Europeans, we can consider extant European ethnic groups being the “multiple species” that can form a pan-European group in a MLS1 mechanism. Pan-European groups can outcompete atomized ethnies via MLS1 mechanisms, suppressing free riding ethnies through successful “force of numbers” as well as using already existing rational mechanisms for approaching the free riding problem. This MLS1 mechanism can lead to a future, more integrated MLS2 pan-European group evolutionary strategy, compared to the weak and not very cohesive MLS2 “Western” organism extant today. Although I do not support the idea of European panmixia, and such does not have to follow from pan-Europeanism, any integration that does occur would be of the potentially more positive nature explained above, and certainly more stable and superior to mechanisms of MLS1 leading to MLS2 including genetically divergent racial groups.
In summary, the theoretical outline for multispecies MLS, particularly the MLS1 and MLS2 paradigms, can apply to multiethnic MLS as part of a pan-European approach.
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