More than a century ago, William Morton Wheeler proposed that social insect colonies can be regarded as superorganisms when they have morphologically differentiated reproductive and nursing castes that are analogous to the metazoan germ‐line and soma. Following the rise of sociobiology in the 1970s, Wheeler's insights were largely neglected, and we were left with multiple new superorganism concepts that are mutually inconsistent and uninformative on how superorganismality originated. These difficulties can be traced to the broadened sociobiological concept of eusociality, which denies that physical queen–worker caste differentiation is a universal hallmark of superorganismal colonies. Unlike early evolutionary naturalists and geneticists such as Weismann, Huxley, Fisher and Haldane, who set out to explain the acquisition of an unmated worker caste, the goal of sociobiology was to understand the evolution of eusociality, a broad‐brush convenience category that covers most forms of cooperative breeding. By lumping a diverse spectrum of social systems into a single category, and drawing attention away from the evolution of distinct quantifiable traits, the sociobiological tradition has impeded straightforward connections between inclusive fitness theory and the major evolutionary transitions paradigm for understanding irreversible shifts to higher organizational complexity. We evaluate the history by which these inconsistencies accumulated, develop a common‐cause approach for understanding the origins of all major transitions in eukaryote hierarchical complexity, and use Hamilton's rule to argue that they are directly comparable. We show that only Wheeler's original definition of superorganismality can be unambiguously linked to irreversible evolutionary transitions from context‐dependent reproductive altruism to unconditional differentiation of permanently unmated castes in the ants, corbiculate bees, vespine wasps and higher termites. We argue that strictly monogamous parents were a necessary, albeit not sufficient condition for all transitions to superorganismality, analogous to single‐zygote bottlenecking being a necessary but not sufficient condition for the convergent origins of complex soma across multicellular eukaryotes. We infer that conflict reduction was not a necessary condition for the origin of any of these major transitions, and conclude that controversies over the status of inclusive fitness theory primarily emanate from the arbitrarily defined sociobiological concepts of superorganismality and eusociality, not from the theory itself.

中文翻译：

---

超有机体和种姓分化是不归路：主要的进化转变如何在翻译中丢失

一个多世纪以前，威廉·莫顿·惠勒 (William Morton Wheeler) 提出，当群居昆虫群落具有类似于后生动物种系和体细胞的形态分化的生殖和哺乳等级时，它们可以被视为超有机体。随着 1970 年代社会生物学的兴起，Wheeler 的见解在很大程度上被忽视了，我们留下了多个新的超有机体概念，这些概念相互不一致，并且无法提供关于超有机体起源的信息。这些困难可以追溯到更广泛的真社会性的社会生物学概念，该概念否认物理上的蜂后-工人种姓分化是超生物群落的普遍标志。与魏斯曼、赫胥黎、费舍尔和霍尔丹等早期进化博物学家和遗传学家不同，谁开始解释获得未交配的工人种姓，社会生物学的目标是了解真社会性的进化，这是一种涵盖大多数合作育种形式的宽泛的便利类别。通过将不同的社会系统归为一个类别，并将注意力从不同的可量化特征的进化上转移开，社会生物学传统阻碍了包容性适应理论与主要进化转变范式之间的直接联系，以理解向更高组织复杂性的不可逆转转变. 我们评估了这些不一致累积的历史，开发了一种共同原因方法来理解真核生物层次复杂性中所有主要转变的起源，并使用 Hamilton' s 规则来论证它们具有直接可比性。我们表明，只有惠勒对超有机体的原始定义才能明确地与从依赖环境的生殖利他主义到蚂蚁、圆蜂、胡蜂和高等白蚁中永久未交配种姓的无条件分化的不可逆进化转变相关联。我们认为，严格的一夫一妻制父母是所有过渡到超有机体的必要条件，尽管不是充分条件，类似于单合子瓶颈是复杂体细胞在多细胞真核生物中收敛起源的必要但非充分条件。我们推断，减少冲突不是任何这些重大转变起源的必要条件，