Generalized Fitness: What Selection Has Been Scoring All Along

Fitness is the most successful idea in biology, and the Price equation is its most general accounting:given a population, a fitness currency, and a trait, Price partitions the change in that trait intoselection and transmission with complete generality, at any level and on any substrate. Thispaper does not dispute that accounting. It asks the question Price brackets and must takeas given, the question of which currency and which unit selection has actually been keepingits books in. The standard answer, differential reproductive success of heritable variants, isexactly right where reproduction is the dominant way a thing persists, and it leaves a long listof cases biology handles one stipulation at a time: the sterile worker whose colony outlives herby decades, the celibate founder whose order outlives him by twelve centuries, the constitutionthat organizes two hundred million lives without being inherited by any of them, the tree and itsfungal partner running one metabolism across the chromosomal divide. We propose that thecurrency selection scores is not reproductive success but Potential Order (PO): the per-resourcecapacity of an arrangement, in the environment it actually inhabits, to evolve toward greaterresistance to decay and to spread that capacity across instances. Reproductive fitness is PO seenthrough the narrow window of heritable replication. The proposal is continuous with a traditionthat has already widened fitness from reproduction to persistence, and it adds three thingsthat tradition lacks: a per-resource rate rather than a raw count, so efficiency enters; a productof a per-instance functional component and a distributional component, so neither unrealizedbrilliance nor realized emptiness scores; and a clean separation between PO, the criterion theanalysis tests against actual persistence, and P̂O, the agent’s own structurally embedded andfallible estimate of it, which is what lets the framework predict rather than redescribe andwhat dissolves the circularity that haunts fitness’s successors. We read cosmic history as threeoverlapping phases sorted by how order chiefly accumulates, give institutions and symbioses atreatment that matches how they actually work, resolve a set of boundary cases where existingframeworks give opposite answers, and stake the account on a pre-registered test: that joint-POpredicts the survival of religious orders better than founder lineage or founder prestige alone.Two further tests, built to read the framework’s recursive loops at one and two levels, separateit from prestige-bias and additive accounts.