ResultsTasks, domains, and comparability across speciesEach informational task identified in this study is defined by the type of situation that reliably precedes ESR-relevant behavior change, rather than by specific behaviors, anatomical structures, or presumed internal mechanisms. Tasks are therefore indexed to domains of informational flow—environmental states, free-moving entities, perception shaping, group structure, and symbolic systems—rather than to taxon-specific repertoires.
This operationalization allows lineages with radically different morphologies, ecologies, and biological substrates to be compared on a common footing: by the kinds of situations that force them to change what they are doing. In the sections that follow, we summarize each task, its corresponding domain of informational control, and the characteristic behavior-change patterns through which that control becomes observable.
Group 1 — Task 1: Binary (0–1) orientation — Environmental controlDomain of informational relevance: Environmental states and fragments.
The immediate physical setting and its fragments: gradients, shelter/exposure, substrate changes, microclimate, barriers, vibration fields, current/flow direction, etc.
Shared capacity (potential)
Species in Group 1 can detect and control changes in this domain. Without being struck or forced, they register changes in the immediate setting that reliably precede behavior change. In this sense, features such as light, temperature, substrate, cover, and moisture function as informational cues.
Foundational decision layer — options to be chosen
Choosing among simple, binary alternatives: favourable / not favourable; safe-enough / not safe-enough; stay / leave; process / pause.
Behavior change patterns (characteristic switches)
These lineages reliably perform controlled shifts such as: move → stay, hold → relocate, drift → anchor, activate → deactivate. Each is a spatial or exposure alternative (here vs. there; now vs. not-now) deployed in service of energy, safety, or reproduction. Accordingly, at this level, informational control is identified solely through observable selection among viable behavioral continuations—expressed through regulated changes in timing, rate, direction, or engagement—rather than through direct physical forcing or invariant coupling, in accordance with the general inclusion and exclusion criteria defined above.
Clarifying the boundary between Task 1 and non-task processes
Below Task 1 lie physical and chemical processes that change state only through direct interaction or force, without the capacity to select among alternatives. Such processes do not produce behavior change in response to environmental change, and therefore cannot act to align conditions with Energy, Safety, or Reproduction. Task 1 marks the minimal transition at which a system can regulate its own behavior in response to changing surroundings in order to sustain itself over time.
Decision cue — what this task is really asking
Informational question the organism is tackling:
“Should I stay or should I move?”, “Is this state safe enough to continue?”, “Do I activate or suspend what I’m doing?”
What this task does not require
It does not require tracking another moving organism, signalling to an audience, coordinating with group members, or following a learned social rule. This is not coalition behavior, display strategy, or symbolic interpretation. It is direct regulation of local conditions.
Representative lineages
All extant taxa in the dataset express this capacity at full strength—from prokaryotes to vertebrates. Every living species is coded BCBS_1 = 1100 for Task 1. At the same time, there is a subset of lineages whose survival appears to rely on this task and essentially this task alone—bacteria, many unicellular eukaryotes, plants, fungi, and comparable forms.
Group 2 — Task 2: Distal engagement choice — Free-moving-entity controlDomain of informational relevance: Free-moving entities at a distance
Motile organisms in the surroundings that can approach, flee, pursue, threaten, court, or otherwise act independently (prey, predators, rivals, mates, intruders).
Shared capacity (potential)
Species in Group 2 can control behavior in response to these free-moving agents at range. They do more than notice “the environment changed” (Task 1). They track a particular moving other, treat it as meaningful “for me, now,” and select among structured options: close in, shadow, confront, evade, hide, or ignore. In other words, they can make decisions about that moving entity before direct contact.
Foundational decision layer — options to be chosen
Distal agent-level alternatives: prey vs. predator vs. rival vs. mate vs. irrelevant intruder.
Once an entity is differentiated in this way, a corresponding class of behavior change becomes available.
Behavior change patterns (characteristic switches).
These lineages reliably show dyadic, target-directed switches such as: forage → flee, patrol → pursue, approach → veer off, stalk → abandon, intercept → break contact, chase → avoid, trail → dodge. Each is a B1→B2 that is about a specific moving other, not about the backdrop.
Decision cue (what this task is really asking)
Informational question the organism is tackling: “What do I do about that moving entity at a distance?” This is already qualitatively different from Task 1. The organism is no longer only regulating itself against a setting (“Is this place okay?”). It is regulating itself in relation to an independently moving agent at range; it is “I must do X about that mover.”
Clarifying the boundary between Task 2 and Task 1
In Task 1, behavior change is selected to regulate the organism’s relation to its immediate setting (stay/leave, activate/suspend) without reference to any specific moving other. In Task 2, behavior change is selected in relation to a particular free-moving entity, requiring the organism to track, engage, avoid, or disengage from that entity at a distance. Put simply, Task 1 governs whether and how I adjust to the situation; Task 2 governs what I do about that mover.
What this task does not require
- Perception-shaping or audience design (that is Task 3).
- Coalition, role, or recruitment logic (that is Task 4).
- Formal rule-following, symbolic media, or cumulative culture (that is Task 5).
Clear, stable Group-2 exemplars in the dataset include
- Many solitary and reef-associated fishes (including many sharks and predatory teleosts) that patrol, detect, trail, lunge, or disengage purely on the trajectory/speed/proximity of another swimmer, but do not routinely use adaptive camouflage, display choreography, or signalling to shape the other’s perception.
- Turtles and many salamanders/newts that react to detected movers (approach or withdraw) but do not regularly modulate what the other sees.
- Non-social, visually hunting insects, non-displaying lineages such as many dragonflies/damselflies and tiger beetles, and host-seeking dipterans such as mosquitoes and horse flies, and other species that track, close in, or break off based on the motion of a target, but do not, as a lineage-level routine, add perception-shaping displays.
Representative lineages
In coding terms, Group-2 species are those with:
- BCBS_1 = 1100 (full Task-1: Binary environmental control), and
- BCBS_2 = 2100 (full Task-2: free-moving-entity control).
Transitional 2→3 cases (e.g. strong but not universal signalling/decoy/display in some contexts) are coded separately and do not alter the five-group scaffold.
Group 3 — Task 3: Perception-shaping in context — Perception-shaping & signalling controlDomain of informational relevance: Context, contextual cues, signals, and audience perception
The domain here is not “the world out there” (Task 1) and not “a mover out there” (Task 2), but the perceptual channel between organism and observer. The organism adjusts what others perceive—via display, concealment, mimicry, exaggeration, posture, coloration, movement pattern, timing, or signal emission—in order to alter the other’s interpretation before contact.
Shared capacity (potential)
Species in Group 3 can control how they appear to others, actively manipulate others’ perception of themselves or the situation, and, when needed, steer observers toward a particular interpretation or direction of action. They do more than regulate themselves against the backdrop (Task 1) or decide how to engage a moving other (Task 2). They can modulate how they are detected through the visual, auditory, or chemosensory channels of another organism. That includes attraction (“notice me as a mate / as dangerous / as cooperative”), deception (“do not notice me,” “misread me,” “treat me as something I am not”), and direction/steering (“move where I point you”). The key is that the organism’s behavior change is directed at altering what is perceptually available to others.
Foundational decision layer — options to be chosen
Given a situated audience (predator, prey, mate, rival, intruder), select among: attract vs. camouflage; show vs. hide; exaggerate vs. downplay; signal vs. stay silent; mimic vs. reveal true form; lure closer vs. push away; direct this way vs. that way.
Behavior-change patterns (characteristic switches)
These lineages reliably perform BCs of the form: neutral posture → threat display; cryptic stillness → courtship dance / color flare; foraging posture → warning exaggeration (spines out, body inflation, wing spread); background-match camouflage → sudden contrast flash (startle, misdirection); ordinary movement → luring movement (prey-attraction feint, brood-defense decoy); freeze → motion-jerk to trigger pursuit; ordinary presence → directional cueing (gestures/movements to steer conspecifics).
Decision cue (what this task is really asking)
Informational question the organism is tackling: “What do I make you think I am right now?” rather than only “What do I do about you?” It also covers questions of influence, such as: “How can I act so that your behavior changes the way I need?” and “How can I present myself so you interpret this situation in the direction I want?” That is the diagnostic leap from Task 2 to Task 3. In Task 2, the organism chooses how to deal with an agent. In Task 3, the organism chooses how the agent will read the situation. Importantly, the same individual can alternate hide → advertise → hide again, depending on who is present and what is at stake (mate nearby vs. predator nearby vs. offspring nearby). That context-sensitivity of presentation is the signature of Task-3 control.
Clarifying the boundary between Task 3 and Task 2
In Task 2, behavior change is selected to regulate how the organism engages a free-moving entity (approach, avoid, pursue, disengage). In Task 3, behavior change is selected to regulate how that entity interprets the situation, by selectively shaping what it perceives about the organism or the context. Put simply, Task 2 governs my action relative to you — “W
hat I do about you”; Task 3 governs your interpretation because of me — “W
hat I make you think is happening”.
What this task does not require
- It does not require coalition or shared role structure (Task 4). All of this can be done dyadically or one-to-many.
- It does not require symbolic rule systems, language, money/tokens, or maps (Task 5).
- It does not count always-on morphology as Task 3. Aposematic coloration that is permanently visible is a carrier, not a choice. Task-3 evidence requires selectable presentation: show / withhold / transform based on context. This avoids the slide from “feature visible” to “feature chosen.”
Representative lineages
- Many reptiles and amphibians that switch between crypsis and display (cryptic posture → sudden throat fan, color flush, or body inflation to deter or attract).
- Numerous birds and fishes that perform courtship displays, alarm postures, or fin/spine exaggerations on demand, not continuously.
- Insects that deploy deimatic/startle flashes, wing-spot reveals, or mate-attraction signalling only in specific contexts—not as an always-on message.
Coding interpretation
Group-3 species express:
- BCBS_1 = 1100 (Task-1: Binary orientation — Environmental control)
- BCBS_2 = 2100 (Task-2: Distal engagement choice — Free-moving-entity control)
- BCBS_3 = 3100 (Task-3: Perception-shaping & signalling control)
Lineages with BCBS_3 = 3066 (strong but not universal Task-3) are coded as Transitional 2→3 or 2–3 span in Appendix A. These transitional cases are important: they show that Task-3 style perception-shaping emerges in partial, context-bound form before it becomes a universal, load-bearing strategy for the lineage.
Group 4 — Task 4: Coalition alignment — Group-dynamics controlDomain of informational relevance: A structured group of independent organisms
The domain here is not a single mover (Task 2) and not just an audience to manage (Task 3), but a multi-individual system in which members are interdependent: they recruit help, share risk, delegate responsibility, allocate roles, maintain or shift hierarchy, defend together, raise young together, enforce tolerance or status, or suppress rivals.
Shared capacity (potential)
Species in Group 4 can control behavior in relation to changes in group dynamics—coalitions, alliances, rank, joint projects. This includes both collaborative and competitive moves inside the group: coordinating movement or attack; supporting an ally and not a rival; tolerating one juvenile and rejecting another; exchanging roles and sharing responsibilities; defending a nest or den as a joint project; recruiting others for mobbing or hunting; deferring to rank when it is strategically necessary. In other words, individuals act in ways that make sense only relative to a live social structure.
Task 4 groups a diverse set of behaviors not because they are superficially similar, but because they share the same underlying control problem. In all Task-4 cases, behavior change is selected with respect to a structured, multi-individual system in which outcomes depend on roles, relationships, and coalition state rather than on a single other agent or audience. Actions such as alliance support, cooperative defense, parental provisioning beyond the dyad, rank-dependent tolerance or aggression, and recruitment during mobbing differ in form, but converge in that their appropriateness depends on who is involved, how individuals are related, and how the group is currently organized. This distinguishes Task 4 from Task 3, where behavior is conditioned on shaping how others perceive the situation, and from Task 5, where behavior is regulated by abstract, symbolic conventions. The apparent heterogeneity of Task-4 behaviors therefore reflects the breadth of group-structured contexts rather than a lack of specificity in the task itself.
Foundational decision layer — options to be chosen
Load-bearing choices for group maintenance, parental success, offspring survival, or individual access to resources/mating, such as: collaborate vs. compete; recruit vs. do not recruit; help vs. defeat; protect vs. harass; share vs. take; follow vs. lead; reinforce hierarchy vs. challenge it; defend this offspring/resource vs. yield; intervene for A against B vs. stay neutral; admit this individual vs. exclude that one; stay with the group vs. split off.
Behavior-change patterns (characteristic switches)
These switches are routinely observed as:
- forage solo → join coordinated hunt/defense
- ignore conspecific → intervene on its behalf (alliance support)
- tolerate proximity → chase/expel after rank breach
- rest quietly → alarm-call and recruit a mob against a predator or rival group
- tend own young → provision or guard non-offspring in the same unit
- withdraw from fight → re-enter when an ally is threatened
- These are not just “approach/avoid a mover” (Task 2). They are role-aware engagements: you do X because that one is your ally, your cub, a tolerated subordinate, or a coalition partner against an outgroup.
Clarifying the boundary between Task 4 and Task 3
In Task 3, behavior change is selected to influence the perception of an observer or audience, typically in dyadic or one-to-many contexts. In Task 4, behavior change is selected relative to a structured group, where actions depend on roles, alliances, coordination, and competition within a multi-individual system. Put simply, Task 3 governs how I appear to others — “What do I make you think I am?”; Task 4 governs how I act within a coalition — “What do I do given who we are to each other?” Task 3 manipulates perception; Task 4 navigates relational structure.
Decision cue (what this task is really asking)
Informational question the organism is tackling: “Who is us and who is them right now?”, “What position do I take inside this live social structure?”, “What action keeps this arrangement working for me (and mine) in this moment?”, and “What is my role in keeping this structure working for me and mine?” That is qualitatively different from Task 3 (“what do I make you think I am right now?”). Task 3 is performance for an audience. Task 4 is navigation of a network.
What this task does not require
- It does not require formal, externalized symbolic systems (Task 5). Coalition alignment can run without language, writing, maps, money, or law codes.
- It does not require explicit rule-teaching; much of it is enacted through practice, reinforcement, and expectation.
- It does go beyond simple proximity or shoaling. Mere aggregation (schooling fish, loose herds with no role structure) does not count as Task 4. Task 4 requires patterned role-taking, coalition logic, or coordinated joint action where individual fates are entangled.
This distinction matters: being near others is not the same as structuring a coalition with them. For that reason we reserve full Task-4 for systems in which we can see collaboration and competition operating together, and in which role taking, coalition choice, or support decisions are themselves behaviorally selectable by the individual (join/withhold, back A against B, tolerate X but not Y), rather than fixed by developmental caste or chemistry. Highly coordinated insect or fish systems are therefore coded 3→4 or 4-partial, when coordination is strong but coalition recombination or role reassignment is limited at the individual level.
Representative lineages
- Mammals very often express Task-4 style control. Because mammalian life histories are scaffolded by juvenile group interaction, provisioning, protection, and rank/tolerance management, removal of this capacity would severely constrain mammalian life histories as we know them.
- Many cooperative or family-living birds (some corvids, parrots, cooperative breeders) that jointly defend territories, provision young beyond the breeding pair, and recruit group members in alarm/mobbing contexts.
- Some fishes and insects with role differentiation, coordinated defense, or shared nest/colony maintenance. In the dataset, these often sit in 3→4 or 4-partial slots when coordination is strong but internal competition, negotiated alliances, or true coalition choice is still limited. Full Task-4 is reserved for cases where coalition alignment and differentiated roles are both central and behaviorally selectable.
Coding interpretation
Stable Group-4 species express:
- BCBS_1 = 1100 (Task-1: Binary orientation — Environmental control)
- BCBS_2 = 2100 (Task-2: Distal engagement choice — Free-moving-entity control)
- BCBS_3 = 3100 (Task-3: Perception-shaping & signalling control)
- BCBS_4 = 4100 (Task-4: Coalition alignment — Group-dynamics control)
Species that show coalition-like organization but not universally (e.g. BCBS_4 = 4066) appear as Transitional 3→4 in Appendix A. These transitional spans (3–4, 4-partial) are exactly where we see cooperative hunting, communal defense, extended parental care, or group-structured rank negotiation present, but not yet obligatory for all members of the lineage.
Group 5 — Task 5: Rule-guided abstraction — Formal symbolic systems controlDomain of informational relevance: Formalized symbolic systems and abstract conventions (“Symbolic–Sapient”)
Population-level systems of shared symbols—language, number, writing, money/tokens, mapped space, explicit norms, ritual contracts, durable collective memory—that can prescribe action and coordinate expectations even when the relevant individuals or objects are not directly present.
Shared capacity (potential)
Species in Group 5—in the present dataset, at full maturity humans only—can control behavior using abstract, conventional symbols and structured rule systems. Behavior can now be selected and switched not only because “the environment changed” (Task 1), or “that agent is doing something I must address” (Task 2), or “I want to shape how you see me” (Task 3), or “our coalition is in state X so I take role Y” (Task 4), but because “a shared symbolic rule says we now do Z,” or “the map/plan/story says go here,” or “this token changes ownership/obligation,” or “this declaration updates the social state.” In other words, behavior change can be driven by symbols that stand for absentees, elsewheres, and non-immediate states of affairs.
Foundational decision layer — options to be chosen
Given a shared symbolic frame, the organism can decide to:
- follow an instruction, policy, law, promise, or norm that is only symbolically present;
- act on a plan encoded in language or marks (“meet at sunrise at that rock”);
- accept or transfer obligations via a token (currency, promissory object, seal, record);
- update group state by declaring something in language (“we are at peace now,” “you are in/out”);
- generate novel combinations (new sentences, new marks, new routes) and have them understood as binding by others. These are not just “make ally / don’t make ally.” They are “instantiate a convention, then behave because of that convention.”
Behavior change patterns (characteristic switches)
Routinely observed human-style B1→B2 include:
- idle → comply based on an instruction, rule, or agreement (no immediate physical force);
- local foraging → coordinated task-execution based on a shared, verbally stated plan;
- hold resource → transfer resource because of an abstract token (“this paper now means it’s yours”);
- short-horizon action → long-horizon investment because of a symbolic narrative, model, schedule, or map;
- spontaneous motion → ritualized behavior because “this ceremony/norm now says we do X.”
- In all such cases, a symbolic frame can trigger or suppress action even when no immediate cue, rival, mate, ally, or predator is present.
Clarifying the boundary between Task 5 and Task 4
In Task 4, behavior change is coordinated through live group dynamics, roles, and relationships among present individuals. In Task 5, behavior change is coordinated through abstract, shared symbolic systems that can prescribe action independent of immediate presence, perception, or interaction. Put simply, Task 4 governs alignment through social structure; Task 5 governs alignment through formalized rules and symbols.
Decision cue (what this task is really asking)
Informational question the organism is tackling: “How do I act now because of a shared symbol/rule that is not physically here?”, “How do I use our symbolic system to align, coordinate, and bind others (and myself), “How do I use our symbolic system to maintain coordinated expectations across time and space?”
This is qualitatively beyond coalition. In Task 4, alignment depends on who is present and what the coalition is currently doing. In Task 5, alignment can depend on a rule that outlives the situation and can be invoked later by anyone who knows the system.
What this task does not reduce to
- Not just rich signalling or ritual display. Many birds, cephalopods, reptiles, and mammals have elaborate, context-sensitive signalling—that is Task 3 (and, in social settings, Task 4). Task 5 requires conventions that can be followed, referenced, revised, and transmitted as systems, not just emitted moment by moment.
- Not just social coordination. Many mammals and birds coordinate hunts, raise young cooperatively, or enforce rank—that is Task 4 (Group-dynamics control). Task 5 requires explicit, shared symbols that can organize behavior beyond immediate presence or immediate memory.
- Not “an animal in a lab can press a symbol.” We explicitly distinguish an individual’s lab-trained potential from a population-level practice. Full Task-5 requires that the symbolic system is normal, entrenched, and socially scaffolded for that lineage outside the lab.
This last boundary matters. Comparative cognition has clearly shown that great apes, corvids, parrots, cetaceans, and elephants can acquire symbols, transmit practices, and use context-sensitive signals. What we do not yet see outside humans is a population-level, obligatorily shared, and consciously revisable symbolic system that organizes day-to-day coordination across time and space. For that reason we code these lineages as Task-5 partials (5033, 5050, 5066) and place them in transitional 4→5 slots, rather than in the full Group-5 class.
Representative lineages
Humans are the only lineage assigned full Task-5 control (BCBS_5 = 5100), because routine human life depends on language, shared fictions, money/tokens, mapped abstractions, explicit norms and laws, and consciously revisable social contracts.
Some other vertebrates (great apes, corvids, parrots, cetaceans, elephants) exhibit partial Task-5 signatures in restricted contexts—symbol learning, proto-conventional calls, tool traditions with social transmission. In the dataset these are coded with partial maturities (e.g. 5033, 5050, 5066) and placed in Transitional 4→5, not in full Group-5.
Task 5 is defined by control over formal symbolic systems that regulate behavior through shared conventions rather than immediate context, presence, or direct interaction. This includes population-level use of symbols that persist across time and space, prescribe obligations, and coordinate action among individuals who may not be co-present. On this definition, humans are the only lineage in the present dataset that exhibits full, routine control of Task 5 as a load-bearing feature of daily life. Importantly, this does not imply an absence of Task-5-related capacities in other species. Several non-human lineages show partial or context-limited signatures of symbolic mediation, such as socially transmitted conventions, learned symbolic associations, or proto-normative signals. These cases are explicitly coded as partial Task-5 expression rather than excluded. The distinction between partial and full control is therefore empirical rather than definitional: full Task-5 control is assigned only where symbolic systems function as a primary, population-wide regulator of behavior, not merely as an individual or experimental capability.
Summary of task structure and constraintsAcross the full dataset of 1,530 species, task control patterns exhibit two consistent empirical constraints. First, we did not observe stable cases in which a lineage routinely expresses control over a later task while lacking reliable control over any earlier task in the sequence. For example, no species shows regular Task-3 (perception-shaping) control without also expressing Task-2 (distal engagement) control, and no species shows Task-4 (coalition alignment) control without Tasks 1–3 in place. Second, among lineages expressing multiple tasks, the internal ordering of tasks is conserved: when several tasks are present, they appear in the same ordinal sequence across taxa. We did not observe alternative or scrambled combinations such as [Task-1, Task-3, Task-4] or [Task-2, Task-4] in routine, species-typical behavior. Transitional cases occupy only adjacent spans [e.g., 2→3, 3→4, 4→5], indicating partial or context-limited emergence of the next task rather than bypassing or reordering. These constraints are descriptive properties of the dataset and hold across major clades, ecological niches, and social systems represented here.
Building on these observed constraints, the five domains of informational control sort lineages into five major groups.