Table 1.
Examples from vertebrates that form multi-level societies and engage in tasks with individuals in different social units. Social units are termed as is in the original studies.
|
definition of social units and tasks performed within... |
|||||
|---|---|---|---|---|---|
|
species |
number of social levels |
...social level 1: |
...social level 2: |
...social level 3: |
... social level 4: |
|
3 or 4 |
household: male–female complementarity and sex division of labour. |
cluster: social units formed by extended family. Assistance from kin and intergenerational division of labour. |
camp: cooperation with both related and unrelated individuals. Selection of foraging partners. |
between-camp visits: information sharing and cultural innovations. |
|
|
4 |
OMU: one male unit which hosts multiple females. Breeding unit. |
clan: two to three OMUs led by kin males, observable during resource scarcity. Foraging as OMUs or clans. |
band: multiple clans together, like troops in olive baboons. Communal sleeping, movement coordination, some affiliative interactions. |
troop: two bands sharing sleeping sites. Predator detection and defence, but no other group tasks. |
|
|
geladas [30] (Theropithecus gelada) |
4 |
OMU: one male unit which hosts multiple females and potentially also a few follower males. Breeding unit. AMG: all-young-male units are an alternative first-level structure. |
team: aggregation of two or more first-level units that associate with each other at least 90% of the time. Tasks not specified. |
band: collection of first-level units that spend between 50 and 90% of their time together. Communal sleeping and foraging, not more specific tasks. Like band in hamadryas baboons. |
community: the set of units with overlapping home ranges that are found together < 50% of the time. Tasks not specified. |
|
(Papio papio) |
3 |
reproductive unit: one male, one to several females, young and many secondary males. |
party: three to five reproductive units. Foraging, socializing, collective movement. |
gang: several parties together. Predator detection and defence in communal sleeping sites. |
|
|
3 |
OMU: one male unit which hosts multiple females. Breeding unit. |
band: social units formed by different OMUs. Males within bands likely defend females from bachelor males. |
troop: several bands together. Females disperse between troops to breed. |
||
|
(Loxodonta africana) |
up to 6 |
breeding females with calves are listed as the first level. We kept the paper’s definition for the first level here, but we would not characterize mothers with calves as a distinct social level. |
family: small size compared to third and fourth levels. Not affected by seasonality. Raising offspring and coordinating movement. |
bond/kinship social units: affected by seasonality. Predator defence, territoriality, knowledge sharing and rearing of young. |
clans: broader compared to third but functional differences have not been identified. |
|
Indo-Pacific bottlenose dolphins of Shark Bay [24–26] (Tursiops aduncus) |
3 |
first-order alliance: duos or trios of males consorting females in a cooperative manner. |
second-order alliance: pool for forming first-order alliances. Socializing and supporting second-order allies to defend females. Stable membership. |
third-order alliance: individuals from two or more second-order alliances come together and fight against rival males from neighbouring third-order alliances. |
|
|
3 to multiple |
permanent social units: contain around 11 females and immatures from unrelated matrilines. Caring for offspring. |
temporary intermediate social units of multiple first-level units of the same cultural clan: defence from killer whales. |
clans: can contain up to 20 000 individuals. They have culturally determined vocalizations and distinct movement patterns. |
there is a big difference in size from the second- to the third-level units and the literature suggests that intermediate levels might be missing. |
|
|
(Malurus cyaneus) |
3 |
cooperatively breeding unit: individuals that assist a breeding pair to raise a brood of young. |
supergroup: two neighbouring breeding units that merge stably. Tasks not specified. |
community: emerges from repeated interactions between three to four breeding units and supergroups. Likely cooperative defence against predators, and communal defence against competitors. |
|
|
(Manorina melanophrys) |
3 |
cooperatively breeding unit: individuals that assist a breeding pair to raise a brood of young. |
coterie or clan: occupying a discrete area within the colony. May contain one or more breeding pairs and nonbreeders. Members associate preferentially with each other, and helpers may assist more than a single pair within a coterie. |
colony: a geographically discrete aggregation of between 20 and 200 individuals that together inhabit and communally defend an area against avian intruders and predators. |
|
|
vulturine guineafowl (Acryllium vulturinum) |
3 |
cooperatively breeding units and stable clusters of males: breeding pairs can be stable from one breeding season to the next and raise young cooperatively with specific non-parents. Clusters of males (potentially highly related) remain stable across years. |
groups: can contain from 13 to 65 individuals that forage, decide collectively and move as single entities. |
stable between-group associations: driven by social preferences as well as by resource abundance and distribution. They share communal roosts and information. They also form supergroups and travel together to rarely visited areas. |
|
|
cooperatively breeding cichlid fish [47–49] (Neolamprologus pulcher) |
3 |
cooperatively breeding group: individuals assist a breeding pair to raise a brood of young. |
colony: neighbouring breeding groups jointly defend against predators. |
aggregation: feeding assemblies of members from different breeding groups exchanging social information by sporadic interactions. |