Cited by 235Primate Behavior and EcologyNeuroendocrine regulation and behaviorAnimal Behavior and Reproduction

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Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees

Open access

Wittig, Roman M., Crockford, Catherine, Deschner, Tobias, Langergraber, Kevin E., Ziegler, Toni E., Zuberbühler, Klaus·2014·Proceedings of the Royal Society B: Biological Sciences

DOI 10.1098/rspb.2013.3096PMC PMC3906952

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Abstract

Humans excel in cooperative exchanges between unrelated individuals. Although this trait is fundamental to the success of our species, its evolution and mechanisms are poorly understood. Other social mammals also build long-term cooperative relationships between non-kin, and recent evidence shows that oxytocin, a hormone involved in parent–offspring bonding, is likely to facilitate non-kin as well as kin bonds. In a population of wild chimpanzees, we measured urinary oxytocin levels following a rare cooperative event—food sharing. Subjects showed higher urinary oxytocin levels after single food-sharing events compared with other types of social feeding, irrespective of previous social bond levels. Also, urinary oxytocin levels following food sharing were higher than following grooming, another cooperative behaviour. Therefore, food sharing in chimpanzees may play a key role in social bonding under the influence of oxytocin. We propose that food-sharing events co-opt neurobiological mechanisms evolved to support mother–infant bonding during lactation bouts, and may act as facilitators of bonding and cooperation between unrelated individuals via the oxytocinergic system across social mammals.

Checked 2026-10-09 · 1 reference not checked yet

AI snapshot

Written from the paper’s full text.

TL;DR

In wild chimpanzees, urinary oxytocin was higher after food-sharing events than after social feeding without sharing, and higher after food sharing than after grooming. The authors propose that food sharing co-opts lactation-related oxytocin mechanisms to support bonding and cooperation among kin and non-kin.

Key findings

Urinary oxytocin after food sharing was significantly higher than after non-sharing social feeding: mean log10 OT 1.407 ± 0.102 versus 0.884 ± 0.066 pg mg⁻¹ crea; F₁,₇₉ = 9.623, p = 0.003.
Food sharing was associated with higher urinary oxytocin than grooming: mean log10 OT 1.41 ± 0.10 versus 0.99 ± 0.05 pg mg⁻¹ crea; F₁ = 16.496, p < 0.001; a within-subject subset also showed this difference (n = 9, Wilcoxon exact p = 0.035).
Within food-sharing samples, oxytocin did not differ significantly by kin/non-kin, bond/non-bond, giving/receiving, meat/other food, or sex combination; models 2a and 2b did not explain variance better than null models. Food sharing with bond partners had the highest levels (1.42 ± 0.11 pg mg⁻¹ crea), significantly higher than grooming with bond partners (1.13 ± 0.07), but not significantly higher than food sharing with non-bond partners (1.36 ± 0.23).
The study observed 42 food-sharing events in 2009; urine samples were obtained after 33 food-sharing and 46 social feeding events. Shared foods were meat (46%), fruit (30%), rotten wood pith (18%), and honey (6%).

Methods

Observational field study of the Sonso community, Budongo Forest, Uganda, January 2009–July 2010; 79 urine samples from 26 chimpanzees (10 adult females, 3 subadult females, 6 adult males, 7 subadult males; mean 3.0 ± 1.97 samples per chimpanzee).
Urine was collected 15–60 min after single food-sharing events or after 1 h of social feeding without sharing; oxytocin was measured by enzyme immunoassay and expressed as pg mg⁻¹ creatinine.
Relationship quality was assessed with the Composite Relationship Index from focal/all-occurrence sampling; kinship was determined from faecal DNA; dominance was based on pant-grunts. Five generalized linear mixed models tested log10-transformed urinary oxytocin with subject, partner, and/or dyad identity as random factors.

Limitations

Stated: models 2a and 2b had small sample sizes (n = 33), which may have reduced power and caused instability; there were relatively few food-sharing samples between non-bond partners; the relationship between central and peripheral oxytocin release remains unclear; and whether food sharing has a stronger bonding effect than grooming remains to be tested.
Evident from the design: the study is correlational and observational, so it cannot establish causality between food sharing and oxytocin release; oxytocin was measured after events without baseline samples; and results come from one wild chimpanzee community, limiting generalization.

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Registry

Crossref · 10.1098/rspb.2013.3096

Cites

87 of 89 references verified0 citation errors · 1 not checkable

Statistics

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Claims the paper makes

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A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Subjects showed higher urinary oxytocin levels after single food-sharing events compared with other types of social feeding, irrespective of previous social bond levels.”
- Also stated in: § (b) Link between food sharing and urinary oxytocin levels · plain
  “Urinary oxytocin concentrations after social feeding events were significantly higher with food sharing than without (F 1,79 ¼ 9.623, p ¼ 0.003, table 3 and figure 1; mean log 10 OT share + s.e.¼ 1.407 + 0.102 pg mg 21 crea versus mean log 10 OT non-share + s.e.¼ 0.884 + 0.066 pg mg 21 crea).”
- Also stated in: § 4. Discussion · boosted
  “Our results provide empirical evidence that food-sharing events in chimpanzees are associated with significantly higher urinary oxytocin levels than non-sharing social feeding events.”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > Abstract
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A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Also, urinary oxytocin levels following food sharing were higher than following grooming, another cooperative behaviour.”
- Also stated in: § (c) Comparison of urinary oxytocin levels following either food-sharing or grooming events · plain
  “Urinary oxytocin levels following food-sharing events (mean log 10 OT + s.e.¼ 1.41 + 0.10 pg mg 21 crea) were significantly higher than following grooming events (mean log 10 OT + s.e.¼ 0.99 + 0.05 pg mg 21 crea; table 5a and figure 2a).”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > Abstract
Load-bearingStrength:plain· not yet calibratedCausal:correlational· not yet calibrated
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passes

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Probably not a substantial claim. 32
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In the paper
“Also, it should be noted that in contrast to the Gombe field site (Tanzania) where the harassment avoidance hypothesis was examined [55], in Budongo we only observed low and not high harassment levels.”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > 4. Discussion
What the sources say
- Not checked yetcites [55] source
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In the paper
“food share 1 9.623 0.003 share 0.427 3.102 0.003”
- Also stated in: § 4. Discussion · plain
  “sex of subject 1 0.127 0.725 male 20.043 20.357 0.725 age of subject 1 ,0.000 0.989 adult 20.002 20.014 0.989 monopolizability of food 1 1.522 0.221 high 0.171 1.234 0.221”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > 4. Discussion
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In the paper
“Physiologically, oxytocin acts in the brain by reducing anxiety and fear [19,20], enhancing social memory [16][17][18]21] and activating neural reward circuits [21].”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > 1. Introduction
What the sources say
- Not checked yetcites [19] source
- Not checked yetcites [20] source
- Not checked yetcites [16][17][18]21]
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Strength:plain· not yet calibratedCausal:direct· not yet calibrated
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Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
In the paper
“Such a link is also evident in chimpanzees, where grooming among bonding partners is associated with higher urinary oxytocin concentration than grooming between non-bonding partners [15].”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > 1. Introduction
What the sources say
- Not checked yetcites [15] source
Strength:plain· not yet calibratedCausal:correlational· not yet calibrated
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Probably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“When centrally administered, oxytocin enhances both affiliation with bonding partners and aversion to strangers in unrelated female voles [14].”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > 1. Introduction
What the sources say
- Not checked yetcites [14] source
Strength:plain· not yet calibratedCausal:direct· not yet calibrated
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Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
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In the paper
“Additionally, in humans, exogenously administered oxytocin enhances the encoding of social memory, both of positive [30] and negative [31] events, suggesting that higher oxytocin levels improve memorization of social interactions.”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > 1. Introduction
What the sources say
- Not checked yetcites [30] source
- Not checked yetcites [31] source
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In the paper
“In humans and other social mammals, the hormone oxytocin plays a central role in facilitating bonding between mother and offspring, and between mating partners [16][17][18].”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > 1. Introduction
What the sources say
- Not checked yetcites [16][17][18]
Strength:plain· not yet calibratedCausal:direct· not yet calibrated
Tests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passes
Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
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In the paper
“Recent studies show that, in addition to humans, other social mammals form cooperative relationships between unrelated adults, which can last over months or years (chimpanzees [7], baboons [8,9] and feral horses [10]).”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > 1. Introduction
What the sources say
- Not checked yetcites [7] source
- Not checked yetcites [8] source
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None of the sources cited for this claim have been checked yet.
In the paper
“If exogenously administered, oxytocin enhances cooperative behaviour [22,23] between related animals, and behaviours associated with trust [24][25][26] and generosity [27] between unrelated humans.”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > 1. Introduction
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- Not checked yetcites [22] source
- Not checked yetcites [23] source
- Not checked yetcites [24][25][26]
- Not checked yetcites [27] source
Strength:plain· not yet calibratedCausal:direct· not yet calibrated
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Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“In humans, the concentration of plasma oxytocin is related to the perceived closeness to [28] or support received by [29] a partner, suggesting a relation between higher oxytocin levels and stronger social bonding.”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > 1. Introduction
What the sources say
- Not checked yetcites [28] source
- Not checked yetcites [29] source
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In the paper
“Food sharing in chimpanzees, however, is not restricted to meat but has also been reported with non-meat foods, for example honey [52], or large fruits such as Carica papaya [57] or Treculia africana [52].”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > 1. Introduction
What the sources say
- Not checked yetcites [52] source
- Not checked yetcites [57] source
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Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
In the paper
“To compensate for variation in urine concentration, we measured creatinine (crea) levels in each sample [66] and expressed all oxytocin values as pg mg 21 crea.”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > (c) Urine sampling and oxytocin extraction
What the sources say
- Not checked yetcites [66] source
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Probably not a substantial claim. It may state two separate claims as one.
In the paper
“In both conditions, samples were not collected if grooming or copulation also occurred within 60 min prior to urination, as both of these behaviours are likely to independently increase urinary oxytocin levels [39].”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > (c) Urine sampling and oxytocin extraction
What the sources say
- Not checked yetcites [39] source
Load-bearingStrength:hedged· not yet calibratedCausal:conditional· not yet calibrated
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In the paper
“Variables did not exhibit problems of collinearity [68] (Kendall's t and Spearman's r , 0.7 in all cases).”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > (f ) Statistics
What the sources say
- Not checked yetcites [68]
Strength:plain· not yet calibrated
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Probably not a substantial claim. The paper’s conclusions may not depend on it.
None of the sources cited for this claim have been checked yet.
In the paper
“Significantly, peripheral measures of oxytocin from plasma or urine correlate positively with psychological and behavioural patterns, such as aversion reduction in male mice [19], social contact in marmosets [40], lactation in rhesus macaques [41], rates of affiliative behaviours in pair-bonded tamarins [42] and grooming in closely bonded chimpanzees [15], indicating that peripheral oxytocin level”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > 1. Introduction
What the sources say
- Not checked yetcites [19] source
- Not checked yetcites [40] source
- Not checked yetcites [41] source
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In the paper
“As in some species, low testosterone is linked to nurturance (i.e.behaviours that involve gentle warm contact with others) [70] and high oxytocin is linked to social bonding [70], and together these results suggest that food sharing might provide the optimal conditions for social bonding (see [70]).”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > 4. Discussion
What the sources say
- Not checked yetcites [70] source
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In the paper
“Food sharing between unrelated individuals outside the mating context is rare but has been reported for humans, vampire bats, chimpanzees and bonobos [13,46,48,49,52].”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > 1. Introduction
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- Not checked yetcites [13] source
- Not checked yetcites [46] source
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- Not checked yetcites [52] source
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Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“In most species, food sharing occurs between kin, such as parents provisioning their offspring (birds [44], mammals [45,47,49]), or between mating partners (insects [44], birds [43], mammals [45,47,50], humans [51]).”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > 1. Introduction
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In the paper
“In chimpanzees and vampire bats, food sharing typically occurs between individuals that also engage in other cooperative behaviours at higher rates, such as grooming and providing coalitionary support [13,49,[52][53][54].”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > 1. Introduction
What the sources say
- Not checked yetcites [13,49,[52][53][54]
Strength:plain· not yet calibratedCausal:correlational· not yet calibrated
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In the paper
“Methodologically important is the fact that oxytocin not only acts within the brain, but is also released into the peripheral circulation before being excreted in urine, although the exact nature of the relationship between central and peripheral releases have yet to be clarified [34][35][36][37][38][39].”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > 1. Introduction
What the sources say
- Not checked yetcites [34][35][36][37][38][39]
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None of the sources cited for this claim have been checked yet.
In the paper
“Indeed, the possibility that food sharing may serve several functions in chimpanzees seems supported by several studies [61,72].”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > 4. Discussion
What the sources say
- Not checked yetcites [61] source
- Not checked yetcites [72] source
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None of the sources cited for this claim have been checked yet.
In the paper
“Also, as oxytocin is thought to act on positive feedback circuits [16,17,21], sharers may experience a mutual increase in positive attitude [15,79,80], resulting in the promotion of stronger social bonds and longer-term benefits of more frequent cooperation [23,26,80].”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > (b) Oxytocin circuitry co-opted for non-kin cooperation hypothesis
What the sources say
- Not checked yetcites [16] source
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None of the sources cited for this claim have been checked yet.
In the paper
“This positive feedback circuitry is considered a key mechanism for bond formation between mother and infant [34,75].”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > (a) Food-sharing -lactation hypothesis
What the sources say
- Not checked yetcites [34] source
- Not checked yetcites [75] source
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In the paper
“In some mammals, maternal care and provisioning of food, however, aids infant survival beyond the age of lactation [75].”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > (a) Food-sharing -lactation hypothesis
What the sources say
- Not checked yetcites [75] source
Load-bearingStrength:plain· not yet calibratedCausal:direct· not yet calibrated
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None of the sources cited for this claim have been checked yet.
In the paper
“Such links between food sharing and long-term benefits have been described on a behavioural level for chimpanzees reciprocally sharing meat [7,78], and exchanging meat for sex [50] or agonistic support [54].”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > (b) Oxytocin circuitry co-opted for non-kin cooperation hypothesis
What the sources say
- Not checked yetcites [7] source
- Not checked yetcites [78] source
- Not checked yetcites [50] source
- Not checked yetcites [54] source
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Tests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passes
It may state two separate claims as one.
In the paper
“suggesting that both perceived the interaction positively [70].”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > 4. Discussion
What the sources say
- Not checked yetcites [70] source
Load-bearingStrength:hedged· not yet calibratedn = 33
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None of the sources cited for this claim have been checked yet.
In the paper
“It is thought that the interface between oxytocin and the dopaminergic reward system may contribute to the success of mother-infant and pair-bonding processes [18,[75][76][77].”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > (a) Food-sharing -lactation hypothesis
What the sources say
- Not checked yetcites [18] source
- Not checked yetcites [75] source
- Not checked yetcites [76] source
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Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passes
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None of the sources cited for this claim have been checked yet.
In the paper
“This link between food sharing and oxytocin found in chimpanzees may also be relevant for humans [70], where pro-social behaviour has often been linked to food sharing and provisioning [46,81,82].”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > (b) Oxytocin circuitry co-opted for non-kin cooperation hypothesis
What the sources say
- Not checked yetcites [70] source
- Not checked yetcites [46] source
- Not checked yetcites [81] source
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Probably not a substantial claim. It may be too vague to check against evidence. The paper’s conclusions may not depend on it.
None of the sources cited for this claim have been checked yet.
In the paper
“and is well known to be connected to peripheral and central oxytocin release in mammals [34,74].”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > (a) Food-sharing -lactation hypothesis
What the sources say
- Not checked yetcites [34] source
- Not checked yetcites [74] source
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In the paper
“In mammals, lactation has been described as the primary form of food sharing [73]”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > (a) Food-sharing -lactation hypothesis
What the sources say
- Not checked yetcites [73] source
Load-bearingStrength:plain· not yet calibrated
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passes
Probably not a substantial claim.
None of the sources cited for this claim have been checked yet.
In the paper
“What makes food sharing in chimpanzees unusual in the animal kingdom is that, as well as occurring between kin [52] and in sexual contexts [50], food sharing also occurs between non-kin adults in non-sexual contexts [13,15,52,53].”
§ Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees > (b) Oxytocin circuitry co-opted for non-kin cooperation hypothesis
What the sources say
- Not checked yetcites [52] source
- Not checked yetcites [50] source
- Not checked yetcites [13] source
- Not checked yetcites [15] source
- Not checked yetcites [53] source
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Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passes
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51 of 51

The paper relies on this without support.
Baseline urinary oxytocin levels before food-sharing events do not differ systematically from those before non-sharing social feeding events.
“Our target behaviours were single food-sharing events or 1 h of feeding in the presence of chimpanzees without sharing food.”
The paper relies on this without support.
Unmeasured factors do not confound the association between food sharing and urinary oxytocin levels.
“Thus, our results suggest a direct link between food-sharing events and urinary oxytocin levels.”
The paper relies on this without support.
The food-sharing events observed in the Sonso community are representative of food sharing in wild chimpanzees.
“We therefore hypothesize that food sharing may be a key behaviour for social bonding in chimpanzees.”

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Model 3b behavioural-context test has conflicting degrees of freedom
Minor
The results report F with 1 numerator degree of freedom for the five-context comparison, whereas Table 5b reports 4 degrees of freedom for the same behavioural-context test with F = 9.821.
Model 2a predictors differ between the table caption and results
Minor
The Table 4 caption lists sex combination as a Model 2a predictor, but the Model 2a results list sharing direction instead. The reported three predictor degrees of freedom correspond to the three variables in the results, not to both variables.

2 of 2

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Authors: Wittig, R. M., Crockford, C., Deschner, T., Langergraber, K. E., Ziegler, T. E., & Zuberbühler, K.
Affiliations: Max Planck Society (DE)
Boston University (US)
University of Wisconsin–Madison (US)
University of St Andrews (GB)
Corresponding author: Wittig, Roman M.
Crockford, Catherine
ORCID: Wittig, Roman M.
Crockford, Catherine
Deschner, Tobias
Langergraber, Kevin E.
Ziegler, Toni E.
Zuberbühler, Klaus
Venue: Proceedings of the Royal Society B: Biological Sciences
Publisher: The Royal Society
Published: 2014-03-07
Volume: 281
Issue: 1778
Pages: 20133096
ISSN: 0080-4649 (ISSN-L)
0962-8452
1471-2954
DOI: 10.1098/rspb.2013.3096
PMC: PMC3906952
OpenAlex: W2123761254
Cited by: 235
References: 87
Citations by year: 2026: 4 · 2025: 13 · 2024: 20 · 2023: 11 · 2022: 20 · 2021: 14 · 2020: 17 · 2019: 29 · 2018: 16 · 2017: 28 · 2016: 33 · 2015: 19 · 2014: 8
FWCI: 43.40
Type: journal article
Language: en
Open access: bronze
doi.org
Licence: royalsociety.org/journals/ethics-policies/data-sharing-mining/ (tdm)
Locations: Proceedings of the Royal Society B Biological Sciences (publishedVersion)
Europe PMC (PubMed Central) (publishedVersion)
PubMed Central (publishedVersion)
PubMed (publishedVersion)
Max Planck Digital Library (submittedVersion)
MPG.PuRe (Max Planck Society) (submittedVersion)
Funders: Wenner-Gren Foundation
Royal Zoological Society of Scotland
Leverhulme Trust
British Academy
Max-Planck-Institut für Evolutionäre Anthropologie
National Institutes of Health (P51OD011106)
U.S. Public Health Service (NCRR000167)
National Center for Research Resources
Topics: Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Primate Behavior and Ecology
Life Sciences › Neuroscience › Endocrine and Autonomic Systems › Neuroendocrine regulation and behavior
Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Animal Behavior and Reproduction
Keywords: food sharing, social bonding, chimpanzees, non-kin cooperation, grooming, oxytocinergic system, social evolution, cooperative behavior
MeSH terms: Animals, Pan troglodytes, Cooperative Behavior, Feeding Behavior, Female, Grooming, Male, Object Attachment, Oxytocin

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Food sharing is linked to urinary oxytocin levels and bonding in related and unrelated wild chimpanzees

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