DOI: 10.5061/dryad.kd51c5bhw
title: Social and ecological drivers of behavioural diversity in wild Costa Rican capuchins

description: The extent of behavioural diversity versus synchrony exhibited among
 members of the same social group impacts group cohesion. Understanding
 social and environmental drivers of behavioural diversity has implications
 for understanding the evolution of group living and has been an enduring
 goal of biologists. Here we analyse a 15 years behavioural dataset from
 wild habituated capuchin monkeys to explore social and environmental
 drivers of behavioural diversity. Between 2007-2022, instantaneous scan
 samples were recorded every 30 minutes from 214 individuals across 8
 social groups, ranging in size from 7-35 individuals. We examine social,
 ecological, and demographic predictors of behavioural diversity at the
 group and individual level, with a focus on the impact of group size. We
 find that the behavioural richness and diversity of behaviours were
 greater in large groups compared to small and medium-sized groups, that
 individuals in small groups exhibited higher diversity across the day
 compared to individuals in medium and large groups, and that daily maximum
 temperature and rainfall were associated with decreased behavioural
 diversity at the group level. Overall, group size appears to impact
 important aspects of behavioural diversity within a group and among
 individuals in this socially complex species. The breakdown in behavioural
 synchrony of large groups likely influences movement ecology and social
 relationships and may contribute to fission events that occur at upper
 limits of group size. In the present study, we leverage a large dataset to
 provide new insights into how social and environmental variables can
 influence behavioural dynamics in an intelligent, social mammal. 
description: Detailed account of methods are available in the corresponding
 manuscript. Please see this abridged version of the methods, modified from
 the manuscript: We collected behavioral data on eight socia; group in
 rotations of 2-5 consecutive days, from sunrise to sunset. Each group was
 followed for 1-3 such rotations per month. Inter-observer reliability
 (IOR) is tested during a training period for new researchers, and they
 begin to collect data only once their IOR is &gt;95% with established
 researchers. We conducted instantaneous scan sampling every 30 minutes
 (Martin et al. 1993). For the majority of contact hours, multiple
 researchers (n = 2-5) were present for each scan. For each scan, we
 recorded the instantaneous state behavior for as many individuals as we
 could identify within a window of up to 10 minutes, using an established
 ethogram (Supplemental Table 1A). Within foraging behaviors, we collected
 more specific foraging types (Supplemental Table 1B) as well as the type
 or species of food (when possible) that each individual was foraging on.
 Prior to each scan, each member of the observer team would disperse to
 maximize the possibility of locating each group member as quickly as
 possible. One observer would record each data point, and other observers
 would dictate the behavior of the individual primates, to minimize
 duplicate entries. In the rare cases where duplicate entries were
 recorded, the second entry was removed prior to data analysis. Typically,
 the behaviors of the majority of group members were recorded within the
 first ca. 3 minutes of each scan. During times when the monkeys were
 widely dispersed (e.g., after an intergroup encounter or during inclement
 weather), observers communicated via two-way radio to help ensure that
 data points were recorded as close together as possible, but researchers
 would use the full 10-minute window to locate individuals. Not all monkeys
 in the group were always able to be found in each scan. We have taken
 steps to account for this in our models, as detailed below. Finally, to
 minimize bias in data collection, such as over-representation of more
 centrally located monkeys, we began sampling with different individuals
 each time, and moved our positioning within the group to equalize, as much
 as possible, the coverage and order of recording different group members.
 We additionally collected data on rainfall, temperature, and fruit
 biomass. These variables were z-scored in our analysis. All code to
 reproduce these results is available at
 https://github.com/webbshasta/behavioral_diversity
description: # Social and ecological drivers of behavioural diversity in wild Costa
 Rican capuchins
 [https://doi.org/10.5061/dryad.kd51c5bhw](https://doi.org/10.5061/dryad.kd51c5bhw) ## Description of the data and file structure Our goal was to examine the relationship between group size, behavioral diversity, and agonism. We collected data from 8 social groups of wild, white-faced capuchins at a long-term field site in Costa Rica. Using scan data, we sought to test the Resource Distribution Hypothesis in a wild system. ### Files and variables #### File: df\_census\_filt.csv **Description:** This file is used in analysis to examine group size over time.  ##### Variables * CensusYear: The year the data were collected. * Group: The code for the social group. The names AD, RM, GN, GN2, LV, EX, BC, and CP refer to the social groups in this study.  * CensusMonth: The month in which data were collected as an integer. * Month: The month in which data were collected as a string. * TotalGroupSize: Number of residents of the social group. * AdultFemales: Number of adult females in the social group. * MonthYear: The months and year in which data were collected. * GroupJoinKey: A key made of month, year, and social group used to link to other tables if needed. * lab: The year the data were collected. #### File: df\_combinedInd\_forage\_fruit.csv **Description:** This file is used to test whether group size is related to diversity in fruit foraging. Each of the behavior columns (i.e., all but column 1) represents the number of times that behavior was recorded for a given individual on a given day.  ##### Variables * DateIdInd: Date and individual ID * VFO: Visual foraging behavior * FIN: Foraging for invertebrates (general category) * EFI - ACOL: Extractive foraging for invertebrates from species ACOL * FOT: Forage other (general category) * FIN - CATA: Forage invertebrates (caterpillars) * FFR - XXXX: Forage for fruit, unknown species * EFI: Extractively forage for invertebrates * FFR - SEXC: Forage for fruit, species SEXC * FFR - LCAN: Forage for fruit, species LCAN * FFR - FCOT: Forage for fruit, species FCOT * FIN - INVE: Forage for invertebrates, species INVE * FFR - FUNK: Forage for fruit, species FUNK * FFR - FHON: Forage for fruit, species FHON * FIN - LCAT: Forage for invertebrates, large caterpillar * FOT - MCHI: Forage other, species MCHI * FFR - GAME: Forage for fruit, species GAME * FFR - STER: Forage for fruit, species STER * FIN - SCRU: Forage for invertebrates, small chitinous * FIN - INSE: Forage for invertebrates, insects * FIN - XXXX: For for invertebrates, category unknown * FFR - ZGUI: Forage for fruit, species ZGUI * FFR - BSIM: Forage for fruit, species BSIM * FFR - AACU: Forage for fruit, species AACU  * FFR - ACOL: Forage for fruit, species ACOL * FFR - MCAL: Forage for fruit, species MCAL * FFR - CARG: Forage for fruit, species CARG * FFR - BPLU: Forage for fruit, species BPLU * FOT - VERT: Forage other, vertebrate pray * FOT - XXXX: Forage other, category and species unknown * FOT - ACOL: Forage other, species ACOL * EFI - ECYC: Extractively forage for invertebrates from ECYC species * FOT - INSE: Forage other, insects * FFR - FMOR: Forage for fruit, species FMOR * FIN - LINS: Forage for fruit, species LINE * EFI - XXXX: Extractively forage for invertebrates, species unknown * FFR - SPUR: Forage for fruit, species SPUR * FFR - RMON: Forage for fruit, species RMON * FFR - CAME: Forage for fruit, species CAME * FFR - DDEN: Forage for fruit, species DDEN * FFR - SGLA: Forage for fruit, species SGLA * FFR - FOVA: Forage for fruit, species FOVA * FOT - ECYC: Forage other, species ECYC * FIN - LGRA: Forage for invertebrates, large grasshopper * FFR - MARG: Forage for fruit, species MARG * FFR - DKUN: Forage for fruit, species DKUN * FIN - ANTS: Forage for invertebrates, ants * FFR - AREV: Forage for fruit, species AREV * FOT - BPLU: Forage other, species BPLU * FFR - TOCH: Forage for fruit, species TOCH * FOT - SQUI: Forage other, species SQUI * FIN - STIK: Forage for invertebrates, stick bug * FIN - KATY: Forage for invertebrates, katydid * FFR - DCUN: Forage for fruit, species DCUN * FIN - COCO: Forage for invertebrates, cockroach * FOT - BPIN: Forage other, species BPIN * FFR - KCAL: Forage for fruit, species KCAL * FIN - SGRA: Forage for invertebrates, small grasshopper * FFR - TMAR: Forage for fruit, species TMAR * FFR - MTIN: Forage for fruit, species MTIN * FIN - HIVE: Forage for invertebrates from a hive * FOT - BROM: Forage other, species BROM * FFR - SOBO: Forage for fruit, species SOBO * FIN - SCAT: Forage for invertebrates, small caterpillar * FIN - CRIK: Forage for invertebrates, cricket * FFR - QOLE: Forage for fruit, species QOLE * FFR - GMAC: Forage for fruit, species GMAC * FFR: Forage for fruit, general category * FFR - SPAL: Forage for fruit, species SPAL * FIN - ECYC: Forage for invertebrates, species ECYC * FOT - HIVE: Forage other, hive * FFR - CGUA:  Forage for fruit, species CGUA * FFR - BCRA: Forage for fruit, species BCRA * FFR - LSPE: Forage for fruit, species LSPE * FOT - BSIM: Forage other, species BSIM * FFR - SMOM: Forage for fruit, species SMOM * EFI - AACU: Extractively forage for invertebrates, species AACU * FIN - CICA: Forage for invertebrates, cicada * FFR - ARET: Forage for fruit, species ARET * FFR - JPUN: Forage for fruit, species JPUN  * FIN - ATIB: Forage for invertebrates, species ATIB * FIN - SCOR: Forage for invertebrates, scorpion * FIN - ACOL: Forage for invertebrates, species ACOL * EFI - CRIK: Extractively forage for invertebrates, cricket * EFI - COCO: Extractively forage for invertebrates, cockroach * FFR - DCOS: Forage for fruit, species DCOS * FFR - RGLA: Forage for fruit, species RGLA * FIN - SPID: Forage for invertebrates, spider * FFR - FGOL: Forage for fruit, species FGOL * FFR - TRAC: Forage for fruit, species TRAC * FFR - HRAC: Forage for fruit, species HRAC * FFR - CSYL: Forage for fruit, species CSYL * FOT - BIRD: Forage other, bird * FFR - LSOR: Forage for fruit, species LSOR * FIN - COKR: Forage for invertebrates, cockroach * FOT - CRGU: Forage other, species CRGU * FOT - CMOL: Forage other, species CMOL * FFR - BPIN: Forage for fruit, species BPIN * FIN - BEET: Forage for invertebrates, beetle * FFR - FOBT: Forage for fruit, species FOBT * FFR - XFLE: Forage for fruit, species XFLE * FFR - PCUR: Forage for fruit, species PCUR * FOT - INVE: Forage other, invertebrate * FOT - COCO: Forage other, cockroach * FFR - HCOS: Forage for fruit, species HCOS * FFR - AOCC: Forage for fruit, species AOCC * FOT - LCAN: Forage other, species LCAN * FOT - LCAT: Forage other, large caterpillar * FOT - CATA: Forage other, caterpillar * EFI - CATA: Extractively forage for invertebrates, caterpillars  * FFR - EHAV: Forage for fruit, species EHAV * EFI - GAME: Extractively forage for invertebrates, species GAME * FIN - RTHU: Forage for invertebrates, species RTHU * FIN - SPAV: Forage for invertebrates, species SPAV * EFI - SPAV: Extractively forage for invertebrates, species SPAV * EFI - SSAM: Extractively forage for invertebrates, species SSAM * FFR - AVIN: Forage for fruit, species AVIN * EFI - SGLA: Extractively forage for invertebrates, species SGLA * FFR - RTHU: Forage for fruit, species RTHU * FFR - AEDU: Forage for fruit, species AEDU * FFR - MCHI: Forage for fruit, species MCHI * EFI - LCAN: Extractively forage for invertebrates, species LCAN * FOT - TAME: Forage other, species TAME * FFR - TAME: Forage for fruit, species TAME * FFR - MARB: Forage for fruit, species MARB * FFR - CPAN: Forage for fruit, species CPAN * FFR - CPEL: Forage for fruit, species CPEL * FIN - USPP: Forage for invertebrates, species USPP * FFR - PGUA: Forage for fruit, species PGUA * FFR - USPP: Forage for fruit, species USPP * FFR - PPLA: Forage for fruit, species PPLA * FOT - SMOM: Forage other, species SMOM * FFR - BOCE: Forage for fruit, species BOCE * FIN - THOR: Forage for invertebrates, species THOR * FOT - USPP: Forage other, species USPP * FFR - GULM: Forage for fruit, species GULM * FIN - ORTH: Forage for invertebrates, species ORTH * FFR - APUR: Forage for fruit, species APUR * FFR - SMEX: Forage for fruit, species SMEX * FOT - AMPH: Forage other, species AMPH * FFR - MSEG: Forage for fruit, species MSEG * EFI - USPP: Extractively forage for invertebrates, species USPP * FFR - SHAZ: Forage for fruit, species SHAZ * FFR - ORTH: Forage for fruit, species ORTH * FFR - KFER: Forage for fruit, species KFER * FFR - FCIT: Forage for fruit, species FCIT * FFR - SPAV: Forage for fruit, species SPAV * FIN - MAYA: Forage for invertebrates, species MAYA * FFR - CELA: Forage for fruit, species CELA * FOT - CPAN: Forage other, species CPAN * FFR - ATIB: Forage for fruit, species ATIB * EFI - MAYA: Extractively forage for invertebrates, species MAYA * FFR - OVER: Forage for fruit, species OVER * FFR - HCOU: Forage for fruit, species HCOU  * FFR - MMYR: Forage for fruit, species MMYR * FOT - ATIB: Forage other, species ATIB * FFR - SAPE: Forage for fruit, species SAPE * FFR - BUNG: Forage for fruit, species BUNG * FFR - CMAC: Forage for fruit, species CMAC * FFR - CLAT: Forage for fruit, species CLAT * FIN - CPEL: Forage for invertebrates, species CPEL * FFR - CVIM: Forage for fruit, species CVIM * FFR - DSAL: Forage for fruit, species DSAL * FFR - ACOR: Forage for fruit, species ACOR * EFI - COKR: Extractively forage for invertebrates, cockroach * FFR - SSPI: Forage for fruit, species SSPI * FFR - CBRE: Forage for fruit, species CBRE * FOT - CVIT: Forage other, species CVIT * FFR - CDIV: Forage for fruit, species CDIV * FFR - BCAU: Forage for fruit, species BCAU * FFR - CLUZ: Forage for fruit, species CLUZ * FFR - CVER: Forage for fruit, species CVER * FFR - MNOB: Forage for fruit, species MNOB * FFR - SGLN: Forage for fruit, species SGLN * FFR - SCAN: Forage for fruit, species SCAN * FOT - AHOL: Forage other, species AHOL * FFR - CCAN: Forage for fruit, species CCAN * FFR - PRIV: Forage for fruit, species PRIV * FFR - AHOL: Forage for fruit, species AHOL * FIN - ZSET: Forage for invertebrates, species ZSET * FFR - GLAT: Forage for fruit, species GLAT * FFR - TGLA: Forage for fruit, species TGLA * FFR - LARB: Forage for fruit, species LARB * FFR - CLAN: Forage for fruit, species CLAN * FFR - CMIC: Forage for fruit, species CMIC * FOT - SNAI: Forage other, snail * FFR - MQUI: Forage for fruit, species MQUI * FFR - ESAL: Forage for fruit, species ESAL * EFI - INSE: Extractively forage for invertebrates, insects * FFR - GBA: Forage for fruit, species GBA * FFR - MESC: Forage for fruit, species MESC * FFR - BGUI: Forage for fruit, species BGUI * FFR - PMEX: Forage for fruit, species PMEX * EFI - ANTS: Extractively forage for invertebrates, ants * FFR - CSPI: Forage for fruit, species CPEL * EFI- LGRA: Extractively forage for invertebrates, large grasshoppers * FFL - DAME: Forage for flowers, species DAME * EFI - THOR: Extractively forage for invertebrates from species THOR * FFR - BPOL: Forage for fruit, species BPOL * FFR - SRAD: Forage for fruit, species SRAD * FFR - SLAT: Forage for fruit, species SLAT * FFR - DAME: Forage for fruit, species DAME * FFL - LCAN: Forage for flowers, species LCAN * FOT - CCAP: Forage other, species CCAP * FFL - CMOL: Forage for flowers, species CMOL * EFF - STER: Extractively forage for fruit, species STER * EFF: Extractively forage for fruit  * FFR - SSCH: Forage for fruit, species SSCH * FFL - XXXX: Forage for flowers, species unknown * FFR - CPAP: Forage for fruit, species CPAP * FFL - CFAR: Forage for flowers, species CFAR * EFF - LCAN: Extractively forage for fruit, species LCAN * TotalScans: Total number of scans recorded for a given individual in a day * shannon: Shannon diversity score that ranges from 0-1 * S.obs: Number of distinct behaviors recorded in a given day\ * S.chao1: Chao1 index of richness of behaviours * se.chao1: Standard error of Chao1 index * S.ACE: Adjusted coverage estimator index * se.ACE: Standard error of adjusted coverage estimator index * MonkeyCode: Individual identity of monkey * Date: Date of individual scan * TempMax: Daily maximum temperate degrees Celsius * Rainfall: Daily rainfall in cm * Group.x: Social group of monkeys. Ad, GN, GN2, BC, CP, RM, EX, and LV are each distinct social groups. * IndJoinKey: Key for joining tables of data * DateOfBirth: Date individual was born * Sex: Biological sex of individual monkey * TempMax_z: Daily maximum temperate degrees Celsius, z-scored * Rainfall_z: Daily rainfall in cm, z-scored * AgeYears: Age of monkey in years * AgeClass: Age class of monkey (i.e. infant, small immature, large immature, adult) * GroupJoinKey: Key for joining tables of data * CensusYear: Year of study * CensusMonth: Month of study (numeric) * Month: Month of study (written word) * TotalGroupSize: Total number of monkeys inhabiting the social group  * AdultFemales: Total number of adult females inhabiting the social group * DateBegin: First day of calendar year (used to assess dominance rank) * DateEnd: Last day of calendar year  (used to assess dominance rank) * domLevel: Level of dominance in the hierarchy based on standardized ordinal ranking (i.e. low ranking, mid ranking, high ranking) * biomass monthly kg: Monthly estimate of fruit biomass per hectare of study area, measured in kg * biomass monthly kg z: Monthly estimate of fruit biomass per hectare of study area, measured in kg, z-scored * TotalGroupSize_z: Total number of monkeys inhabiting the social group, z-scored ### File: df\_combinedInd\_general\_fruit.csv **Description:** This file provides the data necessary to examine the relationship between group size and behavioral diversity at the individual level. For behavioral variables, the numbers represent the number of records that individual was seen doing a given behavior on a given day.  ##### Variables * DateIdInd: Date and individual ID * SAC: Social active behavior * RES: Resting behavior * PLA: Play behavior * SDI: Self- directed behavior * TRA: Traveling behavior * SAG: Agonistic behavior * OTH: Other behavior * DRI: Drinking * EXC: Excreting urine or feces * SRE: Social resting behavior * VIG: Vigilance behavior * FORAGE: Foraging behavior * shannon: Shannon index of behaviors * TotalScans: Number of scans recorded per individual per day * S.obs: Number of distinct behaviors observed in a day * S.chao1: Chao1 index of richness of behaviours * se.chao1: Standard error of Chao1 index * S.ACE: Adjusted Chao1 * se.ACE: Standard error of adjusted Chao1 * MonkeyCode: Individual identity of monkeys * Date: Date of data collection * TempMax: Maximum temp in C  * Rainfall: Rainfall in cm * Group.x: Name of social group * MonthYear.x: Month and year in dd-mmm * IndJoinKey: Join key with individual ID and date for joining to other tables, if needed * DateOfBirth: Date of birth for individual study animal * Sex: Sex of animal * Rainfall_z: Rainfall z-scored * AgeYears: Age in years (numeric) * AgeClass: Age class (categorical) * GroupJoinKey: Key with date and social group to join to other tables, if needed * CensusYear: Year  * CensusMonth: Month of year (numeric) * Month: Month of year (string) * TotalGroupSize: Total number of individuals inhabiting the social group * AdultFemales: Number of adult females in the social group * MonthYear.y: Month year * GroupSizeCat: Group size (categorical) * DateBegin: Start of each year (Jan 1) * DateEnd: End of each year (Dec 31) * domLevel: Level of dominance (categorical) * MonthYear: Month year * month_of: Month (numeric) * biomass_monthly_kg: Monthly measure of fruit biomass in kg/hectare * biomass_monthly_kg_z: Monthly measures of fruit biomass in kg/hectare (z-scored) * TotalGroupSize_z: Total group size (z-scored) #### File: df\_prop\_general\_fruit.csv **Description:** This file contains data that can be used to assess the relationship between group size and behavioral diversity and agonism at the group level. ##### Variables * ScanId: Id for each scan per group per day * GroupJoinKey: Key for joining group ID data to other tables * Date: Date of data collection * Time: Time of day * TempMax: Maximum temperature in degrees C * Rainfall: Rainfall in cm * Group: Social group  * NumberMonkeys: Number of monkeys recorded in a given scan * SAC: Social active behavior * RES: Resting behavior * PLA: Playing behavior * SDI: Self-directed behavior * TRA: Traveling * SAG: Agonistic behavior * OTH: Other behavior * DRI: Drinking * EXC: Excreting urine or feces * SRE: Social resting behavior * VIG: Vigilance behavior * FORAGE: Foraging behavior * TotalScans: Number of scans recorded at a given scan period * shannon: Shannon diversity index of scans at group level * S.obs: Number of distinct behaviors observed in a given scan * S.chao1: Chao1 index of behaviors observed in a given scan * se.chao1: Standard error of Chao1 * S.ACE: Adjusted coverage estimator index * se.ACE: Standard error of adjusted coverage estimator index * CensusYear: Year (numeric) * Group.y: Social group * CensusMonth: Month of data collection (numeric) * Month: Month of data collection (string) * TotalGroupSize: Total number of individuals in the group * AdultFemales: Total number of adult females in the group * MonthYear: Month and year * PropFemales: Proportion of females in the group * PropMonkeysInScan: Proportion of group that was represented in a given scan * Temp_z: Temperature in degrees C (z-scored) * Rainfall_z: Rainfall in cm (z-scored) * Year: Year * month_of: Month * biomass_monthly_kg: Measure of biomass in kg/ha (measured monthly) * biomass_monthly_kg_z: Measure of biomass in kg/ha (z-scored) * TotalGroupSize_z: Total group size (z-scored) #### File: df\_prop\_forage\_fruit.csv **Description:** This files contains data to assess the relationship between diversity of foraging and group size.  ##### Variables * ScanId: Id of scan (date, time, and social group) * VFO: Visual foraging behavior * FIN: Foraging for invertebrates (general category) * EFI - ACOL: Extractive foraging for invertebrates from species ACOL * FOT: Forage other (general category) * FIN - CATA: Forage invertebrates (caterpillars) * FFR - XXXX: Forage for fruit, unknown species * EFI: Extractively forage for invertebrates * FFR - SEXC: Forage for fruit, species SEXC * FFR - LCAN: Forage for fruit, species LCAN * FFR - FCOT: Forage for fruit, species FCOT * FIN - INVE: Forage for invertebrates, species INVE * FFR - FUNK: Forage for fruit, species FUNK * FFR - FHON: Forage for fruit, species FHON * FIN - LCAT: Forage for invertebrates, large caterpillar * FOT - MCHI: Forage other, species MCHI * FFR - GAME: Forage for fruit, species GAME * FFR - STER: Forage for fruit, species STER * FIN - SCRU: Forage for invertebrates, small chitinous * FIN - INSE: Forage for invertebrates, insects * FIN - XXXX: For for invertebrates, category unknown * FFR - ZGUI: Forage for fruit, species ZGUI * FFR - BSIM: Forage for fruit, species BSIM * FFR - AACU: Forage for fruit, species AACU  * FFR - ACOL: Forage for fruit, species ACOL * FFR - MCAL: Forage for fruit, species MCAL * FFR - CARG: Forage for fruit, species CARG * FFR - BPLU: Forage for fruit, species BPLU * FOT - VERT: Forage other, vertebrate pray * FOT - XXXX: Forage other, category and species unknown * FOT - ACOL: Forage other, species ACOL * EFI - ECYC: Extractively forage for invertebrates from ECYC species * FOT - INSE: Forage other, insects * FFR - FMOR: Forage for fruit, species FMOR * FIN - LINS: Forage for fruit, species LINE * EFI - XXXX: Extractively forage for invertebrates, species unknown * FFR - SPUR: Forage for fruit, species SPUR * FFR - RMON: Forage for fruit, species RMON * FFR - CAME: Forage for fruit, species CAME * FFR - DDEN: Forage for fruit, species DDEN * FFR - SGLA: Forage for fruit, species SGLA * FFR - FOVA: Forage for fruit, species FOVA * FOT - ECYC: Forage other, species ECYC * FIN - LGRA: Forage for invertebrates, large grasshopper * FFR - MARG: Forage for fruit, species MARG * FFR - DKUN: Forage for fruit, species DKUN * FIN - ANTS: Forage for invertebrates, ants * FFR - AREV: Forage for fruit, species AREV * FOT - BPLU: Forage other, species BPLU * FFR - TOCH: Forage for fruit, species TOCH * FOT - SQUI: Forage other, species SQUI * FIN - STIK: Forage for invertebrates, stick bug * FIN - KATY: Forage for invertebrates, katydid * FFR - DCUN: Forage for fruit, species DCUN * FIN - COCO: Forage for invertebrates, cockroach * FOT - BPIN: Forage other, species BPIN * FFR - KCAL: Forage for fruit, species KCAL * FIN - SGRA: Forage for invertebrates, small grasshopper * FFR - TMAR: Forage for fruit, species TMAR * FFR - MTIN: Forage for fruit, species MTIN * FIN - HIVE: Forage for invertebrates from a hive * FOT - BROM: Forage other, species BROM * FFR - SOBO: Forage for fruit, species SOBO * FIN - SCAT: Forage for invertebrates, small caterpillar * FIN - CRIK: Forage for invertebrates, cricket * FFR - QOLE: Forage for fruit, species QOLE * FFR - GMAC: Forage for fruit, species GMAC * FFR: Forage for fruit, general category * FFR - SPAL: Forage for fruit, species SPAL * FIN - ECYC: Forage for invertebrates, species ECYC * FOT - HIVE: Forage other, hive * FFR - CGUA:  Forage for fruit, species CGUA * FFR - BCRA: Forage for fruit, species BCRA * FFR - LSPE: Forage for fruit, species LSPE * FOT - BSIM: Forage other, species BSIM * FFR - SMOM: Forage for fruit, species SMOM * EFI - AACU: Extractively forage for invertebrates, species AACU * FIN - CICA: Forage for invertebrates, cicada * FFR - ARET: Forage for fruit, species ARET * FFR - JPUN: Forage for fruit, species JPUN  * FIN - ATIB: Forage for invertebrates, species ATIB * FIN - SCOR: Forage for invertebrates, scorpion * FIN - ACOL: Forage for invertebrates, species ACOL * EFI - CRIK: Extractively forage for invertebrates, cricket * EFI - COCO: Extractively forage for invertebrates, cockroach * FFR - DCOS: Forage for fruit, species DCOS * FFR - RGLA: Forage for fruit, species RGLA * FIN - SPID: Forage for invertebrates, spider * FFR - FGOL: Forage for fruit, species FGOL * FFR - TRAC: Forage for fruit, species TRAC * FFR - HRAC: Forage for fruit, species HRAC * FFR - CSYL: Forage for fruit, species CSYL * FOT - BIRD: Forage other, bird * FFR - LSOR: Forage for fruit, species LSOR * FIN - COKR: Forage for invertebrates, cockroach * FOT - CRGU: Forage other, species CRGU * FOT - CMOL: Forage other, species CMOL * FFR - BPIN: Forage for fruit, species BPIN * FIN - BEET: Forage for invertebrates, beetle * FFR - FOBT: Forage for fruit, species FOBT * FFR - XFLE: Forage for fruit, species XFLE * FFR - PCUR: Forage for fruit, species PCUR * FOT - INVE: Forage other, invertebrate * FOT - COCO: Forage other, cockroach * FFR - HCOS: Forage for fruit, species HCOS * FFR - AOCC: Forage for fruit, species AOCC * FOT - LCAN: Forage other, species LCAN * FOT - LCAT: Forage other, large caterpillar * FOT - CATA: Forage other, caterpillar * EFI - CATA: Extractively forage for invertebrates, caterpillars  * FFR - EHAV: Forage for fruit, species EHAV * EFI - GAME: Extractively forage for invertebrates, species GAME * FIN - RTHU: Forage for invertebrates, species RTHU * FIN - SPAV: Forage for invertebrates, species SPAV * EFI - SPAV: Extractively forage for invertebrates, species SPAV * EFI - SSAM: Extractively forage for invertebrates, species SSAM * FFR - AVIN: Forage for fruit, species AVIN * EFI - SGLA: Extractively forage for invertebrates, species SGLA * FFR - RTHU: Forage for fruit, species RTHU * FFR - AEDU: Forage for fruit, species AEDU * FFR - MCHI: Forage for fruit, species MCHI * EFI - LCAN: Extractively forage for invertebrates, species LCAN * FOT - TAME: Forage other, species TAME * FFR - TAME: Forage for fruit, species TAME * FFR - MARB: Forage for fruit, species MARB * FFR - CPAN: Forage for fruit, species CPAN * FFR - CPEL: Forage for fruit, species CPEL * FIN - USPP: Forage for invertebrates, species USPP * FFR - PGUA: Forage for fruit, species PGUA * FFR - USPP: Forage for fruit, species USPP * FFR - PPLA: Forage for fruit, species PPLA * FOT - SMOM: Forage other, species SMOM * FFR - BOCE: Forage for fruit, species BOCE * FIN - THOR: Forage for invertebrates, species THOR * FOT - USPP: Forage other, species USPP * FFR - GULM: Forage for fruit, species GULM * FIN - ORTH: Forage for invertebrates, species ORTH * FFR - APUR: Forage for fruit, species APUR * FFR - SMEX: Forage for fruit, species SMEX * FOT - AMPH: Forage other, species AMPH * FFR - MSEG: Forage for fruit, species MSEG * EFI - USPP: Extractively forage for invertebrates, species USPP * FFR - SHAZ: Forage for fruit, species SHAZ * FFR - ORTH: Forage for fruit, species ORTH * FFR - KFER: Forage for fruit, species KFER * FFR - FCIT: Forage for fruit, species FCIT * FFR - SPAV: Forage for fruit, species SPAV * FIN - MAYA: Forage for invertebrates, species MAYA * FFR - CELA: Forage for fruit, species CELA * FOT - CPAN: Forage other, species CPAN * FFR - ATIB: Forage for fruit, species ATIB * EFI - MAYA: Extractively forage for invertebrates, species MAYA * FFR - OVER: Forage for fruit, species OVER * FFR - HCOU: Forage for fruit, species HCOU  * FFR - MMYR: Forage for fruit, species MMYR * FOT - ATIB: Forage other, species ATIB * FFR - SAPE: Forage for fruit, species SAPE * FFR - BUNG: Forage for fruit, species BUNG * FFR - CMAC: Forage for fruit, species CMAC * FFR - CLAT: Forage for fruit, species CLAT * FIN - CPEL: Forage for invertebrates, species CPEL * FFR - CVIM: Forage for fruit, species CVIM * FFR - DSAL: Forage for fruit, species DSAL * FFR - ACOR: Forage for fruit, species ACOR * EFI - COKR: Extractively forage for invertebrates, cockroach * FFR - SSPI: Forage for fruit, species SSPI * FFR - CBRE: Forage for fruit, species CBRE * FOT - CVIT: Forage other, species CVIT * FFR - CDIV: Forage for fruit, species CDIV * FFR - BCAU: Forage for fruit, species BCAU * FFR - CLUZ: Forage for fruit, species CLUZ * FFR - CVER: Forage for fruit, species CVER * FFR - MNOB: Forage for fruit, species MNOB * FFR - SGLN: Forage for fruit, species SGLN * FFR - SCAN: Forage for fruit, species SCAN * FOT - AHOL: Forage other, species AHOL * FFR - CCAN: Forage for fruit, species CCAN * FFR - PRIV: Forage for fruit, species PRIV * FFR - AHOL: Forage for fruit, species AHOL * FIN - ZSET: Forage for invertebrates, species ZSET * FFR - GLAT: Forage for fruit, species GLAT * FFR - TGLA: Forage for fruit, species TGLA * FFR - LARB: Forage for fruit, species LARB * FFR - CLAN: Forage for fruit, species CLAN * FFR - CMIC: Forage for fruit, species CMIC * FOT - SNAI: Forage other, snail * FFR - MQUI: Forage for fruit, species MQUI * FFR - ESAL: Forage for fruit, species ESAL * EFI - INSE: Extractively forage for invertebrates, insects * FFR - GBA: Forage for fruit, species GBA * FFR - MESC: Forage for fruit, species MESC * FFR - BGUI: Forage for fruit, species BGUI * FFR - PMEX: Forage for fruit, species PMEX * EFI - ANTS: Extractively forage for invertebrates, ants * FFR - CSPI: Forage for fruit, species CPEL * EFI- LGRA: Extractively forage for invertebrates, large grasshoppers * FFL - DAME: Forage for flowers, species DAME * EFI - THOR: Extractively forage for invertebrates from species THOR * FFR - BPOL: Forage for fruit, species BPOL * FFR - SRAD: Forage for fruit, species SRAD * FFR - SLAT: Forage for fruit, species SLAT * FFR - DAME: Forage for fruit, species DAME * FFL - LCAN: Forage for flowers, species LCAN * FOT - CCAP: Forage other, species CCAP * FFL - CMOL: Forage for flowers, species CMOL * EFF - STER: Extractively forage for fruit, species STER * EFF: Extractively forage for fruit  * FFR - SSCH: Forage for fruit, species SSCH * FFL - XXXX: Forage for flowers, species unknown * FFR - CPAP: Forage for fruit, species CPAP * FFL - CFAR: Forage for flowers, species CFAR * EFF - LCAN: Extractively forage for fruit, species LCAN * TotalScans: Total number of scans recorded for the group at a given scan time * shannon: Shannon diversity score that ranges from 0-1 * S.obs: Number of distinct behaviors recorded in a given scan * S.chao1: Chao1 index of richness of behaviours * se.chao1: Standard error of Chao1 index * S.ACE: Adjusted coverage estimator index * se.ACE: Standard error of adjusted coverage estimator index * GroupJoinKey: Key to join data tables * Date: Date of study  * Time: Time of day in hh:mm:ss * TempMax: Maximum temperature recorded on a study day in degrees Celsius * Rainfall: Total rainfall recorded on a given study day in cm * Group: Social group of monkeys. AD, GN, LV, GN2, BC, CP, EX, and RM are each distinct social groups. * NumberMonkeys: Number of monkeys recorded in a given scan. * Year: Year of study * CensusMonth: Month of study (numeric) * Month: Month of study (written word) * TotalGroupSize: Total number of monkey that inhabit the social group * AdultFemales: Total number of adult female monkeys that inhabit the social group * MonthYear: Month of year (dd-mmm) * PropFemales: Proportion of group that is adult females * PropMonkeys: Proportion of monkeys represented in a given scan * Temp_z: Maximum temperature recorded on a study day in degrees Celsius, z-scored * Rainfall_z: Total rainfall recorded on a given study day in cm (z-scored) * biomass_monthly_kg: Measure of biomass in kg/ha (measured monthly) * biomass_monthly_kg_z: Measure of biomass in kg/ha (z-scored) * TotalGroupSize_z: Total number of monkeys in a social group, z-scored NA - Not applicable ## Code/software Analyses for this project were run using R version 4.3.0. The following packages are required: vegan, lubridate, dplyr, tidyr, nlme, ggplot2, sjPlot, ggpubr, fuzzyjoin, car, jcolors, and glmmTMB. ## Access information Other publicly accessible locations of the data: * NA Data was derived from the following sources: * NA