DOI: 10.5061/dryad.4b8gthtmw
title: Social rank, female competition for food, and behavioural time-budgets in a temperate primate multi-level society

description: Competition within primate groups often translates to a social hierarchy,
 with high-rank individuals gaining privileged access to resources,
 especially food. Golden snub-nosed monkeys Rhinopithecus roxellana live in
 a multi-level society, with multiple one-male units (OMUs), each
 containing a single adult male and several females, forming a
 breeding-band. High-rank males have increased access to high-value foods
 and spend less time feeding and more time being groomed by females,
 potentially enhancing social cohesion within their OMUs. The adults of
 each OMU mainly feed and socialize together, with food competition
 predominantly acting at the OMU level. We thus predicted that adult
 females by association attain the rank and feeding privileges of their OMU
 leader males, and make similar time-budget trade-offs. By
 food-provisioning a wild breeding-band during winter and spring, when
 natural foods are abundant or limited, respectively, we found that females
 of high-rank OMUs ate more provisioned foods at higher rates, especially
 during winter when provisioned foods had increased value. In winter,
 females of low-rank OMUs fed for longer on natural foods and females of
 high-rank OMUs longer on provisioned foods. Females of high-rank OMUs
 spent longer being groomed by other OMU members, especially during winter.
 Our results are consistent with females attaining the feeding privileges
 of their OMU leader male, enhanced during winter due to increased value of
 provisioned foods, high thermal demands, and reduced natural food
 availability. Importantly, the feeding privileges attained by females of
 high rank OMUs were less pronounced than those previously found for leader
 males, possibly due to higher tolerance between females of different OMUs.
 We suggest that behavioral time-budget effects of food competition in
 female R. roxellana enhance cohesion in high-rank OMUs, contributing to
 OMU integrity and, hence, the social structure of this multi-level
 society.
description: Field study of female-female competition in wild golden monkeys.
 The effects of seasonality and social rank were tested. Data are arranged
 in order to be analysed in R using mixed-models.
description: **Spreadsheet column headings** ** ** ** Foodstack data sheet** ** **
 **This spreadsheet is made up of values taken in spring and winter by
 recording the same group of monkeys. The values have also been split
 between two food types, provisioned and natural. This is to enable the
 comparison of food type and season. However, this results in data from the
 same individual females being included because measurements were taken
 from the same monkeys in spring and winter, and because they eat both
 provisioned and natural food on the same day. These data for the same
 monkeys have thus been ‘stacked’ on top of each other. This has been
 accounted for in our statistical analysis by using ‘individual’ as a
 random factor** ** ** **‘Day’ of data recording reflects the same ‘date’
 in each dataset. ‘Rank’ is the OMU leader male rank at the time the data
 were recorded.** ** ** ***Column headings and what they mean.***  
 **Number** – row number. To enhance data sorting and management.  
 **Season** – winter or spring. This column is used to differentiate
 between seasons.   **Date** – the date on which the data for the row were
 recorded.   **Day** – from the first through to the last, this is day in
 words. The total equals the number of days data were recorded. This
 enables ‘day’ to be used as a random factor in mixed models.   **Am temp**
 – air temperature before morning data recording.   **Pm temp** – air
 temperature before afternoon data recording.   **Observer** – the student
 who recorded the data for the particular row.   **Individual** – the
 individual adult female whose behaviour is being recorded for this row of
 data.   **OMU** – the OMU (individual OMU leader male) to which the
 individual female belongs   **OMU size** – the number of adult females
 within the foacl female’s OMU at the time of sampling   **Rank** – the
 rank on the OMU leader male to which the individual female belongs (as
 determined by David’s score data).   **Food** - the food the data for the
 particular row the female was consuming during the time of recording,
 either provisioned or natural. The same individual eats both provisioned
 and natural foods each day so the data have to be partitioned between the
 two food types. We therefore have two data rows for each individual for
 each recording day, one for provisioned food and the other for natural
 food.   **Feeding** – the time in hours the particular individual spent
 feeding on both provisioned and natural foods.   **Tottime** – the total
 time the individual had its behaviour recorded on the day of recording.  
 **Foodtime** – the time the individual spent feeding on the food type in
 ‘food’. Food time for ‘provisioned’ + foodtime for ‘natural’ therefore =
 ‘feeding.’   **Food_g** – the mass in g of food consumed by the individual
 during the day of recording for the particular food type indicated by
 ‘food.’ So we have two measures for each recording of an individual
 monkey, one for provisioned and another for natural.   **Food_jk **–
 energy intake for the food type ingested as designated in ‘food.’ There
 are thus two measures each on different rows per monkey, one for
 provisioned and another for natural.   **Propfeedtime** – the proportion
 of time spent feeding on food of the particular food type designated in
 ‘food’ of the total time spent feeding for that day’s recording:
 Foodtime/feeding. Propfeedtime for provisioned + propfeedtime for natural
 therefore = feeding.   **Feedrate** – the amount of foodg (of the type
 designated in ‘food’) consumed per unit time, out of the total time the
 monkey was feeding that day: food_g/feeding.   **Propfeedtimearc** –
 arcsine square root transformation of ‘propfeedtime.’   **Logfood g** –
 log(10) values of Food_g   **Logfeedrate** – log10 transformation of
 ‘feedrate.’   **Feedmin** – total time spent feeding ‘feeding’ in minutes.
 This creates a vector of integers.   **Foodmin** – total time spent
 feeding on a food type, either provisioned or natural, ‘foodtime’ in
 minutes.   **Totmin** – total daily recording time ‘tottime’ in minutes.  
     **Time budget data sheet** ** ** **The data concern different
 behaviours exhibited by focal female individuals during each day’s data
 recording.** **Number** – row number. To enhance data sorting and
 management.   **Season** – season data were recorded either spring or
 winter   **Date** – date when data sere recorded   **Morning temperature**
 - air temperature in the morning before recording began   **Sunset
 temperature** – temperature in afternoon   **Temperature difference** –
 difference between morning and sunset temperature   **Observer** – code
 for the person who recorder the values for this row of data  
 **Individual** – the individual female whose behaviour has been recorded  
 **Sex** –The sex of the focal individual, here (AF) all are female  
 **Unit** – the code for the OMU leader male of the OMU to which the focal
 female ‘individual’ is a member.   **OMU size** – the number of adult
 females within the same OMU as the focal female   **Rank** – OMU leader
 male rank as determined by David’s score: rank at the time data were
 recorded   **Feeding** – the time in hours the focal monkey spent feeding
 during that day’s recording.   **Not_feed** – the time spend during the
 day’s data recording by the focal monkey on activities other than feeding.
 Feeding + not_feed therefore = total observation time.   **Prop_feed** –
 the proportion of all time the focal monkey spent feeding; prop_feed =
 Feeding/(feeding + not_feed)   **Asin_feed** – prop_feed transformed to
 acrsine square root   **Moving** – time spent moving in hours  
 **Not_move** – time spent on all activities other than moving  
 **Prop_more** – proportion of all time spent moving   **Asin_move** –
 arcsine square root transformation of prop_move   **Resting** – time spent
 resting in hours   **Not_rest** – time spent on all activities other than
 resting   **Prop_rest** – proportion of all time recorded that day spent
 resting   **Asin_rest** – arcsine square root transformation of prop_rest
   **Groomer** – time spent grooming other individuals (usually within the
 same OMU)   **Not_groomer** – time spent on other activities other than
 grooming other individuals   **Prop_groomer** – proportion of all time
 spent grooming other individuals   **Asin_groomer** – arcsine square root
 transformation of prop_groomer   **Groomed** – time individual spent being
 groomed by another individual   **Not-groomed** – time spent on activities
 other than being groomed by another individual ** ** **Prop_groomed** –
 proportion of all time being groomed by another individual  
 **Asin_groomed** – arcsine square root transformation of prop_groomed  
 **Attack** – proportion time spent involved in attacking other OMUs  
 **Not_attack** – time spent not attacking other OMUs   **Prop_attack** –
 proportion of all time spent attacking other OMUs   **Asin_attack** –
 arcsine square root transformation of prop_attack   **mating** –
 proportion time spent mating   **Not_mating** – time spent not mating  
 **Prop_mating** – proportion of all time spent mating   **Asin_mating** –
 arcsine square root transformation of prop_mating   **embracing** –
 proportion time spent embracing or being embraced by other individuals  
 **Not_embracing** – time spent not embracing or being embraced by other
 individuals   **Prop_embracing** – proportion of all time spent embracing
 or being embraced by other individuals   **Asin_embracing** – arcsine
 square root transformation of prop_embracing ** ** **Social+** - time
 spent engaged in social behaviours (grooming, being groomed, and
 embracing). NB mating not included as time is minimal.   **Not_social** –
 time spent on behaviours spent on non-social activities: i.e. tottime –
 social+ **Prop_non_social** – proportion of all time spent on non-social
 behaviours   **Asin_non_social** – arcsine square root transformation of
 prop_non_social   **Prop_social** – proportion of all time spent on social
 behaviours   **Asin_social** – arcsine square root transformation of
 prop_social   **Tottime** – time in hours spend recoding the focal monkey
 that day   **Feed_prov** – time in hours spent feeding on provisioned food
   **Feed_nat** – time spent feeding on natural food   **Prop_prov_feed** –
 proportion of tottime spent feeding on provisioned food   **Pr_feedarc** –
 arsine square root transformation of prop_prov_feed   **Prop_nat_feed** –
 proportion of tottime spent feeding on natural food   **Nat_feedarc** –
 arsine square root transformation of prop_nat_feed   **Prov_feed** –
 duplicate of feed_prov   **Nat_feed** – duplicate of feed_nat