DOI: 10.5061/dryad.c866t1ggk
title: Multimodal communication of adult, subadult, and infant plains zebras (Equus quagga)

description: Juveniles occupy a different social niche than adults, engaging in a
 smaller diversity of social contexts and perceiving greater social risks.
 Either or both of these factors may influence the form communication takes
 in immaturity and its developmental trajectory. We investigated the
 relative influence of these social forces on the development of multimodal
 communication in plains zebras (Equus quagga). Juveniles possessed smaller
 repertoires than adults, with lower combinatorial flexibility and greater
 stereotypy, particularly for signals used in submission. When interacting
 with adults, juveniles used a larger fraction of their repertoire, but
 with reduced combinatorial flexibility. The usage of a contextually
 flexible signal, “snapping”, also shifted across development, beginning as
 a stereotyped, submissive signal before diversifying into the full range
 of adult usage.
description: <strong>Study species and
 subjects</strong>   We
 conducted our study on a population of ~2,000 wild plains zebras living in
 Ol Pejeta Nature Conservancy of Laikipia, Kenya (0°00 N, 36°56 E). This
 population experiences natural predation and is not provisioned. Our
 sample comprised 22 focal harems with an average harem size of 5 members
 (2-13 individuals). To supplement our sample size for the immature age
 class, we included additional data collected ad libitum from a subset of 5
 of the 22 followed harems followed between July 9<sup>th</sup>
 and August 9<sup>th</sup>, 2019. All other methods were the
 same across all sample periods. This inclusion allowed us to double our
 sample of immature individuals (2019 sample = 10). We are able to
 individually identify zebras based on their unique stripes. All research
 was conducted with the permission of Ol Pejeta Nature Conservancy and the
 Kenyan Ministry of Wildlife (NACOSTI/P/21/11647), and was in compliance
 with the guidelines of the Princeton University Institutional Animal Care
 and Use Committee (IACUC 1835F).  
 <strong>Observations and collection of
 data</strong>   Data
 collection occurred daily between July 13<sup>th</sup> and
 September 30<sup>th</sup>, 2021 and June
 13<sup>th</sup> and August 12<sup>th</sup>, 2022
 on the Eastern Sector of Ol Pejeta Conservancy. During this period, we
 conducted multiple focal follows of each harem, with a median inter-follow
 interval/harem 14.2 ± 6.6 days (range 2-35 days), allowing us to capture a
 range of behaviors from each individual. Focal follows occurred between
 08:00-16:30, during which time we followed two harems a day, one in the
 morning block, and another in the afternoon block. Each focal harem was
 continuously followed for 2-4 hours. Within each observation period, a
 single individual from the harem was selected as the focal animal, to be
 followed for two consecutive 30 minute blocks before a new focal
 individual was selected. All behaviours, including interactions, were
 captured with video using a Canon Vixia HR R700. Audio was recorded using
 a Sennheiser ME66 directional microphone in a Rode Blimp windshield and a
 Tascam Dr-100MKII Linear PCM recorder sampling at 48kHz.
description: # Multimodal communication of adult, subadult, and infant plains zebras
 (Equus quagga)
 [https://doi.org/10.5061/dryad.c866t1ggk](https://doi.org/10.5061/dryad.c866t1ggk) ## Description of the data and file structure Multimodal communication of adult, subadult, and infant plains zebras (*Equus quagga*) This dataset comprises social interactions engaged in by adult, sub adult (&gt;2 years old, pre-dispersal), and foal (&lt;2 years old) wild plains zebras (*Equus quagga*). Data were extracted from videos collected during focal follows of 22 known focal harems between 2021 and 2022, and data collected ad libitum from a subset of 5 of the 22 followed harems followed between July 9th and August 9th, 2019. Behaviors were coded to define the full suite of visual, acoustic, tactile, and chemical signals that were produced. Data were used to construct co-occurrence networks of multimodal communication. ### Files and variables #### File: Adult\_Juvenile20212022.xlsx **Description of data and file structure** **Adult_juvenile_20212022.** Each row represents an observed configuration of visual, tactile, acoustic, and/or chemical signals seen in a given moment in time. “Begin_Time” and “End_Time designate the time in the video that the given configuration was observed, while”Duration” designates how long the given posture was observed. All three are in “hh:mm:ss” format. The entire body, “Ears”, “Eyes”, “Upper_Lip”, “Corner_of_Lip”, “Jaw”, “Head_Neck_Position”, and “Tail” are represented for each posture except for instances where there were mutually exclusive or redundant states (e.g. “jaw open” and “bite”). All other potential components were coded only as observed, and therefore may not be present in every observation. Empty cells reflect that there was no behavior observed in that component (e.g. no tactile behavior). Within the excel file, a separate tab designates “All” (the entire dataset), “Juveniles” (all non-adults), “Adults Only”, “TwoandOlder”, “FoalsYearlings”. In addition, there are two tabs for interactions between “Juvenile-Juvenile”, and “Juvenile-Adult”. ## Code/software **OntogenyRCode.** R code used to conduct all analyses and construct all visualizations.