The Paper

The published Biology Letters paper here.

Author preprint version of Biology Letters manuscript here.

Electronic supplemental information of Biology Letters manuscript here.

Full Citation 

Bibi, F., Kraatz, B., Craig, N., Beech, M., Schuster, M., and Hill, A. 2012. Early evidence for complex social structure in Proboscidea from a late Miocene trackway site in the United Arab Emirates. Biology Letters, XX:XXX-XXX. doi: 10.1098/rsbl.2011.1185


Click on any of the below thumbnails for high-resolution versions.

Figure 1. Close-up of the central portion of Mleisa 1. A: The kite aerial orthophotomosaic. B: Corresponding colour map of the trackways, with the prints forming each trackway connected by lines. Track sections that were measured and reported in Fig. 2A are connected by solid lines and numbered, with tracks sections in green indicating those used to calculate the minimum herd profile (Fig. 2B). Remaining proboscidean prints are shown in blue (herd) or black (solitary individual in the herd area). A single non-proboscidean trackway is shown in red. The full site aerial photograph and map are presented as Electronic Supplementary Materials.

Figure 2. A: Stride lengths for the solitary and herd trackways. Trackways and areas are numbered according to Fig. 1B. Box plots show median, 1st and 3rd quartiles, and bars reaching the outermost data points within a 1.5 interquartile range. The horizontal grey lines mark the value of each area’s average stride length. Only a single trackway could be traced throughout all measured areas of the herd (in green). In all areas A–D, 8 to 11 individuals of intermediate to large size are recovered. Area C additionally records the presence of a small individual (trackway section 18). B: Estimated body mass profile for the minimum number of measurable Mleisa 1 proboscideans. These comprise the solitary individual (blue), and all individuals from area A plus trackways 32 from area B and 18 from area C (red). Shown also are reported body mass ranges for adult males of Loxodonta africana (La) and Elephas maximus (Em) and adult females of both species [2, 5, 17, 18].

ESM Figure 1. The Mleisa 1 site kite aerial orthophotomosaic. The full resolution orthophotomosaic is permanently archived and viewable at

ESM Figure 2. The Mleisa 1 colour-coded site map.

ESM Figure 3.  Local stratigraphy and sediments at Mleisa 1. A. Local stratigraphy as observed in the site surface and central yardang, and a view of the solitary trackway. B. View to the west showing the trackway level and trackways from within the herd. C. Close-up of one of the better-preserved prints in the trackway.