DOI: 10.25338/b8bk8g
title: Eco-correlates of coloration in primates

description: Primates are noted for their varied and complex pelage and bare skin
 coloration but the significance of this diverse coloration remains opaque.
 Using new updated information, novel scoring of coat and skin coloration,
 and controlling for shared ancestry, we reexamined and extended findings
 from previous studies across the whole order and the five major clades
 within it. Across primates we found (i) direct and indirect evidence for
 pelage coloration being driven by protective coloration strategies
 including background matching, countershading, disruptive coloration and
 aposematism, (ii) diurnal primates being more colorful, and (iii) the
 possibility that pelage color diversity is negatively associated with
 female trichromatic vision; while (iv) reaffirming avoidance of
 hybridization driving head coloration in males, (v) darker species living
 in warm, humid conditions (Gloger’s rule), and (vi) advertising to
 multiple mating partners favoring red genitalia in females. Nonetheless,
 the importance of these drivers varies greatly across clades. In
 strepsirrhines and cercopithecoids, countershading is important; greater
 color diversity may be important for conspecific signaling in more diurnal
 and social strepsirrhines; lack of female color vision may be associated
 with colorful strepsirrhines and platyrrhines; whereas cercopithecoids
 obey Gloger’s rule. Haplorrhines show background matching, aposematism,
 character displacement, and red female genitalia where several mating
 partners are available. Our findings emphasize several evolutionary
 drivers of coloration in this extraordinarily colorful order. Throughout,
 we used coarse but rigorous measures of coloration, and our ability to
 replicate findings from earlier studies opens up opportunities for
 classifying coloration of large numbers of species at a macroevolutionary
 scale.
description: Torso pattern as 0 = Not black-and-white or may be all black or
 all white; 1 = ‘Salt and pepper’ like splotches of black-and-white that
 appear conspicuous; 2 = Transverse stripes of black-and-white; 3 =
 Longitudinal stripes of black-and-white; 4 = A solid block of black/dark
 brown/dark grey on the dorsum with a solid block of white/light
 yellow/light grey on ventrum; 5 = A solid block of white/light
 yellow/light grey on dorsum with a solid block of black/dark brown/dark
 grey on ventrum; 6 = Irregular blocks of black-and-white; 7 = Disordered
 stripes of black-and-white, often resembling contor lines; 8 = Large spots
 of black on white background or vice versa. <br>
 Tail color proportions were scored as 0 = Not black or white (may be grey,
 brown or red); 1 = All black; 2 = Black more than 50% and white less than
 50%; 3 = 50% of tail black and 50% white; 4 = White more than 50% and
 black less than 50%; 5 = All white; 999 = No tail. In tail analyses we
 reformulated these categories as having any white fur (#2-5) or
 not. <br> Total number of distinct color and
 separately shades (Figure S2) were documented on each part of the body
 (with increasing numbers here termed as being ‘more colorful’). Species
 were additionally categorized as having either red (yes or no) and/or dark
 (yes or no) torso coloration (as described in the legends SOM Figures 1
 and 2). Male and female genitalia scores were 0 = not
 different from surrounding pelage, 1 = white, 2 = red or pink, 3 = black,
 4 = yellow, 5 = orange, 6 = blue. For females, we focused on category 2 in
 analyses as the vast majority of colorful female genitals are red or pink.
 Male genitalia are often multicolored so for males we combined categories
 1-6 in analyses. Activity cycle: 1 = Nocturnal, 2 =
 Crepuscular, 3 = Diurnal, 4 = Cathemeral, 12 = Nocturnal/Crepuscular, 13 =
 Diurnal/Crepuscular’ 0 = absence of data. We combined categories 2 and 4
 into a new category diurnality. Group size: 1 = 1
 individual (solitary); 2 = 2 - 5 individuals; 3 = 6 - 25 individuals; 4 =
 26 - 100 individuals; 5 = &gt;100 individuals.
 <br> Social organization: (i) pair bonded; (ii) multimale;
 (iii) fission-fusion; (iv) multimale and multifemale, or fission-fusion;
 (v) multimale, one female, (vi) one male and multifemale, and (vii)
 solitary foraging, (viii) variable, i,e., reported as (i) or (ii), or (i)
 or (vi), or (ii) or (vi), or a  ‘a few males’. We subsequently lumped
 (iii) and (iv) together as ‘fission-fusion societies’. We combined (ii),
 (iii), (iv) and (v) together as ‘multimale societies’ irrespective of
 female number. Multilevel societies from Table S2 in
 Grueter et al (2015). Mating system: (i) monogamous,
 (ii) polygynous, (iii) polyandrous and (iv) promiscuous (also termed
 polygynandrous). Some species displayed more than one mating system ((v)
 monogamous and polygynous, (vi) polygynous and promiscuous, (vii)
 monogamous and polyandrous, (viii) polyandrous and promiscuous, and (ix)
 monogamous and promiscuous) and in these cases, we recorded both. We
 lumped categories (ii), (v) and (vi) together as ‘polygynous’; and
 categories (iv), (vi), (viii) and (ix) together as
 ‘promiscuous’. Hair: “long” or “shaggy” (1) or else as
 short/absent (0). Fur thickness was scored as dense if reported as
 “dense”, “shaggy” or “thick” (1) or as not dense (0). These two variables
 were subsequently combined to generate a new variable thick hair, where
 species that had either long or thick hair were scored as 1, and those
 that had scores of 0 for both were scored as 0.
 <br> Species range overlap. Species were scored as either
 having sympatric congeners (1) or not (0). Shade
 scores: see methods. Visual system: monochromatic,
 dichromatic, trichromatic or polychromatic (i.e., sex-linked alleles in
 females). If a species was trichromatic or polymorphic (for females), it
 was recorded as a 1. If a species was dichromatic or monochromatic, it was
 recorded as a 0. Trichromatic males were scored as 1, otherwise
 0. But please see Methods in the paper
  
description: Contact Tim Caro or Theodore Stankowich for details