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White-faced capuchins primarily eat accomplished bake-apple and invertebrates, but they are accepted to absorb flowers in aerial abundances seasonally (i.e., up to 25% of the annual bulb diet24). Capuchins accept polymorphic colour vision, and acquire three long-to-mid amicableness acute (L/M) cone types with aiguille ashen sensitivities at 532 nm, 543 nm, and 561 nm37. All males and homozygous females are dichromats, admitting heterozygotic females display trichromacy. All behavioural abstracts were calm in accordance with protocols accustomed by the University of Calgary’s Life and Environmental Care Committee (LESACC).

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We acquired behavioural and ashen abstracts from Sector Santa Rosa (SSR), Área de Conservación Guanacaste (ÁCG), Costa Rica. This abstraction was conducted with permission from the Government of Costa Rica, and Área de Conservación Guanacaste administration. SSR is primarily close dry backwoods characterized by able aberration in precipitation. Although there is melancholia aberration in all-embracing annual affluence in SSR, flowers are almost abounding throughout the year, and abounding bulb breed aftermath flowers during the dry division back added capuchin foods are beneath abundant24.

This abstraction combines two acreage seasons: 2007–2008 and 2013–2014. In 2007–2008 we advised four acclimatized capuchin groups (CP, EX, GN, LV) over 13 months amid January 2007 and August 2008. Amid May 2013 and March 2014, we followed three groups (AD, LV, RM) for 8 months. We followed anniversary accumulation for 2–4 after abounding canicule (typically 12 hours/day) for anniversary “cycle”, with one aeon per ages in 2007–2008 (n = 13 cycles) and two per ages in 2013–2014 (n = 13 cycles). Abstraction groups are able-bodied acclimatized and alone monkeys are identifiable. Best individuals had been ahead genotyped for the L/M opsin from begrimed DNA, and we genotyped new individuals afterward the aforementioned protocols14,37. Individuals of anniversary of the six accessible colour eyes phenotypes are represented in our abstraction (Table 1). Researchers and administration accession behavioural abstracts were not abreast of any monkey’s phenotype to abbreviate eyewitness bias.

We calm reflectance spectra from flowers and leaves application a USB 4000 carriageable spectrometer and LS-1 ablaze antecedent (2007–2008) and a Jaz EL-200 spectrometer with a PX pulsed xenon ablaze antecedent (2013–2014), from Ocean Optics Ltd. Both spectrometers apprehend wavelengths amid 200–850 nm, although we use alone readings accordant to the capuchin beheld arrangement (400–700 nm38) in this study. Spectrometers were calibrated application a WS-1 reflectance accepted at the alpha of anniversary altitude session, and every 15 minutes during use to anticipate alluvion over time. Measurements were taken application the reflectance action of Ocean Optics SpectraSuite software (64-bit adaptation 1.6.0.11). To annual for baseline alluvion via aphotic correction, we connected our abstracts by ambience the everyman account aural the 400–700 nm ambit to 0 nm, and adjusting all others accordingly, application Microsoft Access 2010. Five abstracted samples were analyzed for anniversary annual allotment whenever possible. If altered annual genitalia were audibly altered colours to the trichromatic researcher’s eyes (e.g., Centrosema macrocarpum has white/cream coloured petals with a aphotic amethyst center), anniversary colour application was advised as a abstracted sample, as were the altered annual adeptness stages captivated by capuchins. In absolute we analyzed 21 annual genitalia from 14 annual aliment breed (Supplemental Data S1). To simulate a amalgamate abounding accomplishments that would best almost SSR’s forest, we analyzed the bright backdrop of the aerial and lower blade surfaces from these 14 breed added the leaves from an added 14 timberline breed accepted in the SSR forest.

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Predicting the conspicuity of altar to a non-human abbey application abstract animal colour categories is problematic, a shortcoming that can be addressed by modelling the acknowledgment of specific cone sensitivities for anniversary breed or phenotype37,38,39,40. The best ashen sensitivities (λmax) of capuchin opsin pigments accept ahead been bent via barometer the absorbance spectra of pigments reconstituted in vitro37. Application these values, we estimated the absorbance spectra of anniversary of the three beheld pigments application the adapted adaptation of Lamb’s equation41 by giving the λmax amount of anniversary beheld pigment. To quantify how arresting a annual was to anniversary colour eyes phenotype, we acclimated Just Apparent Aberration (JND) modelling37 (Supplemental Data S1). The minimum bright ambit at which two altar can be differentiated as acutely altered colours is authentic as 1 JND7,42, and altar with more college JND array are accepted to be arresting beneath beneath ideal situations. We bent the JND array of anniversary annual ambition adjoin the beggarly blade accomplishments amount for anniversary capuchin colour eyes phenotype. If a flower-leaf allegory denticulate >1 JND for a phenotype it was advised detectable, and array at atomic 1 JND college for one phenotype about to addition was advised advantageous.

To apart appraise our JND modelling after-effects we additionally analyzed our blush abstracts application “Support Vector Machine” (SVM) modelling (Supplemental Data S2), which uses a apparatus learning, leave-one-out access to adumbrate whether a accustomed capuchin colour eyes phenotype should be able to analyze a annual allotment from a amalgamate abounding accomplishments application the colour eyes pathways accessible to that phenotype14 (Supplemental Data S2). In this analysis, anniversary annual is advised and adjourned about to all leaves separately.

We conducted direct browse sampling to actuate if time adherent to annual adeptness use differed amid colour eyes phenotypes43. Every bisected hour, board amid as abounding monkeys as accessible in 10 minutes and recorded their character and behavioural state43,44. If a monkey was foraging, we recorded the blazon of aliment account whenever possible. We calm browse abstracts on ample juveniles, and adults/subadults.

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We additionally calm abstracts application a anatomy of all-occurrences sampling address we accredit to as “flower application visit” (FLPV) sampling, which is advised to abduction rare, cursory behaviours14,24,44,45. We denticulate any foraging by a capuchin accumulation to a annual application as one FLPV. Back an FLPV was noted, assemblage recorded the annual breed actuality captivated and the alone monkeys involved.

Using R software (64-bit adaptation 3.3.0) and the lme4 statistical package46 (v. 1.1–11) we acclimated a ambiguous beeline alloyed archetypal (GLMM) to actuate whether annual burning abundance (the cardinal of browse samples per abstraction aeon in which annual foraging was recorded) was afflicted by colour eyes adeptness (trichromat or dichromat). Since this assay uses calculation abstracts with a aerial cardinal of zeroes, we acclimated a abrogating binomial distribution47. We included all adults, subadults and ample juveniles present for the majority of a acreage year (i.e., at atomic 7 cycles of abstracts calm in a abstraction aeon of 13 cycles, n = 83 individuals). All individuals and abstraction cycles were advised again measures, and the absolute sampling accomplishment per alone was controlled by including an account in the model, application the log of the absolute scans per alone per cycle. We acclimated a minimum acceptation beginning of p < 0.05 for all analyses. We did not ascendancy for the phyletic relatedness of annual breed in our models. However, the 14 breed captivated during our abstraction aeon were not carefully accompanying and were from 10 altered families. Additionally, annual colouration varies appreciably amid carefully accompanying bulb breed as able-bodied as intraspecifically48,49,50; therefore, we do not accept this to be a above agency influencing our conclusions.

To appraise whether trichromats were bigger able to locate baby annual patches, we acclimated a GLMM with abrogating binomial administration to analyze the cardinal of times dichromats and trichromats were empiric in FLPVs. For this analysis, to anticipate the furnishings of challenge antagonism abashing detection-based foraging advantages, alone visits to bulb breed bearing baby annual crops were included. A bulb breed was authentic as a baby application if it alone produced a baby cardinal of flowers that provided a bound and quickly-depleted patch. A absolute of 11 breed ahead articular as capuchin annual foods met these criteria: Bromelia pinguin, Bromelia plumerii, Bauhinia ungulata, Callistemon viminalis, Centrosema macrocarpum, Cordia guanacastensis, Malvaviscus arboreus, Pithecoctineum crucigerum, Stemmedenia obovata, Tabebuia ochracea, and Vachellia collinsii. In this model, the absolute cardinal of “small-patch” FLPVs per alone monkey was the abased variable, with colour eyes phenotype acting as the augur variable. We advised accumulation associates as a accidental aftereffect and acclimated the log of the cardinal of ascertainment hours for anniversary accumulation as an account to ascendancy for differences in sampling effort.

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Foraging differences due to 51 or possessing a long-wavelength allele52 accept been predicted for Neotropical monkeys. To ascendancy for the achievability that foraging differences were (1) -based or (2) consistent from an advantage due to possessing a long-wavelength (561 nm) acute allele behindhand of whether an alone was a dichromat or trichromat, we activated another models for both annual foraging abundance and annual application appointment analyses. First, we activated for differences by comparing macho and changeable dichromats of the 561 nm acute phenotype, which was the phenotype of best (29/31) of the changeable dichromats in our abstraction citizenry during the abstraction period. We additionally ran anniversary archetypal with alone females to affirm any furnishings were not due to colour eyes adeptness differences and weren’t ashamed by . Secondly, we compared dichromatic males of phenotype 532 to those with the 561 phenotype. We begin no cogent differences in foraging ante due to or from possessing the 561 allele, accordingly and allele-specific phenotype was not included in the final models (see Supplemental Data S3 for all models with abstracts subsets).

Behavioural and spectroscopic abstracts acclimated for this arrangement are provided as Supplemental Data S4.

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