1335
Fruit color and contrast in seasonal habitats – a case study from
a cerrado savanna
Maria Gabriela G. Camargo, Eliana Cazetta, H. Martin Schaefer and L. Patrícia C. Morellato
M. G. G. Camargo (gabicamargo@yahoo.com) and L. P. C. Morellato, Depto de Botânica, Laboratório de Fenologia, Grupo de Fenologia e
Dispersão de Sementes, Inst. de Biociências, Univ. Estadual Paulista (UNESP), Avenida 24A 1515, CEP 13506-900, Rio Claro, SP, Brazil.
– E. Cazetta, Depto de Ciências Biológicas, Universidade Estadual de Santa Cruz, Rodovia Ilhéus-Itabuna km 16, CEP 45662-900,
Ilhéus, BA, Brazil. – H. M. Schaefer, Dept of Evolutionary Biology and Animal Ecology, Faculty of Biology, Univ. of Freiburg, Hauptstr. 1,
DE-79104 Freiburg, Germany.
Communication contributes to mediate the interactions between plants and the animals that disperse their genes. As yet,
seasonal patterns in plant–animal communication are unknown, even though many habitats display pronounced seasonal-
ity e.g. when leaves senescence. We thus hypothesized that the contrast between fruit displays and their background vary
throughout the year in a seasonal habitat. If this variation is adaptive, we predicted higher contrasts between fruits and
foliage during the fruiting season in a cerrado–savanna vegetation, southeastern Brazil. Based on a six-year data base of fruit
ripening and a one-year data set of fruit biomass, we used relectance measurements and contrast analysis to show that fruits
with distinct colors difered in the beginning of ripening and the peak of fruit biomass. Black, and particularly red fruits,
that have a high contrast against the leaf background, were highly seasonal, peaking in the wet season. Multicolored and
yellow fruits were less seasonal, not limited to one season, with a bimodal pattern for yellow ones, represented by two peaks,
one in each season. We further supported the hypothesis that seasonal changes in fruit contrasts can be adaptive because
fruits contrasted more strongly against their own foliage in the wet season, when most fruits are ripe. Hence, the seasonal
variation in fruit colors observed in the cerrado–savanna may be, at least partly, explicable as an adaptation to ensure high
conspicuousness to seed dispersers.
Traditionally, fruit colors are thought to represent adap-
tations to seed dispersers, which are regarded as the main
selective pressure on fruit color evolution (Darwin 1862,
Kerner 1895, Willson and Whelan 1990). During the 1980s
and early 1990s, the inluential ‘syndrome hypothesis’
assumed that seed dispersers have strong preferences for
certain colors and that these preferences inluence the
evolution of fruit coloration (Janson 1983, Gautier-Hion
et al. 1985). Although some studies reported color prefer-
ences of fruit consumers (Puckey et al. 1996, Siitari et al.
1999, Whitney 2005), these preferences were often transient
(Schmidt and Schaefer 2004), and most studies did not
support the syndrome hypothesis owing to inconsistent
color choices (Willson et al. 1990, Willson and Comet 1993,
Traveset and Willson 1998, Schmidt et al. 2004).
Alternatively to color preferences, diurnal seed dispersers
might select fruit colors by consistently detecting a color
more easily (Kerner 1895). his hypothesis, termed later
‘contrast hypothesis’ (Schmidt et al. 2004) is consistent
with the main tenet of signal theory: that detectability is
one of the main factors driving the evolution of signals
(Schluter and Price 1993). Supporting the contrast hypo-
thesis, studies have documented that red and black, the
most common colors of bird-dispersed fruits, contrast
more strongly against the background than other fruit
colors (Lee et al. 1994, Schmidt et al. 2004). Furthermore,
the rate at which birds pecked artiicial fruits was a function
of the conspicuousness of these fruits in southeastern
Brazil (Cazetta et al. 2009). hus, if seed dispersers select
fruits mainly on the basis of their detectability, the contrasts
between fruit and background (leaves, bark and structures
associated with the fruit display) as they are perceived by
animals are more important, from an evolutionary perspec-
tive, than the color of a fruit per se (Schmidt et al. 2004,
Melo et al. 2011). here is also evidence that secondary
structures (non-green pedicels, capsules or unripe fruits
associated with the ripe fruits, according to Schaefer and
Schaefer 2007) increase the contrasts of fruit displays
and inluence fruit detection (Willson and Melampy 1983,
Burns and Dalen 2002, Schaefer et al. 2007).
he contrast between fruit displays and their background
are not necessarily constant. In seasonal habitats contrast
can change during the year, e.g. when leaves senesce and
are shed during the dry season. Burns and Dalen (2002)
suggest that temporal changes in the availability of fruits
with diferent colors are explicable by changes in fruit
conspicuousness against seasonal changes in foliage color-
ation. However, Burns and Dalen (2002) characterized fruit
Oikos 122: 1335–1342, 2013
doi: 10.1111/j.1600-0706.2013.00328.x
© 2013 he Authors. Oikos © 2013 Nordic Society Oikos
Subject Editor: Paulo Guimarães Jr. Accepted 21 January 2013