Current Zoology 61 (4): 729–738, 2015
Received Mar. 11, 2015; accepted May 22, 2015.
Corresponding author. E-mail: gjensen@uw.edu
© 2015 Current Zoology
A whiter shade of male: Color background matching as a
function of size and sex in the yellow shore crab
Hemigrapsus oregonensis (Dana, 1851)
Gregory C. JENSEN
*
, Michael S. EGNOTOVICH
School of Aquatic and Fishery Sciences, Box 355020, University of Washington, Seattle, WA 98195, USA
Abstract Juveniles of the shore crab Hemigrapsus oregonensis are highly variable in color, ranging from the typical yellowish-
green of adults to pure white and myriad patterns of white mottling and other disruptive markings, but large individuals with
white coloration appear to be very rare. Using image analysis to quantify the relative “whiteness” of beaches, we sampled crabs
from nine locations in Washington State that varied widely in their amount of shell fragments and other light-colored material.
The total proportion of white individuals in the different locations was strongly correlated to the proportion of white material on
the beaches, but there was a striking difference between sexes. Although white specimens of both sexes declined significantly at
sizes above 10 mm carapace width, white females generally persisted throughout the entire female size range on lighter-colored
beaches while white males larger than 10 mm were virtually absent from all of the sampled populations. Pure white males held on
dark backgrounds in captivity remained white, as they lack the dark chromatophores in their hypodermis needed to change color;
off-white males became darker and in some cases lightened up again when transferred back to a white background. Behavioral
differences between the sexes may result in differential mortality of white individuals by visual predators [Current Zoology 61 (4):
729–738, 2015 ].
Keywords Camouflage, Color, Polymorphism, Disruptive patterns, Crabs
There are a number of different ways in which the
color or pattern of an organism can make them harder
for predators to detect. The most straightforward of
these, color matching, can be effective in homogeneous
habitats but is difficult for motile individuals to achieve
in heterogeneous environments due to the constantly
changing background. In these circumstances two very
different (but not mutually exclusive) strategies are
useful: color polymorphism and disruptive patterning.
Color polymorphism—color variations within a species
—can be the result of pigments incorporated from the
diet (Lindberg and Pearse, 1990) or due to genetic va-
riability (Palmer, 1985). Such variation can make it dif-
ficult for predators to establish a search image (Pietre-
wicz and Kamil, 1979, Kono et al., 1998) and as a result
can be maintained through apostatic selection (Bond
and Kamil, 1998).
Color polymorphism is widespread among inverte-
brates and has been investigated in a number of differ-
ent crab species, including Cancer irroratus (Palma and
Steneck, 2001), C. productus (Krause-Nehring et al.,
2010), Carcinus maenas (Hogarth, 1975, Todd et al.,
2006), Xantho poressa (Bedini, 2002), Pugettia spp.
(Hultgren and Stachowicz, 2008) and Gaetice depressus
(Murakami and Wada, 2015). A general pattern is emer-
ging in which juveniles of large species may exhibit a
range of color variants while adults may be monochro-
matic (Todd et al., 2009), corresponding with ontoge-
netic changes in habitat (Hogarth, 1978) and/or adults
growing too large to be susceptible to fish predators
(Palma and Steneck, 2001, Krause-Nehring et al., 2010).
Conversely, species that never gain a size refuge tend to
retain their polymorphism throughout their life (e.g.,
Xantho poressa; Reuschel and Schubart, 2007, Parax-
anthus barbiger; Manríquez et al., 2008).
Disruptive coloration (sensu Stevens and Merilaita,
2009) also makes organisms difficult to differentiate
from their background, often through the use of false
edges that disguise their true shape (Cuthill et al., 2005;
Stevens and Cuthill, 2006). Disruptive coloration has
rarely been explicitly mentioned or differentiated from
color polymorphism in the carcinological literature
(Krause-Nehring et al., 2010); notable exceptions are
the extensive work involving Carcinus maenas (Todd et
al., 2005, 2006, 2009, Stevens et al. 2014a, b) and work
by Merilaita (1998) on strikingly patterned morphs of
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