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Main » Coloration
Black color
occurs in coloration of many freshwater fish beening relatively constant throughout
the year and cryptic (in such, for example, bottom dwelling fish as wels, Silurus glanis). In this paper, we will consider those frequent cases
when chiefly males of freshwater fish acquire black coloration in the
reproductive period (nuptial melanization).
For desalinated
areas and rivers of the Ponto-Caspian basin, five species of gobiid fish with
nuptial melanization of body are most typical and abundant. Round goby, Neogobius melanostomus, monkey goby, N. fluviatilis, ratan goby, N. ratan, black goby, Gobius niger, and racer goby, Mesogobius gymnotrachelus, are among
them. Gobies are marine origin, but the foregoing species (as well as many
amelanistic species) inhabit desalinated areas and readily migrate into rivers
of the Ponto-Caspian basin where successfully breed (Pinchuk et al., 1985;
Romanesku, 2012). Some other euhaline gobies develop black nuptial coloration,
such as giant goby, Gobius cobitis),
but they avoid oligohaline bays and fresh waters.
In the
reproductive period, gobies acquire conspicuous black coloration practically of
the whole body using such an appearance, on the one hand, to repel rivals (in
this case, black coloration is called threat, or antaposematic) and, on the
other hand, to attract females (Trifonov, 1955; Yankovsky, 1966). Some gobies
acquire color rims on the edge of both dorsal fins, pair pectoral and anal
fins: white in N. melanostomus,
yellow in M. gymnotrachelus and
orange in N. fluviatilis. In accordance with the accepted terminology (Trifonov, 1955; Yankovsky, 1966), these
conspicuous color signs are called gamosematic, bacause these signs appear in
males when they prepare the nests, indicating in this way on their readyness to
breed, and disappear in males when they begin to protect the nests with laying
eggs.
Spawning
and nest guarding in N. melanostomus are
well documented (Meunier et al., 2009).
In general,
round goby, N. melanostomus, and other gobies are rather visually guided
fish with the specialized chemoreceptory channel. N. melanostomus respond poorly to the odors of lake whitefish (Coregonus) tissues, crushed dreissenids
and fish eggs (Sreedharan
et al., 2009; Yavno & Corkum, 2011). At least Sreedharan et al. (2009) do
not recommend to use food-baited traps to control the spread of these fish.
According to Rollo et al. (2007), N. melanostomus have well developed vocalization and
are attracted by conspecific calls in both laboratory and field trials.
Nuptial
melanization is also found in such fish of the North American ichthyofauna as dirty
darter, Etheostoma olivaceum, Gila topminnow, Poeciliopsis
occidentalis, and Olympic mudminnow, Novumbra
hubsi (Kodric-Brown,
1998). Coloration of males in brook stickleback, Culaea
inconstans (Ward & McLennan, 2006), mosquitofish, Gambusia holbrooki (Horth, 2004) and
Amur sleeper, Percottus glehni (Tsepkin, 1977), are other examples
of nuptial melanization.
Interestingly,
in mosquitofish, G. holbrooki, black males have advantages and
disadvantages to more common silver rivals. On the one hand, largemouth bass, Micropterus salmoides, crayfish (Procambarus) and dragonfly larvae (Libellulidae) prefer, as natural predators, silver males (Horth, 2004), that is black
males are under less predation pressure. According to Taylor et al. (1996), on the
other hand, females of G. holbrooki prefer silver males and even can avoid
black males.
Basic References
Horth L.
2004. Predation and the
persistence of melanic male mosquitofish (Gambusia
holbrooki). Journal of Evolutionary Biology 17,
672-675
Kodric-Brown
A. 1998. Sexual
dichromati
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Conspicuous coloration of males occurs
in many species of fish, birds and other animals being advantageous in
attracting potential mates. Although bright colors can entice females of the
same species, these colors may also attract predators. Female choice for bright
males and an enhanced risk of predation for bright males are both well
documented in numerous works (see Dill et al., 1999, and references therein).
Because
fresh waters are optically turbid in comparison with pure sea waters, curves of
photopic spectral sensitivity in freshwater fish are strongly displaced to the
red part of the spectrum. For example, the maximum of spectral sensitivity in
threespined stickleback, Gasterosteus
aculeatus, is near 605 nm (Rowe et al., 2004). It means that for eyes of freshwater fish red and orange colors are brighter than
all other equipower monochromatic colors, and this feature occurs in nuptial signaling
coloration.
The nature
allows males of freshwater fish to practice several strategies to find
trade-offs between conspicuousness for sexual mates and crypticity for
potential predators, including plasticity in nuptial color development (e.g., Endler,
1983; Candolin, 1998; Ruell et al., 2013). In this context, an ability to develop bright
red and orange colors in the under less illuminated parts of the fish’s body,
in conformity with the theory of color countershading in fresh waters, is the
primary.
Indeed, red
and orange colors occur in breeding males just in the under parts of their
bodies such as breast, ventral part, belly and the lower fins. An important
role of this elements in nuptial coloration of males is documented in guppy, Poecilia reticulata (Endler, 1983;
Kodric-Brown, 1985), and other species of genus Poecilia, threespined stickleback, G. aculeatus (Rowe et al., 2004), European bitterling, Rhodeus sericeus (Candolin & Reynolds, 2001), and in other spesies of genus Rhodeus, as well as in males of other freshwater
fish. In some cases red and orange colors occur in the upper most illuminated
parts of the fish’s body, but patterns of this type will be considered
separately.
According
to Kodric-Brown (1998),
breeding males with red
fins occur in many families of North American freshwater fish, including
minnows (Cyprinidae), suckers (Catostomidae), killifish
(Fundulidae), sunfish (Centrarchidae), darters (Percidae) aa well as cichlids
(Cichlidae).
Generally,
bright red and orange colors are conspicuous at the shot distance for sexual
mates, but are cryptic in the countershading complex at the longer (that is optically
thick) distances for potential predators. According to Evans & Norris (1996),
red pigmentation of the fish’s body cannot be assessed accurately under green
light or hereof if viewed through the water column, as the natural green
filter.
Vorobyev et
al. (2001) demonstrate how fish can see other fish through the water column.
Basic References
Candolin U. 1998. Reproduction under predation risk and the
trade-off between current and future reproduction in the threespine stickleback. Proceedings of the Royal Society, Biological Sciences 265, 1171-1175
Candolin U., Reynolds J.D. 2001. Sexual signaling in the European
bitterling: females learn the truth by direct inspection of the resource. Behavioral Ecology 12, 407-411
Dill L.M.,
HedrickA.V., Fraser A. 1999. Male mating strategies under predation risk: do females call the shots? Behavioral Ecology 10, 452-461
Endler J.A. 1983. Natural and sexual
selection on color patterns in poeciliid fishes. Environmental Biology of Fishes 9, 173-190
Evans M.R.,
Norris K. 1996.
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