We
sometimes encounter two very similar fish species in similar places. When the
two species are difficult for the novice to differentiate, we further wonder
“How are the species reproductively isolated?”
Ernst Mayr, a leading evolutionary biologist of the 20th
century, classified reproductive isolating mechanisms into pre-zygotic (act before fertilization) and post-zygotic (act after fertilization). In the case of sympatric species, behavioral
isolation may involve distinctive behaviors designed to increase contact
between breeding individuals of the same species. Post-zygotic mechanism are less efficient for reproductive
isolation. Although, we don’t often
observe these isolating behaviors, there must be some subtle cues present
during breeding. Fish use pheromones
(smell), vision, and sound for species recognition and reproductive isolation.
Yes, fish make sounds that travel well underwater. We’ve known this for a long
time (Myrberg et al. 1965; Gerald 1971). Listen to a few fish making sounds here.
Recent
studies confirm that many fish species communicate via sound during courtship. Typically
these are low-frequency sounds produced by males when close to females, while
other fish make long-distance sounds to advertise their location. Black Drum Pogonias cromis can communicate up to
30–100 m (Amorim et al. 2015). Goby
males emit sounds from their nests to attract mates (Malavasi et al. 2008). It
may not sound like singing to humans. But the male fish sings in order to
attract a mate. There is enough information in the sound signals that the
female can assess size and condition of potential mates. In a recent paper, Eva-Lotta Blom and
coauthors studied two species of gobies (Gobiidae) to examine visual and
auditory cues during courtship. Two species, Common Goby Pomatoschistus microps and the Sand Goby Pomatoschistus minutus, are members of the species-rich family
Gobiidae. Gobies are small, benthic
fishes with modified pectoral fins fused together, two dorsal fins, and often
distinctive color patterns. Many are popular aquarium specimens favored for
their colorful patterns or behaviors. See gallery of gobies.
The
Common Goby and the Sand Goby have sympatric distributions. During breeding
they are found in close proximity, often competing over the same nest resources.
To study the role of vision and sound in reproductive isolation, Blom and
coauthors simultaneously recorded sounds with a hydrophone and visuals with a
camcorder of breeding individuals under controlled lab conditions. What they
learned was that females like males that sing. In addition, there were visual cues that
differed between species so that a female Common Goby would not be fooled by
the behaviors of a male Sand Goby.
Common Goby (left, by Michel Barrabes) and Sand Goby (right, by Mazzun Tar-Ramel) |
Differences
in courtship behaviors and cues also help explain why the two morphologically
and ecologically similar species select different habitats for breeding. The courtship sound of the male Common Goby
has shorter duration than the one of the Sand Goby. Male
Common Goby swim faster than male Sand Goby during courtship displays. Finally, only the female Sand Goby display
black eyes during courtship. Nests of Common Gobies are more prevalent in
shallow, soft bottom areas whereas Sand Goby nests use Mya clams in deeper, sandy beaches. The habitat differences
influence sound transmission.
Oscillograms
of representative sounds illustrate the distinctness of sounds of (A) male Common Goby and (B) male Sand Goby. (Blom
et al. 2016)
|
During
breeding, visual cues are typically expressed by males as breeding
coloration. In the Sand Goby, the female
display of black eyes acts as a declaration of intent that the female is ready
to mate. Female Sand Gobies are able to change eye coloration, sometimes
gradually, other times quickly, even within seconds. The
black eyes are not conspicuous in males. In experimental aquaria, Olsson et al. (2017) observed
that occurrence of female displaying dark eyes are more likely in those females
close to spawning readiness, as measured by body roundness.
Bar chart illustrating the relationship between female roundness and frequency of dark eyes (black line). Frequency of dark eyes are illustrated in dark grey bars. (Olsson et al. 2017). |
As the studies of gobies illustrate, females prefer males that
sing and can distinguish sounds for closely related species in order to isolate
species reproductively. The study raises
questions about how males learn to sing and whether and how noise from human
activities affect signals during breeding. But now you know why the male goby sings.
References
Amorin, M.C.P., R.O. Vasconcelos, and
P.J. Fonsesa. 2015. Pages 1-33 in F.
Ladich, editor. Sound Communication in
Fishes. Springer.
Blom,E-L., I. Mück, K. Heubel and O.
Svensson. 2016. Courtship sound and associated behaviours of two sympatric
marine Gobiidae species – Pomatoschistus
microps and Pomatoschistus minutus.
Environmental Biology of Fish 99: 999–1007.
Gerald, JW.1971. Sound production
during courtship in six species of sunfish (Centrarchidae). Evolution 25:75–87.
Malavasi S, S. Collatuzzo, and P. Torricelli. 2008.
Interspecific variation of acoustic signals in Mediterranean gobies
(Perciformes, Gobiidae): comparative analysis and evolutionary outlook. Biological Journal of the Linnean Society
93:763–778.
Myrberg AA Jr, E. Kramer E, and P.
Heinecke. 1965. Sound production by cichlid fishes. Science 149:555–558.
Olsson, K.H., S. Johansson, E-L. Blom,
K. Lindström, O. Svensson, H. Nilsson Sköld and C. Kvarnemo. 2017. Dark eyes in
female sand gobies indicate readiness to spawn. PLoS One 12: e0177714 http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0177714