Salamanderfish Lepidogalaxias salamandroides (Mees 1961) Photo by Gerry Allen |
-Burrowing & Aestivation
In order to utilize a habitat that dries out, a fish must
have a way of getting through this hostile period until wetter times arrive.
This ability is not unheard of in the fish world: some killifish, for instance,
lay eggs that can survive (and may even require) a period of dormancy in dry
conditions. Others, like the large familiar lungfishes, burrow into the ground
and aestivate. The Salamanderfish falls into this second category. These
remarkable little fish move below the surface of the sand and leaf litter in
search of substrate hydrated by ground water, and, remarkable for a fish of
this size, they have been recovered up to 60cm (about 2 feet) below the
surface! Interestingly they are incredibly quick to re-emerge, and when their
habitat was experimentally rehydrated with water from a fire truck, fish were
captured as little as 8 minutes later (Berra and Allen 1989). The ability to survive
dry periods and rapidly resume normal activity is of huge benefit to a fish
that inhabits such a seasonally hostile environment.
Salamanderfish in process of burrowing. Photo by Auscape. |
-Cutaneous respiration
Another incredible adaptation, related to aestivation but
significant enough to mention separately, is the ability of this species to
perform cutaneous respiration. In what must have been a delicate task,
researchers separated the head and gill structures from the rest of the body by
putting a “collar” around the fish and measured oxygen and CO2
levels while the fish was out of water. They were able to determine that L. salamandroides is capable of
considerable gas exchange through the skin. Surprisingly, however, they also
determined that when out of water, this fish does not produce extra mucous or
have any other apparent mechanism to prevent dessication, so while they can
breathe out of water, they must stay moist in order to survive any prolonged
period on land or in the substrate (Martin et al. 1993). Another interesting
aspect of this is that, unlike many other aestivating fish species, the
Salamanderfish does not have the ability to survive hypoxic water conditions
(Berra & Allen, 1995) and has no accessory breathing apparatus in the swim
bladder or gills (Berra et al. 1989). The reason for this
may be because the typical habitat of this fish is physically predisposed to
gas exchange, being relatively shallow pools of water with relatively large
surface area. Except under the influence of extreme amounts of microbial oxygen
usage, water in this situation would likely contain plenty of dissolved oxygen
by default. One possible benefit of cutaneous respiration when the fish is not
able to maintain water balance, may be forays above the waterline to forage on
insects. This has not yet been studied in this species, but it is within the
realm of possibility, as the Mangrove Killifish (Kryptolebias marmoratus Poey 1880), a small estuarine species of
similar habitat and ability, has been shown to actively feed above the
waterline (Pronko et al. 2013).
-Internal fertilization
Back in the water, in good conditions at the appropriate
time of year, it’s time for the Salamanderfish to breed. In yet another plot
twist, this fish defies the odds yet again by practicing internal fertilization.
Lepidogalaxias salamandroides males
possess a modified anal fin with a scaly sheath that serves as an intromittent
organ, while females have ciliated ducts connecting to the ovaries and are
capable of storing sperm. Strangely, there has not been observed to be any
courtship display, but instead the male approaches a female and rolls her so he
can position his anal fin and scaly sheath adjacent to her vent. The scaly
sheath structure apparently secretes a kind of adhesive mucous that connects
the mating pair (with a researcher even noting that when lifted from the water
they remained attached!)(Pusey and Stewart 1989). The function or
history of internal fertilization can only be theorized at this point. This
species has undergone dramatic changes in taxonomic placement, most recently being
placed in the basal position of euteleosts (Li et al. 2010), and therefore it is
difficult to make connections to where this may have arose in the evolution of
this species. Its mode of fertilization is quite unlike any other teleost (Pusey and Stewart 1989), making it difficult to
ascertain the origin. It has been theorized that internal fertilization evolved
as a response to the often highly acidic conditions in which this species
lives, which is a hostile environment for sperm, or that this (and the mucous
adhesion which subsequently serves to form a “plug”) arose as a result of sperm
competition (Pusey and Stewart 1989). This subject requires
additional study to elucidate the evolutionary details of this process.
-Bone and skull adaptations –
neck bending
When
one first observes the Salamanderfish, most of these interesting facets are not
readily apparent. One thing that does, however, immediately grasp the attention
is this fish’s amazing (in the fish world) ability to turn its head. This fish
is capable of moving its skull directionally both side-to-side and up-and-down
as much as 90 degrees (Berra and Allen 1989). This is possible
because the distance between the back of the skull and the cervical vertebrae
is relatively large, allowing an enhanced degree of flexibility. Besides this
increased flexibility potentially aiding in the ability to burrow, this fish
lacks typical musculature surrounding the eye, preventing the Salamanderfish
from moving the eye within its socket (Mcdowall and Pusey 1983). Thus, the ability to
bend the neck may be of crucial importance during foraging and feeding,
allowing the fish to lie nearly motionless on the bottom while scanning the
environment for prey. Despite being such a small fish, L. salamandroides has a formidable array of teeth, which may assure
the consumption of any prey captured by the fish. The reinforced, wedge-shaped
skull and largely reduced ribs may be adaptations that additionally enhance
burrowing by decreasing drag and increasing flexibility, aiding in travel
through the substrate (Berra and Allen 1989). For a demonstration of the neck-bending ability of this
fish, see video.
Neck bending Salamanderfish. Photo by Tim Berra |
With a huge variety of specialized adaptations, the Salamanderfish
(Lepidogalaxias salamandroides Mees
1961) is a truly fascinating example of a fish living where a fish shouldn’t
really be. A curious suite of characters have led this species to drive
taxonomists crazy, with its placement on the evolutionary tree changing often
since its discovery as ichthyologists struggle to determine where it belongs. This
animal has proven intensely fascinating some researchers who continue to
unravel the mysteries of its uniqueness. In the meantime, while we are
struggling to understand it, the Salamanderfish will continue to eke out a
living in one of the most hostile environments known to fish-kind.
References
Berra, T., and G. Allen. 1989. Burrowing, emergence, behavior, and
functional morphology of the Australian salamanderfish, Lepidogalaxias
salamandroides. Fisheries 2415(May 2014):37–41.
Berra,
T. M., D. M. Sever, and G. R. Allen. 1989. Gross and Histological Morphology of
the Swimbladder and Lack of Accessory Respiratory Structures in Lepidogalaxias
salamandroides , an Aestivating Fish from Western Australia. Copeia
1989(4):850–856.
Li, J.,
R. Xia, R. M. McDowall, J. A. Lopez, G. Lei, and C. Fu. 2010. Phylogenetic
position of the enigmatic Lepidogalaxias salamandroides with comment on the
orders of lower euteleostean fishes. Molecular Phylogenetics and Evolution
57(2):932–936.
Martin,
A. K. L. M., T. M. Berra, and G. R. Allen. 1993. Cutaneous Aerial Respiration
during Forced Emergence in the Australian Salamanderfish, Lepidogalaxias
salamandroides. Copeia 1993(3):875–879.
Mcdowall,
R. M., and B. J. Pusey. 1983. Lepidogalaxias Salamandroides Mees – a
Redescription, With Natural History Notes. Records of the Western Australian
Museum 11(1):11.
Pronko,
A. J., B. M. Perlman, and M. a Ashley-Ross. 2013. Launches, squiggles and
pounces, oh my! The water-land transition in mangrove rivulus (Kryptolebias
marmoratus). The Journal of Experimental Biology 216(Pt 21):3988–95.
Pusey,
B. J. 1990. Seasonality, aestivation and the life history of the salamanderfish
Lepidogalaxias salamandroides (Pisces: Lepidogalaxiidae). Environmental Biology
of Fishes 29(1):15–26.
Pusey,
B. J., and T. Stewart. 1989. Internal fertilization in Lepidogalaxias
salamandroides mees (Pisces: Lepidogalaxiidae). Zoological Journal of the
Linnean Society 97(1):69–79.
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