The text below is
quoted from the article 'Pipefish, Museums, Marine Naturalists and Fish
Conservation'
in the Marine Life Society of South Australia
2003 Annual Journal no. 13 (Browne 2003)
The reproductive biology of the Syngnathids
is particularly interesting as the males brood the eggs. Seahorses have
well developed brood pouches, seadragons have brood patches, and pipefish
have brood patches which are enclosed to varying degrees. The brooding of
eggs by males means that acceptance of the female by the male is a
limiting factor in conferring genes to the next generation. This infers
that the females would be advantaged by competing for males by
ornamentation, pairing, courting or aggression, and all these activities
have been observed
.

Courting is normal before mating in the Syngnathids, with complex courting
rituals a pre-requisite to mating. In South Australia, females of both
the Wide-bodied Stigmatopora nigra and Spotted pipefish court the males
by displaying their chests which are barred. The chest of the Wide-bodied
pipefish may be bright red. The South Australian Deep-bodied pipefish,
Kaupus costatus has the greatest difference of form between males and
females of any pipefish. The females are flattened sideways to display
bright red and blue bars. Pairing is common in seahorses with pairs observed
over long periods. However, pairing does not always mean fidelity as a
wide range of mating patterns are documented in the Syngnathids, including
genetic monogamy (faithful pairs) in a seahorse, and polygynandry (more
than one female mate) and polyandry (more than one male mate) in pipefish
(Jones and Avise 2001).
Recent studies using molecular techniques to elucidate paternity have
shown that polyandry is common in some pipefish species. Moreover, in
these species the intensity of sexual selection on females rivals that
of any other animals (Jones et al 2001). This, and more females than males,
with some females never reproducing in some species, results in very different
behaviour between males and females; the males are faithful and the females
are dedicated, if promiscuous, partners. Male pipefish reject courting
females other than their partner, maintaining the pair bond over seasons.
This fidelity, and studies showing that even in primitive pipefish where
external fertilization occurs, eggs exposed on the brood patch were all
fertilized by the tending male, show the evolution of enclosed brood pouches
is not a response to cuckoldry by sneaker males (McCoy et al 2001).

Some have suggested
that species which live amongst shelter which could brush away the eggs
would have brood pouches. However, brood pouches are found in seahorses
which do not violently interact with substrates, and brood patches in
seadragons which have similar behaviours. Why then do advanced pipefish
have a brood pouch? In these species, the embryos are attached to a placenta-like
tissue which seals the pouch folds. The most apparent reason would be
to reduce predation. However, no evidence exists of this. One study showed
that within this enclosed pouch concentrations of salt were lower than
in seawater (Watanabe et al 1999), perhaps reducing the energy expenditure
from the egg needed for osmoregulation resulting in fitter larvae from
similar sized eggs. If this is the case it is a further transfer of reproductive
effort from the female to the male.
Mate guarding by females has been suggested as the main mechanism for
maintenance of monogamy in males of pipefish; males losing their partners
re-mate within a few days. Females will mate with other males besides
their mate during a breeding episode (McCoy et al 2001). However, for
unknown reasons widowed females remain unmated for a considerable period
(Matsumoto and Yanagisawa 2001). Females in their quest to reproduce with
as many males as possible have larger home ranges and are more active
in courtship displays (Matsumoto and Yanagisawa 2001). Some interesting
benefits have been shown from competition for partners in the pipefish.
Broods from preferred matings when either males or females were allowed
to choose a partner are superior at escaping predators and grow faster
(Sandvik et al, 2000).
The number of eggs in most species of pipefish is not documented. A number
of males from the South Australian Museum carried eggs or had mature brood
pouches. This has enabled the first tabling of the fecundity of many species.
A surprising find was that most species of pipefish only had between 20-30
eggs. This is in contrast to many seahorses which lay hundreds of eggs.
In the Wide-bodied pipefish, reproduction had been shown throughout the
year. However, there were few species with enough specimens over the seasons
to show seasonality from the South Australian records. In many other species,
the presence of brood pouches showed few males and in some species no
males were found. Brood pouches could be subjected to seasonal variation
or these samples could include only female or juvenile pipefish. If this
is the case, other un-sampled habitat may be needed for reproduction.
Further knowledge will be gained on reproductive status of individuals by
their dissection to show the sex and the maturity of gonads
.
Although in many pipefish, hatching time and the period between batches
is not known, hatching time generally varies from 10-30 days. Pipefish
can have several broods in a season, with non-brooding intervals as short
as a few days (Matsumoto and Yanagisawa 2001). Many species have been
shown to reproduce throughout the year (Howard and Koehn 1985). However,
even closely related species may vary in respect to seasonality. The most
likely reason for seasonality is variation of the potential adult food
and larval supply.
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