Water movement activating fragmentation: a new dispersal
strategy for hydractiniid hydroids
Giorgio Bavestrello*, Stefania Puce*, Carlo Cerrano
O
, Laura Castellano
P
and Attilio Arillo
O
*Istituto di Scienze del Mare dell'Universita© di Ancona,Via Brecce Bianche, 60131Ancona, Italy.
*E-mail: bavestrello@pocsi.unian.it.
O
Dipartimento per lo studio delTerritorio e delle sue Risorse dell'Universita© di Genova,Via
Balbi 5, 16126 Genova, Italy.
P
Acquario di Genova, Area Porto Antico, 16100 Genova, Italy
The fragmentation process in Hydractinia pruvoti Motz-Kossovska, 1905) which live on shells inhabited by hermit crabs is
activated by the intensity of water movement that induces a rapid and abundant production of propagules by transversal
¢ssion. Being produced by all kinds of polyps in the polymorphic colony, fragments are very di¡erent from each other both in
shape and size. After their liberation, fragments ¢rst settle on the bottom, then dedi¡erentiate and produce a network of
hydrorhizae from which new polyps arise.
Asexual reproduction is prevalent in hydroids and takes
place by means of di¡erent strategies. Cloning can be produced
accidentally due to injury of the colony, physiologically by
budding, or by the autotomy of di¡erent portions of the colony.
Budding is the most common mechanism of asexual reproduc-
tion among hydroids. Buds generally arise from hydranths,
branches or stolons producing both medusae and/or polyps.
Stolonization is the formation of stolons from hydrorhizae or
hydrocauli which grow and attach to another substratum near
the parent colony and then segregate to form a new colony. In
addition to the stolons, other fragments of colonies, such as frus-
tules and free hydranths, may detach by autotomy giving rise to
propagules with very di¡erent degrees of specialization for a
review see Gravier-Bonnet,1992).
In this work we report on the water activated fragmentation
process of Hydractinia pruvoti with a description of the fate of
their released fragments.
Our original colony, living on the shells of Monodonta turbinata
inhabited by the hermit crab Clibanarius erythropus, was collected
by SCUBA divers at a depth of 10m along the coast of Alassio
Ligurian Sea) in September 1998. They were reared in an
aquarium ¢lled with ¢ltered natural seawater and fed daily with
Artemia salina nauplii.Within a month, several colonies arose on
the carbonatic fragments of the bottom. These colonies 50^55
polyps each), which were individually reared in cups of 250ml,
were used for the water movement experiments.
In our colonies polyps arose from a creeping hydrorhiza
composed of a confused arrangement of coalescent stolons.Three
main kinds of polyps were present: i) gastrozooids, 1^2 mm in
height, with 5^10 ¢liform tentacles; ii) gonozooids, 1^2mm in
height, characterized by a single, very extensible tentacle
0.5^2mm), arising laterally at the base of a vestigial hypostome.
Each gonozooid bore 3^5 medusoids in the middle of its body;
iii) dactylozooids, made up of a spherical body 0.5mm in
diameter) which directly arises from the hydrorhiza and bears, at
its apex, a long tentacle very similar to that of the gonozooids.
Conical spines 0.2^0.3mm high were present too Figure 1A).
The occurrence of anomalous gastrozooids with two tentacular
rings and two hypostomes Figure1B) was also observed.
When mature medusoids Figure 1C) only male specimens
in our material) detach from their gonozooids, they vigorously
swim for several days before the gamete release. They show four
well-developed radial canals ending in the tentacular bulbs on
the circular canal.The gonads are concentrated in a large masse
around a non-functional manubrium.
During the ¢rst phase of rearing, it was observed that
increased water turbulence in the aquarium produced a
dramatic increase in the number of detached fragments so it was
decided to test colonies in water arti¢cially circulated.The water
movement was obtained by means of a moving plane on which
cups containing the hydroid colonies were placed. The rotary
movement of the plane was adjusted in order to produce turbu-
lence causing oscillation of the polyps without changing the
position of the entire colony to avoid any mechanical injuries.
Plotting the average number of fragments released vs the
time of induced turbulence, the maximum production of ten
propagules per day per 100 polyps was achieved. Colonies
maintained in calm water produced fragments at a rate of about
0.5^1/d per 100 polyps. The main section of fragmentation were
on the long tentacles of gonozooids and dactylozooids 48%)
followed by gastrozooids 40%) and the body of gonozooids
12%). The ¢rst propagules settled four days after the start of
fragmentation despite the continuous water movement, while,
on the other hand, the fragmentation process continued for
some days after the cessation of the water oscillation. Fifteen
days after the start of the experiment seven days after the stop
of water movement) the success of propagules settlement was
very near 100%.
The fragmentation process stems from a transversal ¢ssion
acting at di¡erent levels both in gastro- Figure1D) gono- and
dactylozooids which produce fragments of variable shape
Figure 1E), while their basal portions remain adherent on the
original substratum producing new hypostomes and tentacles
Figure 1F). Fragments of gastrozooids were composed of almost
entire polyps or mere tentacle rings which surrounded the
hypostome.The gonozooids which had detached from the mother
colony immediately released mature medusoids, and proceeded
to undergo either body or long tentacle fragmentation, giving
origin to planula-like propagules. The fate of the medusoids
released during the fragmentation process di¡ers according to
their developmental stage: mature medusoids actively swim and
spawn, while immature medusoids undergo a regressive
Journal of the Marine Biological Association of the United Kingdom 2000)
J. Mar. Biol. Ass. U.K. 2000), 80,361^362
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