P. S. Z. N. I: Marine Ecology, 5 (2): 9>118 (1984) Accepted: 7.7.

1983
0 1984 Paul Parey Scientific Publishers, Berlin and Hamburg
ISSN 0173-956YInterCode: MAECDR

The Ecology of Marine Hydroids

and Effects of Environmental factors:
A Review
F. BOERO

Istituto di Zoologia dell’Universit8, Via Balbi 5 , 1-16126 Genova, Italy

With 17 figures

Key words: Hydroids, ecology, environmtntal factors, adaptations.

Abstract. The history of the study of hydroid ecology is briefly outlined, pointing out the major
methodological innovations which have contributed to the development of ecological research in the
last thirty years. The influence of the major ecological factors on hydroid ecology and biology is
synthesized, taking into account: substratum, water movement, light, salinity, sedimentation,
exposure to air, temperature, food availability and pollution. Besides affecting the species composi-
tion of the hydroid community, these factors also influence the morphology and general biology of
the individual species. The adaptations and reactions of hydroids to different intensities of the
various environmental factors and to their .combinations are reported.

Problem
When, in the early 1950s, R.RIEDLdemonstrated the efficiency of SCUBA
diving techniques for marine biology research, he chose an as yet unexplored
environment: marine caves. Here he studied in detail the ecology of the hydroid
population. The work of RIEDL(1959) is not only of scientific but also of
historical value.
In the same period in which RIEDLcarried out this work, J. PICARD studied
hydroid ecology and systematics, introducing new classification criteria for both
species and ecological associations (PICARD, 1951a, b; 1952, 1956, 1958).
In the 1950s 0.JSINNE (1956 a, b; 1957, 1958) dealt with hydroid ecology in
the North Sea, supporting laboratory studies with in situ observations and
demonstrating experimentally the effects of salinity and temperature on some
hydroid species.
While REDL showed the absolute necessity of in situ ecological studies,
PICARDclearly indicated that systematics and ecology had to become more
integrated. KINNEfurther demonstrated that in situ observations can be control-
led and confirmed with laboratory experiments.

U.S. Copyright Clearance Center Code Statement: 0173-9565/84/0502-0093$02.50/0

94 BOERO

Earlier observations on hydroid ecology, however, can be found as far back

as the last century. ALLMAN(1871-72) proposed a bathymetric zonation of
British hydroids, grouping the species according to their ecological preferences.
MOTZ-KOSSOWSKA (1903) and HALLEZ(1905 a, b) realized that water movement
produces morphological variations in hydroids and that, in some cases, species
considered as being different are merely ecological forms of the same species.
BILLARD (1904) devoted part of his studies to the influence of environmental
factors on hydroid morphology. PRENANT & TEISSIER (1924) studied the ecology
of the Roscoff hydroids, providing abundant information on the individual
species and their associations as well as offering interpretations about the
influence of physical factors on the zonation of the hydroid population.
These forerunners had a scant following and for a long time hydroid ecology
was almost completely neglected. The notes that systematic specialists put
before morphological descriptions were the only available ecological informa-
tion and usually reported only date, locality and depth of the collection.
The utilization of hydroids as laboratory animals has led, since the beginning
of this century, to numerous contributions to the knowledge of the biology of
those species which are easily reared. These studies, however, rarely began with
observations under natural conditions, so that such research fell within the field
of pure physiology rather than ecology.
In the last thirty years ecological studies on hydroids have received remark-
able attention and the results have brought our knowledge to a sufficiently
advanced stage.

Fig. 1. Detail of an experimental plate immersed since 40 days. Agluopheniu tubiformis is colonizing
the margin, passing from another plate where it is dominant. Clytia hemkphericu has settled all over
the plate.
Hydroid ecology and environmental factors 95
The importance of hydroids in ecological theory is due to a number of
characteristics of this group, which also make it particularly suited for more
general investigations, both in sifu and in the laboratory.

Colonization and choice of the substratum

Owing to their rapid settling and quick growth, hydroids are among the first
metazoans to colonize virgin substrates (HuvB, 1953; SENTZ-BRACCONOT, 1966)
(Fig. 1). They are then often substituted by other more massive and stoutly
constructed organisms such as algae, sponges, polychaetes, cirripedes, bryo-
zoans, molluscs and ascidians. At this stage secondary settlement occurs, with
epibiosis and recolonization of the substratum at a superior level (Fig. 2). Some
hydroids are capable of strenuously defending their territory (Fig. 3): SUTHER-
LAND & KARLSON (1977), in their study on Beaufort fouling, report Hydructinia
echinatu (FLEMING) as being the organism most resistent to overgrowth by other
species competing for space. FRADETTE & BOURGET (1981) found that Obeliu
longzssima (PALLAS) is the dominant species in the benthic epifauna of the Saint
Lawrence maritime estuary and gulf. This dominance was explained by the work
of STANDING (1976) on another species of this 'genus, 0. dichotomu. With
experiments involving the local removal of colonies, he demonstrated that 0.
dichoroma (L.), prevents or delays the settlement of other organisms on
experimental surfaces.

Fig. 2. Three steps in the colonization of

a horny sponge. Hydroids are among the
first settlers. When the surface of the
sponge is almost completely covered hy
other epibionts, hydroids settle on them
(redrawn after RIEDL,1966).
96 BOERO

Fig. 3. Aglaophenia tubiformis, Aglaophenia ocrodonia, Dynamena disiicha covering an artificial

substratum immersed for 6 years.

In the epiphytic community of Fucus serratus (L.), Dynamena pumila KRAUSS

seems almost immune from interspecific competition for space: the vital parts of
the colony are borne on erect stems which are rarely overgrown by epizoites
(STEBBING, 1973).
During the development of a hard-bottom benthic community hydroids settle
precociously and, if overwhelmed in the competition for space, can settle again
on both the dominant vegetal and animal species.
Hydroids rarely settle on shallow soft bottoms, although with increasing
depth they play a more and more important role and form large communities
below 40-50 m. Hydroids living on soft bottoms can reach considerable dimen-
sions. The solitary species Branchiocerianthus imperator (ALLMAN) reaches a
maximum height of two meters in deep-sea trenches.
Many hydroid species have evolved more or less exclusive preferences for
other organisms as a substratum. DALES(1957) reports twenty-eight hydroid
species which have relations with gorgonaceans, pennatulaceans, polychaetes,
hermit-crabs, parasitic copepods, bivalves, gastropods, tunicates and fishes.
Other species live exclusively in sponges or on hydroids, bryozoans, echino-
derms and reptiles. Strict associations between hydroids and algae or sea grasses
also exist. MILLARD (1973), introducing the concept of auto-epizoism, describes
the modified habit of hydroids which settle on conspecifics. These relationships
show a complete range from epiphytism and epizoism to inquilinism, commen-
salism, mutualism and parasitism.
Hydroid ecology and environmental factors 97
The choice of the substratum occurs during the larval (or planula) stage, or
during the propagation by propagula, stolons, etc. DONALDSON (1974) studied
the attachment of Proboscidactyla flavicirrata BRANDT planulae to the pinnulae
of their sabellid host. The planulae pass from the pinnulae to the rim of the tube
when the worm withdraws. Nematocysts play a very important role during the
attachment of Proboscidactyla planulae.
NISHIHIRA (1973) studied the settlement of certain hydroid species on littoral
algae. He showed that planulae of the various species differ in their behaviour in
that they choose only particular parts of certain algae.
KATOet al. (1961) investigated the epiphytic hydroids of Sargassum. They
found that Clyria volubilis (L.) usually settles on the middle regions of the
branches. When Obelia dichotoma is present, it takes the place of C. volubilis.
The latter species is then to be found on more proximal and distal portions of
the thallus of Sargassurn. This phenomenon, known as coaction, was then
studied under controlled conditions and a hierarchy of overgrowth among
hydroids was determined (KATOet al., 1962, 1963, 1967).
The settlement of hydroid planulae on algal substrata is promoted or inhi-
bited by substances produced by the plants. From Sargassum extracts KATOet al.
(1975) isolated the substance which promotes the settlement of Coryne uchidai
STECHOW.
On the other hand, tannin-like substances from the growing tip of Sargassum
kill hydroid colonies (SIEBURTH & CONOVER, 1965). As a result hydroids are
absent in the distal regions of the Sargassum thallus (CONOVER & SIEBURTH,
1964).
Margelopsis haeckeli HARTLAUB has a very peculiar life-cycle; it passes the cold
season as a cyst. WERNER (1963 a) has experimentally demonstrated that the
presence of more or less suitable substrata determines both the form of the cysts
and their attachment.
Hydroids can also be hosts for other organisms. The epifauna of Nernerte.uk
antennina (L.) consists of 180 species (HUGHES,1975). ROUNDet al. (1961)
report an equally abundant inventory of epibionts on Aphisbetia (Sertularia)
operculata (L.)*. LAGARDBRE & TARDY(1980) have described relationships
between hydroids and polychaetes. In these situations the polychaetes that have
settled on hydroids (of which they are commensals) assault the predators of
their hosts (Fig. 4).
REES& ROA (1966) briefly discuss a system of asexual reproduction which
could ensure the propagation of the species when substrates suited for the
attachment of the planula are unavailable. Some medusae, in fact, asexually
produce polyps or gonothecae from which other medusae are liberated. In these
cases the polyp phase lives on the medusa from which it has originated.

The ecological significance of the reproductive strategies

of hydroid polyps
All hydroid species possess a polyp phase. Some also have a medusa phase,
considered by some authors as the adult and by some others as a simple carrier
* When a species is reported by the cited author under a name not retained as valid, it will be named
with the correct name and, in parenthesis, with the name used by the author.
98 BOERO
Fig. 4. The fauna associated with Ectopleu-
ra dumortieri. The polychaete worm (Syl-
lidae, Aufolyfinae), living in tubes attached
to Ecfopleura stems, feed on the prey at-
tached to the tentacles of the hydroid. They
also attack the nudibranch molluscs (i. e.
Cumanotus cuenofi) which feed on Ecro-
pleura, causing their detachment from the
hydranth (redrawn from LAGARD~RE & TAR-
DY, 1980).

of sexual products (see BOUILLON, 1981). The presence or absence of a medusa

stage must be of ecological significance. The possibility of colonizing new sites
varies greatly according to the lifespan of the planktonic stage of the cycle.
Some hydroid polyps, e. g. Eudendrium, release planulae which remain
attached to the mother colony by a thread of mucus. The thread breaks off when
the planula reaches the bottom (WASSERTHAL & WASSERTHAL, 1973). In this way
the species occupy very definite sites and an induced gregariousness between the
mother colony and its offspring occurs. HUGHES(1977) has found a similar
system of planula release in Nemertesia antennina. In this case, however, the
mucus thread allows the planula to reach locations further from the mother
colony (5-50 m). Instead of falling immediately on the sea bottom the planulae
can be transported by currents via elongation of their mucus threads.
Other species release free planulae which crawl on the substratum rather than
swim. This is the case in many plumularians and sertularians. As in Euden-
drium, the occupied sites are very close to the mother colony.
Swimming ciliated planulae are present in many athecates and thecates. They
can live in the plankton for several days and are capable of colonizing quite
distant sites.
Hydroid ecology and environmental factors 99
The behaviour of planulae has been studied in only relatively few species
(DONALDSON, 1974; %TO et al., 1975; NISHIHIRA, 1967 a, b; 1968 a, b; SPINDLER
& MULLER,1972; TITELBAUM, 1966; WILLIAMS, 1965, 1966), some of which are
gregarious.
The release of actinulae occurs in many tubulariid species. The settlement of
actinulae usually takes place after a brief period. Some species, e. g. Tubularia
crocea ( L . A ~ ~ s s r z can
) , shed their hydrants, which also live free in the
plankton for long periods and then release actinulae (RUNNGER, 1969).
The production of medusoids (free-living, swimming structures deprived of a
digestive system and which carry the sexual products) enable several species,
e. g. Halocordyle disticha (GOLDFUSS),Campanufaria integra MCGILLIVRAY,
Stylactis spp. , Lytocarpus phifippinus (KIRCHENPAUER) , t o spread over short
distances.
Free medusae are the most complicated structures present in hydroid life-
cycles. They actively swim and can survive for several months, feeding on other
organisms. The possession of a medusa stage has the advantage of increased
dispersal. The disavantage is that the medusae can be taken by currents to areas
where no possibilities for planulae settlement or gamete interaction exists.
Some hydroids (e. g. Velelfa, Porpita, Margelopsis) have planktonic polyps
and thus avoid or reduce any sessile phase in their life-cycles.
The choice of a certain reproductive strategy, however, does not seem to play
a very important role in the colonization possibilities of a species, even in very
specialized habitats. Hydractinia (fixed gonophores), Stylactis (medusoids) and
Podocoryne (free medusae) settle on gastropod shells still inhabited by the
mollusc or used by hermit crabs. The three very different reproductive strategies
are all perfectly adapted for the occupation of very similar ecological niches.
Carveia grisea (MOTZ-KOSSOWSKA) (fixed gonophores) and Bougainuiflia ramo-
sa (VANBENEDEN) (free medusae) are very abundant on the big Eudendrium
spp. colonies in the Mediterranean. They live near each other and are often
mixed. If not fertile they are very similar in appearance. The colony structure
and the form of the polyps are so similar that it is unlikely that they occupy
different ecological niches. Once again opposite reproductive strategies give
very similar results.
Both in athecates and thecates reproduction by medusae or fixed gonophores
occur in parallel in many families. The medusa stage has been completely
abandoned in the families which are currently considered as the most evolved
among hydroids (Plumufariidae and Sertulariidae). These families are very
successful in all marine habitats. The loss of the medusa stage, then, actually
appears to be a great advantage from the point of view of general ecology.
The absence of a medusa stage has probably a positive selection value in
many plumularians and sertularians. The enormous number of species in these
families and the great number of endemisms indicate that the populations of the
various species are restricted to specific areas. Once colonization has occurred,
these “conservative” species remain in the geographical and environmental area
where their success is “guaranteed”. On the other hand species with a medusa
stage are often cosmopolitan and, in many cases, can colonize very different
habitats. They are opportunistic and adapted to face very different environmen-
tal conditions. The Campanulariidae, with the genera Clytia and Obelia, are a
100 BOERO

good example of this tendency (see CORNELIUS,

1975, 1982 for a detailed
review).

Water movement
REDL (1959, 1966, 1971) showed, with in situ observations, that planar (i. e.
plumose) hydroid species (such as Aglaophenia, Plumularia, Halopteris, etc. )
orientate their colonies in order to he perpendicular to the dominant direction
of water movement (Fig. 5 ) .
These species are very frequent at every depth and can be used as recorders of
the mean conditions of water movement. SVOBODA (1976) has, under controlled
conditions, verified RIEDL’Sobservations. In his zonation system RIEDLdistin-
guishes three critical depths, related to the quality of water movement. The first
critical depth is the boundary between the surf zone and the zone of a
predominantly oscillating water body. The second critical depth is the boundary
between the oscillating water body and the unidimensional flow zone along a
steep coast. The third critical depth is the boundary between the unidirectional
(parallel to the coast) streaming water body and the multidirectional flow
possible over level bottoms. In correspondence with the critical depths the
orientation of planar hydroids (and of planar colonies in general) changes from
(a) no orientation in the surf zone to (b) parallel to the water line in the
oscillating zone (Fig. 6) to (c) perpendicular in longshore currents to (d) bush-
like or with movable fans on the level bottoms where longshore and tidal
currents can produce water movement from any direction.
Surface Surt Pandular mowmunt

Fig. 5 . Four types of planar hydroid positions (a: general view; b: detailed view) individuate four
forms of water movement (surface movement, surf, pendular movement, orbital movement) in the
first seven meters of depth (redrawn after RIEDL,1971).
 Hydroid ecology and environmental factors 101
The importance of water movement is easily understood because a high water
exchange around the colony allows for a greater oxygen and food supply; at the
same time it will diminish the possibility of excessive sediment deposition on the
colony itself.

Fig. 6 . Agfuopheniu ocfodontu at 0.5 m depth, where water movement is pendular. The cofonies are
all consistently oriented perpendicularly to the direction of water movement.

movement
~ O Q in cmlsec

Fig. 7. Ecological distribution of hydroid species in a marine cave. Seven hydroid-zones are
distinguished and related to the decreasing intensity of water movement in the cave (redrawn after
RIEDL,1966).
102 BOERO

RIEDL(1959) distinguishes seven hydroid-zones characterized by different

water movement conditions in marine caves, where hydroid zonation should be
more influenced by water movement than by light (Fig. 7).
Water movement causes considerable resistence problems to the delicate
hydroid colonies. Those near the surface tend to diminish their dimensions and
increase the number of rings on their stems, thus becoming more elastic. Those
colonies which attain a large size at low depths (i. e. Hulocordyle) have very
flexible stems and stout hydrorhizae , with which they tenaciously cling to the
substratum. The feeding strategies of Halocordyle are best adapted to take
advantage of turbulent or oscillatory water movement conditions. The feather-
like colonies of Hulocordyle are very flexible at their base and, under the
influence of water movement, “beat” the surrounding space in a fanlike
manner. This increases their chances of capturing prey. In this respect they can
be considered intermediate between active and passive filter-feeders.
The morphological variations caused by water movement have been
described for various species since the beginning of the century. Recently BOERO

-
1 cm

Fig. 8. Modification of hydroid ecology

by water movement. Ventromma
halecioides - A: specimens with fascicu-
lated hydrorhizae and stems in calm wa-
ter movement conditions (gonothecae
on stems). B: specimens with fascicu-
lated hydrorhizae in turbulent water
movement conditions (gonothecae on
hydrorhizae). Plumularia obliqua - C :
specimens from 1m depth. D:specimen
from 30m depth. E: hydrotheca from
1m depth (with septum). F: hydrotheca
from 30m depth (without septum). (A,
B redrawn after BOERO,1981 a; C-F re-
drawn after BOERO,I981 b).
Hydroid ecology and environmental factors 103
(1981a, b) showed, in Ventrumma halecioides (ALDER)and Plurnularia obliqua
(JOHNSTON), important structural and morphological variations between superfi-
cial colonies and those settled at greater depths (Fig. 8).
Hydroids climb on other organisms with increasing depth. They are elevated
above the bottom, taking advantage of better trophic conditions due to the
increased water movement around their colonies (Figs. 9,10,11). This is termed
“acrophily”.
The reproductive strategies of some hydroid species can also undergo modifi-
cations as water movement varies. STEFANI(1959) reports that, in the Gulf of
Cagliari, the same population of Campanularia integra (= caliculata) produces
fixed gonophores in sheltered zones, while colonies settled in exposed sites
produce free medusoids. Tubularia crocea, by autotomy, liberates hydranths
that can sexually reproduce even when detached from the colony. In this way
the species, thanks to water movement, increases its dispersion possibilities
(RUNNGER, 1969).
In general, as already stated by ALLMOJ (1871-72), with increasing depth and
a decrease in light and water movement, species with fixed gonophores are more
abundant while those producing medusae tend to disappear.

Light
Hydroids sensitivity towards water movement has been widely demonstrated
and quantified, while it is not completely clear what the direct influence of Light
is, even though many observations confirm the importance of this factor in
hydroid biology.
MCDOUGALL (1943) has shown, in Tubularia crocea and Halocordyle (= Pen-
naria) tiarella (AYRES), the damaging or inhibiting effects of ultraviolet light on
the metabolism and growth of young colonies. Tubularia crocea cannot survive
exposition to various intensities of ultraviolet light (=UGH, 1929). ROSS (1964)
has demonstrated a more intense growth in Eudendrium racernosum (CAVOLIKT)
when exposed to light. Bougainvillia ramosa, on the contrary, has a more
intense regenerative activity in the dark (PEEBLES, 1900).
THORSON (1964) reports positive phototropism in hydroid larvae. WILLIAMS
(1965) observes the same phenomenon in Claw multicornis (= squamata)
(FORSKAL) planulae. In the phase preceeding settlement, however, the photo-
tropism becomes negative. In most cases only the behaviour of planulae of
species living in or near the turbulent zone has been studied. These planulae
must reach superficial sites and then settle on the substratum. In this case
photophily is without any doubt the most common system used for reaching
optimal settling sites. On the other hand this does not mean that such species
have evolved this behaviour to colonize zones with certain light conditions.
Light has no influence on the behaviour of Tubularia larynx (ELLIS &
SOLANDER) and Nernertesia antennina planulae (PYEFINCH & DOWNING, 1949;
HUGHES, 1977).
Photophilous and skotophilous populations are well known, each formed by
different species mixing where light undergoes sudden variations. The internal
mechanism determining these phenomena in every species remains unclear.
Fig. 9 . Serntlarella crassicaulis at 4 m depth, settled directly on the sea bottom.

Fig. 10. Sertularella crassicaulis at 10m depth, settled along the stem of Eudendrium glomeratum.
Fig. 11. Sertularella crassicnulis at 20m depth, settled on the top of Errnicella singularis.

Fig, 12. Paracoryne huvei in the mid-littoral zone, completely out of the water (Photo R. CASTANEO).
106 BOERO

The presence of zooxanthellae in many hydroids makes them very sensitive to

light variations, although studies attempting to demonstrate the mechanism of
this sensitivity ar far from being complete. It is still difficult to understand the
photophily of some species deprived of zooxanthellae and the moderate
skotophily of many species with zooxanthellae. Photophily and skotophily are
probably often masked by rheophily and rheophoby.
The presence in marine caves of species also common outside at superficial
levels (RIEDL,1959) shows that hydroids are not very sensitive to light dimin-
utions. Not a single hydroid species typical of or exclusive in marine caves is
known. Unlike sponges, bryozoans and serpulids, hydroids almost never col-
onize the deepest and darkest parts of marine caves. In such caves, however,
environmental conditions are formed by a complex integration of factors which
can only be discriminated with difficulty.
Examples of distributions sharply mfluenced by light are nevertheless avail-
able. Synthecium evansi (ELLIS& SOLANDER) is typical of coralligenous bottom
and of the rhizomes of Posidonia oceanica (L.) DELILEmeadows, where both
light intensity and water movement are feeble. S. evansi can also occur at l m
depth in rock fissures where water movement is intense but where light is very
weak. The ecological preferences of this species are, then, principally charac-
terized by a sharp skotophily.
Hydroids have also been found in the deepest oceanic bottoms in absolute
darkness (VERVOORT, 1966) and, in general, they tend to avoid well illuminated
sites.
Species living directly exposed to sunlight, such as Paracoryne huvei PICARD
(Fig. 12), are also known. This species colonizes the intertidal zone of the

0 B

Fig. 13. Diagrammatic scheme of the life cycle

of Paracoryne huvei throughout the year. The
species passes the summer months under a cyst
stage (redrawn after BOUILLON,1975).
Hydroid ecology and environmental factors 107
Mediterranean, but is present only in winter when sunlight is reduced (Fig. 13)
(BOUILLON, 1975).
Light has a noticeable effect on hydroid reproduction. BAKER(1936) and
BALLARD (1924), for instance, have demonstrated that the alternation of light
and darkness can initiate the release of sexual products in Hydractinia and
Halocordyie (Pennaria). The anthomedusa Polyorchis karafutoensis KISHI-
NOUYE releases its gametes only if it is subject to alternate conditions of light and
darkness; no sexual activity occurs if light or darkness are continuous (ZEN,
1963).
GOY(1973) reports that the liberation of medusae in Gonionemus suvaensis
A. AGASSIZ & MAYERis more intense under a full moon and that this
phenomenon is probably common to all hydromedusae. Previous observations
by ORTON (1920) and ELMHIRST (1925) tend to confirm this hypothesis, even
though the information available from the literature thus far comprises too small
a number of species to allow a generalization.

Salinity
For marine animals salinity in its normal range does not represent an ecological
factor; variation therefore does not influence population biology. Marked
changes in salinity occur only under certain circumstances (brackish waters,
tropical inner reefs, heavy rains, etc.), and can affect the biology of the species
and their populations.
KINNE(1956 a, b; 1957, 1958), studying Cordylophora caspia (PALLAS),has
demonstrated a sharp morphological and structural variability both of hydranths
and of colonies in relation to sharp salinity changes (Fig. 14). As salinity
decreases, the specific richness of the hydroid fauna tends to decrease gradually
(Fig. 15). Along salinity gradients the typical marine species disappear and are
substituted by euryhaline species or by those adapted to low salinities (THIEL,
1970; CALDER,1976; MORRI& MARTINI,1981).
As is the case for light, the influence of salinity is probably masked by other
factors such as water movement. Brackish waters often have conditions of
reduced water movement. Waters with variable salinity often have strong
currents and a strong water exchange. In these situations water movement could
be of considerable importance.

Sedimentation
The influence of this factor is strictly linked to the inclination of the substratum
and to water movement. The species living in zones where water movement is
turbulent or oscillatory use no particular system to avoid sediment covering
because they are continuously “cleaned” by moving water. As depth increases
hydroids solve the sedimentation problem in two ways. The smaller species
settle on upright organisms and are thus elevated above the sediment stratum.
Other species develop tree-like and thin colonies, obtaining the same results and
offering scant space for the accumulation of sediment.
 108 BOERO
Fig. 14. Cordylophora caspia.
Shapes of colonies and polyps at
different salinities and, for
polyps only, temperatures. The
form of the colonies has been
observed at 20°C (redrawn after
KINNE,1958).

2O0C IOT 20% 1o0c 20°C IOOC

Fresh water 15 %o S 30%o S
? Lafoeina tenuis
Agalaaphenia pluma
Sertularella ellisi
Obelia dichotama
Obelia geniculata
Obelia bidentata
Eudendrium sp.

Ventromma halecioides

Fig. 15. Penetration of hydroids in the

Orbetello Lagoon (Central Italy) from
the two principal sea-water inlets (re-
drawn after MORRI& MARTINI,1981).
Hydroid ecology and environmental factors 109
These strategies are not only useful in avoiding sediment covering, but also
effect a greater water exchange around the colony. Consequently the trophic
situation of the colonies is improved. OSMAN(1977) has observed that Tubularia
larynx retains sediment around its hydrorhizae, thus preventing the settlement
of other species.

Exposure to air
Some hydroid species have colonized the intertidal zone and face the problem of
drying due to exposure to air. PRENANT & TEISSIER (1924) described a number of
systems employed by hydroids to survive this danger. Similar systems have been
described for the hydroids of the Mediterranean mussel belt (BOERO,1981 a). In
general, when exposed to air, hydroids use two strategies to avoid desiccation:
the formation of bushy clumps which retain water, or settlement on algae to
remain in contact with surfaces which retain water.
Paracoryne huvei, a Mediterranean species living in the intertidal zone,
succeeds in preventing desiccation by forming encrusting colonies with dense
clumps of polymorph polyps. This species has also developed a life-cycle with
winter activity peaks, when water movement is stronger and sunlight weaker; it
passes the warm season with cyst stages (BOUILLON, 1975) (Figs. 12, 13).

Temperature - Seasonality
While the preceeding factors mainly influence the spatial distribution of hydroid
populations, temperature plays an important role in determining its composition
in time (i. e. cold and warm seasons). Seasonal phenomena cause a continuous
change in the composition of hydroid populations which, in one and the same
zone, have very different features in the various periods of the year.
A s for the Mediterranean area, the research of BOERO& FRESI(in prepara-
tion) provides evidence for sharp differences between winter and summer
populations (Fig. 16). These seasonal differences are almost negligible in the
first half meter of depth, where a few specialized species form a compact group
which is stable in time. This is followed by an intermediate zone between 0.5
and 5-6 m, where both superficial and deep features are present. From 5-6 m to
20m a sharp division between winter and summer populations occurs. In
summer, hydroids are influenced by the presence of large algae. In winter large
colonies of Eudendrium glomeratum PICARDand E. ramosum (L.) occupy the
spaces used by algae in summer, forming a facies. The alternation between
summer algae and winter hydroids suggests that, in this case, a biotic component
(presence-absence of macroalgae) also plays an important role in determining
the development of the Eudendrium facies. From October-November an
epizoic population settles on the large winter Eudendrium; it reaches its peak in
December-January and then declines to disappear towards the end of April
(Fig. 17). Many hydroid species contribute to this epigrowth.
CALDER(1971), HAMOND (1957) and WEDLER(1975) have studied hydroid
seasonality in the Chesapeake Bay, on the British and the Colombian coasts,
 110 BOERO

-+ 9 J -3
lo

0.5-5
’ 3

f9 6

0-0.5 5-1 0

respectively. WEDLER, in particular, observed that the variations in the hydroid

population are related not only to temperature but also to the rainy season.
Along the Colombian coast these variations are more evident at the superficial
levels while there is greater stability with increasing depth; here colonies attain
greater dimensions and have longer life-spans and reproductive cycles.
Temperature is the regulating factor in the cycle of Paracoryne huvei (BOUIL-
LON, 1975). The temperature increase favours the autotomy of the hydranth of
Tubularia crocea, initiating a special process of propagation (RUNNGER, 1969).
Laboratory experiments on some hydroid species show that temperature
directly influences nutrition and assimilation of food (KINNE,1957; KINNE&
PAFFENHOFER, 1965;PAFFENHOFER, 1968).Other researchers (MOORE, 1939;KINNE
1956a, b; WERNER,1956, 1958, 1962, 1963b) report about the effects of
temperature on the life-cycles of certain hydroids and hydromedusae. Usually
there are optimal temperatures for sexual reproduction. Growth, on the other
hand, can continue indefinitely at temperatures higher or lower than those
optimal for reproduction, within certain limits. Growth can also be interrupted,
with reduction of the colonies, while waiting for better conditions.
BROCK(1975) has found that Laomedea (as Campanularia)flexuosa ALDER
shows endogenous rhythms of circannual periodicity that “anticipate seasonal
variations in the coastal water temperature of the species natural environment”.
Seasonal species (i. e. species with different growth and reproductive rates in
different seasons) are apparently adapted to the variability of their environ-
ment, anticipating the changes occurring in the environmental factors.

Food availability and food chains

Food availability greatly influences the development of hydroid colonies. This
phenomenon has been studied in the laboratory by BERIULL(1950) and
CROWELL (1957). Undernourished colonies of Laomedea (as Campanularia)
show a normal stolon growth, while stem growth is lacking. This hydroid, then,
occupies more horizontal space and terminates vertical growth. This phenome-
non also occurs in normally fed colonies as a “regression cycle”, i. e. old
hydranths degenerate and others are produced peripherally. STREHLER &
CROWELL (1961) have described the regression cycle of species belonging to
several hydroid families. They interpreted this as a strategy “to place the feeding
Hydroid ecology and environmental factors 111
Fig. 16. History of a hydroid

“a,.
11 community during one year at
five different depth ranges in the
Portofino Promontory (Ligurian
Sea). Ordination model of a PC
Analysis in the plane of the two
Principal Components (tPC 1 =
24.98; tPC2 = 11.42). Numbers
indicate months (after BOERO&
15-20 76 FRESI, in preparation).

10-15

organs (hydranths) in the most efficient conditions (peripherally) during periods

of low food supply”. The regression cycle - according to STREHLER & CROWELL
(1961) - “serves the colony by automatically forcing it in effect to evaluate in a
primitive way and on a regular schedule the state of its nutrition”.
When the environmental conditions are particularly favourable hydroids can
become enormously abundant, assuming a very important role in food chains.
Main Russian authors (TURPAEVA et al., 1977; MARFENIN& BURYKIN, 1979;
LETUNOV & MARFENIN, 1980; SIMKINA,1980) have studied the diet of common
and abundant species along the Soviet coasts, among them Garveia franciscana
(Perigonimus megas) (TORREY) and Dynamena pumila, and have evaluated their
role as predators of coastal plankton. With few exceptions (e. g. CHRISTIENSEN,
1967), the importance of these phenomena has been neglected until now by
other than Russian researchers. This field, once studied on a wider scale, will
reveal new and unsuspected information about the role of hydroids in the

Fig. 17. Two stages of the life cycle of

Eudendrium rameum. A t the beginning
of the cycle only few epizoites are pre-
sent. After several months the epibionts
have nearly covered the entire colony,
the polyps of which are degenerated
(redrawn after RIEDL,1966).
112 BOERO
energetic transfers between plankton and benthos. The big “prairies” of Nemer-
tesia and Thecocarpus on muddy bottoms and the big assemblages of Euden-
drium on rocky bottoms probably consume large quantities of energy. Nemer-
tesia and Eudendrium, in particular, display sudden growth and short life-cycles,
implying high energy demand during those times.
Hydroids enter also food chains as a food source for nudibranchs, crusta-
ceans, fishes, etc.

Pollution
Information regarding the effects of pollution on hydroids is fragmented in the
enormous literature on fouling. MILLARD (1959), MONTANARI & RELINI(1970),
CHIMENZ-GUSSO & RIVOSECCHI-TARAMELLI (1975) have studied the composition
and distribution of the hydroid fauna in harbour waters with different degrees of
pollution. Only some genera are very resistant in non-natural water conditions.
Tubularia is one of the most characteristic and BARNES (1948) reports it as one
of the organisms more resistant to poisons such as copper, widely employed in
anti-fouling paints. PYEFINCH & DOWNING (1949) report that copper and mercury
stimulate the settlement of larvae of Tubularia larynx. Subinhibitory levels of
toxic substances, with a process called hormesis, have stimulatory effects on the
growth rates of various organisms (STEBBING, 1982). STEBBING (1981) has studied
hormesis in Laornedea (as Campanularia) flexuosa. An increase in growth is
present independent of the type of toxic or other inhibitory agents used (copper,
cadmium, tributyl tin fluoride, reduced salinity) and is due to the influence of
these agents on a general control mechanism.
In noxious conditions (e. g . high concentration of copper) Laomedea flexuosa
increases the production of gonozooids (STEBBING, 1980). This phenomenon is
explained by STEBBING as an adaptation of the species where, in general, “the
planktonic phase in the life cycle is initiated by conditions that are unfavourable
for the sessile parent colony. This appears to be an adaptive response favouring
the survival of the parent colony’s genes, by the dispersal of the medusae in the
plankton”.
Those species typical of fouling and generally polluted waters are cosmopoli-
tan and well adapted to great variations of environmental factors. ‘The hydroid
population in harbour waters is much less species-rich than those of non-
polluted waters. Seasonal cycles, reproductive periods and depth preferences,
however, still remain evident in the various species.
Depending on the geographical region, certain species can become dominant
in a particular season.
As MILLARD (1959) observes, the more evolved families (Plurnulariidae and
Sertulariidae) contribute virtually nothing to fouling. The more primitive
families are, on the other hand, well represented.
CLAREet al. (1971) relate the formation of detached spherical colonies of
Sarsia tubulosa (M. SARS)and Tubularia larynx to the presence of inhibiting
factors (probably pollutants) affecting the first stages of development of the
planulae.
In general, when the waters reach a high degree of pollution, hydroids
disappear almost completely.
Hydroid ecology and environmental factors 113

Indicator species
The great plasticity of hydroid populations, both in time and space, makes this
group a faithful recorder of environmental conditions. In fact, modifications in
the composition and distribution of the hydroid population, together with the
morphological modifications of the most tolerant species, are strictly related to
variations of the environmental factors.
The sensitivity of hydroids to different intensities of light and water move-
ment has led MERGNER (1977), after a careful study of the hydroid ecology in
coralline reefs (MERGNER, 1972), to propose some species as ecological indi-
cators. He has chosen stenoecious species of sufficiently large dimensions which
are easily identifiable in situ. Of a total of thirty-seven species considered, seven
are excellent indicators exhibiting different combinations of the effects of the
two factors. The ecology of these species can be used to characterize all the
“structural zones and many biophysiographical zones between the coralline
lagoon and the external reef”.
The hydroid population of the leaves of Posidonia oceanica (a common
Mediterranean sea grass) undergoes noticeable modifications as depth in-
creases. Some species, strictly epiphytic on marine phanerogames, have a
distinct superficial distribution (BOERO,1981b). This has been interpreted by
BOERO as a proof of the validity of the theory of DENHARTOG (1973) according
to which Posidonia oceanica is a superficial species living also at deeper levels
owing to the last rise in the sea level. In this case the hydroid population is a
good ecological indicator, and is much more sensitive than Posidonia itself.
Those species characteristic of sea grasses, in fact, remain only at the more
superficial levels which are optimal for both themselves and Posidonia.
Hydroid species which are easily reared have been successfully used as test
animals in the evaluation of the effects of pollutants on marine organisms
(KARBE,1972).

Conclusions
The role of hydroids in benthic biocoenoses is of primary importance. An
increase in work on this topic will make it possible to better understand the true
position of the group in the energetic balances of ecosystems and in the
distribution of populations along gradients of environmental factors. The
studies carried out thus far in this field have already indicated the direction for
further research.
It must also be remembered that our systematic knowledge is still insufficient
and that the identification of species is based on characters which are not always
natural (BOERO,1981c).
Ecologists very often identify their species from obsolete monographs that
require radical revisions. This leads to the propagation of imprecise and often
false information on the distribution and the ecology not only of hydroids, but
also of nearly all the other invertebrate groups.
Systematists, on the other hand, are nearly always completely unaware of the
ecology of the species, which are frequently described from preserved material.
114 BOERO

Some monographs have recently appeared (e. g. BRINCKMANN-VOSS, 1970;

CORNELIUS, 1975, 1979,1982; MILLARD,1975; SVOBODA, 1979; KUBOTA,1983) in
which systematics and ecology are integrated to form a complete picture of the
morphology and biology of the treated species. As for the medusa generation,
the monograph of RUSSELL(1953) contains much ecological information.
The conceptions of RIEDL,PICARDand KINNE are then still valid: direct
observation, the integration of systematics and ecology, and the experimental
verification of in situ observations are the indispensable tools for a satisfactory
knowledge of the structure and function of marine ecosystems.

Summary
Hydroids are heavily influenced by ecological factors: Some species show very
specialized adaptations to particular environmental conditions, other species
can live under very different ecological conditions. The hydroid population as a
whole can be used for describing the overall ecological features, especially on
hard substrata. Some hydroids have been used as indicator species for describ-
ing the intensities of single parameters such as water movement or for more
general indications on the ecology of a defined zone. The qualitative importance
of hydroids within benthic communities is already well established, but only few
studies have been done on their quantitative importance and especially on their
role in marine food chains.

Acknowledgements
I wish to thank Dr. P. CORNELIUS, London, Dr. L. ROSSI,Torino, Dr. B. WERNER, Hamburg, for
reviewing the manuscript; Dr. J. Om, Wien, for advice on RIEDL’Scritical depths; Prof. M. SARA,
Genova, for stimulating the writing of this paper.

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