Professional Documents
Culture Documents
1983
0 1984 Paul Parey Scientific Publishers, Berlin and Hamburg
ISSN 0173-956YInterCode: MAECDR
With 17 figures
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.
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.
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
-
1 cm
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
0 B
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).
Ventromma halecioides
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
“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
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
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.
References
ALLMAN, G . , 1871-72: A Monograph of the Gymnoblastic or Tubularian Hydroids: Vol. 1-2. Ray
Society, London.
BAKER,E., 1936: Photoperiodicity in the spawning reaction of Pennaria fiarella Mc Cr. Proc.
Indiana Acad. Sci., 45: 251-252.
BALLARD, W., 1942: The mechanism of synchronous spawning in Hydractinia and Pennaria. Biol.
Bull., 82: 329-339.
BARNES, H., 1948: Studies on anti-fouling compositions. IV. The relationships between leaching
rate, copper loss and anti-fouling performance under raft and service conditions. J. Iron Steel
Eng., 1948: 175-185.
BERRILL, N., 1950: Growth and form in calyptoblastic hydroids. 11. Polymorphism within the
Campanulariidae. J. Morphol., 87: 1-26.
BILLARD, A., 1904: Contribution A I’Ctude des hydroides (Multiplication, rkgkneration, greffes,
variations). Ann. Sci. Nat. (Zool.), 20: 1-251.
BOERO,F., 1980: Life cycles of hydroids and hydromedusae: some cases of difficult interpretation.
Mem. Biol. Mar. Oceanogr., 10 (Suppl.): 141-147.
- -, 1981 a: Osservazioni ecologiche sugli idroidi della fascia a mitili della Riviera Ligure di
Levante. Cah. Biol. Mar., 22: 107-117.
- -, 1981b: Systematics and ecology of the hydroid population of two Posidonia oceanica meadows.
P. S . Z. N. I: Marine Ecology, 2 (3): 181-197.
Hydroid ecology and environmental factors 115
BOUILLON, J., 1975: Sur la reproduction et I'tcologie de Puracoryne huvei, PICARD(Tubuiaroidea -
Athecutu - Hydrozou - Cnidaria). Arch. Biol., 86 (1):45-96.
- -, 1981: Origine et phylogenese des cnidaires et des hydropolypes-hydromeduses. Ann. SOC.R .
Zool. Belg., 111: 45-56.
BRINCKMANN-VOSS, A., 1970: AnthomedusuelAthecutue (Hydrozou, Cnidariu) of the Mediterra-
nean. Part I: Capitata. Fauna e Flora del Golfo di Napoli, 39: 1-96.
BROCK,M., 1975: Circannual rhythms - 111. Rhythmicity in the longevity of the hydranths of the
marine cnidarian, Cumpunuluriu flexuosu. Comp. Biochem. Physiol., 51 A: 391-398.
CALDER, D., 1971: Hydroids and Hydromedusae of Southern Chesapeake Bay. Va. Inst. Mar. Sci.,
Spec. Pap. Mar. Sci., 1: 1-125.
- -, 1976: The zonation of hydroids along salinity gradients in South Carolina estuaries. In: G. 0.
MACKIE(Ed.), Coelenterate Ecology and Behavior. Plenum Press, New York: 165-174.
CHIMENZGusso, C. & E. RIVOSECCHI TARAMELLI, 1975: Idroidi del porto di Civitavecchia. Boll.
Pesca Piscic. Idrobiol., 30 (1): 111-125.
CHRIsnENsEN, H., 1967: Ecology of Hydructinia echinatu. I. Feeding biology. Ophelia, 4 (2):
245-275.
CLARE,J., D. JONES& A. O'SULLIVAN, 1971: On the occurrence of detached spherical colonies of
the hydroids Sursiu rubulosu and Tubularia larynx in Morecambe Bay. J. Mar. Biol. Assoc.
U. K., 51: 495-503.
CONOVER, J. & J. SIEBURTH, 1964: Effect of Surgussum distribution on its epibiota and antibacterial
activity. Bot. Mar., 6: 147-157.
CORNELIUS, P., 1975: The hydroid species of Obelia (Coelenterata, Hydrozoa: Campanufariidae),
with notes on the medusa stage. Bull. Br. Mus. (Nat. Hist.) Zool., 28 (6): 249-293.
- -, 1979: A revision of the species of Sertuluriidue (Coelenterufa:Hydroida) recorded from Britain
and nearby seas. Bull. Br. M U ~(Nat. . Hist.) ZOO~., 34 (6): 243-321.
- -, 1982: Hydroids and medusae of the family Curnpunuluriidue recorded from the eastern North
Atlantic, with a world synopsis of genera. Bull. Br. Mus. (Nat. Hist.) Zool., 42 (2): 37-148.
CROWELL, S., 1957: Differential response of growth zones to nutritive level, age and temperature in
the colonial hydroid Campanularia. J. Exp. Zool., U 4 (1): 63-90.
DALES,R., 1957: Interrelations of organisms. A. Commensalism. In: J. HEDGEPETH (Ed.), Treatise
on marine ecology and paleoecology. Geol. SOC.Am., New York: 391-412.
DONALDSON, S . , 1974: Larval settlement of a symbiotic hydroid: specificity and nematocyst
responses in planulae of Probosciductylu fluvicirrufu. Biol. Bull., 147: 573-585.
ELMHIRST, R., 1925: Lunar periodicity in Obelia. Nature, 116 (2914): 35g3.59.
FRADETTE P. & E. BOURGET, 1981: Groupement et ordination appliques B I'ttude de la rtpartition de
I'tpifaune benthique de I'estuaire maritime du golfe du Saint-Laurent. J. Exp. Mar. Biol.
Ecol., 5 0 133-152.
GOY, J., 1973: Conionemus suvuemis: structural characters, developmental stages and ecology.
Publ. Set0 Mar. Biol. Lab., 20: 525-536.
HALLEZ,P., 1905a: Rhtotropisme de quelques hydroides polysiphonks. C. R. Acad. Sci., 141:
727-730.
- - , 1905 b: Rheotropisme de quelques hydroides monosiphones et de Bugulu. C. R. Acad. Sci.,
141: 84Q-843.
HAMOND, R., 1957: Notes on the Hydrozou of the Norfolk coast. J. Linn. SOC.London, Zool., 43:
294-324.
HARTOG,C. DEN,1977: Structure, function and classification in seagrass communities. In: C. P.
McRov & C. HELFFERICH (Eds.), Seagrass Ecosystems. Marcel Dekker, Inc., New York and
Basel: 89-121.
HUGHES,R., 1975: The distribution of the epizoites on the hydroid Nemertesiu antenninu (L.). J.
Mar. Biol. Assoc. U. K., 55 (2): 275-294.
--, 1977: Aspects of the biology and life-history of Nemertesia anfenninu (L.) (Hydrozoa:
Plumulariidue). J. Mar. Biol. Assoc. U. K., 51: 641-657.
Huv6, P., 1953: Compte rendu prkliminaire d'une expkrience de peuplement de surfaces
immergkes. Recl Trav. Stn Mar. Endoume, 3: 9-59.
KARBE,L., 1972: Marine Hydroiden als Testorganismen zur Priifung der Toxizitat von Abwasser-
stoffen. Die Wirkung von Schwermetallen auf Kolonien von Eirene viridulu. Mar. Biol., 12
(4): 316-328.
116 BOERO
KATO,M., E. HIRAI& Y. KAKINUMA, 1962: Laboratory experiments on the interspecific relationship
in the colony formation of some hydrozoan species. Bull. Mar. Biol. Stn Asamushi, Tohoku
Univ., 11: 87-89.
- -, E. HIRAI& Y. KAKINUMA, 1963: Further experiments on the interspecific relation in the colony
formation among some hydrozoan species. Sci. Rep. Tohoku Univ., Ser. 4 (Biol.), 29:
317-325.
- -, E. HIRAI& Y. KAKINUMA, 1967: Experiments on the coaction among hydrozoan species in the
colony formation. Sci. Rep. Tohoku Univ., Ser. 4 (Biol.), 33: 359-373.
--, K. NAKAMURA, E. HIRAI& Y. KAKINUMA, 1961: The distribution pattern of hydrozoa on
seaweed with some notes on the so-called coaction among hydrozoan species. Bull,. Biol. Stn
Asamushi, Tohoku Univ., 10: 195-202.
KATO, T., A. KUMANIRENG, I. ICHINOSE, Y. KATAHARA, Y. KAKINUMA, M. NISHIHIRA & M. KATo,
1975: Active components of Sargassum torfile affecting the settlement of the larvae of Coryne
uchidai. Experientia, 31: 433.
KINNE,O., 1956a: Uber den EinfluB des Salzgehaltes und der Temperatur auf Wachstum, Form
und Vermehrung bei den Hydroidpolypen Cordylophora caspia (PALLAS), Afhecata, Clavidae.
Zool. Jahrb., Abt. Allg. Zool. Physiol. Tiere, 66: 565-638.
- -, 1956b: Zur Okologie der Hydroidpolypen des Nordostseekanals. Z. Morphol. Okol. Tiere, 45:
217-249.
- -, 1957: Uber den EinfluB von Temperatur und Salzgehalt auf die Kopfchenform des Brackwas-
serpolypen Cordylophora. Zool. Anz. (Suppl. Bd.), 20: 445-449.
- -, 1958: Uber die Reaktion erbgleichen Coelenteraten-Gewebes auf verschiedene Salzgehalts-
und Temperaturbedingungen. 11. Mitteilung iiber den EinfluB des Salzgehalts auf Wachstum
und Entwicklung mariner, brackischer und limnischer Organismen. Zool. Jahrb., Abt. Allg.
Zool. Physiol. Tiere, 67: 407-486.
- - & G. PAFFENHOFER, 1965: Hydranth structure and digestion rate as a function of temperature
and salinity in Clava rnulticornis (Cnidariu, Hydrozoa). Helgol. wiss. Meeresunters., 12:
329-341.
KLUGH,A , , 1929: The effect of the ultra-violet component of sunlight on certain marine organisms.
Can. J . Res., 1: 1OC-109.
KUBOTA, S . , 1983: Studies on life history and systematics of the Japanese commensal hydroids living
in bivalves, with some reference to their evolution. J. Fac. Sci., Hokkaido Univ., Ser. 6
(Zool.), 23 (3): 296-402.
LAGARDBRE, F. & J. TARDY,1980: Un facits d'epifaune nouveau: la faciCs a Ectopleura dumorfieri
(VANBENEDEN) et Elecfra pilosa (LINNB).Faune associte, cartographie et Cvolution saison-
nitre. Cah. Biol. Mar., 21: 265-278.
LETUNOV, v . & N. MARFENIN, 1980: Some characteristics of feeding behaviour in winter colonies of
Dynamenapumilu under various temperature regimens. Biol. Nauki, 6 (1): 51-55 (In Russian).
MARFENIN, N. & Y. BURYKIN, 1979: The growth of colonies of Dynametiapumila (L.) as a function
of the quantity of food received (Hydrozoa, Sertulariidae). Vestn. Mosk. Univ. Ser. 16, Biol.,
1: 61-68 (In Russian).
MCDOUGALL, K., 1943: Sessile marine invertebrates at Beaufort, North Carolina. Ecol. Monogr.,
13: 321-374.
MERGNER, H., 1972: The influence of several ecological factors on the hydroid growth of some
Jamaican coral cays. Proc. Symp. Corals and Coral Reefs 1969. Mar. Biol. Assoc. India:
275-290.
- -, 1977: Hydroids as indicator species for ecological parameters in Caribbean and Red Sea coral
reefs. Proc. 3rd Int. Symp. Coral Reefs, Miami: 119-125.
MILLARD, N., 1959: Hydrozoa from ships' hulls and experimental plates in Cape Town docks. Ann.
S. Afr. Mus., 45 (1): 239-256.
- -, 1973: Auto-epizoism in South African hydroids. Publ. Set0 Mar. Biol. Lab., 20 (Proc. second.
Int. Symp. Cnidaria): 23-34.
- -, 1975: Monograph on the Hydroida of Southern Africa. Ann. S. Afr. Mus., 68: 1-513.
MONTANARI, M. & G. RELINI,1970: Fouling di zone inquinate. Osservazioni nel Porto di Genova:
Idroidi e Ascidiacei. Pubbl. Stn. Zool. Napoli, 38 (Suppl.): 34-54.
MOORE,J., 1939: The role of temperature in hydranth formation in Tubularia. Biol. Bull., 76 (1):
104-107.
MORRI,C. & F. MARTINI, 1981: Gli idroidi della laguna di Orbetello. Quad. Lab. Tecnol. Pesca, 3 (I
Suppl.): 305-313.
Hydroid ecology and environmental factors 117
MOTZ-KOSSOWSKA, S., 1903: Sur I’action morphogtne de I’eau en mouvement sur les hydraires. C.
R. Acad. Sci., 137: 863.
NISHIHIRA,M., 1967a: Observations on the selection of algal substrata by hydrozoan larvae,
Sertularella miurensis, in nature. Bull. Mar. Biol. Stn Asamushi, Tohoku Univ., 13: 35-58.
- -, 1967b: Dispersal of the larvae of a hydroid Sertularella miurensis. Bull. Mar. Biol. Stn.
Asamushi, Tohoku Univ., 13: 49-56.
- -, 1968a: Experiments on the algal selection by the larvae of Coryne uchidai STECHOW (Hy-
drozoa). Bull. Mar. Biol. Stn Asamushi, Tohoku Univ., 13: 83-89.
- -, 1968b: Brief experiments on the effect of algal extracts in promoting the settlement of the
larvae of Coryne uchidai STECHOW (Hydrozoa). Bull. Mar. Biol. Stn Asamushi, Tohoku Univ.,
13: 91-101.
- -, 1973: Ecological distribution of epiphitic hydrozoa with special reference to Sertularella
miurensis. Publ. Set0 Mar. Biol. Lab., 20: 401-418.
ORTON,J., 1920: Sea-temperature, breeding and distribution in marine animals. J . Mar. Biol.
Assoc. U. K., 12 (2): 339-365.
OSMAN,R., 1977: The establishment and development of a marine epifaunal community. Ecol.
Monogr., 4 7 : . 3 7 4 3 .
PAFFENHOFER, G., 1968: Nahrungsaufnahme, Stoffumsatz und Energiehaushalt des marinen Hydroid-
polypen Clava rnulficornis. Helgol. wiss. Meeresunters., 18: 1-44.
PEEBLES,F., 1900 Experiments in regeneration and in grafting of Hydrozoa. Arch. Entwicklungs-
rnech. Org., 1 0 435-488.
PICARD, J., 1951a: Les hydraires des formations coralligtnes des c6tes Franpies de la MCditerranCe.
Vie Milieu, 2 (2): 254-261.
- -, 1951 b: Note sur les hydraires littoraux de Banyuls-sur-mer. Vie Milieu, 2 (3): 338-349.
- -, 1952: Les hydrozoaires des herbiers de Zosttractes des cdtes FranGaises de la Mtditerrante.
Vie Milieu, 2 (Suppl.): 217-233.
- -, 1956: Les especes et formes Miditerraneennes du genre Serfularella. Vie Milieu, 7 (2):
258-266.
- _ , 1958: Origines et affinittes de la faune d’hydropolypes (Gymnoblastes et Calyptoblastes) et
d’hydromeduses (Anthomtduses et LeptomCduses) de la Mediterranee. Rapp. Proc. Verb.
Reunions Comm. Expl. MCdit. N. S., 1 4 187-199.
PRENANT, M. & G. TEISSIER, 1924: Notes tthologiques sur la faune marine sessile des environs de
Roscoff: Cirripedes, Bryozoaires, Hydraires. Trav. Stn. Biol. Roscoff, 2: 1-49.
PYEFINCH, E. & F. DOWNING, 1949: Notes on the general biology of Tubuluria larynx ELLISand
SOLANDER. J. Mar. Biol. Assoc. U. K., 28: 21-44.
REES,W. & E. ROA,1966: Asexual reproduction in the medusa Zanclea implexa (ALDER).Vidensk.
Medd. Dan. Naturhist. Foren., Khobenhavn, 129: 39-41.
RIEDL,R., 1959: Die Hydroiden des Golfes von Neapel und ihr Anteil an der Fauna unterseeischer
Hohlen. Ergebnisse der Osterreichischen Tyrrhenia-Expedition 1952. Teil 16. Pubbl. Stn Zool.
Napoli, 30 (Suppl.): 589-755.
- -, 1966: Biologie der Meereshohlen. Paul Parey, Hamburg und Berlin.
- -, 1971: Water movement. 5. 3. Animals. In: 0. KINNE(Ed.), Marine Ecology, Wiley Inter-
science, London: 1123-1156.
Rossr, L., 1964: Fattori ecologici ed accrescimento in colonie di Eudendrium racemosum (GMELIN).
Boll. ZOO^., 31 (2): 891-905.
ROUND,F., J. SLOANE,F. EBLING & J. KITCHING, 1961: The ecology of Lough h e . X. The hydroid
Sertularia operculata (L.) and its associated flora and fauna: effects of transference to sheltered
waters. J. Ecol., 49: 617-629.
RUNNGER, D., 1969: Autotomy in Tubularia crocea and its ecological and physiological significance.
Pubbl. Stn Zool. Napoli, 37: 95-139.
RUSSELL,F., 1953: The Medusae of the British Isles. Cambridge University Press, Cambridge.
SENTZ-BRACONNOT, E., 1966: Donntes tcologiques sur la fixation d’invertkbrts sur des plaques
irnmergCes dans la rade de Villefranche-sur-mer. Int. Rev ges. Hydrobiol., 51 (3): 461-484.
SIEBURTH, J. & J. CONOVER, 1965: Sargassum tannin: an antibiotic which retards fouling. Nature,
208: 52-53.
SIMKINA, R. G., 1980: A quantitative feeding study of the colonies of Perigonimus megas (Hydroida,
Bougainvillidae). Zool. Zh., 59 (4): 500-506 (In Russian).
SPINDLER, K. & W. MULLER,1972: Induction of metamorphosis by bacteria and by a Lithium-pulse
in the larvae of Hydractinia echinata (Hydrozoa). Wilhelm Roux’ Arch., 169: 271-280.
118 BOERO
STANDtNG, J., 1976: Fouling community structure: effects of the hydroid Obelia dichofoma on larval
recruitment. In: G . MACKIE(Ed.), Coelenterate Ecology and Behavior. Plenum Press, New
York: 155-164.
STEBBING, A,, 1973: Competition for space between the epiphytes of Fucus serratus. L. J. Mar. Biol.
ASSOC.U. K., 53: 247-261.
- -, 1980: Increase in gonozooid frequency as an adaptative response to stress in Cumpanularia
jlexuosa. In: P. TARDENT & R. TARDENT (Eds.), Developmental and cellular biology of
Coelenterates. ElseviedNorth-Holland Biomedical Press, Amsterdam: 27-32.
- -, 1981: Hormesis - Stimulation of colony growth in Campanularia flexuosa (Hydrozoa) by
copper, cadmium and other toxicants. Aquat. Toxicol., 1: 227-238.
- -, 1982: Hormesis - The stimulation of growth by low levels of inhibitors. The Science of the
Total Environment, 22: 213-234.
STEFANI, R., 1959: Sulla variabilith ecologica di un idrozoo (Campanularia caliculafuHINCKS). Boll.
ZOO^., 26 (2): 115-119.
STREHLER, B. & S. CROWELL, 1961: Studies on comparative physiology of aging. I. Function vs. age
in Campanularia jlexuosa. Gerontologia, 5: 1-8.
SUTHERLAND, J. & R. KARLSON, 1977: Development and stability of the fouling coinmunity at
Beaufort, North Carolina. Ecol. Monogr., 47: 425-446.
SVOBODA, A., 1976: The orientation of Aglaophenia fans to current in laboratory conditions
(Hydrozoa, Coelenterufu).In: G . MACK~E (Ed.), Coelenterate Ecology and Behavior. Plenum
Press, New York: 41-48.
- -, 1979: Beitrag zur okologie, Biornetrie und Systematik der mediterranen Aghophenia-Arten
(Hydroidea). Zool. Verh., 167: 3-114.
THIEL, H., 1970: Beobachtungen an den Hydroiden der Kieler Bucht. Ber. Dtsch. Wiss. Komm.
Meeresforsch., 21 (1-4): 474-493.
THORSON, G., 1964: Light as an ecological factor in the dispersal and settlement of larvae of marine
bottom invertebrates. Ophelia, 1: 67-208.
TITELBAUM, M., 1966: Behavior and settling mechanism of planulae of Hydructinia echinata. Biol.
Bull., 131: 410-411.
TURPAEVA, E., M. GAL’PERIN & R. SIMKINA, 1977: Food availability and energy flux in the mature
colonies of the hydroid polyp Perigonirnus megas KINNE.Okeanologiya, 17 (6): 1090-1101 (In
Russian).
VERVOORT, W., 1966: Bathyal and abyssal hydroids. Galathea Report, 8: 97-174.
WASSERTHAL, L. & W. WASSERTHAL, 1973: Okologische Bedeutung der Schleirnsekretion bei der
Planula-Larve der Hydroidengattung Eudendrium. Mar. Biol., 22: 341-345.
WEDLER, E., 1975: okologische Untersuchungen an Hydroiden des Felslitorals von Santa Marta
(Kolumbien). Helgol. wiss. Meeresunters., 27, 324-363.
WERNER, B., 1956: Uber die entwicklungsphysiologischeBedeutung des Fortpflanzungswechsels der
Anthomeduse Rafhkea octopuncfafaM. SARS.Zool. Anz., 156: 159-177.
- -, 1958: Die Verbreitung und das jahreszeitliche Auftreten der Anthomeduse Rathkea ocfopunc-
tata M. SARSsowie die Temperaturabhangigkeit ihrer Entwicklung und Fortpflanzung. Helgol.
wiss. Meeresunters., 6 137-170.
- -, 1962: Verbreitung und jahreszeitliches Auftreten von Rathkea octopuncfafa(M. SARS)und
Bougainvillia superciliaris (L. AGASSIZ),(Athecafa, Anthomedwae). Ein Beitrag zur kausalen
marinen Tiergeographie. Kiel. Meeresforsch., 18: 55-66.
- -, 1963a: Effect of some environmental factors in differentiation and determination in marine
Hydrozoa, with a note on their evolutionary significance. Ann. N.Y. Acad. Sci., 105 (8):
461-488.
- -, 1963 b: Experimentelle Beobachtungen uber die Wirksamkeit von AuRenfaktoren in der
Entwicklung der Hydrozoen und Erorterung ihrer Bedeutung fur die Evolution. Veroff. Inst.
Meeresforsch. Brernerhaven, 3: 153-177.
WILLIAMS,G., 1965: Observations on the behaviour of the planulae larvae of CIuva squamatu. J.
Mar. Biol. Assoc. U. K., 45: 257-273.
- -, 1976: Aggregation during settlement as a factor in the establishment of coelenterate colonies.
Ophelia, 15: 57-64.
ZEN,N., 1963: The early development of the Anthomedusa Polyorchis karafufoensis KISHINOUYE.
Annot. Zool. Jpn., 36: 187-193.