J .

Fish B i ~ l(1974)
. 6, 119-133

Biosociology and ecology of pomacentrid fishes around the

Sinai Peninsula (northern Red Sea)
L. FISHELSON, AND A. AVIDOR*
D. POPPER
Department of Zoology, Tel Aviv University, Tel Aviv, Israel
(Received 2 August 1973)
Pomacentrid fishes inhabiting the coral formations along the Red Sea coast of Sinai
Peninsula form specificgroups of species on various habitats. Ainphirionbicinctus, Dascyilus
ariianus, D. marginatus and Pumacentpus tvichoirrus are typical for the backreef lagoon;
Pomacentrus sulfuveus, Abudefduf melanopus and A. melas are common along the rearreef
wall; Pomacentrus albicaudatus, P. tripunctatus, Abudejduf lacrymatus, A . leucozona and
A. unnulatus dominate the coral reef table. The forereef wall is occupied by Abudefduf
soxatilis, A. sexfasciatus and A . leucogaster. The lower part of the forereef and coral
knolls along it are inhabited by groups of Abudefduf azysroiz, Chromis dimidiatus, C.
caeruleus, C. ternatensis and of the Dascyllus trimaculatus. Among the fishes observed a
part are living in groups or schools, whereas others are typical solitary, permanent territory
holders. Most of them are polygamous, breeding in colonies, or solitary in nests, guarded
by males. The variation in feeding behaviour as well as in sociobiology, enables the high
diversity of these fishes on various parts of the habitat.

I. INTRODUCTION
Among the fish families found along the coral and rocky environments of the tropical
seas the family Pomacentridae seems to be the most conspicuous one, occurring in
almost all habitats and in a variety of forms. They have attracted the attention of
ichthyologists and ethologists from almost all countries bordering tropical seas and so
a great number of investigations have been published about them. Most of these
papers deal with the reproductive behaviour of various species (Reese, 1964), while
others deal with their sound production (Garnaud, 1957; Albrecht, 1969; Schneider,
1964; Myrberg, Brahy & Emery, 1967), their distribution (Monro, 1955; Bardach,
1958; Schultz, 1960; Smith, 1960) and feeding behaviour (Hiatt & Strasburg, 1960;
Eibl-Eibesfeld, 1962; Hobson, 1965; Randall, 1967). From the Red Sea, pomacentrid
fishes are first mentioned in the works of Forski3 (1775), Ruppell (1828), Klunzinger
(1870) and Gunther (1880). The behaviour of various species occurring in the
northern Red Sea have been discussed in papers of Gohar (1948), Abel (1961) and
Fishelson (1964, 1965, 1968, 1970). Most of the publications mentioned deal with
various aspects of the biology of single species, and almost no information was pub-
lished dealing with general aspects of sociobiology and the ecodistribution of groups
of pomacentrid-species occurring in a given locality.
This paper summarizes observations and experiments on pomacentrid fishes carried
out over 20 years of work on coral reef ecology of the northern Red Sea. During these
investigations special emphasis was given to the social structure and behaviour of
single species, as well as to the link of various fish species among them and to a certain
type of habitat.
II. METHODS
The observations were made in the very shallow subtidal waters to a depth of 40 m. They
were carried out during trips around the Sinai Peninsula from the north of the Gulf of
Aqaba, around Sinai, to the north of the Gulf of Suez. Two expeditionswere also made to the
Dahlak Archipelago of thc southern Red Sea (Oren, 1962; Lewinson & Fishelson, 1967).
* Partly sponsored by the Smithsonian Institntion Grant, SFC-7-0074.
119
120 L . F I S H E L S O N , D. POPPER A N D A . A V I D O R

Most extensive and frequent observations were performed in the vicinity of Eilat harbour in
the Gulf of Aqaba. During the underwater observations notes were taken on plastic PVC
sheets, and animals were phtographed with Rollimarine and Nikonus cameras. For motion
pictures a Bolex l6mm camera with underwater housing was used. In addition to field
observations, various species of pomacentrids were collected and observed in captivity in the
aquaria of the Department of Zoology, Tel Aviv University. Several specimens of almost
each pomacentrid species were dissected and their stomach contents noted, for information
on food habits.

In. DESCRIPTION OF THE STUDY AREAS

The coral reef tracts along the shallow waters of the Sinai Peninsula form several
main habitats very different in their topography and distribution (Fishelson, 1968, and
in preparation). Starting from Eilat harbour in the northern part of the Gulf of Aqaba
and extending approximately 30 km to the south, the corals form typical reef crests,
separated from the shore by a backreef channel (lagoon) 10-15 m wide and 1.0-1.5 m
deep at high tide (Fishelson, 1971). In this habitat the first small growths of corals are
found in the very shallow subtidal, usually represented by the colonies of Stylophora
pistillata (Loya, 1972; Fishelson, 1973). From here on, dispersed coral heads are
found all over the sandy bottom of the lagoon. The actual crest forms a relatively
wide belt, frequently interrupted by crevices and passages that connect the lagoon
waters with the forereef and open sea. The upper part of the crest forms horizontal
platforms 5-40 m wide, extensively eroded and rich in cracks and crevices. Compared
to the 3.0-6.0 m high forereef, the backreef wall of these crests is only 1.0-1.5 m high
and much poorer in coral cover. The corals dominating the forereef slope are species
of the genera Acropora, Pocillopora and Porites; and the h yd rocoralline species
Millepora dichotarna and M. platyphylla. The richest coral growth is in the uppermost
fringe of the forereef. Seawards the coral crest descends into a flat sandy platform on
which large coral pinnacles and patches are found. These are 2.5-5 m high and look
like submerged islets, composed of the same coral elements as the coral crests. These
pinnacles are adjacent to the forereef wall or may be located farther away into the sea.
The investigations on coral-species diversity in this region (Loya, 1972) showed that
the coral crests are exceptionally rich in coral species that form a very complex
environment serving coral dwelling fishes and invertebrates.
An entirely different habitat starts 30-40 km south of Eilat, and is found along the
shore of the Gulf of Aqaba to its southern end. In this habitat, instead of a shore
channel, coralogenous or beach-rock platforms are found that extend all along the
subtidal area. These more or less horizontal tables of 10-120 m in width, slant slightly
seawards. From such places the coral growths take over, forming a dense and rich
cover over the descending substrate. In this way again a well developed coral fringe
and forereef are formed that provide a complex environment for fish settlement.
The third and most extreme coral habitat is found only in several isolated places,
where the deep sea is very close to the shore line. The organogenic beach rock in such
Iocalities forms a narrow belt 2-0-6.0m wide. All the coral formation extends on
the vertical slope, forming only the forereef habitat. Such places are constantly
agitated by wave motion, the coral heads are strong, but dispersed, and the surface of
the rocky substrate is exposed.
In the Gulf of Suez, along the western Sinai Peninsula, a different coral habitat is
found; close to the most southern point at Ras Muhamed the coral crests resemble
those of the northern part of the Gulf of Aqaba. Northward, the Gulf of Suez
 BIOSOCIOLOGY AND ECOLOGY OF POMACENTRID FlSHES 121

becomes shallow and the coral formations occur in forms of iieogenic growths
over sandstone banks, patches and platforms. In many cases, these grow far out in the
Gulf, without any connection to the shoreline. The shape of those coral habitats is
elongate or rounded, with gradually descending outer parts. In most cases, the bulk
of the platforms is formed by sandy beach-rock. The coral covers found here are
usually poor in species, large areas being covered by a single species of the genus
Acropora (A. hemprichi, and A . microphthalma) and Stylophora pistillata.

DISTRIBUTION OF POMACENTRID FISHES

The highest coral species diversity was found in the coral crest of the northern Gulf
of Aqaba. Here also was found the most complex distribution of pomacentrid species.
Along cross sections that extend from the shore line over the coral crest, to the reef
edge, a total of 20 species of fish from this family was noted. These fishes form groups
of species in each zone of the habitat (Fig. 1). The anemonefish, Amphiprion bicinctus
(Riippell) is found in the shallowest waters close to the shoreline [Plate I(a)]. Juvenile
clown-fishes are found in the upper subtidal, hiding among arms of the host sea-
anemone Radiunthus koseirensis. Occasionally also single juvenile specimens of
Dascyllus trimaculatus (Ruppell) occur with them together in the same host. The
living coral colonies of Stylophora pistillata, usually found in the slightly deeper
subtidal, and across the lagoon, are inhabited by schools of Dascyllus aruanus (L.) and
D. rnarginatus (Ruppell). These three Dascyllus (Cuvier) species and the anemonefish
with their host-anemones (Fishelson, 1965) are found all across the reef, extending
seawards to a depth of 15 m. It seems that the factor which limits their distribution is
the availability of Stylophora corals and sea-anemone hosts.
The typical species of the backreef wall facing the lagoon are Abudefduf melas (Cuv.
et Val.), A. melanopus (Bleeker), and Pomacentrus sulfureus (Klunzinger) [Plate I(b)].
The latter two species are usually found on places covered by large colonies of the
soft coral, Lithophytum arboreum. Another species that occurs here is Pomacentrus
iripunctatus (Cuv. et Val.). The optimal habitat of this last species is the horizontal
surface of the coral table. Hiding in crevices and holes, are also found Abudefdqf
lacrymatus (Quoy & Gaimard) and occasionally Pomacentrus albicaudatus (Bosch.-
Salv.) and A. leucozona (Bleeker). In crevices like this also dwell young specimens of
A . melas. All of these species that occur and dwell on the coral tables are rather
small forms, the length of adults being 80-100 mm total length.
Entirely different are the pomacentrid species assembled along the forereef wall,
especially on its upper part. Here the dominating species are Abudefduf saxatilis
(L.) [Plate I(e)], A . sexfasciatus (Lac.), and at some places also A. sordidus (Forskid).
These species are large, the adults being 100-250 mm total length. On the lower zones
of the forereef the dominant pomacentrids are schools of Abudefduf azysron (Bleeker)
and Chromis dimidiatus (Klunzinger), while A . leucogaster (Bleeker), also occur,
especially on places rich in colonies of the fire coral, Millepora dichotoma, among
whose plates they hide [Plate T(f)]. In the same habitats large heads of Acropora corals
are populated by schools of Chromis caeruleus (Cuv. et Val.). Mixed with those three
gregarious living species, are also found Pomacentrus albicaudatus, P. sulfureus and
occasionally Dascyllus trimaculatus. Around coral pinnacles, the pomacentrid
populations are similar to those found on the nearshore lying reef: on top of the
pinnacles are found species that dwell on the reef table, and around the pinnacles
especially prominent are the schooling Abudefduf azysron [Plate II(a)] and Chromis
122 L. FISHELSON, D . POPPER AND A . AVIDOR

dimidiatus [Plate II(b)]. Also groups of Dascyllus trinzaculatus are occasionally found
in this habitat [Plate II(c)J. In some places, mixed with groups of A. uzysron groups
of Chromis ternatensis (Bleeker) also occur.
The above distribution of various pomacentrid species illustrates the ranges in
which the animals are found most of the time, especially during their reproduction
cycle. Occasionally, during various activities they may leave these places, wandering
further afield. Individuals of Abudefduf saxatilis, A . melus, and A. leucogaster
encountering the forereef fringe, during high tides invade the lagoon, collecting food.
As the sea-level descends, they swim back to their usual places. Especially mobile are
juvenile individuals of A. saxutilis, which may be found along the shore line and even
within the tidal rock-pools. When the open surface waters are rich in plankton,
pomacentrids of the tables as well as those from the forereef, ascend together and
swim out to feed, on such occasions frequently mixing with fishes from other families.
The lagoon habitat is replaced farther south along the Gulf of Aaqba by a beach-
rock table and the assemblage of pomacentrid species occurring here becomes scarcer.
From a point approximately 30 km south of Eilat, on the beach rock platform, the
dominating species are Abudefduf leucozonu, A. lacrymatus and A . annulatus (Peters).
Scattered on the table are also found specimens of Pomaceiztrus tripunctafus. The
upper part along the forereef wall is again dominated by A. saxatilis, mixed with A.
sexfasciatus.
In such habitats are also found single individuals of A . sordidus [Plate II(d)] the
rarest Abudefduf species in the Gulf. In only one locality was a group of 18 specimens
of this fish observed. Usually, only single A . sordidus are found dispersed along the
holes of the most agitated part of the fringe. Close to Eilat single specimens of A .
sordidus were observed beneath large beach rock plates found within the lagoon.
The lower part of the forereef region along the beach-rock platforms is also occupied
by schools of Abudejduf azysron, Chrornis dimidiutus, and occasionally also C.
caeruleus. In the localities where the corals form only steep forereefs, being a narrow
fringe along the rocky shore, the smallest number of pomacentrid fishes is found. At
such places the first to disappear are the platform dwelling species but those inhabiting
forereefs remain common, with Abudefduf saxatilis and A. sexfasciatus dominating.
Along the reef fringe and slightly below, along the coral wall, groups of A . azysron
and the rarer A. sordidus occur.
A . saxatilis is the dominant species on the northern reefs of the Gulf of Aqaba,
whereas A . sexfasciutus replaces it towards the southern end of the Gulf, and remains
the dominant one in the Gulf of Suez.
In this Gulf, the large schools of pomacentrid fishes, so typical of the Gulf of
Aqaba, are no longer observed. Fishes, such as Abudefduf saxatilis; A. sexfasciatus
and A. azysron occur here singly or form groups of several species only. This decline
in fish-species diversity observed in the Gulf of Suez seems to be in accordance with
the decline of coral-species diversity on the shallow coral tables, as compared to those
in the Gulf of Aqaba (unpublished observations).
Species that remain numerous along the coral platforms of the Gulf of Suez are
Dascyllus aruanus and D. marginatus; these form colonies between branches of the
corals Styfophorapistiflata and S. wehlsi [Plate II(e)]. The most common fish species
dwelling on the coral platforms are Abudefduf lacrymatus, A. Ieucozona and A.
annulatus. Although no quantitative data are available, it seems that along the coral
crests within the Gulf of Aqaba, a constant increase of the coral reef fish species is
Knoll Fore reef Table Rearreef Lagoon

Legend

P tripunctatus A saxatiles

A lacrymatus A bicinctus
f9@
P albicaudatus

P sulfureus

D marginatus a
A leucogaster

D trrrnaculatus

A azysron
*a
Ch. caeruleos D aruanus
@
FIG.1. Distribution of pomacentrid fishes in various zones of the coral reef in the Gulf of Aqaba. Ch dimidiatus
 BIOSOCIOLOGY A N D E C O L O G Y O F P O M A C E N T R I D FISHES 123

observed towards the north, including pomacentrids. This seems to be associated with
the gradual increase of complexity in the coral formation growing along the shore line.
The opposite is observed along the Gulf of Suez: toward the north of the Gulf, a
constant decrease in the living coral cover is evident, along with a decrease in poma-
centrid fishes.
SOCIOLOGY OF RED SEA POMACENTRIDAE
All the species observed could be divided into two main categories according to
their tendency to aggregate : gregarious species and solitary species.
GREGARIOUS POMACENTRIDAE
Here are included species that spend at least a part of their life-cycle in groups, in
which there is some kind of affinity among the individuals. The size of such groups
may vary between several to several thousand individuals and may include animals of
different sizes and ages. The gregarious pomacentrid fishes can again be divided into
schooling fishes and shoal-forming species.
(a) Schooling fishes
Schooling fishes are those that during various steps of their behaviour, swim
strongly unidirectional and perform stereotypic movements, synchronically
repeated by most members of the group. The best example of a schooling pomacentrid
fish is Chromis caeruleus. These stationary fish hover above and inhabit large heads of
Acropora corals. The most striking behaviour of such a school is the synchronized
ascent and descent of the group out of and into their coral. During these movements
they resemble a flock of birds in that they act as an organized unit: all of them rise
together to feed and when alerted escape downwards together. Descent was observed
to occur spontaneously without any visible external stimuli. Such a school forms also
a permanent reproductive unit, in which agonistic behaviour is observed only during
courtship. During reproductive periods the fishes are close together and individual
territories are established on the coral surface by males. Parts of those surfaces form
the nests on which the eggs are spawned by the females, and which are guarded by
males until the fry hatch (Swerdloff, 1970). A social structure like this is found also in
DuscyZlus aruunus and D.marginatus (Fishelson, 1965). Groups of 10-20 individuals
of these species inhabit isolated coral heads and remain together in such numbers
even though ' empty ' corals are present in the vicinity [Plate II(e)]. Also in these
species, as in Chromis, synchronized up and down swimming, or so-called signal
jumps (Abel, 1960) could be observed, especially during reproduction. These two
species also form mixed schools and in such cases they act as one homogeneous unit.
Dascyllus species seem to be more aggressive in their interactions than C . caeruleus.
Each individual defends its site among the branches of the occupied coral against
other members of the group. This agonism seems to prevent crowding in small coral
heads and keeps the Dascyllus school small.
(b) Shoal-forming species
This category includes species living in groups, in which the individual swims in any
direction and behaves more or less independently, moving around only partially in
accordance with other members of the group. The largest shoals are those of Abudefduf
mxatilis and A. sexfusciatus. As described for A . suxatilis (Fishelson, 1970) most
shoaling species form loose small bands within the group, that may split and aggregate
repeatedly. While feeding they disperse and move in various directions, but never
perform synchronized spontaneous actions as do schooling fishes. At night each fish
124 L. F I S H E L S O N , D . P O P P E R A N D A . A V I D O R

looks for its own place to hide and rest, and each morning they gather again, forming
small groups that merge to form large shoals.
According to the relation to the surrounding substrates all the gregarious species could be
divided into two groups. The typical schooling species are strictly stationary. Their
home range always includes a solid object, that forms the centre of their activity. Such
an object is a permanent coral head, rock, or floats constructed by man. Schools of
Abudefduf azysron, Chromis dimidiatus, together with schools of fishes from other
families (Anthias (=Frenzia) squamipinnis (Peters), Pempheris oualensis Cuv. and Val.)
form multispecific assemblies that act as one uniform society (Fishelson, Popper &
Gunderman, 1971; Popper & Fishelson, 1973). Such groups of species remain for
years together at the same place [Plate III(a), (b)]. This was confirmed for a group of
adult individuals of Dascyllus trimaculatus marked with dorsal tags and studied
near Taba (Eilat) [Plate IV(b)]. These tagged fish, which were recaptured at least
twice a month, were found around the same coral pinnacle ;for almost two years.
None of them were observed to move to the surrounding coral structures. The
whole group spent the night between branches of a single head of Acropora coral.
Such stationary groups of fishes are typical around coral pinnacles, as well as on
appreciated places along the forereef slope. Also stationary are small schools of
Dascyllus aruanus and D. marginatus, as well as the more numerous Chromis cacruleus.
The first two species often occur together in one coral head, although among the
coral branches they act individually, each defending his site. When swimming
out around the coral they behave as typical homogeneous schools. Chromis caeruleus
forms much larger colonies and usually does not mix with other colonial species.
The juvenile fishes of those schooling species are also found on the same sites as
the adult fishes. In most cases such groups of young fishes have been observed
to form some kind of sub-schools within the schools of adults. Those juveniles are
more attached to the rocky shelter and while the school of adults swims out for
feeding, the juveniles form clumps that remain closer to the substrate. They are
also the first to disappear when alarmed. Shoaling fishes are usually the wanderers
among the pomacentrids. Of those, Abudefduf saxatilis and A . sexfasciatus are the
typical vagabonds moving along the coral walls, entering and swimming out from
various caves and lagoons. Their swimming, although occurring together, is defin-
itely less organized than that of the schooling species (Fishelson, 1970). Such
wandering groups of Abudefduf usually attract groups of other fish species, and in
such a manner, multispecific assemblies are formed that for a short time act together.
The juveniles of wanderers behave like the adults except they are found in more
sheltered places, among rocks or in caves and crevices. Juveniles of A . saxatilis
attaining 30-50 mni total length emerge from cover and rising in the water join shoals
of the adult fishes, usually forming the ' tail ' of a moving group.
Solitary pomacentrids. This category includes fishes found always loosely distributed
over their habitats, where each specimen acts individually, usually close to a chosen
shelter. In such species agonistic behaviour is much more pronounced than in those
living in groups. The extreme solitary species are Pomacentrus albicaudatus, P.
tripunctatus, P. sulJureus, Abudefduf leucozona, A . inelas and Chromis ternatensis. Of
these species, P. sulfiweus and A. melas are open water fishes, that occur along the
shoreward and seaward coral rcef fringes. Only rarely may individuals of these species
be observed to swim higher than 0-5-0-7 m above the substrate; most of the time they
remain very close to their retreats. The other solitary species are inhabitants of
PLATE1. (a), A pair of Amphipvion biciticrus in their host anemone (c. of natural size); (b), Pomu-
cetitvus src/fureus (c. 4 of natural size); (c), Pomacetitvus albicaudutus ( c . f of natural size);
(d), Abude/ihj leucozotzu (c. 4 of natural size); (e), a shoal of Abudefduf saxatilis along the
forereef fringe (black fish at the bottom is A . melus); (f), Abudefduf leucogasrev (close) and
A. uzyrson, along the forereef wall.
 11. (a), Abudefdufazysron (c. natural size); (b), Chromis dimidiatus (c, natural size); (c), a group
PLATE
of Duscyllus trimaculatus; ( e ) , Dascyllus aruanus among branches of the Stylophora coral
colony; (d), Abudefduf sordidus (c. 4 of natural size).
PLATEI 11. Synchronized behaviour of an interspecific group of pornacentrids. (a), dispersion during
feeding; (b), spontaneous escape descent toward the shelter.
PLArE IV. (a), Communal spawning of Dascyllus orwanus (c. natural size); (b), male Dascyllus trimu-
crrlutus guarding the nest (with a tag on its back) (c. 4 natural size); (c), male Abudefduf leuco-
gcrster guarding the spawn on a plate of Millepora (c. 1 natural size); (d), spawn with embryos
of Abudefduf azysrun (c. x 25).
 B I O S O C I O L O G Y A N D E C O L O G Y OF P O M A C E N T R I D F I S H E S 125

crevices and holes within the coral reef structure. During the day they swim close to
the rocky substrate, entering and emerging from the holes. Within their range a
certain hole or crevice serves as the centre of the home or territory, and around this
spot the fish activity is the most prominent. During low tide, these hole-dwelling
solitary fishes are almost the only pomacentrids found on the reef table.
Among these fishes, Pomacentrus albicaudatus [Plate Z(c)] seems to be the most
euryeucous one, occurring from the subtidal to deep water. This species is also the
most aggressive one, attacking intruders viciously. In captivity, even in large tanks, it
was impossible to keep two adult fishes together.
All other solitary species are less aggressive and most of them are found on coral
tables and pinnacles, as well as on sand banks of the Gulf of Suez covered by corals.
Juvenile specimens of solitary fishes behave similarly to the adults-they hide in
crevices usually in shallow water and are also very aggressive in their behaviour. It
seems that the range of their movements is more restricted than the adults.
Intermediate social t-ypes. Among the various pomacentrid species, a special behav-
ioural pattern iq displayed by the anemone fish, Amphiprion bicinctus: the adults are
strictly stationary, living in pairs within one, or rarely two, sea anemones [Plate I(a)].
The hosts are the centre of their territory, and if such a host moves over the rock,
changing place, the fishes move together thus altering their home range (Fishelson,
1964). The distribution and number of fishes is virtually in accordance with the
distribution and number of available sea anemones. The juvenile fishes of this species
behave differently from the adults: they are usually found in groups of 5 to 20 indi-
viduals in one sea anemone, usually the shallow water species Radianthus koseirensis.
Upon reaching the length of 30-50 mm the fish disperse, forming pairs that occupy
large anemones. The anemone fishes, during their life span, begin as juveniles in
shoals and end as monogamous pairs, each of which with territorial behaviour. The
example of A. bicinctus leads to a group of species in which the group or solitary mode
of life is not so precise. Here belong Abudefduf leucogaster and Dascyllus trimaculatus,
species that demonstrate intermediate elements of social organization, in which
schooling and solitary behaviour occur together. A. leucogaster usually occupies gaps
among raised plates of the fire coral Millepora dichotoma, which grows principally
along the forereef region [Plate IV(c)]. In such a habitat each fish occupies a separate
gap which is used as shelter during alerts and serves as the resting place during the
night. They defend those places against individuals of their own species as well as
other fishes. At various times, as the fish rise above the coral colonies, they usually
aggregate, forming groups of 10 to 60 individuals. Such temporary units may move
together even to a distance of 5-20m from their territories. On returning to the
substratum, they switch back to territorial behaviour. Individuals of Dascylfus
trimaculatus are found scattered along the various zones of the reef, occurring with
other fishes. As mentioned earlier, juveniles often occupy sea anemones together with
A . bicinctus. Adult fishes are occasionally found in groups of 10 to 200 individuals
that demonstrate shoaling behaviour, but more frequently they are found swimming
singly along and within the reef. Elements of such transitional behaviour were also
observed in Abudefduf melas and A. lacrymatus. On several occasions fishes of those
species were observed to join groups of passing A. saxatilis, but as soon as the group
passes beyond their territories the joiners separate and descend to their homes.
Feeding behaviour. As evident from data published by Hiatt & Strasburg (1960)
Hobson (1968) & Randall (1967), most of the pomacentrid species that have been
126 L. FISHELSON, D . POPPER AND A . AVIDOR

investigated were found to be grazers or browsers that feed on a variety of food

organisms. Only a small number feed on planktonic animals.
The northern Red Sea pomacentrids, according to observations on feeding in
nature and in captivity, can be divided into three groups: one feeding on benthic
organisms, the other on plankton feeders and the third one with a mixed diet. The
most typical for the first group are Pomacentrus albicaudatus, P. trichourus Gunther,
P. sulfureus, Abudefduf leucozona, A. lacrymatus and probably A. sordidus. Specimens
of all those species are frequently observed to move close to the rocky surface and feed
directly on it. The stomach contents of P. trichourus consisted of algae, ostracods,
demersal fish eggs, small blennies and fragments of sponges. P. sulfureus was found
to feed mainly on algae, harpacticoid copepods, small polychaetes, and larvae of the
marine chironomid, Caelia marina. This last organism was found to hide and grow
among algae. The stomach contents of A . sordidus are composed of rock fragments,
nudibrancli molluscs, polychaetes and algae. Planktonic organisms and floating
organic material serve as food for Abudefduf saxatilis, A . sexfasciatus, A . leucogaster,
A. azysron, Chromis dimidiatus and C. caeruleus. While feeding, these fishes ascend
in the surrounding waters, often rising several metres above their rocky habitat. This
is especially prominent when southern currents occur and plankton masses approach
the coral region along the shore line. In such instances, shoals of A. saxatilis and A .
sexfasciutus mix with schools of Caesio lunare (Cuv. & Val.) and Acanthopagrus
bifaciatus (Forskil), all feeding together. At the same time while swimming reotac-
tically, schools of A. azysron, C. dimidiatus and C. coeruleus spread out in the water
collecting food [Plate Ill(a)]. The stomachs of these fishes were found to contain
large amounts of mysid shrimps, calanoid copepods, arrow worms, pelagic poly-
chaetes and hydromedusae. On one occasion the stomachs of several specimens of
A . sexfasciatus were found packed with salps and pelagic amphipods.
The following species feed on a mixed diet : Pomacentrus tripunctatus, Amphiprion
bicinctus, Dascyllus trimaculatus, D. aruanus and D. marginatus. When plankton
is abundant, these fishes rise in the water and collect food like the typical plankton
feeders. They may even join other fish feeding in the same way. In other instances,
when plankton is scarce, these fishes are observed near their shelters, collecting food
from the rocky surfaces and above the sediment. The stomachs of P. tripunctatus
were found filled with algae, rhizomes of hydrozoan colonies, as well as with pelagic
mysid shrimps and polychaetes. D. trimaculatus was found to feed on harpacticoid
and calanoid copepods, benthonic isopods and planktonic fish eggs. The stomachs
of D. marginatus and D. aruanus contained filaments of algae, salps, benthic copepods
and tunicates. Amphigrion was found to feed on various fouling organisms, amphipods
and planktonic organisms.
Reproduction. The reproductive behaviour of pomacentrid fishes has been described
in captivity and nature. Common to all the species investigated, is the development
of territoriality, accompanied by specific courtship behaviour, especially of males.
The pattern of these activities seems to be more or less uniform, beginning with the
establishment of nesting places, their defence, signal swimming and leading the
females to spawn. Subsequently, the eggs are guarded by the territory holding male
until hatching. Most of the species observed in the northern Red Sea are polygamous,
the only known exception being Amphiprion bicinctus, that for years live as mono-
gamous pairs. In this case, both of the partners take care of the eggs and defend them.
All other species, as far as is known, pair only for the time needed for spawning.
 B I O S O C I O L O G Y A N D E C O L O G Y O F P O M A C E N T R I D FISHES 127

These polygamous fishes may be divided into colonial breeders and solitary breeders.
Dascyllus aruanus and Abudefduf saxatilis (Fishelson, 1964, 1971) are the typical
colonial breeders, but they differ in the duration of the bond between the sexes. A
breeding colony of D. aruanus consists of several fishes to several tens of individuals
and is one indivisible territorial unit restricted to a single coral head. In such a
colony, all the fishes (especially the males) know each other and will bite and chase
away any new intruder. Being together permanently reduces the agonistic behaviour
during courtship and spawning to a set of ritualistic postures. This enables each male
to hold his nesting site among the coral branches. When spawning begins, the borders
of such individual nest sites are often abolished [Plate IV(a)], and two or three neigh-
bouring males may together fertilize the eggs produced by several females (Fishelson,
1964). After such dense spawning, no limits are found between one batch of eggs and
another, and in this case each male guards and defends a part of the communal layer
of eggs. This type of group reproduction has also been observed in Chromis caeruleus.
In A . saxatilis the reproducing individuals represent a fraction of a large, mobile
shoal o f fishes. The localization and establishment of a breeding colony is a temporary
act and is started by one or several ripe males: these individuals choose the site for a
temporary colonization on a rocky surface and their individual territories are estab-
lished by antagonistic encounters among the pretenders. These territories are often
set well apart from each other. The fighting among these males is very vigorous in the
beginning of colonization, as new fishes try to take a part in this, but after the individual
nesting places are with spawn, the aggressiveness decreases, and is succeeded by a
more peaceful coexistence of males defending their respective nests. The hatching of
larvae ends the existence of a colony, and the now free male joins the wandering fishes.
This type of reproduction was also observed for A. sexfasciatus, and presumably
occurs also in A . sordidus. Abudefduf azysron and Chromis dimidiatus seem to be
intermediate in reproductive behaviour between the permanent colony breeders such
as D.aruanus and the temporary colonial breeders such as A . saxatilis. These species
are stationary schooling fishes, but instead of occupying an isolated coral head, they
are usually found along vertical walls of forereefs and of pinnacles. During repro-
duction each male occupies and defends his nest on the rocky surface inhabitated by
all of the group. Usually those nests are separated from each other and thus resemble
the pattern described for the temporary colony holders. Here each male guards his
spawn [Plate IV(a)] but remains in the vicinity with the other fishes after the young
hatch.
Solitary breeders. This group includes all the pomacentrid species that are dwellers of
holes and crevices of the coral framework, especially the platform surfaces. Some of
them belong to the genus Abudefduf (Zeucozona, lacrymatus, annulatus) and others to
the genus Pomacentrus (albicaudatus, tripunctatus and trichourus).
In those species, within a territory that is sometimes 2-4 m in diameter, each male
occupies a nest hole, actively defending it against conspecific males. Such nest holders
perform courtship signal movements, leading receptive females to the nest for spawn-
ing. After mating with one female, the male leads another to the nest, if more are
available. Such guarding and nesting may continue throughout the season, usually
from March/April till September, with some decline in July. This pattern of repro-
ductive behaviour occurs also in Pomacentrus sulfureus and Abudefduf melas, species
which perform solitary courtship motions along the coral reef fringe.
Somewhat different are the breeding habitats of Abudefduf leucogaster and Dascyllus
128 L . F I S H E L S O N , D . P O P P E R A N D A. A V I D O R

trimucuZutus. The first species breeds normally in groups on dead bases of vertically
rising plates of MilJeporu [Plate IV(c)]. Only exceptionally were single males found to
breed beneath plates lying on the bottom. Each male has his side of a plate on which
he guards the spawn, and so two males may breed on either side of the same plate,
not intefering with each other. Such sites can be used by the same male for several
years (Dafni, pers. comm.). As for D. triuiaculutus, each male chooses his nest
without concern for other males. The nests may be found on vertical cliffs, small
stones and rocks [Plate IV(b)]. Acting individually close to its nest, the male performs
courtship behaviour, producing sounds and signal swimming (Shpanier & Fishelson,
in prep.). On one occasion when a group of 180 individuals was observed, the males
were seen to establish their nests well dispersed from each other, in a radius of approxi-
mately 5-7 m from the main group. From time to time each of the nest holders was
observed to swim into the main aggregation and after a while lead a female toward
their territory. Among such performing males almost no aggressive encounters were
observed. Contrary to this, in captivity, D.trimuculatus showed persistent aggressive
behaviour like other Duscyllus species. In this species, as in many others, the transfer
into the limited space of aquaria triggers agonistic behaviour. Biting and wounding
seen in aquaria were never observed in nature.

IV. DISCUSSION
The distribution pattern of animals in any ecosystem evolves as the end product
of an evolutionary process under particular environmental conditions. The major
factors that govern such developments are environmental, among them, habitat
complexity (MacArthur & MacArthur, 1961 ; Smith, 1972), ecosystem predictability
(Slobodkin & Sanders, 1969) and the particular physico-chemical properties of the
habitat (tropics versus non-tropics as presented by MacArthur, 1964, and others).
Other important factors in this context are of biological nature, the tolerance of the
animals to environmental extremes (McIntosh, 1963); the decrease in competition
(Dice, 1952; Klopfer, 1962) and development of interspecific social relations such as
mimetic and symbiotic associations (Wickler, 1969; Springer & Smith-Vaniz, 1972;
Slobodkin & Fishelson, in press). The intergroup structures like those observed in
birds (Davis, 1962; Crook, 1962; Lack, 1965; Murray, 1971) seem to be the most
complex one observed, that favours animal diversity in chosen places. The primary
factor that governs the distribution of pomacentrid fishes in the northern Red Sea
seems to be the complexity of the habitat. The reefs of the northern part of the Gulf
of Aqaba, along the Sinai Peninsula, are the most diverse and complex, when com-
pared with those of other localities (Loya, 1972; Fishelson, 1973). Only here the
various elements of a reef, as backreef lagoon, tablereef platform, and forereef region
are found in one locality. Only in such places could all the 24 species of pomacentrid
fishes, mentioned in this investigation, and inhabitors of those various sub-habitats,
be observed. As the foliage layer diversity is reflected in the bird species (in the sense
of MacArthur, Rechner & Cody, 1966) so the coral subhabitats influence fish species
diversity. Because of this, with the decrease in coral reef diversity as observed in the
southern parts of the Gulf of Aqaba, and especially in the Gulf of Suez (Fishelson, in
prep.) a comparable decrease in the pomacentrid fish diversity is observed. So for
example, in localities where no lagoons are found, the colonies of Duscyllus spp.
disappear, normally inhabiting here isolated Stylophoru coral heads. Also less
frequently found are the sea-anemone fishes, Amphiprion bicinctms. Similarly in places
 B l O S O C l O L O G Y A N D E C O L O G Y O F P O M A C E N T R I D FISHES 129

with a narrow, belt-like coral platform, crevice dwelling species ( P . albicaudutus,

A . annulatus, A. leucozonu) are very few or almost absent. Steep forereef walls,
partly bare, without coral growths, will also be poor in fish species, especially in the
school-forming Chromis (C. dimidiatus and C. caeruleus); as well as Abudefduf
azysron.
Such specific habitat-connected distribution of pomacentrid fishes points to a
certain type of specific utilization of the coral reef. As food resources are practically
unlimited, mostly because the major portion, plankton, is provided by the open sea,
the main factor of this utilization seems to be the availability of space (refuges) acting
against predation (see also Smith & Tyler, 1972). This leads to a socio-ecological
distribution of available space among the shelter seeking fishes. Such properties, as
mentioned by Tobach & Schneirla (1968), must have acted in different ways as factors
of natural selection, with high survival values. As open-water fishes among our
pomacentrids are very few (Abudefduf SaxatiEis being the only example) it seems that
the general evolutionary trend has been towards restricted movement and constant
shelter use. The most simple from this point of view, are the solitary species, that are
persistently territorial, and use individual shelters that are protected against con-
specific and even non-specific fishes (Reese, 1964; Low, 1971). Among our fishes, P.
ulbicaudatus was observed to attack strongly P. tripunctatus and vice versa; P. tui-
punctutus was also observed to chase away A . lacrynzatus. As the morphological
pattern of those fishes are very similar, one is coping with misdirected aggression, as
in birds (Murray, 1971) based on lack of recognition. This occurs when similar species
compete for space optimal for both of them.
In those solitary species of pomacentrids the factors that promote dispersion are
individual distances, territorial aggression, escape and avoidance tendency. Also in
those fishes, as in birds (Crook, 1962) the nests are cryptic and the home range
(feeding space) small. Thus the population distribution of such persistent territorial
pomacentrids will be the function of nest-site selection in relation to active competition
and predation. These circumstances should limit the population to small or moderate
numbers. In fact, on a platform of 100 square metres 8 specimens of P. tripunctatus
and 6 specimens of A . lacrymatus were observed, whereas among the social porna-
centrids hundreds can be found on 2 or 3 square metres. One of the important
ecological results of the solitary territorial intra- and interspecific behaviour, will be
the persistent numerical stability of the population in a specific habitat. This will
effect the survival rate as well as the ability of sexes to find each other. The picture
changes almost entirely as one considers the social living pomacentrids. As mentioned
by Wynne-Edwards (1963) each group of animals ' automatically constitutes a
society ' and natural selection occurred between such competing social structures
' favouring the more efficient variants '. The factors that promote congregation seem
to be social and sexual tendency coupled with decrease in aggression and common
habitat preference.
Abudefdufsuxatilis illustrates the transitional type of fish (Fishelson, 1970) in which
short time territoriality during reproduction occurs within dominating school
formation, with school-properties as described by Shaw (1965) and Cullen, Shaw &
Baldwin (1965). As in some of the Ploceine-birds, the gregarious behaviour in A.
suxatilis constitutes an important part of its daily-night changeover system (term by
Collette & Talbot, 1972). During this the school acts as a unit and like in insect social
bonds (Schneirla & Rosenblat, 1961) this limits the individual action of each single
130 L. F I S H E L S O N , D . P O P P E R A N D A. AVIDOR

fish to a minimum. Such behaviour disappears towards sunset, when each individual
begins to act solitarily, searching for a refuge in which to spend the night. The
survival value of such daily schools seems to be very high (Zweifellos-according to
Eible-Eibesfeldt, 1962) making it difficult for a predator to take aim at a single
victim. In the case of A. saxatidis the population size seems to be controlled by
predation during the daytime and not by space-availabjlily of nocturnal hiding places.
In an area of reef platforms, that theoretically could hide several thousands of A .
saxatizis, a population of 800 to 900 fish was observed during the course of several
years. It seems that most of the predation occurs during the juvenile stages of their
development. It seems that something like this should be found in A. sexfasciatus,
A . abdominalis, and presumably in Chromis multilineata, described by Myrberg,
Brahy & Emery (1967). As this type of behaviour was observed in A . saxatilis from
rhe smallest observable individuals, it seems that they are dominated by internal
innate factors.
The next stage of more complex social structure is found in DascyZlus aruanus D.
marginatus and Chromis caeruleus. Here the group forms a permanent unit, stationary
on a chosen coral head. The persistent interrelation between the individual of such a
bond, that includes feeding around the coral, and the attachment to a common shelter
(coral) plays the major role in the development of reciprocal processes and mutual
stimulations. The most interesting thing in such colonies is the decrease in aggression
among the members that permits the high compatibility of individuals. In such a way
20 to 25 DascyElus could be found in a coral head of one quarter cube metre, where
100 to 120 Chromis caeruleus are observed to inhabit a single Acropora-coral colony
of 1 m in diameter and 40 cm thick. Such density consequently led to the communal
spawning and fertilization of eggs (Fishelson, 1964; Sale, 1971). In DascylZus aruanus,
it was also observed that the males communally guard the egg layers till hatched. This
type of fish behaviour resembles the observations on communal nests of some of the
Anis cuckoos, as described by Davis (1942).
The highest and most complex level of social organization is observed in the inter-
specific school of fishes, formed by pomacentrid and non-pomacentrid species. Such
groups involve : Abudejduf azysron ; Dascyllus trimaculatus; Clzropnis dimidiatus; C .
caeruleus, and the serranid Anthias squamipinnis. In such cases the mutual stimulative
processes are formed on the intergeneric and even interfamilial basis. As mentioned
by Popper & Fishelson (1973) such groups of species show intergroup behaviour
patterns resembling those described for colonies of birds. In these cases not only can
single individual fishes not act individually-even single-species groups are strongly
limited in their acting. In such cases of inter-group socialization, a mutual defence
(pro-survival) mechanism developed, that includes in it elements of mimetic behaviour
[Plate 10(a, b)]. No doubt such levels of group socialization could develop only on
the basis of a common ecological-evolutionary, long-term process typical for most of
the coral ecosystems.
V. SUMMARY
(1) On the coral formation of the Red Sea coast of the Sinai Peninsula, the poma-
centrid fishes form assemblies of species typical for each zone of the habitat. The
diversity of pomacentrid fish species seems to be primarily correlated with the diversity
of the coral formation.
(2) Within the backreef channel developed especially in the northern part of the
 BIOSOCIOLOGY A N D ECOLOGY O F P O M A C E N T R I D FISHES 131

Gulf of Aqaba, the most typical species are Amphiprion bicinctus, Dascyllus aruanus,
D. marginatus and Pomacentrus trichourus.
(3) Pomacentrus sulfureus, Abudefdyf melanopus and A , melas are the typical
species along the rearreef wall.
(4) Within crevices of the reef table dwell Pomacentrus alhicaudaius, P. triyunctatus,
Abudefduf lacrymatus, A . leucozona and A. annulatus.
(5) Along the upper part of the forereef the dominant pomacentrid groups are A .
saxatilis, A . sexfasciatus and A. leucogaster. A. sordidus also occurs here but is not
common.
(6) The lower forereef and coral pinnacles are inhabited by stationary schools of
A . azysron, Chromis dimidiatus, C. caeruleus, C. ternatensis and occasionally groups of
Dascyllus trimaculatus.
(7) Among the fishes observed, a part live gregariously, forming well-organized
groups (Abudefduf azysron, Chromis dimidiatus, C. caeruleus, Dascyllus aruanus and
D. marginatus) which are permanently stationary on chosen sites. Others form shoals
(Ahudefduf saxatilis, A. sexfasciatus, A . sordidus, and partly Dascyllus trimaculatus)
which are mobile along and among the corals.
(8) The small coral table crevices are occupied by A. lacrymatus, A. leucozona,
Pomacentrus albicaudatus, P. trichourus; these are solitary fish, holding permanent
territories in this habitat.
(9) Amphiprion bicinctus and Abudejduf leucogaster possess an intermediate type
of social behaviour; during their lives they are partly gregarious and partly solitary.
(10) Some of the fishes investigated feed on benthic organisms, including algae and
animals, whereas others feed on planktonic organisms, A third group of species was
found to have a mixed diet from the benthos and plankton.
(I 1) Most of the pomacentrids observed are polygamous : only Ainphiprion
bicinctus is monogamous, and1 ives in pairs. The gregarious fishes breed in colonies,
whereas the solitary species have males which guard separate nests.
(12) The various sociobiological types found in the pomacentrid fishes enables
comparison with similar phenomena described for birds, and other animals.

The authors wish to thank Drs V. S. Springer, E. Clark and G. Allen for their very valuable
remarks on the manuscript. Thanks are also due to M. S. Shaefer for the preparation of
Fig. 1, as well as to the many students and colleagues whose discussions stimulated this
work.

References
Abel, E. F. (1960). Zur kenntnis des Verhaltens und def Okologie von Fischen an Korallen-
riffen bei Ghardaqa (Rotes Meer). 2. Morph. Okol. Tiere 49, 430-503.
Abel, E. F. (1961). Freiwasserstudien uber des Fortpflanzunyerhalten von Monchfishes
Chromis chromis L., einem vertreter der Pomacentriden in Mittelmer. Z . Tierpsychol.
18,441-449.
Albrecht, H. (1969). Behaviour of four species of Atlantic Damselfishes from Colombia,
South America (Abudefduf suxutilis, A. tuuvus, Chromus multilineatu, C. cyanea,
Pisces, Pomacentridae). 2. Tierpsychol. 26, 662-676.
Bardach, J. E. (1958). On the movement of certain Bermuda reef fishes. Ecology 39,139-140.
132 L. F I S H E L S O N , D. P O P P E R A N D A. A V I D O R

Collette, B. B. & Talbot, F. H. (1972). Activity patterns of coral reef fishes with emphasis on
nocturnal-diurnal changeover. In Results of the Tektite Program: Ecology of Coral
Reef Fishes (Ed. Collette, B. B. & Earle, S. A.), Nat. Hist. Mus. Sci. Bull. (Los Angeles)
14, 98-124.
Crook, J. H. (1962). The adaptive significanceof avian social organization. Symp. zool. SOC.
London 14, 181-218.
Cullen, J. M., Shaw, E. & Baldwin, H. A. (1965). Methods for measuring the three-dimen-
sional structure of fish schools. Anim. Behav. 13, 534-543.
Davis, D. E. (1942). The phylogeny of social nesting habits in Crotophaginae. Rev. Bid. 17,
115-134.
Davis, D. E. (1962). An inquiry in the phylogeny of gangs. In Roots of Behaviour (Ed. Bliss,
E. L.), pp. 316-320. New York: Harper and Brothers Publ.
Dice, L. R. (1952). Natural Communities. Michigan: University of Michigan Press.
Eibl-Eibesfeld, I. (1962). Freiwasserbeobachtungen zur Deutung des Schwarmverhaltens
verschiedener Fische. Z. Tierpsychol. 19, 165-182.
Fishelson, L. (1964). Observations on the biology and behaviour of Red Sea coral fishes.
Bull. Sea Fish. Res. Stn Israel 37, 11-26.
Fishelson, L. (1965). Observation and experiments on the Red Sea Anemones and their
symbiotic fish Amphiprion bicinctus. Bull. Sea Fish. Res. Stn Israel 39, 3-1 6.
Fishelson, L. (1968). Marine Biological and Oceanographical Research in ihe Red Sea.
Final Report for the Office of Naval Research (mimeographed).
Fishelson, L. (1970). Behaviour and ecology of a population of Abudefdufsaxatilis (Poma-
centridae, Teleostei) at Eilat (Red Sea). Anim. Behav. 18,225-237.
Fishelson, L. (1971). Ecology and distribution of the benthic fauna in the shallow waters of
the Red Sea. Mar. Biol. 10, 113-133.
Fishelson, L. (1973). Ecology of coral reefs in the Gulf of Aqaba (Red Sea) influenced by
pollution. Oecologia (Berl.) 12, 55-67.
Fishelson, L., Popper, D. & Gunderman, N. (1971). Diurnal cyclic behaviour of Pempheris
oualensis, Cuvier and Valenciennes (Pempheridae, Teleostei). J. Nat. Hisf. 5,503-506.
ForskB1, P. (1775). Descriptiones animalium avium, piscium, amphibiorum, insectorum,
vermium; qua1 in itinere orientali observavit : 1-19-XXXII, 1-164. Hafniae.
Garnaud, J. (1957). Ethologie des Dascyllus trimaculatus (Rupp.). Bull. Inst. Oceanogr. 54,
1-10.
Gohar, H. A. F. (1948). Commensalism between fish and anemone (with a description of the
eggs of Amphiprion bicinctus). Publs. mar. biol. stn. Ghardaga (Red Sea) 6, 3544.
Giinther, A. C. L. G. (1880). An Introduction to the Study of Fishes. Edinburgh: Adam.
Hiatt, R. V. & Strasburg, D. W. (1960). Ecological relationship of the fish fauna on coral
reefs of the Marshall Islands. Ecol. Monogr. 30, 65-127.
Hobson, E. S. (1968). Predatory behaviour of some shore fishes in the Gulf of California.
Res. Rep. U.S. Fish. Wildl. Serv. 74.
Klopfer, P. H. (1962). Behavioral Aspects of Ecology. New Jersey: Prentice-Hall.
Klunzinger, C. B. (1870). Synopsis Der Fische des Rothen Meeres mit einer Einleitung von
W. Klausewitz. Repr. 1964, Weinheim: J. Cramer.
Lack, D. (1965). Evolutionary ecology. J. appl. Ecology 2,247-255.
Lewinsohn, Ch. & Fishelson, L. (1967). The second Israel South Red Sea Expedition, 1965
(General Report). Israel J. Zool. 16, 59-68.
Loya, Y. (1972). Community structure and species diversity of hermatypic corals at Eilat,
Red Sea. Mar. Biol. 13, 100-123.
Low, R. M. (1971). Interspecific territoriality in a pomacentrid reef fish Pomucentrus fiavi-
cauda Whitley. Ecology 52, 648-654.
MacArthur, R. N. & MacArthur, J. (1961). On bird species diversity. Ecology 42, 594-598.
MacArthur, R. H. (1964). Environmental factors affecting bird species diversity. Am. Nut.
98, 387-398.
MacArthur, R., Rechner, H. & Cody, M. (1966). On the relation between habitat selection
and species diversity. Amer. Nut. 100, 319-325.
McIntosh, R. P. (1963). Ecosystems, evolution and selectional patterns of living organisms.
Am. Scien. 51, 246-267.
 B I O S O C I O L O G Y A N D E C O L O G Y O F P O M A C E N T R I D FISHES 133

Monro, I. S. R. (1955). The Marine and Freshwater Fishes of Ceylon. Canberra: Dept. of
External Affairs.
Murray, B. G. jr. (1971). The ecological consequences of interspecific territorial behavior in
birds. Ecology 52,414-423.
Myrberg, A. A. jr., Brahy, B. D. & Emery, A. R. (1967). Field observations on reproduction
of the Damselfish Chromis multilineata (Pomacentridae) with additional notes on
general behaviour. Copeia 819-827.
Oren, 0. H. (1962). The Israel South Red Sea expedition. Nature, Lond. 194, 1134-1137.
Popper, D. & Fishelson L. (1973). Ecology and behavior of Anthius squamiphinnis (Petters,
1855) (Anthiidae, Teleostei) in the coral habitat of Eilat (Red Sea). J. exp. 2001. 184,
409-424.
Randall, J. E. (1967). Food habits of reef fishes of the West Indies. Studies in tropical
oceanography 5, 665-847.
Reese, E. S. (1964). Ethology and marine zoology. In (Ed. Barnes, H.), Annual Review
Oceanography and Marine Biology, Vol. 2, pp. 445-488. London: George Allen and
Unwin.
Riippell, E. S. (1828). Atlas zu der Reise im nordlichen Africa. Fische des Rothen Meeres.
144 S., 35 Taf., Frankfurt a.M.
Sale, P. F. (1971). The reproductive behaviour of the pomacentrid fish Chromis caeruleus.
Z. Tierpsychol. 29, 1 5 6 164.
Schneirla, T. C. (1960). Instinctive behaviour, maturation, experience and development.
In Perspectives in Psychological Theory (Eds Kaplan, B. & Wagner, S.). New York:
Inter. University Press, 303-334.
Schneirla, T. C. & Rosenblat, L. (1961). Behavioral organisation and genesis of social bond
in insects and mammals. J. Orthopsychiat. 31, 223-253.
Schneider, H. (1964). Bioakustiche Untersuchungen an Anemonenfishen der Gattung
Amphiprion (Pisces). Z . Morph. Okol. Tiere. 53,453-457.
Schultz, L. P. (1960). Fishes of the Marshall and Marianas Island. Smithsonian Inst., U.S.
National Museum Bull. 11. 202.
Shaw, E. (1965). The optomotor response and the schooling of fish. Spec. Publs int. Commn
N.W. Atlant. Fish. 6, 753-759.
Slobodkin, L. B. & Sanders, H. (1969). On the contribution of environmental predictability
to species diversity. In Diversity and Stability in Ecological Systerrrs. Brookhaven
Symp. Biol. 22, 82-95.
Slobodkin, L. B. & Fishelson, L. (in press). The effect of Labroides dirnidiatus on the point
diversity of fishes on the reef front at Eilat. Am. Nut.
Smith, C. L. & Tyler, J. C. (1972). Space resource sharing in a coral reef fish community. In
Results of the Tektite Program: Ecology of Coral Reef Fishes (Eds Collette, B. B. &
Earle, S. A.). Nat. Hist. Mus. (Los Angeles) Sci. Bull. 14, 125-170.
Smith, F. E. (1972). Growth by intussusception. Trans. Conn. Acad. Arts Sci. 44, 307-335.
Smith, J. L. B. (1960). Coral fishes of the family Pomacentridae from the Western Indian
Ocean and the Red Sea. Rhodes Univ. Ichthyol. Bull., Grahamstown 19, 317-349.
Springer, V. G. & Smith-Vaniz, W. F. (1972). Mimetic relationship involving fishes of the
family Blenniidae. Smith Contr. Zool., Washington 112, 1-36.
Swerdloff, S. N. (1970). Behavioral observations on Eniwetok Damsel fishes (Pomacentridae;
Chromis) with special reference to the spawning of Chromis caeruleus. Copeia 371-374.
Tobach, E. & Schneirla, T. C. (1968). The biopsychology of social behavior of animals. In
Biological Basis of Pediatric Practice (Ed. Cooke, R. E. & Levin, S.), pp. 68-82.
New York: McGraw-Hill.
Wickler, W. (1969). Mimicry in tropical fishes. Phil. Trans. R. SOC.251, 473-474.
Wynne-Edwards, V. C . (1963). Intergroup selections in the evolution of social systems.
Nature, Lond. 200, 623-626.