Estuarine, Coastal and Shelf Science (1996) 42, 583595

Population Dynamics and Overgrowth of

the Sea Fan Eunicella cavolini
(Coelenterata: Octocorallia)

Markus G. Weinbauera and Branko Velimirov

Institut fr Allgemeine Biologie, Universitt Wien, Schwarzspanierstr. 17,
A-1070 Vienna, Austria
Received 21 October 1994 and in revised form 14 February 1995

Keywords: gorgonians; population dynamics; recruitment; mortality; overgrowth; Mediterranean Sea

The population dynamics of Eunicella cavolini Koch was investigated from
198991 at three study sites in the Bay of Calvi (Corsica, France). Population
density was highest in the channel (149180 colonies m "2), intermediate at the
wall (5466 colonies m "2) and lowest at the boulders (1519 colonies m "2).
Mortality, expressed as percentage of colonies that died within 1 year of
investigation, did not differ between the study sites and showed large variability
(83267%). Expressed as percentage of new colonies on total colony numbers, recruitment was highest in the channel (117201%), intermediate at the
wall (74120%) and lowest at the boulders (083%). A growth formula for
E. cavolini developed in an earlier study (Velimirov, 1975) was shown to be
suitable for the investigated study sites. The estimated maximum age was 21
years. Age frequency distribution did not vary strongly between subsequent
years during the 3-year investigation. However, the age structure differed
strongly between study sites. Overgrowth of colonies was highest at the
boulders, intermediate at the wall and lowest in the channel. Overgrowth
increased with colony age until the age of 4 years. Mortality of E. cavolini is
? 1996 Academic Press Limited
mainly due to detachment and overgrowth.

Introduction
Compared to other sessile marine animals, little is known of the population dynamics of
octocorals. Benayahu and Loya (1981, 1984a,b, 1985) investigated the life history of the
soft coral Xenia macrospiculata Gohar from the Red Sea; some information is available
on Australian soft corals (Dinesen, 1985; Farrant, 1987) and Caribbean as well as
Californian gorgonians (Grigg, 1975, 1977; Gotelli, 1988; Lasker, 1990). Demographic
models have been established for Caribbean gorgonians (Gotelli, 1991; Lasker, 1991)
and the importance of clonal propagation in population dynamics has been investigated
(Lasker, 1990). Weinberg (1978) studied the life cycle of the gorgonian Eunicella
singularis Esper in the Mediterranean Sea and there are data on the population structure
a

Present address: National Research Center for Biotechnology, Division Microbiology,

Mascheroder Weg 1, D-38124 Braunschweig, Germany.

02727714/96/050583+13 $18.00/0

? 1996 Academic Press Limited

584

M. G. Weinbauer & B. Velimirov

of Corallium rubrum L. (Garca-Rodrguez & Mass, 1986; Abbiati et al., 1993). Life
history tables have been constructed for only two gorgonians and one soft coral (Grigg,
1977; Farrant, 1987). Causes of mortality in octocorals, an important population
parameter, are detachment, abrasion, burial, overgrowth and predation. However, there
are only a few studies investigating quantitatively the mortality of gorgonians. Mass
mortality of gorgonians has been also reported (Woodley et al., 1981; Yoshioka &
Yoshioka, 1987; Garzn-Ferreira & Zea, 1992).
Eunicella cavolini Koch is one of the most abundant gorgonian corals in the
Mediterranean Sea (Weinberg, 1978). This species is azooxanthellate and clonal
propagation is not reported. Velimirov (1975) established a growth curve for E. cavolini,
and the orientation of the fan to the water movement and the morphological variability
have been studied (Velimirov, 1973, 1976; Russo, 1985; Weinbauer & Velimirov, 1995).
The causes of mortality have not been investigated. A preliminary investigation of the
population structure of E. cavolini showed densities of up to 180 colonies m "2
(Velimirov & Weinbauer, 1992). However, the population dynamics have not been
studied. Thus the aims of the present study were: (1) to determine population
parameters of E. cavolini from different study sites, and (2) to determine the amount of
overgrowth.

Material and methods

Study sites
In the investigated depth range, E. cavolini grows on vertical rock substratum characterized by coverage of brown algae (Cystoseira spp.) and various sessile invertebrates,
such as sponges, bryozoans and tunicates. Colonies were counted and measured at the
following sites in the Bay of Calvi (Corsica, France) near Stareso (Figure 1): (1) a
slightly overhanging wall in a large underwater channel at about 25 m depth; (2) a steep
wall from 16 to 28 m depth; and (3) a group of large boulders between 10 and 15 m
depth. In the following text, these study sites are termed channel , wall and
boulders . In the channel and at the wall, two transects 25 cm wide and 3510 m long
were set up on each rock face. The size, growth characteristics and position of the
colonies were recorded. The position of a colony was determined as distance to the
transect line, neighbour colonies and recognizable structures such as small crevices, tiny
holes or other sessile organisms. This allowed the recognition of individual colonies in
the subsequent years. Measurements of parameters was made in August 1989, August
1990 and August 1991. In 1990, two additional areas with colonies at the boulder field
were found and included in the present study, in order to increase the number of
colonies investigated at the boulders.

Age and growth

Total branch length (TBL) was determined using two methods. During a study on the
morphology of E. cavolini (Weinbauer & Velimirov, 1995), fan-like colonies were
collected in the Bay of Calvi. Colonies were dried and photocopied, and the TBL was
determined. The rectangularized fan surface area (RFSA) was determined from photocopies of the collected colonies and was defined as the length (distance between the two
most distant parts of the colony) multiplied by the mean of three measurements made at
equidistant positions at a right angle to the length. The RFSA was regressed against

585

Gorgonian population dynamics

9W

845' W

Wall

4235' N

Channel
Boulders

Calvi

Stareso
Corsica

Punta
Revellata

42N

Figure 1. Study sites of Eunicella cavolini in the Bay of Calvi.

TBL. For very small colonies or where no fan had developed, TBL was determined
directly by measuring each branch length separately with sliding calipers.
The TBL of each colony was converted to age by using the growth equation developed
for E. cavolini:

t is colony age (in years) and Nmax is the largest colony at the study sites (Velimirov,
1975). The maximum TBL of a colony was 278 cm at the investigated study sites.
An attempt was made to test the growth equation for the investigated study site. In
1990, the TBL of 38 colonies was determined in the channel and at the boulders by
measuring each branch length separately. In 1991, the measurements were repeated and
the growth increment was recorded. Unfortunately 14 of the 38 colonies died during
the investigation period. All colonies older than age class 4 (1990) were completely
overgrown with epibionts or were torn off in 1991. Thus data on growth of older
colonies obtained from the channel and the wall during an earlier study (Velimirov &
Weinbauer, 1992) and from non-fan-like growth forms were included. The TBL of all
these colonies was determined by measuring each branch length separately, and this
measurement was repeated in the next year. Since it is not possible to determine the age
of colonies without information on their growth, the following assumption was made. If
the growth equation from the earlier study is also true for the investigated area, then (1)
the age of a colony can be inferred from the size, and (2) the growth and thus size
increment in the following year can be predicted. Thus, if the predicted size does not
differ from the size actually measured in the following year, then the application of the
growth equation was correct and the growth equation can be used to estimate the age of
the colonies.

586

M. G. Weinbauer & B. Velimirov

T 1. Population density and overgrowth of Eunicella cavolini, and depth ranges and
relative light intensity at the various study sites
Density (colonies m "2)

Study
site

Depth
(m)

Relative
light
intensity

Overgrowth

1989

1990

1991

Channel
Wall
Boulders

2327
1628
1015

10
37
49

Low
Intermediate
Strong

159
66
19

180
63
18

149
54
15

Relative light intensity is given as proportion of the light intensity of the least
illuminated study site (channel). Ranking of study sites by overgrowth is based on
Figure 7.

Light intensity
On several occasions, the light intensity was determined at two positions at each of the
various study sites by using a LI-COR Model 185 equipped with an irradiance probe.
Light intensity was determined between 1130h and 1230h at the three study sites and
measured by placing the sensor towards the surface, the bottom, the right and left, and
horizontally towards the water column. This multiple measurement was performed since
light intensity at the study site is not only influenced by the amount of light transmitted
vertically through the water column, but also by reflection from the bottom or by shading
due to overhanging rocks. By calculating mean values from the different orientations, it
was possible to rank the study sites by light intensity.
Overgrowth
After the RFSA was determined, the area of the fan in which overgrown branches could
be observed was estimated. This area was expressed as percentage of the RFSA and
termed overgrowth . When the TBL of colonies was determined directly, the total
length of the overgrown branches was measured simultaneously and expressed as
percentage. Thus, an overgrowth of 50% means that 50% of the branches of a colony are
overgrown by epibionts. Colonies with 100% overgrowth, i.e. dead colonies, were
recorded but not included in the calculation of the mean overgrowth. The mean
overgrowth was calculated separately for various age classes (1, 2, 3, 4 and >4 years).
The dominant epibionts (if present) on the single colonies were also recorded.
Results
Characteristics and population density of study sites
Data on the characteristics and population density of study sites are summarized in
Table 1. Study sites were ranked according to the light intensity as follows: low
(channel), intermediate (wall), high (boulders). Population density was related to light
intensity; highest densities were found at the lowest light intensities. Highest gorgonian
density was found in the channel (149180 colonies m "2) and the lowest density at the
boulders (1519 colonies m "2), whereas the population density was intermediate at the
wall (5466 colonies m "2). Since density varied by 27% in the channel, 28% at the wall
and 31% at the boulders over all investigated years, the density can be considered as
comparatively stable.

587

Gorgonian population dynamics

T 2. Annual recruitment and mortality of Eunicella cavolini at the various study

sites from 1989 to 1991
Channel

Recruits
Deaths
Total

Wall

1989

1990

1991

316

+72
"29
359

+35
"96
298

Boulders

1989

1990

1991

289

+34
"40
283

+18
"57
244

1989

1990

1991

31

+2
"3
30

+2
"8
24

Data are given as total investigated colony numbers. Recruits are given as number of
new colonies, and deaths as number of colonies that could not be found in the
subsequent year.
T 3. Colony numbers, recruitment, death and overgrowth of Eunicella cavolini at
two additional sites at the boulders

Site
1
2

Age
(years)
04
>4
04
>4

Total colony numbers

1990

1991

9
20
4
20

7
20
4
18

Recruits
1991

Deaths
1991

2
2
0
2

Overgrowth (%)
1990

1991

198
250
n.d.
n.d.

178
283
n.d.
n.d.

n.d., not determined. Recruits are given as number of new colonies, and deaths as
number of colonies that could not be found in the subsequent year.

Recruitment and mortality

The number of recruits was equal or even higher in 1990 than in 1991, whereas the
number of deaths was lower in 1990 than in 1991 at all study sites lower in 1990 than
in 1991 (Table 2). Thus there was a slight decrease in the total numbers of colonies from
1990 to 1991 in all sub-populations. Also, at two additional sampling sites at the
boulders, investigated from 1990 to 1991, more deaths than recruitment occurred
(Table 3). Expressed as percentage of new colonies on total colony numbers, recruitment varied between 117 and 201% in the channel, 7412% at the wall and 6783%
at the boulders (Table 2). Mortality, expressed as percentage of colonies that died within
1 year of investigation, varied between 92 and 267% in the channel, 134201% at the
wall and 97267% at the boulders. At the additional study sites at the boulders,
recruitment varied between 0 and 74%, and deaths between 83 and 138% (Table 3).
Determination of total branch length
The non-linear regression between RFSA and TBL of E. cavolini showed a slightly better
curve fit (R2 =09) than the linear regression (R2 =0876), especially for small colonies
(Figure 2). As small size classes are more abundant and more dynamic members of the
population than larger ones (Velimirov & Weinbauer, 1992; this study), non-linear
regression was used for the calculation of the TBL from the RFSA. The TBL of any fan
was obtained by using the regression equation: TBL (cm)= "230+152 RFSA
(cm2)"00017 RFSA2.

588

M. G. Weinbauer & B. Velimirov

400
2

Total branch length (cm)

y = 2.30 + 1.52x 0.0017x

R 2 = 0.900
300

200

100
y = 19.04 + 1.02x
2
R = 0.876

100

200
RFSA (cm2)

300

400

Figure 2. Correlation of the rectangularized fan surface area (RFSA) with the total
branch length in Eunicella cavolini.

Testing the growth equation

The growth curve derived from an earlier study (Velimirov, 1975) was compared with
the growth of colonies from all investigated study sites (Figure 3). Since the observed size
of colonies was close to the expected size in all age classes (Figure 3), it can be assumed
that the growth equation can be used for all investigated study sites to convert total
branch length into age.
Age frequency distribution
The age frequency distributions showed that the most abundant age classes were 14 at
the channel (585&97%), 24 at the wall (554&142%) and 58 at the boulders
(669&725%; Figures 46). At the additional sampling site at the boulders, most of the
colonies belonged to the size class >4 years (Table 3). Although the age frequency
distribution was very similar between the wall and the channel, more older colonies were
present in the channel (Figures 4,5). The oldest colonies were estimated to be 21 years.
Overgrowth
Figure 7 shows that the overgrowth generally increases with age until size class 4. In
colonies older than 4 years, the degree of overgrowth generally did not increase at the
wall and the boulders; in the channel, overgrowth increased slightly from age class 4 to
colonies >4 years. In most size classes, colonies showed the lowest degree of overgrowth
in 1989. When averaging the overgrowth for the age class 4 years (age classes with the
strongest overgrowth), the boulder colonies showed the highest degree of overgrowth
(28%), the channel colonies the lowest (13%), and the wall colonies an intermediate
value (20%). ANOVA showed that these differences were significant (F=536; P<005,

589

Gorgonian population dynamics

250

Total branch length (cm)

200

150

100

50

6
7
Age (years)

10

11

12

Figure 3. Testing the growth equation for Eunicella cavolini. The growth curve was
derived from Velimirov (1975) and compared with data on colonies from the channel,
the wall and the boulders, and three colonies investigated in the channel and at the wall
during an earlier study (Velimirov & Weinbauer, 1992). One colony was omitted from
this earlier study, since it was strongly affected by overgrowth. Note that the size of
1-year-old colonies was derived from the recruits, i.e. from new colonies. Error bars
indicate standard deviations. , Growth equation; -, Velimirov and Weinbauer
(1992); ,, channel; /, wall; 0, boulders.

post hoc P<005). Moreover, at the additional study sites at the boulders, investigated
from 1990 to 1991, a 2528% overgrowth of colonies >4 years was found. Younger
colonies had an overgrowth of 1820% (Table 3).
Discussion
The age of living gorgonian colonies can be estimated by colony height, stem diameter
or total branch length (Grigg, 1974; Velimirov, 1975; Lasker, 1990; Mistri &
Ceccherelli, 1993). Mistri and Ceccherelli (1994) showed that stem diameter of fans and
colony height are better correlated to age than total branch length in the Mediterranean
gorgonian Lophogorgia ceratophyta L. However, since stem thickness of E. cavolini
increases with the intensity of water flow (Velimirov, 1976), this parameter might result
in a bias of the age composition. Colony height would seriously underestimate age in
colonies that are composed of two fans (typical for the second critical depth; Velimirov,
1976) or in bush or dish forms, since growth in width is important in these colonies. The
observed size (total branch length) of E. cavolini colonies matched the expected size at all
sites when applying the growth formula (Figure 3; Velimirov & Weinbauer, 1992). This
showed that total branch length and the growth equation developed for E. cavolini can be
used for the estimation of colony age in this environment.
Differences in age structure (Figures 46) and population density (Table 1) between
study sites comparable to those found for E. cavolini are also known for other octocorals

590

M. G. Weinbauer & B. Velimirov

30
30
25
20
20
15
15
10
10

Age frequency (%)

Age frequency (%)

25

5
5
0
0
1

1989
3

7
9
Age
11
(yea 13 15
rs)
17

1990
19

1991
21

Figure 4. Agefrequency distribution of Eunicella cavolini in the channel for 1989, 1990
and 1991.

(e.g. Birkeland, 1974; Grigg, 1975; Farrant, 1987; Gotelli, 1988; Mitchell et al.,
1993). The fact that mortality was roughly the same at all study sites (83267%),
whereas recruitment was highest at the channel (117201%), intermediate at the wall
(74120%) and lowest at the boulders (083%), indicates that larval settlement and
survival of recruits might be important for the differences of population density
between study sites. Moreover, since reproduction is probably a function of age (or
size) (Wahle, 1983), the absence of large colonies (>10 year; Figure 6) and the low
population density at the boulders (Table 1) might result in low reproduction. Since
fertilization is unlikely at high turbulences (Denny & Shibata, 1989), shallow colonies
on boulders (Table 1) may have lower reproductive success and rely on imported
larvae from other sites compared to populations at greater depths (channel and wall).
In addition, the reproductive success of the strongly overgrown colonies at the
boulders (Figure 7) might be low, since the production of gonads is reduced in
colonies that are strongly affected by overgrowth (Prager & Velimirov, in prep.). Thus,
differences of reproduction could also explain the differences of population density
between study sites.
In the channel, the youngest colonies were the most abundant (Figure 4), indicating
that recruitment and mortality were balanced. At the two other sites, age structure was
skewed to older colonies, since the most abundant colonies were found in age classes
24 years (wall) and 58 years (boulders; Figures 5, 6). Production of larvae or
survival of recruits might differ strongly between subsequent years, and age classes
with numerous colonies might represent years of high larval production or high

591

Gorgonian population dynamics

30
30
25
20
20
15
15
10
10

Age frequency (%)

Age frequency (%)

25

5
5
0
0
1

1989
3

7
9
Age
11
(yea 13 15
rs)
17

1990
19

1991
21

Figure 5. Agefrequency distribution of Eunicella cavolini at the wall for 1989, 1990
and 1991.

survival rates. In Briareum asbestinum Pallas, it was shown that reproductive success
was low in a year when colonies showed a low growth rate (Brazeau & Lasker, 1992).
Thus, larval production may differ between years within a habitat, if factors such as
overgrowth change inter-annually (compare Figure 7), since a significant amount of
energy might be directed to reparative growth instead of reproduction. Moreover,
Grigg (1975) suggested that small changes in larval mortality of gorgonians would
cause large changes in recruitment due to high intrinsic rates of natural increase.
This could also be a reason for the observed differences of age structure between
study sites.
The most important reasons for gorgonian mortality are strong water movements
causing abrasion or detachment, sedimentation, overgrowth and predation (e.g. Kinzie,
1973; Birkeland, 1974; Grigg, 1977). Sedimentation is unlikely to cause mortality of
E. cavolini at the investigated sites, since colonies grow on vertical walls and thus are
protected from burial by sediments. Predation by invertebrates, such as ovulid snails and
amphipods (Salvini-Plawen, 1972), does not seem to be very important for the mortality
of E. cavolini, since signs of snail feeding could be detected in only a few colonies, and
other predators were never observed. Feeding on gorgonians by fish has not been
reported from the Mediterranean Sea. Moreover, it has been suggested that grazing is
probably more important in terms of tissue removal than in terms of mortality
(Birkeland, 1974). On the other hand, damage caused by predation might provide sites
where epibionts could begin to grow.

592

M. G. Weinbauer & B. Velimirov

30
30
25
20
20
15
15
10
10

Age frequency (%)

Age frequency (%)

25

5
5
0
0
1

1989
3

7
9
Age
11
(yea 13 15
rs)
17

1990
19

1991
21

Figure 6. Agefrequency distribution of Eunicella cavolini at the boulders for 1989,

1990 and 1991.

Although no attempt was made to determine quantitatively which group of organisms

was responsible for overgrowth, it was obvious that algae were the dominant epibionts on
the colonies. The high frequency of overgrowth in colonies 4 years (Figure 7) was
possibly a reason for gorgonian mortality at the study sites. In all investigated years and
at all the study sites, completely overgrown axial skeletons were found.
Detachment by weakening of substratum due to boring organisms is a major cause for
gorgonian mortality (Kinzie, 1973; Birkeland, 1974; Yoshioka & Yoshioka, 1991).
However, detachment of colonies from the substratum is more likely in this habitat,
since colonies of E. cavolini grow on solid rock. Colonies of E. cavolini respond to
exceptionally strong current velocities simply by bending over and periodically keeping
the fans parallel to the water flow (Velimirov, 1973). Overgrowth might not only cause
mortality by completely overgrowing colonies; overgrowth might also affect the flexibility
of colonies (Jeyasuria & Lewis, 1987; Esford & Lewis, 1990) thus facilitating detachment, or deterring optimal feeding currents (Sponaugle, 1991; Sponaugle & LaBarbera,
1991) thus decreasing fitness of colonies. Of the 38 colonies tagged in 1990, three were
completely overgrown and 11 could not be found. This indicates that detachment (about
80% of mortality) is more important than overgrowth (about 20% of mortality).
However, it is not known whether some colonies were overgrown before they were
detached or whether overgrowth caused detachment.
In the present study, the population parameters of E. cavolini were investigated in
different habitats. The study sites differed strongly with respect to population density,
recruitment, and overgrowth of the colonies. The age structure showed strong

593

Gorgonian population dynamics

30

(a)

20

10

0
30

>4

>4

3
Age (years)

>4

(b)

Overgrowth (%)

20

10

0
50

(c)

40

30

20

10

Figure 7. Age-specific overgrowth of Eunicella cavolini at the various study sites for
1989 (open bar), 1990 (stippled bar) and 1991 (solid bar). (a) Channel, (b) wall,
(c) boulders.

differences between habitats, but was comparatively stable within a habitat over the time
period investigated. Overgrowth and detachment were identified as the most important
causes for mortality of E. cavolini.

594

M. G. Weinbauer & B. Velimirov

Acknowledgements
This work was financed by G. Bonifacio (President of Stareso) and Stareso SA.
Furthermore, we thank A. Thiery, F. Wernerus and D. Bay for boat and diving
assistance. We are grateful to B. Riegl and three anonymous reviewers for valuable
comments on the manuscript.
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