Professional Documents
Culture Documents
Population Dynamics Growth and Energetics of Jelly
Population Dynamics Growth and Energetics of Jelly
Population Dynamics Growth and Energetics of Jelly
net/publication/250215444
CITATIONS READS
144 236
3 authors, including:
Some of the authors of this publication are also working on these related projects:
All content following this page was uploaded by Hans Ulrik Riisgård on 04 June 2014.
ABSTRACT Seasonal abundance and growth of the scyphomedusa Aurelia a u n t a were studied during
1991 and 1992 In the Inner part of a small shallow fiord, Kertinge Nor, Denmark U u n n g both years,
mass occurrence of A a u n t a was observed, and a maximum value of ca 300 ind m-3 was measured In
Apnl 1992 Growth was poor in the fiord and the maximum mean umblella diameter of medusae was
only 54 i 12 mm a n d 37 + 15 mm In 1991 and 1992 respectively Growth, clearance, lngestlon and
resplratlon were measured at different prey concentrations in the laboratory, a n d a n energy budget
was estabhshed Ephyrae (4 mm in diameter] were offered rotiferas (Brachjonus plicatllls) in concen-
'
trations ranging from 7 to 13000 ind I Clearance of ephyrae decreased 30 to 5 0 % with increasing
prey concentration, and a curved relationship between prey concentration a n d lngestlon w a s found
' ',
A maximum speclflc growth rate of 0 2 d was measured In the laboratory at 400 B plicatilis 1 corre-
spondlng to 60 pg C 1.' In Kertinge Nor, the growth rate never exceeded 0 1 d ' The average
'
zooplankton concentration In Kertinge Nor was only 5 p g C I and it could therefore b e concluded that
A aunta was food hmited in the fiord
O Inter-Research 1994
mar. Ecol. Prog. Ser. 105: 9-18, 1994
MATERIALS AND METHODS In order to measure the mean size of the popula-
tion, live scyphomedusae caught in Kertinge Nor
Study area and sampling site. All sampling of Aure- were measured within 2 h after sampling. Jellyfish
lia aurita was done in Kertinge Nor and Kerteminde were placed with the dorsal side down to flatten out
Fjord, a shallow fjord located on the east coast of the umbrella, and the interradial diameter was
the island of Fyn, Denmark (Fig. 1). The inner part, measured. The arithmetical means and standard
Kertinge Nor, covers an area of 5.8 km2 and has an deviations were calculated. The biomass of Aurelia
average depth of 2 m and a maximum depth of 3 m. aurita medusae in Kertinge Nor was estimated from
Mean residence time of water is approximately 2 mo. the biometric relationships in medusae ranging in
Salinity varies between 14 and 22%0 and water tem- umbrella diameter (D, mm) from 10 to 50 mm. Wet
perature between about 2°C (February) and 22°C weight (Ww,,, mg), dry weight (Wdry= W mg), and
(July). The outer located Kerteminde Fjord (2.8 km2) ash-free dry weight (AFDW, mg) were determined
has a maximum depth of 8 m and a mean residence as a function of bell diameter (Fig. 2) as described
time of water of 1 to 2 wk. Variations in salinity and by Bamstedt (1990). The carbon content of the
temperature are the same as those for Kertinge Nor. medusae was calculated from the dry weight carbon
Population dynamics. Medusae of Aurelia aurita relation given by Schneider (1988): 1 mg dry weight
were sampled approximately every 2 wk from June to = 0.051 mg C for medusae > 2 cm and 0.071 mg C for
December 1991 and from February to September 1992 medusae < 2 cm.
at 7 stations in Kertinge Nor and at 2 stations in Kerte- The instantaneous specific growth rate (p, d-') of
minde Fjord (Fig. 1). Sampling was done using a 0.6 m Aurelia aunta in Kertinge Nor was calculated from the
diameter plankton net with a mesh size of 500 p m The formula:
net was equipped with a 0.5 1cod end and the medusae
p = [In (W,/%)]t-' (1)
were sampled in the daytime by hauling the net at a
fixed depth of 1 m below the water surface. During where and = mean body mass (dry wt) of
sampling, the speed of the boat was 2 to 3 knots, and medusae on Day 0 and Day t respectively. From
the amount of water passing through the net was February to May when new ephyrae were produced
measured with a 'Hydrobios flowmeter'. The volume of continuously, U, was calculated from the dry weight of
water filtered ranged from 3 to 10 m3 at each sampling the largest medusa present on each sampling date.
station depending on the density of medusae. Current Medusae dry weights were calculated from the regres-
measurements inside and outside the net demonstrated sion of umbrella diameter versus dry weight (Fig. 2)
that there was no significant clogging or overflow from given by the equation:
the frontal opening of the net. The abundance of
medusae at each station was estimated as the number
of individuals caught divided by the volume of water Water samples (9.9 1) for zooplankton analysis were
filtered. collected every 2 wk on 2 stations in Kertinge Nor
during 1991 and 1992 by using a 3.3 1 Heart Valve
Water Sampler. Water from 0.5, 1 and 1.5 m below
the surface was pooled and filtered through a 20 pm
plankton gauze. Retained organisms were preserved
in Lug01 solution for later analysis.
l0
RESULTS 0
J F M A M J J A S O N
Month
Population dynamics
Fig. 3. (A) Aurelia aunta. Population density of jellyfish col-
The first ephyrae of the new generation of Aurelia lected at Stns 1 to 7 in Kertinge Nor in 1991 and 1992. Mean
aurita in Kertinge Nor and Kerteminde Fjord ap- population densities f SD and biomass of jellyfish in the
whole fjord system are shown in Table 1. (B) Mean diameter
peared in February (Fig. 3A, Table 1). During the of jellyfish at Stns 1 to 7 in 1991 and 1992. Representative
following months the number increased remarkably SD is shown for 2 dates (C) Instantaneous specific growth
in both 1991 and 1992, and A. aunta obtained a rate of jellyfish at Stns 1 to 7 in 1991 and 1992. (D) Biomass of
maximum density of ca 300 ind. m-3 in April 1992 zooplankton in Kertinge Nor 1991 and 1992
(Fig. 3A). In 1991, the number of medusae decreased
from June and throughout the rest of the season, The growth pattern of Aurelia aunta in Kertinge
while the density in 1992 decreased from late April. Nor was different in 1991 compared with 1992
On all sampling dates, the density of medusae in (Fig. 3B, C). During 1991, the growth rate was poor
Kertinge Nor (Stns 1 to 7) was higher than in Kerte- until early August, when a sudden increase in both
minde Fjord (Stns 8 & 9) (Table 1). diameter and specific growth rate was observed
Olesen et al.: Dynamics and growth of Aurelia aunta 13
Table 1. Aurelia aul-ita. Mean population density f SD and 1992. A close relationship between zooplankton bio-
biomass of jellyfish in Kertinge Nor (Stns 1 10 7) and Kerte- mass and growth of Aurelja aurjta was observed
minde Fjord (Stns 8 & 9, see Fig. 1) in 1991 and 1992. The Fig. 3B, C, D). The maximum in zooplankton
mean population density in Kertinge Nor in 1991 and 1992 is
shown in Fig. 3 biomass in July 1991 (Fig. 3D) was followed by an
increased growth of A. aurita (Fig. 3B, C). Similarly, a
Date Density Biomass minor increase in zooplankton biomass during May
(ind. m-3) c 1-l) 1992 resulted in increased growth rates of the jellyfish
Stns 1 to 7 Stn 8 Stn 9 Stns 1 to 7 (Fig. 3C).
Data from gut content analysis are shown in Table 2.
1991
+
248 292 170.0
Although only a small number of medusae were ex-
3 Jun - -
17 Jun 231 f 136 14 8 171.2 amined, it is evident that rotifers made u p a major part
1 Jul 107 f 127 6 0 93.4 of Aurelia aurita's food on 11 March 1992 and 10 June
15 Jul 81 f 73 <1 0 74.3 1992.
29 Jul +
55 19 7 0 71.2 The biomass of Aurelia aurita in Kertinge Nor
22 Aug 38 f 27 <l 0 251.2
2 Sep 45 f 26 22 0 351.0 obtained a maximum of 350 and 450 yg C 1-' in Sep-
19 Sep 29 + 17 0 0 133.4 tember 1991 and June 1992 respectively (Table 1 ) .The
8 Oct 19 i 16 4 <l 55.2 carbon biomass of A. aurita exceeded that of the zoo-
22 Oct 10 f 12 <l <l 25.5 plankton in Kertinge Nor during most of the year, in
27 Nov 2 f l 0 <l 2.3
some periods even by several orders of magnitude
1992
<1
(compare Fig. 3D, Table 1).
13 Feb 1 f 0.3 1 <0.01
24 Feb 5f4 3 0 <0.01
11 Mar 79 f 29 4 2 0.2
26 Mar 169f104 71 23 0.9 Laboratory experiments
10 Apr 186+87 37 5 3.9
23 Apr 304 f 129 55 13 29.7
11 May +
216 125 2 2 67.6 Feeding and growth
26 May 245 f 251 118 46 118.4
l 0 Jun +
74 37 30 2 109.5 Small medusae (D, = 4 mm) showed rapid growth
25 Jun 1 3 4 f 110 46 <l 450.0 when rotifers were offered in the concentration range
19 Jul 41 k 21 1 1 122.8
20 Aug 39 f 22 1 0 73.8 130 to 13 000 ind. 1-' (Fig. 4A. Table 3). During the 10 d
24 Sep 22 f 27 1 0 24.2 incubation period, the umbrella diameter increased
from approximately 4 to 9 mm at the highest prey
concentrations, corresponding to a n increase in dry
(Fig. 3B, C). A maximum in the umbrella diameter weight from 0.1 to 0.9 nlg (Table 3). The weight
was observed in early September 1991 where a increase was slightly lower with lower concentrations
mean diameter of only 54 mm was measured of prey organisms. Medusae with an initial diameter
(Fig. 3B). In late August 1991,
reached a nlaxlmum of 0.07 d-'. In
1992, a maximum of p = 0.09 d-' was Table 2. Aurelia aurita. Average number of prey items f SD in gut content
found in early J~~~ ( ~ i 3~ ~. with ) a of ~ n e d u s a efrom Kert~ngeNor, sampled in 1992 during daytime, and on
16 October 1991 at various times during a 24 h period. n : no. of medusae
maximum mean diameter of 37 mm examined. D: mean diameter f SD of medusae
in late June (Fig. 3B).
The following zooplankton groups
Date Rotifer Tintin- Gastropoda Harpac- Polychaeta b n
were found in situ [mentioned in order nids larvae ticoida larvae (mm)
of their contribution ( % ) to the average
total zooplankton biomass through 1991 1991
and 19921: Rotatona (54.6) > Harpacti- 16 Oct
08:OO h - - - 5.8 f 4 - 34 + 12 10
coida (16.2) > Chaetopoda (14.1) > 13:30 h - - - <1 - 30k11 25
Calanoida (3.6) > Gastropoda (3.2) > 16:45 h - - - 3.3 f 2 - 28 k 10 10
Lamellibranchia (0.15) > Cirripedia (2.1) 21:45 h - - - 2.6 f 3 - 31 k 13 10
> Cladocera (1.9) > Cyclopoida (1.3). 1992
Total biomass of zooplankton in Ker- llMar 14+1 122 0 0 0 2.5 f 0.6 17
tinge Nor is shown in Fig. 3D. During 3 A p r 1 + 1 1 1 2 5 2 + 2 <1 0 3.5 i 0.8 6
llMay 2 + 4 2+2 I f 1 <1 <l 13.2 i 4 9
the 2 yr, the average zooplankton 10Jun 8 i 7 1+1 4f3 1+2 0 17.9 f 5 8
biomass was low with a mean value of 19 Jul <1 <1 I f 1 3f2 <l 29.2f1 5
6.5 yg C 1-' in 1991 and 4.7 ~g C 1-' in
Mar. Ecol. Prog. Ser. 105: 9-18, 1994
Table 3 Atirelia aurita. Prey organism (Branchionus pl~catilis]concentrations (C) in growth experiments of t days duration with
-dry weight (W,),
2 size groups of jellyfish with initial umbrella diameter (Do= 4 or 10 mm), initial body and final size D, and
on Day t; all values are mean f SD; p: instantaneous specific growth rate, [In (W,/W,) t-l; R,: individual respiration rate
measured on Day t; R,,: weight-specific respiration rate, R,/wf;NGE: net growth efficiency = p/(p + R,,)
-
C t 4 61 WO F P R1 R, NGE
(in l ) (d) (mm) (mm) (mg) (mg) (d-l) (pm01 Oz d-' (J d-' mg-l) (%)
rnedusa)
DISCUSSION
T
10 - Population dynamics
v v v
C
-E 8 - 0 0 0
0 The abundance of Aurelia aurita in Kertinge Nor was
E 8 extremely high during 1991 and 1992. The average
.-
k
a,
6 -
v
O + + + + +
+ density was ca 220 ind, m - 3 from late March to late
May 1992 with a maximum density of approximately
.- 5 C 13
0 4 - &
2 -
,
, P g 300 ~ n d m-"Fig.
. 2A), which is almost 7 times hlgher
than the maximum density reported in the literature:
10 ephyrae m-3 in Urazoko Bay, Japan (Yasuda 1968);
15 ephyrae m - 3 in the Gullmar Fjord, Sweden (Hern-
0 -- A roth & Groendahl 1983) and 44 ephyrae m - 3 in Elefsis
B Bay, Greece (Papathanassiou et al. 1987). The reason
for the large number of medusae in Kertinge Nor is not
;
R 0.2 -
-9
a, fr
l
clear, but the relatively protected fjord may support
high densities of scyphistomae. Groendahl(1988)found
evidence that scyphistomae of A. aurita are very rare
outside protected fjords and lagoons. The abundance
9
of scyphistomae on ceramic settling plates in the inner
U
O, "'-1 part of Kertinge Nor was estimated to be approxi-
-
.-
.
U
0.0 -
,
1 mately 316000 m-' in mid-September 1992, while the
a number was significantly lower in the outer fjord
m
(56 000 m-') (pers. obs.).The higher density of scyphis-
tomae and medusae (Table 1) in the inner part of the
n fjord, and the small individual size of medusae seem to
vI
c I
Indicate a local homogeneous population.
-
I
25 - New ephyrae emerged from February to April in 1991
and 1992 (Fig. 3). A late winter, early spring liberation of
U A
- 20-
A ephyra has been described by many authors (Palmen
A
c A 1945, Moller 1980a, Van der Veer 1985, Papathanassiou
-
E 1987).In contrast, Hernroth & Groendahl(1985) reported
1 5 - 0 0
a,
U 0
that the main period of ephyrae release in the Gullmar
C
Fjord, Sweden, was from October to November. Newly
10-
-a, released ephyrae sank rapidly towards deeper water
0 8 where they wintered in a diapause-like state before as-
5 -
cending towards the surface layers in April. We have no
evidence of such a temporal linkage between release
0 A
and appearance of ephyrae, but a new autumn genera-
0 500 1000 l500 V 3 6000
~ ~9000~ 12000
3000
tion seems unlikely in Kertinge Nor, as no ephyrae were
n found in the autumn in spite of well wind-mixed water.
The decreasing number of medusae during summer tion of approximately 400 Brac.hoionus plicatilis 1-'
in Kertinge Nor (Fig. 3A) may be explained by outdrift and above (Fig. 4B), which corresponds to 60 pg C
of medusae to the open sea. In both years, a positive 1-l On the average, the carbon concentration of zoo-
growth was observed In periods of decreasing num- plankton in Kertinge Nor was only 6.5 and 4.7 pg C
bers, and mortality due to food limitation seems 1-' in 1991 and 1992 respectively (Fig. 3D), and the
therefore to be an unlikely explan.ation during those growth of 4 to 9 mm medusae in Kertinge Nor was
periods. Further, mortality in Aurelia aurita is usually therefore severely food limited. However, this con-
low before maturation (Moller 1980a). clusion can also be extended to other size medusae
in Kertinge Nor by calculating the lowest prey con-
centration, different sized medusae need to attain
Growth rates and energetics maximal growth: using the respiration versus body
mass relationship found for Aurelia aurita by Larson
The growth of Aurelia aurita in 1991 and 1992 was (1986), and assuming a NGE of 35% (Fraser 1969,
very poor in Kertinge Nor (Fig. 3). The specific this study), and an assimilation efficiency of 0.9
growth rate never exceeded 0.1 d-l, and the final (Purcell 1983), the ingestion for a 20 mm medusae
umbrella diameter was only 54 and 37 mm in 1991 is found to be 2.1 J d - ' medusa-' corresponding to
and 1992 respectively (Fig. 3B). These figures are 320 B. plicatilis d-l medusa-l. Clearance of a 20 mm
considerably lower when compared to values ob- medusa feeding on B. plicatilis = 2364 m1 d-l
tained in similar studies. Thus, Moller (1980a) found medusa-I (Olesen unpubl.), and hence the prey con-
a monthly increase of 41 mm in the diameter, corre- centration needed for maximal growth is 135 B. pli-
sponding to a specific growth rate of 0.18 d - ' in an catilis 1-' corresponding to 20 pg C 1-l. The value for
A. aurita population in Kiel Bight (Germany), where a 30 mm medusa is 1 4 pg C 1 - l . These values are
the medusae obtained a maximum size of 197 mm. likely to be underestimated because the respiration
The growth was even more pronounced in Gullmar rate used in the calculations was not obtained from
Fjord, Sweden, where a specific growth rate of jellyfish growing at maximal rate. In conclusion,
0.23 d-l was reported by Hernroth & Groendahl there is little doubt that A. aurita is food limited most
(1983) and in the Western Wadden Sea, where a of the season in Kertinge Nor, and this explains the
maximum specific growth rate of 0.2 d - ' was calcu- poor growth observed.
lated from data of Van der Veer (1985) The maximum growth rate obtained in laboratory
In the laboratory experiment, the specific growth experiments (Fig. 4B) is close to the maximum
rate was determined only for jellyfish with initial sizes growth rate observed in natural populations in other
of 4 and 10 mm, while the growth rate in Kertinge Nor systems. In nature, however, this growth rate is
was obtained for jellyfish of varying sizes (2 to 54 mm). generally only obtained for short periods, and food
It is, however, valid to compare the maximal growth limitation of Aurelia aurita may therefore be preva-
rate found in the laboratory (0.22 d-l) directly with the lent in nature.
maximal growth rate of 0.09 d - ' estimated in Kertinge Fraser (1969) calculated a net growth efficiency of
Nor (Fig. 3 C ) , if the specific growth rate of Aurelia 37 % in Aurelia aurita, which is close to the value of
aurita is constant with body mass. A constant instanta- 35% obtained in this study. From the energy budget
neous growth rate at increasing body mass depends established in the present work it was calculated that
upon a linear relationship between body mass (Wdry) respiration constituted about 66% of assimilation,
and metabolic rate (M),that is b = 1 in the allometric and production constituted 34 % of assimilation at a
relationship M = aWdrYb(see e.g. Jmgensen 1990). maximum growth rate. In Kertinge Nor, however,
Larson (1987) measured respiration rate as a function growth rdte of A . aurita was less than 0.1 d-l dunng
of body mass in 10 different hydromedusae species 1991 and 1992 (Fig 3C), and NGE therefore never
and 2 scyphomedusae species, and found that most exceeded 24 % (Table 3), indicating that the medusae
b-values were not significantly different from 1.0, at the low food conditions channelled more than
and recently Uye (Hiroshima University, pers. comm.) 76% of the assimilated food energy into respiration
found a h-value = 1.0, over a wide size range of A. and less than 24 % into somatic growth. During 1982
aurita medusae. Thus, it may be assumed that labora- when the abundance of medusae was high in the
tory growth data on A . aurita can be compared to Kiel Bight, Schneider (1989) also found that 80% of
growth data from Kertinge Nor, and therefore, it can the assimilated food energy was used for respiration
be concluded that A. aurita does not utilize its growth while 18% was used for somatic growth. These
potential in Kertinge Nor. figures were 61 and 35%, respectively, at a sig-
The maximal growth rate (of 4 to 9 mm medusae) nificantly lower density of medusae and higher prey
in the laboratory was attained at a prey concentra- levels.
Olesen et al.: Dynamics and growth of Aurelia aurita 17
Heinbokel, J . F., Coats, D. W.. Henderson. K. W., Tyler, M. A. Papathanassiou, E., Panayotidis, P., Anagnostaki, K. (1987).
(1988). Reproduction rates and secondary production of Notes on the biology and ecology of the jellyfish Aurelia
three species of the rotifer genus Synchaeta in the estuar- aurita L. in Elefis Bay (Saronikos Gulf, Greece). Mar. Biol.
ine Pontomac River. J. Plankton Res. 10: 659-674 8: 49-58
Hernroth, L.. Groendahl, F. (1983). On the biology of Aurelia Purcell, J. E. (1983).Digestion rates and assimilation efficien-
aurita (L.) 1. Release and growth of Aurelia aurita (L.) cies of siphonophores fed zooplankton prey. Mar. Biol. 73:
ephyrae in the Gullmar Fjord, Western sweden, 257-261
1982-1983. Ophelia 22: 189-199 Riisgard, H. U,,Randlsv, A., Kristensen, P. A. (1980).Rates of
Hernroth, L., Groendahl. F. (1985). On the biology of Aurelia water processing, oxygen consumption and efficiency of
aurita (L.). 2. Major factors regulating the occurrence particle retention in veligers and young post-metamorphic
of ephyrae and young medusae in the Gullmar Fjord, Mytilus edulis. Ophelia 19: 37-47
Western Sweden. Bull. mar. Sci. 37(2): 567-576 Schneider, G. (1988).Chemische Zusarnrnensetzung und Bio-
Jnrgensen, C. B. (1990) Bivalve filter feeding: hydrody- masseparameter der Ohrenqualle Aurelia aurifa. Helgo-
namics, bioenergetics, phys~ologyand ecology. Olsen & lander Meeresunters. 42: 319-327
Olsen, Fredensborg Schneider, G. (1989). Estimation of food demands of Aurelia
Kramp. P. L. (1961). Synopsis of the medusae of the world. aurita medusae populations in the Kiel BighVWestern
J . mar. biol. Ass. U.K. 40: 1-469 Baltic. Ophelia 31(1): 17-27
Krumbach, T. (1925).Syphozoa. Handb. 2001. 1: 522-686 Stoecker, D , Michaels, A. E , Davies, L. H. (1987). Grazing
Larson, R. J. (1987).Respiration and carbon turnover rates of by the jellyf~sh, Aureha aurita, on 'microzooplankton.
medusae from the NE Pacific. Comp. Biochem. Physiol. J. Plankton Res. 9: 901-915
87A1: 93-100 Szyper, J. P. (1989). Nutritional depletion of the aquaculture
Lindahl, O., Hernroth, L. (1983). Phyto-zooplankton commu- feed organisms Euterpina acutifrons, Artemia sp. and Bra-
nity in coastal waters of Western Sweden - an ecosystem chionusplicatilisduring starvation. J. Wrld Aquat. Soc. 20:
off balance? Mar. Ecol. Prog. Ser. 10: 119-126 162-169
Moller, H. (1980a) Population dynamics of Aurelia aurita Van der Veer, H W., Oorthuysen, W. (1985). Abundance,
medusae in Kiel Bight, Germany (FRG). Mar. Biol. 60: growth and food demand of the scyphomedusa Aurelia
123-128 aurita in the Western Wadden Sea. Neth. J . Sea. Res.
Moller, H. (1980b). Scyphomedusae as predators and food 19(1):38-44
competitors of larval fish. Meeresforsch. 28: 90-100 Yasuda, T.(1968). Ecological studies on the jellyfish Aurelia
Palmen, E. (1954). Seasonal occurrence of ephyrae and subse- aurita In Urazoko Bay, Fukui Prefecture. 2. Occurrence
quent instars of Aurelia aurita (L.) in the shallow waters of pattern of the ephyrae. Bull. Jap. Soc. Sci. Fish. 34:
Tvarminne, S. Flnland. Arch. Soc. Vanamo 8: 122-131 983-987
This article was submitted to the editor Manuscript first received: March 14, 1993
Revised version accepted: December 6, 1993