Journal of Experimental Marine Biology and Ecology 436-437 (2012) 5055

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Journal of Experimental Marine Biology and Ecology

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The effects of food and temperature on settlement of Aurelia aurita planula larvae and
subsequent somatic growth
Clare N. Webster , Cathy H. Lucas
Ocean and Earth Science, National Oceanography Centre Southampton, University of Southampton Waterfront Campus, European Way, Southampton, SO14 3ZH, UK

a r t i c l e i n f o a b s t r a c t

Article history: The occurrence and magnitude of coastal scyphozoan jellysh blooms can only be predicted with knowledge
Received 28 February 2012 of how the environment inuences every stage of the life cycle. This study provides the rst information on
Received in revised form 28 August 2012 the effects of food and temperature on the settlement of Aurelia aurita planulae. Planulae from mature medu-
Accepted 29 August 2012
sae collected in southern England (50 48 3.86 N, 1 25 23.06 W) were used for experiments. They were
Available online 23 September 2012
introduced to six different treatments: three temperatures (6 C, 12 C, 18 C) at both high and low food
Keywords:
feeding regimes. Although settling rate was faster at higher temperatures, signicantly fewer planulae settled
Aurelia aurita at the highest temperature. Four types of asexual reproduction were observed in the high food treatments at
Jellysh higher temperatures. Temperature had a signicant positive effect on somatic growth of scyphistomae. Food
Planula larvae had a signicant positive effect on the somatic growth of those with four and more tentacles, and the inter-
Scyphistoma action of the two had a signicant positive effect on those with eight and sixteen tentacles.
Scyphozoa 2012 Elsevier B.V. All rights reserved.
Settlement

1. Introduction
Recent studies have suggested that jellysh populations are on the
rise (Daskalov et al., 2007; Purcell et al., 2007), with the abundance
and distribution of several populations linked to climate cycles
(Lynam et al., 2011). Spatial and temporal distribution of medusae ulti-
mately depends on the location of the scyphistomae and the recruit-
ment of their ephyrae to adult medusae (Decker et al., 2007). Most
work, apart from that on the more well-known medusa stage, has
been experimental work on cultured scyphistomae from aquariums or
collected from the eld. That research predominantly looks at the rela-
tive signicance of temperature and salinity effects on asexual repro-
duction, usually with the consequences for strobilation (Willcox et al.,
2007; Purcell et al., 2009; Han and Uye, 2010) and ephyrae (Widmer,
2005). Exceptions and contradictions can be found in results from
both salinity and temperature studies on Aurelia aurita (Custance,
1967; Olmon and Webb, 1970; Kakinuma, 1975; Willcox et al., 2007).
In some studies food has been found to have a more signicant effect
than temperature or salinity on both the somatic growth and asexual
reproduction rate of scyphozoan polyps (Keen and Gong, 1989;
Stampar et al., 2008), whereas other studies have shown food levels
to be the most signicant inuence on somatic growth (Willcox et al.,
2007, 2008), budding rate (Ma and Purcell, 2005) or strobilation
(Purcell et al., 1999). This is difcult to quantify as perhaps surprisingly,
Corresponding author at: Pelagic Ecology and Research Group, Gatty Marine Labora-
tory, School of Biology, University of St Andrews, Fife, KY16 8LB, Scotland, UK. Tel.: +44
7590689688.
E-mail address: cnw3@st-andrews.ac.uk (C.N. Webster).
food level has received less attention than either temperature or salini-
ty. Other factors that have also been shown to be signicant on scyphis-
toma growth and asexual reproduction are density (Willcox et al.,
2008), predation (Hernroth and Grndahl, 1985; Grndahl, 1988b)
and light intensity (Custance, 1967).
Unlike scyphistoma studies, research on planula settlement is not
dominated by work carried out on Aurelia aurita. So far, studies on
planula settlement in Semeaostomeae have investigated the response
of Cyanea capillata planulae to environmental inuences (Brewer, 1976;
1989; Dolmer and Svane, 1993; Svane and Dolmer, 1995; Holst and
Jarms, 2007; Holst and Jarms, 2010), substrate preferences in Aurelia
aurita, Cyanea lamarckii, Chrysaora hysoscella, Cyanea capillata (Holst
and Jarms, 2007; Hoover and Purcell, 2009) and the effects of bacteria
on the induction of settlement of A. aurita (Kroiher and Berking, 1999).
Importantly, this highlights the dearth of information concerning the en-
vironmental inuences on the rate of planula larvae settlement and
metamorphosis; yet this information is crucial as settlement activity in-
uences the abundance and distribution of scyphistomae and thus
recruitment to the adult medusae population.
This study presents the rst data on the effects of temperature on
A. aurita planula larvae settlement. The aim of this study was to rear
A. aurita from planula larvae in the laboratory exposed to different
temperature and feeding regimes. There are two hypotheses. Firstly,
that temperature rather than food will have a signicant effect on
planula settlement, as measured by time taken to settle and/or settle-
ment success, due to their lecithotrophic nature. Secondly, that both
temperature and food will have a signicant effect on asexual repro-
duction and somatic growth of any subsequent scyphistomae. There-
fore, the null hypotheses are that neither temperature nor food will

0022-0981/$ see front matter 2012 Elsevier B.V. All rights reserved.
http://dx.doi.org/10.1016/j.jembe.2012.08.014
 C.N. Webster, C.H. Lucas / Journal of Experimental Marine Biology and Ecology 436-437 (2012) 5055
have any signicant effects on planulae settlement or subsequent
growth.
2. Material and methods
Thirty mature female medusae of Aurelia aurita with visible planulae
were carefully collected from a large aggregation surrounding the oat-
ing pontoons of Bucklers Hard Marina on the Beaulieu River, which con-
nects to the Solent across from the north western shore of the Isle of
Wight (50 48 3.86 N, 1 25 23.06 W). Concurrent temperature and
salinity measurements were taken using a YSI multiparameter probe.
The medusae were put into individual containers and were returned
to the laboratory within the hour.
Running seawater was repeatedly poured over the gonads, and
planula liberation was further encouraged by the gently rubbing the go-
nads between ngers. Planulae were counted out by the hundred, and a
mixture of planulae from different medusae totalling 1600 was put into
each replicate bowl. Three replicate bowls with a volume of approxi-
mately 0.7 L were used for each of the six treatments: high and low
food at 6, 12 and 18 C. High and low treatments were fed 4.5 ml and
1.5 ml of newly hatched Artemia salina nauplii concentrate at a density
of approximately 1000 ml1 every second day. Bowls were set up with
three glass slides balanced across two wires to allow planulae to settle
underneath (Fig. 1). The glass slides had been preconditioned for ve
days in natural seawater to allow a biolm to develop, which enhances
planula settlement (Schmal, 1985; Holst and Jarms, 2007). The bowls
were placed in six aerated tanks (25 cm 15 cm 18 cm), two of
which were placed into each of the three temperature-controlled
water baths (61 cm 31 29 cm) set at each temperature (1 C).
Black lids were placed over the water baths to maintain a constant
light level and seawater was maintained at a constant salinity of 27.
The glass slides were removed every second day and placed in a petri
dish with seawater for a maximum of 10 min so that numbers of each
growth stage of settled scyphistomae could be counted under a dissec-
tion microscope. Once they reached the eight tentacle stage, pencil
marks were made on the other side of the slide so any movement
could be recorded. Every second day after the slides were counted
50% of the water was changed and dead Artemia salina nauplii were re-
moved with a pipette. Slides were photographed as often as possible
(11 times) before feeding using a digital camera mounted on a stereo-
microscope. Counts from image analysis using SigmaScan software
were used for analysis due to the sizeable human error margin in counts
done by eye. SigmaScan was also used to accurately measure the diam-
eter of the mouth disc (excluding tentacles). Total experimental time
was 64 days.
Fig. 1. Experimental set-up to investigate planulae settlement and subsequent somatic
growth consisted of three replicate bowls at each treatment. Each bowl had three
pre-conditioned slides held off the bottom of the bowl by wires.
51
All data were tested for normality using a ShapiroWilk test and any
signicant differences between treatments were analysed with a
two-way ANOVA in order to address the null hypothesis that tempera-
ture will not have a signicant effect on planula settlement. Days when
asexual reproduction occurred were noted. Gain or loss in numbers
after all settlement had ceased were displayed graphically, and were
analysed with a two-way ANOVA. There are two null hypotheses
concerning somatic growth; H1 temperature has no effect on somatic
growth as measured by mouth disc diameter or tentacle number; and
H2 food has no effect on somatic growth as measured by mouth
disc diameter or tentacle number. Size-frequency of scyphistoma
mouth disc diameters were calculated for each treatment and data
were tested using regression analysis. This was followed by several
two-way ANOVA tests comparing the effect of food and temperature
on the number of scyphistomae with certain numbers of tentacles. All
statistics were carried out by the statistical software SPSS.
3. Results
Temperature of surface water on collection of the mature female
medusae was 19 C, and salinity was 33. The medusa bell diameters av-
eraged 47 0.8 mm. Settlement was gregarious; in one main patch in
each treatment at a density of approximately 150 cm2 with the
remaining few evenly distributed across the remaining area at a density
of 2 cm2. 100% of planula settlement was on the undersides of the
slides. After settlement, scyphistomae rst developed a short stalk and
then two tentacles on opposite sides of the calyx, the mouth and two
more tentacles followed, again on opposite sides. Table 1 shows that
the rst occurrences of particular growth stages varied between treat-
ments. Counts of each growth stage on each day when photos were
taken were also different for each treatment (Fig. 2).
Count data of settled planulae in each treatment (Fig. 3) were nor-
mally distributed (ShapiroWilk; p >0.05). Results of the subsequent
two-way ANOVA showed no evidence of a signicant difference in set-
tling numbers between different food levels (F(1,12) =1.2, P= 0.295)
but a highly signicant difference in settling numbers between the
three different temperatures (F(2,156) = 40.47, P =b 0.001). Planulae
were still present until day 14 at 6 C, day 10 at 12 C and day 5 at
18 C, with no difference in different food levels at each temperature.
The rst null hypothesis was rejected and it was concluded that while
planulae took longer settle at 6 C, greater numbers successfully settled
and metamorphosed at lower temperatures.
Four types of asexual budding were observed: Lateral budding;
lateral budding by means of stolons; reproduction from parts of
stolons/stalks; and longitudinal ssion. These were only observed
in the 16 tentacle growth stages in the high food regime at 12 C
(day 33) and 18 C (day 26); with a 210.08 83% increase in num-
bers in the latter treatment. Note that movement of scyphistomae
was recorded in the two treatments where budding occurred, in-
cluding migration to the inside walls of the experimental bowls.
Table 1
Day of planula larvae observations in six different settlement treatments followed by
the rst appearances of various stages of scyphistosoma growth (4, 8, 16 tentacles).
The six treatments were three different water temperatures, 6 C, 12 C and 18 C,
each with high and low feeding regimes. Note that there were three replicates of
each treatment yet the recorded days of presence/absence did not differ.
Developmental 6 C 6 C 12 C 12 C 18 C 18 C
stage
Low High Low High Low High
food food food food food food
Free swimming 014 014 010 010 04 04
planulae
4-tentacle stage 1064 564 457 364 318 364
8-tentacle stage 64 2664 2664 1064 464
16-tentacle stage 5764 5764 2664 864
52
Number of scyphistomae
Fig. 2. Number of Aurelia aurita scyphistomae reaching each tentacle stage in each feeding regime at the three different temperatures. Scyphistomae with no tentacles (A), two
tentacles (B), four tentacles (C), eight tentacles (D) and sixteen tentacles (E). Counts are from the days pictures were taken. Note that error bars have been removed for legibility.
Space for further movement, migration and asexual reproduction
was still available on the nal day of the experiment (day 64).
All low feeding regimes incurred heavy losses: 6 C, 98.8 11%;
12 C, 98.2 10%; 18 C, 100%. High feeding regimes had lower mor-
tality: 6 C, 37.4 14%; 12 C, 32 17%. It was evident from observa-
tion that dead scyphistomae were succumbing to bacterial
degradation. Note that mortality data of 18 C high food treatments
were obscured by the aforementioned large increase in numbers.
Overall these scyphistomae were of small diameter (Fig. 4). Re-
gression analysis of temperature and mouth disc diameter did not
nd a statistically signicant relationship (r 2 = 0.0276, F = 3.010,
P = 0.086) and the null hypothesis could not be rejected; whereas
analysis of the relationship between food level and mouth disc diam-
eter did nd a signicant positive relationship (r 2 = 0.108, F =
12.812, P = b 0.001) and the null hypothesis was rejected. Somatic
growth was also quantied by counting the number of scyphistomae
at each growth stage as indicated by number of tentacles. Two-way
Final number of settled planula larvae
Fig. 3. The total number of Aurelia aurita planula larvae that settled in each treatment.
Day 14 for 6 C, day 10 for 12 C and day 5 for 18 C. Data did not differ signicantly
between high and low feeding regimes at each temperature.
C.N. Webster, C.H. Lucas / Journal of Experimental Marine Biology and Ecology 436-437 (2012) 5055
A B
C D
ANOVA tests revealed that temperature had a signicant positive ef-
fect on the number of scyphistomae at all growth stages, as indicated
by number of tentacles, and food level signicantly positively affect-
ed numbers of scyphistomae at the four tentacle stage and higher.
Their combined interaction had a signicant positive effect on the
numbers of scyphistomae at the eight and sixteen tentacles stages
only (Table 2). For tentacle number representing somatic growth,
both null hypotheses regarding food and temperature were rejected.
4. Discussion
4.1. Planula collection and settlement
Planula larvae stayed near hard surfaces on the bottom or moved
upwards on the sides of the collecting containers, which is behaviour
also reported by Holst and Jarms (2007) in their study on the planu-
lae of ve scyphozoan species, including Aurelia aurita. They report
that A. aurita planulae had the closest contact with the sides of the
container and so had the least free-swimming planulae compared
with the other four species. We hypothesise that this could be an ad-
aptation to avoid rough seas or predation and Holst and Jarms (2007)
suggest that it could be a strategy to reach particular habitats. This
may be part of the reason for the success of A. aurita in exploiting a
wide variety of environmental conditions at different latitudes
around the world (Vagelli, 2007). Indeed, a pertinent area of future
research would be a comparison of the survival time of A. aurita plan-
ulae with other species to investigate whether survival longevity of
planulae is a reection of their time spent being vulnerable in the
water column. In this study, the planulae maintained at 6 C settled
as late as day 14; any that did not settle were not seen alive after
this day. If Aurelia aurita planulae do not settle after a dened time
period they do not survive. Our ndings suggest that this critical
time period is determined by temperature. The two week survival
time is longer than previously reported for A. aurita. For example,
Schneider and Weisse (1985) concluded they could survive for max-
imum of one week at 19.6-21.5 C). The rhizostome Cotylorhiza
tuberculata from the Mediterranean still had planulae present and
presumably motile after 71 days at 12 C which is the lowest
 C.N. Webster, C.H. Lucas / Journal of Experimental Marine Biology and Ecology 436-437 (2012) 5055 53

A B
60

40

20

C D
60
Percent frequency

40

20

E F
60 60

40 40

20 20

0 0
0.0 0.2 0.4 0.6 0.8 1.0 1.2 0.0 0.2 0.4 0.6 0.8 1.0 1.2
Scyphistoma mouth disc diameter (mm)

Fig. 4. Measured mouth disc diameters of scyphistomae in each of the experimental treatments; 6 C low food (A), 6 C high food (B), 12 C low food (C), 12 C high food (D), 18 C
low food (E), 18 C high food (F). Note that due to differences in time of mortality between treatments these data are independent of time.

temperature found in their natural environment (Prieto et al., 2010).
This highlights a key fact the two week survival at 6 C in this
study is not directly relevant to conditions in the Beaulieu River dur-
ing summer although it does show their robustness and probably the
slowing effect of low temperature on their metabolic rate (Schneider
and Weisse, 1985). At 12 C and 18 C, longevity of the planulae was
similar to those reported by Schneider and Weisse (1985). Brewer
(1978) notes that A. aurita planulae take longer to settle in aerated
water than nonaerated water, so this may also have contributed.
Table 2
Five two-way ANOVAs were carried out on the numbers of scyphistomae at each
growth stage in six experimental treatments. Growth stage was categorised by number
of tentacles. Data from three replicates of each of the six treatments were analysed:
temperature (6 C, 12 C, 18 C) had a signicant effect on the number of all growth
stages and food level (high and low) signicantly affected numbers of growth stages
with four tentacles and higher. Their combined interaction had a signicant effect on
the numbers of eight and sixteen tentacles stages only.
Variable Number of tentacles F d.f.
Temperature 2 5.610
4 7.709
8 21.123
16 6.624
Food 2 0.056
4 18.271
8 39.365
16 21.476
Temperature and food 2 1.631
4 0.625
8 21.243
16 6.664
The duration of planulae survival prior to settlement is in agreement
with Ohtsu et al. (2007) who found that lower temperatures increased
the time taken to settle Nemopilema nomurai planula larvae in the Sea of
Japan. More recent experiments on Cotylorhiza tuberculata (Prieto et al.,
2010) found that higher temperature reduced time to settlement al-
though their numbers remained constant at their three highest temper-
atures and lower at their lowest temperature reecting their natural
environment. For A. aurita, it could be an unknown temperature effect
on the bacterial induction cues within the biolm, an effect on metabo-
lism of the planula (Schneider and Weisse, 1985), a confounding effect
of increased predation by the more developed initial settlers in the
higher temperatures. Time spent in the water column poses major
risks for pelagic larvae in the form of predation, transport away from
suitable substrata, and exposure to extreme environmental conditions.
Lecithotrophic larvae could be considered a lower risk strategy than
planktotrophic larvae as they are independent of food supplies and
have shorter development times (Uthicke et al., 2009).
P
4.2. Asexual reproduction
2 0.008
2 0.002
2 b0.001 Four of the nine types of asexual reproduction described by Adler
2 0.004 and Jarms (2009) were recorded only in fully grown scyphistomae in
1 0.814 the high feeding regimes at 12 and 18 C. Reproduction from parts of
1 b0.001
stolons and reproduction by lateral ssion were only observed six and
1 b0.001
1 b0.001 four times respectively. For purposes of discussion the term budding
2 0.210 will be used to refer to both lateral and stolonic budding which were
2 0.541 commonly observed in both treatments and accounted for the vast ma-
2 b0.001 jority of asexual reproduction. By the 1970s most researchers agreed
2 0.003
budding was not a response to unfavourable conditions (Hofmann et
54 C.N. Webster, C.H. Lucas / Journal of Experimental Marine Biology and Ecology 436-437 (2012) 5055
al., 1978). The agreement from studies on the effect of food levels is that
it increases the occurrence of budding (Ma and Purcell, 2005; Hoover
and Purcell, 2009; Han and Uye, 2010) and that scyphistomae
experiencing low levels of food do not bud even when nutrient enriched
(Grassle and Shick, 1979). On the other hand a study on Chrysaora
quinquecirrha reported food effects on ephyrae but not on clonal growth
(Purcell et al., 1999). However, in this experiment food quantity and
type was varied (copepods, Artemia salina nauplii and natural zooplank-
ton) so it is difcult to make robust comparisons without quantifying
the amount of energy provided to each treatment. At very low levels
of food (3.3 mg carbon per polyp) asexual reproduction resorts to
podocyst formation (Han and Uye, 2010). Podocyst formation was not
observed in our study, either because food levels were not low enough,
or because the polyps were too young to produce podocysts as has been
observed in other species (Widmer, pers. comm.).
Increased budding at increased temperatures has been reported in
many other studies (Ma and Purcell, 2005; Willcox et al., 2007; Holst
et al., 2007; Han and Uye, 2010). In contrast, Liu et al. (2009), report in-
creased Aurelia aurita bud production with decreasing temperature.
This could be because of insufcient food at high temperatures
(30 C). It is also possible that because the lowest temperature
(20 C) reected a natural mid-temperature increased bud production
may not have continued at lower temperatures. Increased bud produc-
tion at higher temperatures could be due to enhanced feeding ability
because of faster tentacles (Elliott and Leggett, 1996) and increased in-
gestion rate (Mangum et al., 1972). This might explain why Purcell
(2007) also reported decreased budding at higher temperatures
(15 C) as it was ensured that feeding time was limited (one to three
hours). It is possible that results of studies reporting signicant effects
of temperature on budding are inuenced by food timings (how long
the scyphistomae have to feed) even when given in equal amounts.
Movement of scyphistomae was evident in the higher temperature,
high food treatments where budding took place. Although no move-
ment was observed as it happened, it is likely that the scyphistomae
moved using their stolons as previously described for Aurelia aurita
(Schmal, 1985; Arai, 1997) and Catostylus mosaicus (Pitt, 2000). The
availability of additional substrate for these budding polyps removed
the density-dependent effects of intraspecifc competition for space (Di
Camillo et al., 2010) increasing the robustness of subsequent statistical
analysis. Another method to remove this effect is removal of daughter
scyphistomae after separation from the mother (Purcell, 2007). The
tentative suggestion is that scyphistomae be reclassied as sedentary
benthic organisms rather than the sessile benthic organisms to which
they are currently ascribed (Kawahara et al., 2006; Ishii et al., 2008;
Stampar et al., 2008; Willcox et al., 2008; Holst and Jarms, 2010;
Straehler-Pohl and Jarms, 2010).
4.3. Survival and somatic growth
A simple explanation for the observed mortality is that competition
for food led to some settled scyphistomae not being able to feed. At
high food levels there was plentiful food for all scyphistomae and com-
petition for food did not factor signicantly, however, some competition
especially among scyphistomae in close proximity could account for low
levels of mortality. Temperature was found not to be a signicant inu-
ence on mortality rates by Willcox et al. (2007) but they report a much
decreased mortality rate than this study. Other studies have also shown
higher survival rates of laboratory polyps (Liu et al., 2009) and our rela-
tively high mortalities may be because these were not from a previously
laboratory cultured scyphistoma population or an already established
colony taken from the wild.
This study found small overall size with most in the range of 0.1 mm
to 0.5 mm mouth disc diameter, with even the largest only reaching
1.3 mm. This could be a reection on the age of the polyps, as other
studies that use previously cultured polyps may already have a 1.1 to
1.4 mm calyx diameter from the beginning of the experiment (Han
and Uye, 2010) and can get as big as 4.9 mm (Willcox et al., 2007). In
contrast, Grndahl (1988a) measured his 10 day old scyphistomae at
0.6 mm (including tentacles). There is little comparable data on the
mouth disc diameter of new Aurelia aurita scyphistomae. It could be
simply a phenotypic difference in this population compared to other
populations of A. aurita which may reect plasticity in the species as
they adapt to different environments. This has been documented in
other life stages of A. aurita (Lucas, 2001) and would be a good topic
for further study.
No relationship between mouth disc diameter and temperature
was found, unlike Willcox et al. (2007) and Han and Uye (2010)
who found that at low temperatures polyps tended to be larger than
at high temperatures. They suggest that the scyphistomae were allo-
cating energy towards bud production at warm temperatures rather
than somatic growth, but scyphistomae have a fast ingestion rate
(Schiariti et al., 2008) and in their experiment food was plentiful, so
it is unclear why they would allocate their energy to one or the
other. This study found signicantly increased mouth disc diameters
at the high food levels, data which are in agreement with Han and
Uye (2010). Although photographs were taken before feeding, previ-
ous meals can change the elastic size of the gastric cavity creating a
size independent of actual somatic growth. This could partly explain
the signicant relationship found between food and somatic growth.
For this reason and because asexual reproduction was only observed
at the sixteen tentacle stage, as well as previously documented
non-linearity of similarly aged scyphistomae tentacle number and
physical dimensions (Holst et al., 2007), we suggest that number of
tentacles is maybe a better, and more informative indicator of somatic
growth.
Our analysis of tentacle number data produced a different result
than mouth disc diameter with regards to temperature nding that
there was a signicant effect of temperature on the number of scyphis-
tomae achieving certain growth stages. This signicant effect of temper-
ature has previously been reported for Aurelia sp. somatic growth
(Willcox et al., 2007; Han and Uye, 2010); however, these studies report
colder temperatures as being benecial for somatic growth. This could
be a difference between populations at different latitudes (Tasmania
and southern Japan respectively) and possibly from different habitats
(Albert, 2005). In the case of Han and Uye (2010), it could be because
low experimental temperatures represented medium natural tempera-
tures. Interestingly, food only had a signicant effect on numbers at the
four tentacle stage and higher which is likely due to the increased feed-
ing capabilities that come with having more tentacles to capture prey.
The interaction between food and temperature was only important
when scyphistomae had large numbers of tentacles. This is probably
down to increasing metabolic demand with increasing temperature
coupled with the amount of food required to sustain a larger organism
(Fitt and Costley, 1998). This pattern of increased growth rate with
food ts with what has already been described for Aurelia aurita medu-
sae (Hansson, 1997).
The intriguing lifecycle of scyphozoans and its relationship with envi-
ronmental factors has many questions that need to be investigated before
transposition of the potential effects of climate change on the abundance
and distribution of Aurelia aurita and other scyphozoans.
Acknowledgements
Special thanks to Dr. Jon Houghton and Ph.D. student Nick Flem-
ing of Queens University, Belfast for their attempts at sampling in
Strangford Lough. For advice on sample postage and polyp husbandry I
would like to thank Jamie Craggs of the Horniman Museum, London. I
am indebted to Dr. Sven Thatje and his Ph.D. students Alistair Brown,
Andrew Oliphant, Adam Reed and Katie Smith for allowing me use of
microscope imaging equipment. Many thanks go to the anonymous
reviewers for their constructive comments and advice. [SS]
 C.N. Webster, C.H. Lucas / Journal of Experimental Marine Biology and Ecology 436-437 (2012) 5055
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