University of Southampton 1 James Fay 23415045

Physical Population Dynamics of Aurelia aurita within the River Beaulieu, UK

By James Fay MSci Marine Biology 2013 Supervisor: Dr Cathy Lucas Word count: 9640

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Abstract Aurelia aurita the common moon jellyfish is commonly distributed between 70 oN and 40oS and is the UKs most common jellyfish. The following study looks at the population dynamics within the River Beaulieu, Southampton, with focus on physical factors such as temperature, salinity and tidal cycle. It also considers possible active behaviours that A. aurita may be using to control its population dynamics such as vertical migration, pulsation rate and ability to adapt to changing salinity. Zooplankton was also sampled to look for predatory relationships for the use as a proxy of A. aurita presence. The study was carried out over the spring-neap cycle of June 2012 from the 8th to the 28th. The results found that salinity was a controlling factor upon the distribution of A. aurita and controls its vertical and horizontal position within the river through density driven currents. Rainfall also has a significant effect on their population and forces them to migrate deeper. There were also differences between the neap and spring cycle; where higher densities were seen during spring cycles which are attributed to greater aggregation and smaller bloom size on a spatial scale. The ability of the A. aurita to adapt to changing salinity over the course of a tidal cycle was too slow to keep up with the change which supports salinity as a major controlling factor in its distribution within the Beaulieu. A. aurita also displayed endogenous vertical migration to the tidal cycle which is a mechanism they use to maintain their isolation and position within the River Beaulieu. Pulsation rate alters due to daily vertical migration (DVM) signals, temperature fluxes and prey presence and may be a factor in A. auritas vertical distribution. Comparing the population to other Southampton water populations they are significantly larger with a max bell size of 192mm and an average size of 132.12mm

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Acknowledgements For this study I would like to first thank my supervisor Cathy Lucas who has provided valuable knowledge, expertise and help throughout the course of the project as well as providing transport to and from the sampling site. I would also like to thank Emma Whelan for her help in collecting the field data and analysing the zooplankton microscope samples which took numerous hours. I would also like to thank Chris Sturdy for providing aquarium facilities for the laboratory experiments. I would like to thank the University of Southampton also for the funding needed to carry out the research presented and I would lastly like to thank Natasha Phillips for her recommendations based on her previous study of the River Beaulieu.

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Contents 1.0. Introduction 1.1. 1.2. 1.3. 1.4. 1.5. 1.6. 2.0. Aurelia aurita Distribution of A. aurita Why study A. aurita? The River Beaulieu Aims Hypotheses 6 6 8 9 11 13 13 14 14 15 17 17 17 18 21 22 22 25 28 29 30 30 32 34

Methods and Materials 2.1. 2.2. Study site Data collection

2.2.1. Aurelia aurita field sampling & collection 2.2.2. Zooplankton collection 2.2.3. Temperature and Salinity 2.2.4. Laboratory 2.3. 3.0. Data analysis

Results 3.1. 3.2. 3.3. 3.4. 3.5. Salinity profiles Temperature profiles Zooplankton Aurelia aurita A. aurita and zooplankton interactions

3.5.1. Monthly A. aurita and zooplankton interactions 3.5.2. Daily A. aurita and zooplankton interactions 3.6. Neap Spring comparison

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3.7. 3.8. 3.9. 4.0.

Salinity adaptation A. aurita pulsation rate A. aurita vertical position

38 40 41 43 43 45 46 46 48 48 49 51 52 53 53 61

Discussion 4.1. 4.2. 4.3. 4.4. 4.5. 4.6. 4.7. 4.8. 4.9. Temperature and Salinity Aurelia aurita size, density and food availability A. aurita and zooplankton interactions Aurelia aurita over neap and spring cycles Vertical position Aurelia aurita Pulsation rate Aurelia aurita Limitations Future recommendations Implications

5.0. 6.0.

Conclusions References

Safety forms

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1.0. Introduction 1.1. Aurelia aurita Aurelia aurita, commonly known as the moon jellyfish, is a Scyphozoan of the Cnidarian phylum and was first described by (Linnaeus, 1758). A. aurita is most physically distinguished by eight marginal lobes and four oral arms as long as its umbrella, four gonads and four gastric pouches shown in (figure 1) which also shows the structure of A. aurita (Russell, 1970). However, individuals with more than 4 gonads or gastric pouches can be observed.

Fig1. Subumbrellar view of adult female meduase with brooding pouches (Russell, 1970). A. aurita has a cosmopolitan diet and feeds on other plankton, but is not limited to eating small fish (Lebour, 1922; Orton, 1922; Arai, 1997), it captures prey items using ciliary currents externally and internally, trapping them within mucus (Russell, 1970), whereby nematocysts situated on the oral arms and tentacles capture the prey where they are then passed to the mouth (Costello & Colin, 1994). A. auritas

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locomotion is controlled by contracting their coronal and radial muscles to propel itself through the water column (Russell, 1970). The life cycle of A. aurita has two forms, one as a pelagic medusa typically lasting 3 to 4 months within Southampton (Lucas and Williams, 1994), where it then reproduces sexually to produce planula larvae which settle to form benthic polyps. The second life stage is the benthic polyp where it reproduces asexually or undergoes strobilation to produce many ephyrae which grow into medusae, (figure 2) shows the life cycle of A. aurita (Brusca & Brusca, 2003).

Fig2. Life cycle of A. aurita a) Medusae > b) Fertilised egg > c) Planula larvae > d) Polyp, the scyphistoma > e) asexual reproduction to form new polyps > f) Strobilation to give off ephyrae > g) Ephyrae grows into adult medusa > a) (Brusca & Brusca, 2003). A. aurita have relatively few predators (Parsons & Lalli, 2002) although they are preyed upon by fish and turtles (Purcell and Arai, 2001; Houghton et al. 2006). The physical resilience of A. aurita to temperature and salinity is renowned (figure 3) they

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are a euryhaline and eurythermal species (Miyake et al. 1997; Lucas, 2001). Few controlling factors on their population distribution have allowed for wide and increasing population ranges (Purcell et al. 2007).

Fig3. A. aurita temperature and salinity ranges within different localities (Miyake et al. 1997). 1.2. Distribution of A. aurita A. aurita has a global distribution ranging from 70oN to 40oS (Lucas, 2001) where they are mostly found within coastal and estuarine environments (Lucas, 2001). There have been many studies upon A. aurita populations and factors relating to their distribution and impacts around the globe including, Southampton, UK, (Lucas & Williams, 1994), the North Sea (Hay et al. 1990), Japan (Shoji et al. 2010), Roscoe Bay (Albert, 2009), the Wadden Sea (van der Veer & Oorthusen, 1985) and the Baltic Sea (Schneider, 1989).

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A. aurita also have been said to have a distribution all over the coasts of Britain and is our most common jellyfish according to Russell (1970). However on the Southern coast it has been said that they are endemic to Southampton and the Solent (Lucas & Williams, 1994). Within this study the population to be studied is within Southampton waters within the River Beaulieu which would support the endemic claim, however further information on A. aurita within the English Channel is absent (Lucas & Williams, 1994). The River Beaulieu population may have first become present here through polyps settling upon oyster shells found within the river (Rodhouse, 1979). 1.3. Why study Aurelia aurita? The study of A. aurita is important due to its common distribution and resilience to survive. With reports documenting that in recent years the population of jellyfish is increasing globally (Purcell et al. 2007) they are of growing concern. Particular problems caused as a result of A. aurita include the splitting of fishing nets within japan under high abundances (Purcell et al. 2007). The blocking of sea cooling intakes for power stations which then over heat and are then forced to shut down is also a problem (Purcell et al. 2007). This may be of particular interest to study Southampton populations due to the presence of the Marchwood power station and its cooling intakes within the River Test. Furthermore the cosmopolitan lifestyle has implications towards marine food webs as their feeding has significant impact on the abundance of certain plankton species such as Acartia omorii documented in Omori et al. (1995). A. aurita also prey on high trophic level predators larvae and commercially important fish larvae (Arai, 1997; Purcell et al. 2007), this cosmopolitan lifestyle leads to trophic cascades similar to ones observed in cod stocks with

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combined effects of overfishing (Frank et al. 2011). Within the Baltic Sea several fish stocks such as bonito, mackerel, bluefish and dolphins became severely depleted as a result of A. aurita removing fish food items such as zooplankton which when coupled with overfishing caused trophic cascades (Daskalov, 2002). Further reasons to research A. aurita include the effects of climate change, as increasing sea temperatures could allow for A. auritas distribution to change as their habitable temperature range increases towards the poles and allows for invasion; where other gelatinous creatures such as Salpa Thompsoni are already exploiting this changing environment (Schofield et al. 2010; Aronson et al. 2011). Increasing populations could be aided via translocation within ballast water from ships, and the settlement of polyps onto boat hulls (Graham & Bahya, 2007). With these impacts in mind it is therefore important to study the population dynamics of A. aurita in order to manage their impact to prevent disruptions to commercial activities and to ensure a balanced marine ecosystem persists which will help to maintain biodiversity. The A. aurita population within the River Beaulieu has not been described by published literature, and there is currently only the previous study carried out by Phillips (2012) on this population which did not focus towards physical effects upon the population. This study will aim to further the understanding of the River Beaulieu population with particular focus on physical factors such as salinity and temperature. 1.4. The River Beaulieu The A. aurita population to be studied is found within the River Beaulieu (figure 4). The A. aurita blooms within the river may be attributed to the oyster beds once present here, where their shells now provide substrate for polyps to settle and

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develop (Rodhouse, 1979). The River Beaulieu is an estuarine river which acquires its nutrients from 80km2 the heath and bog lands of the New Forest, UK. The lower estuary to which the A. aurita have access to is approximately 8km in length where mean flow is estimated at 0.63 m3 s1 (Holliday & Liss, 1976; Fang, 1995; Turner et al. 1998), at this site physical mixing with the Solent occurs (Holliday & Liss 1976) and this creates a very dynamic environment within the estuarine river. The Beaulieu is characterised by high dissolved organic matter (DOM) and with notable iron removal from the system (Holliday & Liss, 1976; Howard et al. 1984; Martino et al. 2003). The River Beaulieu is also a popular yachting destination with around 7500 boats visiting each year where if A. aurita populations become too dense could cause problems. The tidal cycle of the River Beaulieu is characterised by a double high tide where the flood tides lasts 6 hours followed by a second high water approximately 2 hours 30 minutes later where after 3 hours 30 minutes the ebb begins and lasts only 2 hours 30 minutes (http://www.beaulieuriver.co.uk/about).

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River Beaulieu

Fig4. Position of River Beaulieu within the UK (Lucas, 1996). 1.5. Aims The aim of this study is to describe the population dynamics of adult A. aurita medusae over the course of a spring and neap tidal cycle with reference to physical conditions such as temperature and salinity. This will be achieved by collecting abundance data for A. aurita over the course of spring and neap cycle at a fixed station to look for differences between the cycles. The abundance data will also be collected to analyse any changes in density of the A. aurita and their potential prey items within the zooplankton. Temperature and salinity measurements will also be

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taken throughout the spring and neap cycle to see how these influence the distribution of the A. aurita. A. auritas swimming behaviour will also be studied within laboratory conditions to look for changes throughout a tidal cycle, in order to assess if the A. aurita are actively controlling their position within the River Beaulieu. To achieve these aims simple hypotheses have been constructed. 1.6. Hypotheses Testable null hypotheses have been constructed for this study which are: 1. A. aurita salinity or temperature physical tolerances do not control its distribution within the River Beaulieu. 2. The A. aurita population distribution within the River Beaulieu does not change between neap and spring tides. 3. A. aurita pulsation rate and vertical positioning behaviour does not change over a tidal cycle.

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2.0.0. Methods & Materials 2.1.0. Study site

Fig5. Aerial photo of the survey area with superimposed track of the zooplankton net trawl, which was collected between 50o 48 02.71N, 1o 25 21.81W and 50o 48 01.36N, 1o 25 19.80W at the River Beaulieu, United Kingdom.

Data was collected from the fuel jetty located at Bucklers Hard, River Beaulieu, shown in figure 5. This is situated mid-way up the river it has depth ranges from approximatley 5m at high tide and 3m at low tide at the sampling location. Although an ideal place to sample from it was also chosen out of convenience due to funding

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limitations and lack of access to a boat which would have been a preferred sampling method. During the sampling period it was also recorded as the wettest and dullest June on record. Particularly on the 11th June as during sampling there was persistent heavy rain throughout the day (http://www.metoffice.gov.uk/climate/uk/2012/june). 2.2.0. Data collection Date 8.6.2012 11.6.2012 13.6.2012 15.6.2012 18.6.2012 20.6.2012 25.6.2012 27.6.2012 Start Time (BST) 10:18 10:33 11:00 13:31 15:20 06:25 10:00 11:18 End Time (BST) 18:36 22:53 19:40 21:11 23:00 18:25 15:20 17:18

Table1. Dates of sampling with time that sampling commenced and time that sampling ceased on each day. 2.2.1. Aurelia aurita field sampling & collection The A. aurita population was sampled using a 100m by 0.202m2 zooplankton net with a cod-end attached to collect specimens; a flow meter was also attached for recording water flow in order to obtain densities per m3 for qualitative sampling. The net was trawled by hand horizontally (figure 6) just below the surface up and down the pontoon once located at bucklers hard, within the River Beaulieu shown in figure 5, trawls were repeated every 20 minutes during the sampling period shown in table1.

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Where each trawl lasted approximately 5 minutes, after each trawl the cod end was removed and emptied into a bucket, A. aurita were counted and removed. Their bell diameter was then measured within a white tray on top of a ruler (figure 6) placing the specimen within the tray provided a contrastable measuring surface allowing for faster and more accurate measuring, whilst also providing a smooth surface which would minimise stress to the individual allowing them to be returned to the river more efficiently after being measured.
a) b)

Fig6. (a) A. aurita specimen measured within a white tray and ruler. (b) Zooplankton net being trawled horizontally. On the 25th A. aurita were also collected quantitatively using a bucket to collect individuals where their bell was measured and then returned. This provided a more complete size frequency distribution of the population whilst also highlighting some limitations in the sampling methods, which will be discussed within the discussion. On the 27th A. aurita were also collected quantitatively alongside the net specimens; these were measured then placed in buckets of seawater to be returned to the laboratory for further analysis of which 11 individuals were retained for this purpose.

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2.2.2. Zooplankton collection Samples of zooplankton were obtained using the same 100 m by 0.202m2 net used to collect the A. aurita. Samples were taken at different stages of the tidal cycle throughout the day. Collection was also based on the predicted A. aurita density where samples were taken at periods that were considered pre-bloom, bloom period, and post-bloom. This was done with the hope of showing A. aurita impact on zooplankton densities. Before each zooplankton sample was taken the net was rinsed thoroughly to remove any organisms from previous trawls that may bias the results. The net was trawled up and down the pontoon once, where after the trawl the net was rinsed with the cod-end attached to wash any animals trapped within the net mesh into the cod-end, after which the cod-end was emptied into a labelled 500ml bottle for further analysis using microscopes within the laboratory. 10% formalin solution was added to the sample bottle in order to preserve the sample. 2.2.3. Temperature & Salinity A temperature/salinity (T/S) probe was used to record temperature and salinity depth profile measurements. However measurements were only recorded from the 15th onwards, due to the equipment being broken on the first measurements on the 8th beforehand. T/S measurements were recorded every 20 minutes throughout the sampling period or 10 minutes after the start of each net trawl. This was done by lowering a T/S probe 1 metre at a time from the river surface until it reached the river bed.

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2.2.4. Laboratory

Fig7. 48inch sampling water tank used for rhythm studies with attached ruler for depth measurement. On the 28th June 2012 A. aurita was analysed within water tanks to look for endogenous rhythms to the tidal cycle, rhythms to dawn/dusk periods and its ability to adapt to changing salinity. To test for endogenous rhythm to tidal cycle and rhythms for dawn/dusk periods a similar method to Mills (1983) was used, where A. aurita were placed in a 48 inch deep tank with a glass front panel and attached ruler to measure depth (figure 7). A. auritas vertical position within the tank was then recorded every 30 minutes from 6am to 11pm. In addition to vertical position, pulsation rate was recorded for each individual by counting their bell pulses for 2 minutes every 30 minutes, 5 individuals were used for this test however 2 individuals were damaged as a result only 3 individuals were tested. Additionally the A. aurita

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were fed at 10:00 with frozen Artemia sp. and at 14:00 with live Artemia sp. to avoid starvation playing a factor within the experiment. The ability of A. aurita to respond to changing salinity was also tested on the 28th June 2012 by placing 6 individuals from a salinity of 31.9%o to lower salinity tanks of 15%o, 20%o and 25%o; 2 individuals were placed in each condition. These conditions were representative of the conditions found within the River Beaulieu. The lower salinity tanks were made by adding distilled fresh water to the 31.9%o salinity water. Once adapted to these salinities the individuals were moved to the other tanks where their response was tested again. The adaptation time, was taken as the point where the A. aurita swimming ability was no longer impaired. When placed into different salinities A. aurita ability to swim is affected; when placed into lower salinities they sink due to being denser. Whereas when placed into higher salinities they then float to the surface as they are less dense. Therefore the point where they can move between the surface and bottom of the water tanks was taken as the point of adaptation to the changed salinity. The time taken for this to occur was recorded. The adaptation was also stopped regardless of successful adaptation after 7 hours, as in natural conditions the tides will have changed and the A. aurita will not be exposed to a changed salinity for longer than 7 hours. As a result the jellyfish were moved to different tanks to adapt to a different salinity.

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Fig8. Zooplankton sample under microscope. Zooplankton samples were analysed using a microscope (figure 8) where 10ml of each 500ml bottle was pipetted into a bogorov-counting chamber after the bottle was well shaken. The 10ml was diluted down to make counting and identifying the plankton sample easier. Zooplankton was identified to class level; due to A. auritas cosmopolitan feeding style, further identification was not necessary. After the abundance per 10ml for the zooplankton was found, the total abundance per m 3 for each sample was obtained by using the following equation taken from Harris et al. (2000):

AT R 0.3 0.2552
A = abundance per 10ml T = total sample 500ml conversion factor, 50 R = flow meter revolutions

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2.3.0. Data analysis Data was analysed using Sigma plot, Minitab, Microsoft Excel statistical software to produce graphical representations of the data to describe the population dynamics of A. aurita and to perform appropriate statistical tests upon the data.

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3.0. Results 3.1. Salinity profiles Salinity data collected from the Beaulieu was pooled together and then represented using contour graphs to show how salinity changes over the course of a tidal cycle, from which there were two observed profiles (figure 9 & 10).

Salinity Depth Profile Over Time 15/06/2012

0
Salinity < 12 12 14 14 16 16 18 18 20 20 22 22 24 24 26 26 28 > 28

-1

-2

Depth

-3

-4

15/06/2012 14:00 15/06/2012 16:00 15/06/2012 18:00 15/06/2012 20:00

-5

Date-Time

Fig9. Salinity depth profile for the 15th June 2012 where sampling started at 13:43 and ended at 21:08 sampling every 20 minutes, blue indicates low salinity, red indicates high salinity. The first profile to be observed (figure 9) shows vertical stratification with overlying freshwater and underlying seawater. As time progresses towards high water the stratified layers become more mixed and salinity increases as sea water moves in on the flood tide. There is also a large range in the salinity change of 15%o over the tidal

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cycle. The observed profile could be attributed to the large amount of rainfall from the 11th June causing a large input of freshwater to the surface waters.

Salinity Depth Profile over time 18/06/2012

0
Salinity < 21 21 22 22 23 23 24 24 25 25 26 26 27 27 28 > 28

-1

-2

Depth

-3

-4

-5 18/06/2012 16:00 18/06/2012 18:00 18/06/2012 20:00 18/06/2012 22:00 Date-Time

Fig10. Salinity depth profile for the 18th June 2012 where sampling started at 15:30 and ending at 22:50 sampling every 20 minutes, blue indicates low salinity, red indicates high salinity. The second observed profile (figure 10) shows a well-mixed vertical profile which was also observed on the 20th, 25th and 27th. Like the stratified profile salinity increases as sea water moves in on the flood tide and approaches high water. However the range in salinity is only 7%o and the surface salinity is constantly above 20%o. There was no heavy rainfall since the 11th June and the change in salinity profile which was persistent for the rest of the month could indicate that the profile seen on the 15th is what occurs due to rainfall and that the 18th shows normal estuarine mixing profiles which are not influenced by rainfall.

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Salinity Depth Profile Over Time 20/06/2012

0 -1 -2 -3 -4 -5
Salinity < 20 20 22 22 24 24 26 26 28 > 28

Depth

12 20 / 6 /0 0 2

00 8:

12 20 / 6 /0 0 2

0 :0 10

12 20 / 6 /0 0 2

0 :0 12

12 20 / 6 /0 0 2 Date-Time

0 :0 14

12 20 / 6 /0 0 2

0 :0 16

12 20 / 6 /0 0 2

0 :0 18

Fig11. Salinity depth profile for the 20th June 2012 where sampling started at 06:35 and ended at 18:35 sampling every 20 minutes, blue indicates low salinity, red indicates high salinity, high tide at 11:06 BST. Sampling also took place over a 12hour tidal cycle (figure 11) where it showed a well-mixed profile similar to (figure 10) and also only ranged by 7%o. However there is a sharper salinity gradient after the double high tides as the salinity changes over a shorter period of time where the fast flowing ebbing tide occurring over a short period causes this sharp change. This rapid change may have effects on the organisms present within the water column. In addition after the first high tide at 11:06 the salinity remains constant until around 15:00 which demonstrates the duration of the high tide and that the ebb tide does not occur for a long period after the high tide.

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3.2. Temperature profiles Similar to the salinity profiles there was also temperature two profiles observed within the River Beaulieu (figure 12 & 13).

Temperature Depth Profile Over Time 15/06/2012

0
Temperature < 15.0 15.0 15.5 15.5 16.0 > 16.0

-1

-2

Depth
-3 -4
15/06/2012 14:00

-5

15/06/2012 16:00

15/06/2012 18:00

15/06/2012 20:00

Date-Time

Fig12. Temperature depth profile for the 15th of June 2012 where sampling began at 13:43 and ended at 21:08 where samples were taken every 20 minutes. The first observed temperature profile observed on the 15th June (figure 12) shows a vertical stratification with warmer surface water and colder underlying seawater the temperature range however is small only changing by 1oC there is also a very small decrease in temperature as high tide approaches as colder water from the sea floods in. Compared to the 18th June (figure 13) the temperature is lower by 2oC this decreased temperature may be due to the large rainfall, which may have decreased the temperature.

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Temperature Depth Profile over time 18/06/2012

0
Temperature < 16.0 16.0 16.5 16.5 17.0 17.0 17.5 17.5 18.0 > 18.0

-1

-2

Depth

-3

-4

18/06/2012 16:00 18/06/2012 18:00 18/06/2012 20:00 18/06/2012 22:00

-5

Date-Time

Fig13. Temperature depth profile for the 18th June 2012 where sampling started at 15:30 and ending at 22:50 sampling every 20 minutes, blue indicates low salinity, red indicates high salinity. The second observed profile (figure 13) of the 18th June shows a well-mixed profile as the same temperature persists to the river bed; there is also a decrease in temperature towards high tide which could be due to tides with cold seawater influx or due to the sun setting and no heat input from sunlight. This profile was also seen on the 20th, 25th and 27th June.

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Temperature Depth Profile Over Time 20/06/2012

0 -1 -2 -3 -4 -5
Temperature < 18.0 18.0 18.5 18.5 19.0 19.0 19.2 19.2 19.5 > 19.5

Depth

12 20 / 6 /0 0 2

00 8:

12 20 / 6 /0 0 2

0 :0 10

12 20 / 6 /0 0 2

0 :0 12

12 20 / 6 /0 0 2

0 :0 14

12 20 / 6 /0 0 2

0 :0 16

12 20 / 6 /0 0 2

0 :0 18

Date-Time

Fig14. Temperature depth profile for the 20th June 2012 where sampling started at 06:35 and ended at 18:35 sampling every 20 minutes, blue indicates low salinity, red indicates high salinity. The well mixed vertical profile was also seen over the 12 hour cycle however the temperature only increases throughout the day with particular increase at the beginning of the ebb tide and the flow of the colder sea water out of the river. Figure 14 demonstrates that the tidal cycle does have an effect on temperature, but also as sunlight would have increased during this period the increase in the temperature could be partially due to sunlight as well. The temperature of the river had also increased further from the 18th June and now had a maximum temperature of 19.5oC compared to 18oC seen on the 18th June.

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3.3. Zooplankton Zooplankton were analysed to class level within the Beaulieu and their densities recorded. Their composition was also analysed (figure 15).
Zooplankton composition per m3

Zooplankton per m3 Copepods Copepod Nauplii

Copepods Copepod Nauplii Tunicates Echinoderms Polychaetes Cirripedes Siphonophores Teleosts Gastropods Decapods Larvacea

3622.08 10153.19 3.19 6.38 135.08 4.58 1.00 2.19 246.45 8.97 0.20

Tunicates Echinoderms Polychaetes Cirripedes Siphonophores Teleosts Gastropods Decapods Larvacea

Fig15. Pie chart illustrating the percentage composition of zooplankton classes found within the River Beaulieu during June 2012 with density per m3 sampled from 250m3 over June. Within (figure 15) the most dominant class was the copepods consisting of the copepod and copepod nauplii groups which contributed 97.12%. The remaining majority consisted of gastropods 1.75% and polychaetes 0.95%. For the 250m3 of water sampled during June the densities of copepods and copepod nauplii were far greater and within the samples were most abundant. Considering A. aurita is a cosmopolitan feeder it would likely feed upon all of the zooplankton, but, due to the high percentage of copepods; their diet most likely would consist of copepods.

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3.4. Aurelia Aurita

Aurelia aurita size frequency distribution
35

30

25

Frequency #

20

15

10

0 60-70 71-80 81-90 91-100 101-110111-120121-130131-140141-150151-160161-170171-180181-190191-200

Umbrella size (mm)

Fig16. Histogram showing A. aurita umbrella size frequency distribution of 140 individuals, of which 130 collected quantitatively and 10 collected qualitatively during June 2012. Of the A. aurita collected (qualitatively 10 and quantitatively 130) umbrella size was measured combining sampling techniques; from this a size frequency distribution graph was constructed (figure 16). The average mean size of the population was 132.171mm with a standard deviation of 24.341. in addition a Shapiro-Wilks test was run to test for normality which it passed with P = 0.854. This indicates that the specimens caught from the population are normally distributed around the mean. In addition to the size of A. aurita caught there were also specimens caught that had 5 or 6 gonads rather than the common 4 gonads. Due to the normal distribution it would also suggest that the data collected is likely reliable and will not have any outlying points as it represents the majority of the population found.

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3.5.1. Monthly Aurelia aurita and zooplankton interactions

3.5 3.0

3 Aurelia aurita /m

2.5 2.0 1.5 1.0 0.5 0.0

8th
14 12 10

11th

15th 18th

20th

25th

27th

Zooplankton /m

8 6 4 2 0

8th
4.0 3.5 3.0

11th

13th

15th

18th

20th

25th

27th

Tidal Height (m)

2.5 2.0 1.5 1.0 0.5 0.0

8th

11th

13th

15th

18th

20th

25th

27th

Fig17. Time series of A. aurita, zooplankton and tidal height for June 2012 showing how densities changed throughout the month and relations between zooplankton and A. aurita.

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Interactions between zooplankton and A. aurita are shown within (figure 17) of which when A. aurita is present zooplankton densities are either low or in decline. This would imply that there is a predator-prey relationship being exhibited, however to test this interaction on a monthly scale statistically would not give a clear reliable result due to limitations in the net sampling reporting a density of 0 qualitatively when no A. aurita were caught, even when they are being caught quantitatively at the same time. In addition there was no A. aurita caught on the 13th June which is after the heavy rainfall from the 11th June and is possibly during stratified conditions shown within the salinity-temperature profiles within (figures 9 & 12). In addition to this the A. aurita density is lower during neap tides than spring tides according to the figure which was tested using Pearsons product moment correlation upon the mean daily A. aurita density and tidal range. This showed significant strong correlation of C= 0.878 with P < 0.005. however statistically there was no significant relationship between zooplankton and tidal range indicating no significant change in their density between neap and spring tides as Pearsons correlation reported C= 0.553 P=0.155.

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3.5.2 Daily Aurelia aurita and zooplankton interactions Looking in more detail between interactions between zooplankton and A. aurita densities per m3 with relation to tidal cycle (figure 18) it is observable that there is a relationship between A. aurita and the zooplankton where when A. aurita is present zooplankton densities are decreasing or are low. Looking at the 15th June the A. aurita only appear just before high tide compared to other days where A. aurita is present which are present for a few hours before high tide. Referring back to the Salinity profiles (figure 9) the salinity is above 20 when the A. aurita is caught when below this no A. aurita was caught. Furthermore where A. aurita persist hours before such as the 18th June the salinity is constantly above 20 (figure 10). Further looking at the 13th June after the heavy rainfall from the 11th June the zooplankton density was also heavily affected and is much lower compared to zooplankton before the rain on the 8th June. Whilst sampling after the rain the colour of the River had also changed colour to a dark brown and was full of silt and debris possibly washed from the surrounding heath and bog land. In addition on the 20 th where sampling continued for 12 hours from low-high-low tide A. aurita were caught after high tide the only time during June, they were caught approximately 5 hours after high tide at which time it is predicted that the ebb tide starts. It is therefore likely that after the initial high tide the A. aurita population stays up river or continues moving up river until the start of the ebb tide after the second high tide.

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Fig18. Zooplankton, A. aurita and tidal height, daily interactions over a tidal cycle.

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3.6. Neap-Spring comparisons

Fig19. A. aurita density around high tide shown for neap-spring tides for June 2012. Comparisons between the sampling that were considered neap tides 11th, 13th, 15th,18th and 27th June were compared against sampling days that were considered spring tides 8th, 20th, 25th June; although no A. aurita were caught on the 13th June. These were first compared with regards to density (figure 19) which was also previously tested within figure 17 which showed a significant result, where spring tides have a higher density than neap tides. This is evident within figure 19. In addition almost all the density measurements recorded are before high tide as A. aurita caught after high tide were apparently absent except on the 20th where they were caught 5 hours after. In addition the majority of the neap densities are clustered

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closer around the high tide mark with the exception of the 18th which is borderline spring tide. With this in mind it would appear that neap tides have a smaller bloom range around the neap tide and do not move as far up or down the river during neap tides compared to spring tides. This is examined in figure 20. The average time A. aurita appeared before high tide was 2 hours for spring tides and 2 hours 5 minutes for neap tides. The average after high tide was 2 hours 50 minutes for spring tides and 1 hour 54 minutes for neap tides; the larger average seen in the neap tides before high tide is higher due to a larger spread of the data particularly from the 18th June whereby the data point (-5.46, 0.1718) within figure 19 skews the mean by 13 minutes, where the mean would be 1 hour 52 minutes without this point. Considering its close proximity to low water this point may be A. aurita caught from the ebb tide as A. aurita was caught just before low water also.

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Fig20. A. aurita time of sampling net capture shown for each day to show how the blooms position within the River Beaulieu changes between neap-spring cycle. The time at which A. aurita were caught during each sampling day in relation to the tidal state has been plotted (figure 20) which shows that the neap sampling days are present closer around the high tide than spring tides are, showing that the A. aurita blooms distribution alters from neap to spring tide. However figure 20 also shows that during neap tides the bloom is more spread and can be seen at a stationary point for a longer period of time than on a spring tide, whereas on the spring tide the bloom can be seen for a shorter period of time. This shows that during a neap tide the size of the bloom is less aggregated and spread across a larger range, whereas

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on a spring tide the bloom becomes more aggregated and its range shrinks. Illustration of how the blooms dynamics change is shown by figure 21.

Neap Bloom, Low density

Spring Bloom, High density

Fig21. Illustration of bloom size between neap and spring tidal cycles. The illustration (figure 21) shows how the density per m3 changes between neap and spring tides and does not represent a larger or smaller population size observed between spring and neap tides; this figure is also supported from the density statistics run within section 3.5.1. in addition although describing the size of the bloom changes occurring over a tidal cycle it cannot show the spatial limits that the A. aurita are found at within the river as current flow data is needed with ground truthing spatial survey to be discussed in future recommendations.

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3.7. Salinity adaptation

Fig21. Time taken for A. aurita to adapt to changes in salinity where the x axis represents the time taken and the y axis is the level of change the individuals were exposed to. It was carried out on the 28th June 2012. Whilst carrying out the experiment testing for adaptation ability to salinity change, the physical form of the A. aurita was changed whereby when placed in lower salinities the umbrella became inverted, whereas placing in higher salinities the umbrella became more spherical and less-saucer shaped this observation has also been previously documented and observed (Thill, 1937; Russell, 1970). From the salinity profiles (figures 9-11) the change in salinity over a tidal cycle ranged from 7-12%o

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over the space of 6 hours (figure 21) shows how long it takes for A. aurita to adapt to similar changes in salinity of which it takes 4hours 47 minutes to adapt to a change of 5 and for larger salinity changes it takes a longer period of time. Linear regression was carried out on this whereby the R2= 0.392, with a line of regression as Salinity change = (Time+2.381)/1.730 indicating weak positive correlation. Considering this the adaptation time of the A. aurita found within the Beaulieu is too slow as within the space of the 4 hours 47 minutes needed to adapt to the small change, the surrounding salinity will have already changed as shown from the salinity profiles.

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3.8. A. aurita pulsation rate

Fig22. Pulsation rate per 2 minutes of A. aurita from 6am to 11am with repeat measurements every 30 minutes on the 28th June 2012, with a superimposed tidal state curve from the River Beaulieu to match the time the experiment was conducted; vertical black lines represent times of feeding. The pulsation rate of A. aurita (figure 22) shows an initial increase in pulsation early within the morning at 6:08 and a decrease in activity from 20:30 onwards this could be the presence of a dawn/dusk rhythm as at these times the sun was rising/setting and the A. aurita would have been exposed to changing levels of light. However there does not seem to be any rhythm related to tidal cycle as mean pulsation rate remains at a steady level, in addition the A. aurita were fed at 10:00 with frozen

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Artemia sp. and 14:00 with live Artemia sp. where the period just before feeding there is a decrease in pulsation rate, however after feeding the pulsation rate increases, pulsation rate from (figure 22) likely does not change as a result of tidal cycle but there may be a relationship between pulsation rate and the presence of food. 3.9. A. aurita vertical position

Fig23. Vertical height of A. aurita within the sampling tank where 0 is surface and 48 is bottom of tank taken from 6am to 11am every 30 minutes on the 28th June 2012, with a superimposed tidal state curve from the River Beaulieu to match the time the experiment was conducted, vertical black lines represent times of feeding.

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Vertical position of the A. aurita within the deep tank simulates water depth at a smaller scale found to that in the Beaulieu but does not expose the A. aurita to tidal cues, therefore testing for an endogenous rhythm with their vertical positioning within the water column. Figure 23 shows that at low tide and shallow depths the A. aurita move to greater water depths. On the other hand as water depth increases and approaches high tide they move to shallower depths, displaying an endogenous rhythm. The large fluctuations in the mean vertical position may be due to the only 3 healthy individuals being retained for this experiment or due to the complexities of a double high tide tidal cycle found within the River Beaulieu. In addition vertical position does not appear related to dawn/ dusk rhythm as there is a decrease in depth at both dawn/dusk points. After feeding the A. aurita live Artemia sp. there was an increase in vertical position of the A. aurita.

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4.0.

Discussion

4.1. Temperature and Salinity on A. aurita The observed temperature salinity profiles, where stratification occurred and reductions in salinity and temperature can be attributed to rain fall from the 11th June where its effects persisted to the 15th June most likely due to the proceeding fresh water runoff from the surrounding heath and bog lands (Katsaros & Buettner, 1969; Holliday & Liss, 1976; Atkinson & Blanton, 1986). This effect is known as estuarine circulation and is expected to occur where inflowing seawater with greater density or higher salinity mixes with less dense less saline fresh water causing vertical stratification as observed (Hansen & Rattray, 1966). Where the well-mixed vertical profiles occur for the remaining sampling days it would imply low mixing using the flow ratio of coastal plains which would be approximately 0.1 for river Beaulieu (Hansen & Rattray, 1966). The well mixed profile of salinity and temperature observable within the River Beaulieu is most likely due to the barrier entrance, found at the mouth, causing restrictive flow between the sea and the river (Hansen & Rattray, 1966). The absence and low abundance of A. aurita seen on the 13th and the 15th could be due to the rainfall as it has been documented that they move to deeper depths when it begins to rain (Claus, 1877; Russell, 1970). Additionally the medusae are scarce or absent when the after effects of the rain have reduced the salinity (Lo & Chen, 2008). Salinity is likely a controlling factor on the distribution of A. aurita found within the Beaulieu. As salinity changes it causes A. aurita to change vertical position as observed in the results during the salinity adaptation experiment. It is also consistent with how the A. aurita were distributed throughout the Beaulieu as when the tide

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begins to ebb and salinity declines at the surface the A. aurita are forced to deeper depths due to higher density limiting them from the surface. This effect is what causes the absence of A. aurita from being detected on the ebbing tide. On the other hand during the flow tide where salinity is increasing and the A. aurita are adapted to less dense waters they are able to swim to the surface allowing for net capture and sampling. The effect of salinity controlling the distribution of A. aurita is also seen within the Black Sea as well as the Beaulieu described in this study (Russell, 1970; Zhong 1988; Mutlu, 2001). Further evidence also supports that salinity controls vertical distribution and horizontal distribution as salinity has been found to account for 20% of spatial variations (Primo et al. 2012). It is yet again documented to occur within Taiwan when salinity dropped below 20, the A. aurita disappeared (Albert, 2011). Based on the influence salinity has regarding vertical distribution it is likely this holds true for horizontal distribution and A. aurita are most likely found within a narrow salinity range where they are aggregated and their position controlled by density interactions causing them to move with the salinity as it changes tidally. This has also been observed within a Danish Fjord (Nielsen et al. 1997). A. aurita caught throughout the sampling period were caught within a salinity range of >20, this effect is particularly noticeable on the 15th where there is the largest change in salinity along the surface over the course of the tidal cycle. It is likely that the A. aurita prefer to remain in a constant salinity which will provide advantages in conserving energy by not having to adapt to new salinities; saving this energy by not adapting to the changed salinity results in altered distributions of A. aurita (Nielsen et al. 1997). Furthermore when salinity did drop below 20 the A. aurita disappeared, where this trend has been similarly observed in Taiwan (Albert, 2011).

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Changes in temperature within the Beaulieu were fairly homogeneous with very small changes in temperature throughout the water column and tidal cycle although temperature is known to have the effect of lowering pulsation rate of A. aurita it is unlikely that temperature is a controlling factor on its distribution (Russell, 1970). 4.2. Aurelia aurita size, density and food availability Compared to other A. aurita populations found within Southampton Water particularly Horsea Lake (near-Portsmouth) and the River Test (see figure 4 for locations), the bell size of A. aurita within the Beaulieu is much larger with a max size of 192mm compared to 150mm in the River Test and 105mm within Horsea Lake (Lucas, 1994; Lucas, 1996). This could be due to a difference in food availability as zooplankton were abundant within the Beaulieu and under the presence of A. aurita were not fully depleted, whereas within the River Test and Horsea Lake they were said to be food limited particularly Horsea Lake (Lucas, 1994; Lucas, 1996). The availability of the food present within the Beaulieu could be due to the large nutrient input from the New Forest with the barrier entrance at the mouth of the river preventing immediate flushing of the nutrients (Howard et al. 1984; Martino et al. 2003). The density measurements of A. aurita for June 2012 peaked just over 3 individuals per m3, which were differing from results obtained in 2011 which showed 270 individuals per m3 during May and 13 individuals per m3 during July however the bell sizes from 2011 were indicated to be smaller than 2012 (Phillips, 2012). Compared to other studies carried out within the UK on A. aurita densities, the densities within this study are consistent with other studies as within Southampton Waters densities have been recorded from 0 8.71 individuals per m3 (Lucas & Williams, 1994; Lucas

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& Williams, 1995). But differ to other UK sites such as Horsea Lake which had a max abundance of 24.9 individuals per m3 (Lucas, 1996). Compared to other A. aurita populations found outside UK waters the Beaulieu density is considerably lower than Denmark, 300 individuals per m3, Norway, 22.3 individuals per m3, Urazoko Bay, Japan, 71 individuals per m3, (Reviewed in Lucas, 2001). 4.3. A. aurita and zooplankton interactions As the ebbing tide population was not found through net sampling due to the effects of salinity, zooplankton density could be used as a proxy for the presence of A. aurita which could help locate the ebbing tide population. The potential for this to be used as a proxy is noticeable particularly within figure 18 on the 8th and 11th June. However, due to limitations in the zooplankton net the relationship between zooplankton density and A. aurita cannot be significantly proven and therefore requires further work. The observed composition of the zooplankton is similar to Southampton waters which have been found to be 95-99% of calanoid copepods compared to 97.12% copepods observed within this study (Lucas & Williams, 1995). Furthermore the predatory impact upon the zooplankton has been described as varied, as a result the use of zooplankton as a proxy for A. aurita distributions may be unreliable due to this varied impact (Lucas & Williams, 1995). 4.4. Aurelia aurita over neap and spring cycles It was statistically proven that the density changes between neap and spring cycles whereby spring cycles are at a higher density and the population becomes more aggregated during a spring cycle and as a result of this the length of the bloom shortens, whereas during a neap cycle the bloom lengthens but becomes less aggregated. The increase in density during spring tides is consistent with previous

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studies (Kaneda, 2007). This aggregation and increase in density could be due to a longer young flood stand on spring tides where the longer pause before incoming tides would allow for A. aurita to aggregate causing higher densities in addition Southampton waters spring tides have a faster freshwater flow =0.34 where neaps = 0.03. This faster flow would cause faster changes in salinity as more water is flushed through the Beaulieu. This would condense the A. aurita as they are more prone to density driven currents than on neap tides (http://www.estuaryguide.net/pdfs/southampton_water_case_study.pdf). Furthermore the increased water flow during spring tides allows for a larger spatial range of the A. aurita particularly if they are distributed by passive currents (Doyle et al. 2007). Within the Beaulieu faster spring currents allow for access to higher parts of the river in addition A. aurita are known to selectively swim at certain stages of the tidal cycle (Hampel et al. 2003; Lucas, 1994). One theory for the observed difference between the spring and neap cycles and the increased aggregation during spring tides is that the A. aurita being sampled are within spawning season, whereby aggregation would yield higher recruitment and more successful reproduction, considering that A. aurita are not usually seen after June within Southampton (Lucas, 1994) the observed aggregation may be due to reproductive strategies and not tidally related (Hamner et al. 1994). Having suggested these reasons there appears to be little other literature to support the increased bloom size during neap tides, there is also little literature demonstrating how the spatial limits within an estuary change from neap to spring for A. aurita, however based on increased currents during spring tides within Southampton Water it is likely that their spatial range is greater than it is during neap tides (http://www.estuary-guide.net/pdfs/southampton_water_case_study.pdf).

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4.5. Vertical position Aurelia aurita A. aurita demonstrated an endogenous tidal rhythm within the laboratory experiment where they migrate to deeper depths to avoid strong surface ebbing currents towards low tide where at depth currents would be slower and less turbid on the river bed which will prevent them from being flushed out of the river this behaviour is also displayed at Roscoe Bay (Albert, 2007; Albert; 2009; Albert; 2011). They migrate to shallower depths during flood tides to make use of the faster flowing surface currents to transport themselves up the river, this vertical migration is a common feature found within plankton as a means of maintaining their position within a river such as the estuarine River Beaulieu (Kopacz, 1994; Kimmerer et al. 1998). Literature and comparison between other studies (Kopacz, 1994; Kimmerer et al. 1998) suggests that A. aurita is a vertical migrator and is further supported within this study and as a result this behaviour would be exhibited within the Beaulieu and therefore likely that A. aurita is actively trying to maintain its position within the river, due to this it has allowed the population to become isolated from the rest of Southampton waters (Mackie et al. 1981; Mills, 1983). It could be this isolation of the population that could contribute towards significant differences in bell size of other A. aurita populations found within Southampton Water. 4.6. Pulsation rate Aurelia aurita Pulsation rate is affected by temperature by lowering it with decreasing temperature (Russell, 1970) where the results show a possible rhythm of dawn/dusk. This could be due to temperature as temperature increases after dawn and decreases after dusk (Russell, 1970). A. aurita have been shown to exhibit daily vertical migration, which would indicate that there could be a dawn dusk signal present where it has

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been said that they do this in order to track copepod prey which supposedly move deeper to avoid the A. aurita. Where it is suggested that A. aurita ascend to the surface at night the opposite is observed within this study and other field observations show the same trend (Reviewed in Albert, 2011). A. aurita migratory behaviours however, are possibly quite common according to Graham et al. (2001). The literature also suggests that the pulsation rate responds to changes in food availability and presence of predators where pulsation increase with the presence of both which was observed with the introduction of live Artemia sp. (Hansson and Kultiner, 1995; Graham et al. 2001; Matanoski et al. 2004). 4.7. Limitations During the study there were some logistical limitations. It was originally planned to carry out a spatial survey to attain the spatial limits of the neap and spring blooms, with current data of the Beaulieu, which due to adverse weather conditions could not be carried out during the sampling period and during the presence of the adult medusa for the rest of 2012. In addition the T/S probe connecting cable to the hand held readout broke on the 8th and could not be used until it was repaired on the 15th. As a result no temperature salinity data could be obtained for the sampling days prior to this. Further problems were encountered with the zooplankton net where the A. aurita were able to avoid the mouth opening, in addition there were net trawls where no A. aurita were caught but they were visibly sighted within the surface waters where the net density would therefore report a density of 0, as a result the net underestimates the density measurements of the A. aurita within the River Beaulieu. Furthermore hand trawling the net along the pontoon was difficult due to water currents pushing

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the net under the pontoon, where also hand trawling may be towed at inconsistent speeds, which may cause inconsistencies in the flow meter. Lastly the net was limited to the surface 1m of the water column which for obtaining the A. aurita density data after high tide is impractical as they move deeper below the nets sampling capabilities, solutions to the majority of these problems could be solved using a boat to trawl the net which can also sample a larger amount of water. In order to prevent the A. aurita from avoiding the net the consideration of a larger mesh size could be used if zooplankton samples are not required. In order to sample at greater depths attaching a weight to the net mouth may solve this, or using vertical net trawls at frequent 5 minute intervals may solve this issue. Other limitations within the study was the fact that there was no replication within the zooplankton microscope samples, which was due to time consumption and 48 samples to process which contained thousands of zooplankton per 10 ml subsample. However, ideally more zooplankton samples could be taken which would allow for a statistical comparison between their density and the A. aurita density providing the underestimations caused from the net are mostly solved. In addition zooplankton identification may always contain sources of human error through misidentification and lack of experience when identifying zooplankton. Inaccurate counts may also arise if the samples are too dense so dilution is needed to avoid this. With regards to the laboratory work carried out on the 28th only 11 A. aurita were collected on the 27th and retained as it was not possible to retain more, where this was the last sampling day also it was not possible to retain more after the 28th for the laboratory work. This was a limitation as only a couple of individuals could be used per experiment resulting in a low sample size. As a result the trends seen from

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the laboratory experiments could be minor trends amplified to major trends due to individual A. auritas influence on the experiment. Therefore larger sampling size for the laboratory experiments is needed to draw more accurate conclusions to definitively prove/disprove and trends seen. Lastly as there was a change after feeding for pulsation rate and vertical position the A. aurita may need more frequent feeding to exclusively look for tidal rhythms. 4.8. Future recommendations Opportunities for future research identified within this study include; obtaining current flow data for the River Beaulieu using an ADCP or current flow meter, and a spatial survey could be carried out to obtain the upper spatial limit of the A. aurita population and the lower spatial limit within the River Beaulieu, which could be achieved by following the population up river in a shallow draft boat using a GPS and frequent net trawls to confirm presence or absence of the population. The tidal data provided from this study would recommend that to find upper river limit, sampling would continue for up to 5 hours after high tide and to find the lower river limit it would be obtained by following the population after the second high tide which occurs approximately 5 hours after high tide where the population would be followed down river. Additionally to this further research could consider extending the sampling period to look at the spring- neap cycle over a period of months to look for seasonal signals as the majority of A. aurita literature focuss towards this research area, undertaking such a project would also solve limitations caused by low sampling size and replication as this could be corrected with such a study. Furthermore additional studies investigating the vertical position, pulsation rate and salinity adaptation of A. aurita using a larger sample size with more frequent feeding

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periods given during the experiments would be ideal. Considering food availability as a new variable is recommended. Additionally the experiments need to be run for longer period of time that persists further into the night to look for a stronger dawn/ dusk signal. Lastly the polyp distribution of the River Beaulieu is poorly understood in terms of its spatial setting and location. The River Beaulieu used to contain oyster fisheries and therefore could contain large amounts of oyster shells providing suitable substrate for A. aurita to settle on (Rodhouse, 1979). An investigation to discover if the A. aurita are settling upon these or using the pontoons located at bucklers hard would be an important step in better understanding the population dynamics of A. aurita located within the River Beaulieu. To achieve this benthic grabs could be used to collect samples at different points along the river or scuba divers could carry out spatial surveys. 4.9. Implications The implications of this study provide description of population dynamics of A. aurita within the River Beaulieu over a short neap-spring tidal cycle; which within current literature has not been addressed at this scale. The literature currently available focuss towards larger timescale studies comparing the population dynamics over seasonal trends. As a result the implications of this study could provide a good baseline to studying A. aurita population dynamics over large tidal cycle periods to look for further neap-spring differences which have been proven within this study to exist. Further implications of this study provide a detailed description of tidal and physical dynamics relating to the A. aurita population within the River Beaulieu where previously no literature has been published on this population. Furthermore

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the following study also adds valuable population dynamics data to the studies carried out within the surrounding Southampton Waters, which will allow for future studies to draw comparisons and assess changes occurring within the environment, food webs and distribution of A. aurita.

5.0.

Conclusions

Null hypotheses (1) can be partially rejected as salinity effects the distribution of A. aurita but there is not enough evidence to reject the null hypotheses for temperature and its effect on A. aurita. Null hypotheses (2) can be rejected as the distribution and dynamics of A. aurita does change between a neap and spring cycle. Null hypotheses (3) cannot be fully rejected as low sampling size limits reliability however, observed results indicate there is a change in behaviour over a tidal cycle and this can be conditionally rejected with further work still required. 6.0. References

Albert, D.J., 2007, Aurelia labiata medusae (Scyphozoa) in Roscoe Bay avoid tidal dispersion by vertical migration, Journal of Sea Research, 57, pp281287. Albert, D.J., 2009, Aurelia labiata (Scyphozoa) jellyfish in Roscoe Bay: Their Spatial distribution varies with population size and their behaviour changes with water depth, Journal of Sea Research, 61, 3, pp140-143.

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Albert, D.J., 2011, Whats on the mind of a jellyfish? A review of behavioural observations on Aurelia sp. jellyfish, Neuroscience & Biobehavioural Reviews, 35, 3, pp474-482. Arai, M.N., 1997, A functional biology of Scyphozoa., Chapman & hall, London. Aronson, R.B., S. Thatje, J.B. McClintock, K.A Hughes, 2011, Anthropogenic impacts on marine Ecosystems in Antarctica. Book Series: Year in Evolutionary Biology. Annals of the New York Academy of Sciences, 1223, pp82- 107. Atkinson, L.P., Blanton, J.O., 1986, Processes that affect stratification in shelf waters, Coastal and Estuarine Sciences, 3, pp117-130. Brusca, R.C., Brusca, G.J., 2003, Invertebrates: Chapter 8 Cnidaria 2 nd edition, Sinnaeur Associates, Inc, Sunderland, MA, pp219-268. Claus, C., 1877, Studien uber Polypen und Quallen der Adria I. Acalephen (Discomedusen), Denkschriften der Kaiserlichen Akademie der Wissenschaften , 38, 64, pp1-11. Costello, J.H., Colin, S.P., 1994, Morphology, fluid motion and predation by the scyphomedusa Aurelia aurita, Marine Biology, 121, pp327-334. Daskalov, G.M., 2002, Overfishing drives a trophic cascade in the Black Sea, Marine Ecological Progress Series, 225, pp53-63. Doyle, T.K., Houghton, J.D.R., Buckley, S.M., Hays, G.C., Davenport, J., 2007, The broad-scale distribution of five jellyfish species across a temperate coastal environment, Hydrobiologia, 579, pp29-39.

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