Quantification of variability in marcofauna of pre-tsunami sedimentary tropical shores of SW Thailand

By James Fay MSci Marine Biology 2012

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Abstract This report aimed to assess the variability of macrofaunal species at the sites: LSon = Laem Son; TND = Thung Nang Dam; KRa = Ko Ra; Kb = Krabi. Off of the coast of SW Thailand where it then quantified this variability by analysing the data using MDS plots, CLUSTER plots of abundances of the macrofauna at each station first with singleton results and then without singleton results. This showed that the removal of singletons decreased the similarity between the sites most likely due to the removal of a species that was common at another site. It also showed that from the MDS plots the site Kb was particularly dissimilar to the other three sites TND, LSon and KRa. The results then looked at the feeding mechanisms of the species to find a small variation between the sites which was attributed to each site having a different habitat. This was also seen in the last result which showed the sites were variable with regards to having vegetation present as only some sites had vegetation present. This was identified to be sea grass. However further analysis showed that the habitats varied even more due to physical factors where internal waves were present and the possibility of patchiness of the vegetation. Furthermore different levels of nutrient inputs influenced the productivity of each site causing more variation. These contributing factors lead to unique habitats at each site which was found to be the main cause in the variation as the habitat influences the species present at the site.

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Contents Front cover Abstract Contents Introduction Methods Results Discussion References Page Page Page 1 2 3

Pages 4-6 Pages 6-7 Pages 7-15 Pages 16-20 Pages 20-23

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Introduction The importance of quantifying variability of an ecological system is an important study which is needed to understand processes which are occurring that cause various patterns to arise. By quantifying this variability it is possible to construct models which can predict patterns and responses of various anthropogenic and natural inputs to a system. From these models it is also therefore possible to form new hypotheses (Ysebaert & Herman, 2002). By quantifying variations it is possible to construct food webs by looking at the interactions between different sites. This also makes it possible to implement management protocols in order to maintain and sustain ecosystems biodiversitys which are of growing interest with the potential to discover new patterns (Landres et al. 1999 & Barrio Frojan et al. 2005). Within this study the area to be studied is a series of sites located off of the coast of SWThailand within the Andaman Sea shown by (figure 1). The reason for choosing this site for quantifying variability is because there is minimal previous work been carried out on this area before (Barrio Frojan et al. 2006). Where understanding the variability of a system is important in understanding its processes and makes it a scientifically important site (Ysebaert & Herman 2002). Furthermore it is an important location for this study to be carried out seeing as it is an important fishing ground and undergoes a biannual monsoon season creating a unique ecosystem through large seasonal variations (Buranapratheprat et al. 2002 & Nielsen et al. 2004). Other reasons for studying the SW Thailand coast is due to its strong tidal actions which makes suspended particulate matter available causing high productivity and a site of biological interest (Umezawa et al. 2009).

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Figure 1. Map to show study area where data was collected at the stations, LSon, KRa, TND and Kb where LSon = Laem Son; TND = Thung Nang Dam; KRa = Ko Ra; Kb = Krabi (Barrio Frojan et al. 2006). From (figure 1) you can see the locations of the sites where the data was collected the sites are spread over 200km of coastline and are geographically separated as a result of this. Along the coastline mangroves and sea grass are present which provide sediment stability and help dampen the effect of tsunamis. All of the sites within the study have a semi-diurnal tidal cycle (Barrio Frojan et al. 2006). The reason for quantifying macrofauna off of the coast of Thailand is because they are a very important part of marine food webs. Where monitoring them is important for monitoring a system (Ysebaert & Herman, 2002). Macrofauna also make good indicator species of changes to a system due to their constant presence within the system due to various macrofaunal species having sedentary lifestyles. Therefore by studying the macrofauna of Thailand it is a good indicator towards changes occurring such as pollution and will also give an idea what
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will be happening to the ecosystem as a whole which will aid in management (Ababio et al. 1999). Macrofauna are also have important ecological effects on the environment through their sediment interactions due to some species having burrowing lifestyles as burrows increase microbial productivity (Kristensen and Kostka 2004). Furthermore macrofauna can also be classified into bio-turbators and bio-stabilisers where they either weaken or strengthen the sediment respectively via burrowing or foraging motions, this has a major impact ecologically as it influences the habitat they live in (Volkenborn et al. 2009). For these reasons this study will concentrate on the variability of macrofauna as they are of important ecological significance. The aims of this study is therefore to look at the variability of macrofaunal species between the sites Kb, LSon, KRa and TND with an emphasis on polychaete species where it is then to identify any variabilitys and differences found between these sites using statistical analysis. The aim is to then quantify the variability in order to understand why these sites differ from each other in order to understand any patterns that may occur. To hypothesis the sites will be varied from one another as they are located within different areas and will have different processes acting upon them. On the other hand the stations within the sites will be less varied as they will have the same processes acting upon them. Methods The data was collected from the methods of (Barrio Frojan et al. 2006). This data was then analysed using Primer where MDS plots and CLUSTER plots were constructed to look at the similarities between each station of the study area. The data was also re-analysed after removing species that only appeared once at a station so the more abundant species could be compared more closely. Furthermore as polychaetes were the primary subject of the study and also most abundantly found the results were also analysed without the other species found so the stations variability can be compared with respect to just polychaetes. Whilst constructing graphs for where singleton species had been removed for polychaete and crustacean species the stress level of the graphs were too high and therefore not reliable enough to be used as a result. In addition to looking at the variability of species found. The feeding mechanisms of each species found were identified and compared using MDS and CLUSTER plots and a stacked bar graph to look at breakdown of feeding types at each station which would reveal variations attributed to feeding. Lastly the seabed

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habitat was identified for each station and compared comparatively within a table in order to identify variation at a habitat level. Results After analysing the results collected off the coast of Thailand the graphs showed the following variations between the different sites of macrofaunal species:
Sp e c i e s _ Ab u n d a n ce

Transf orm: Log(X+1) Resemblance: S17 Bray Curtis similarity

Kb1a_C2 Kb1a_C1 Kb1a_B1 Kb2a_A1 Kb2a_B1 Kb1a_B2 Kb1a_A2 Kb2a_A2 Kb2b_A4 Kb2b_C4 Kb2a_C2 Kb2b_C3 Kb2b_B3 Kb2a_B2 Kb2a_C1 Kb2b_A3 Kb2b_B4 Kb1a_A1

2D Stress: 0.24

Site
TND(N) TND(S) KRa(N) KRa(S) LSon1 LSon2 Kb1a Kb1b Kb2a Kb2b

TND(S)_C2 TND(S)_A1 TND(S)_B1 TND(N)_C3 TND(S)_B4 TND(N)_B3 TND(S)_B3 TND(N)_B4 TND(S)_A2 TND(N)_A3 TND(N)_B1 KRa(N)_C1 KRa(S)_C4 KRa(N)_A1 KRa(N)_B1 TND(N)_C1 KRa(S)_C1 KRa(S)_B2 KRa(S)_A2 KRa(S)_A3 KRa(S)_B3 LSon1_C1 KRa(N)_C4 KRa(N)_C3 KRa(N)_A3 LSon2_C2 LSon2_C3 LSon1_A4 LSon2_B4 LSon2_B1

Kb1b_B3 Kb1b_C3 Kb1b_A3

Kb1b_A4 Kb1b_C4

Similarity
24 40 60

Kb1b_B4

LSon1_C4 LSon1_B2 LSon2_C1 LSon1)_B3 LSon1_C3 LSon2_A1

Figure 2. MDS plot of polychaete and crustacean species abundances at each site including singleton species with similarity rings in (%) to show how similar each site are from each other. From (figure 2) the variation between different sites is very high as they are only 24% similar and there is a larger variation between Kb and the other sites TND, LSon and KRa, therefore a larger difference between them. However the stations at each site were more closely related as they have a higher similarity shown by the 40% similarity ring. This shows that there is still a high amount of variation within each site of the study area. As a result when quantifying the variation there are site specific and station specific variables that need to be explained.

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Sp e c i e s _ Ab u n d a n ce

Transf orm: Log(X+1) Resemblance: S17 Bray Curtis similarity

Kb1a_A1 Kb1a_B1 Kb1a_C2 Kb1a_A2 Kb1a_B2 Kb1a_C1

2D Stress: 0.23

Site
TND(N) TND(S) KRa(N) KRa(S) LSon1 LSon2 Kb1a Kb1b Kb2a Kb2b

Kb2a_A1

Kb2a_A2 Kb2a_C2 Kb2a_B1 TND(S)_C2 TND(S)_A1 TND(S)_B1 TND(N)_B3 TND(N)_C3 TND(S)_B4 TND(S)_B3 TND(S)_A2 TND(N)_B4 TND(N)_A3 TND(N)_B1 KRa(S)_C4 KRa(N)_C1 TND(N)_C1 KRa(S)_C1 KRa(N)_B1 KRa(S)_B2 KRa(S)_A3 KRa(S)_A2 KRa(S)_B3 KRa(N)_C4 KRa(N)_C3 Kb2a_B2 Kb2b_B3 Kb2b_C3Kb2a_C1 Kb2b_C4 Kb2b_B4 Kb2b_A4 Kb2b_A3 Kb1b_B4 Kb1b_B3 KRa(N)_A1 Kb1b_C3 Kb1b_A4

LSon1_C1 LSon2_C2 Kb1b_A3 Kb1b_C4

Similarity
24 40 60

LSon2_B4 KRa(N)_A3 LSon2_B1 LSon2_C3 LSon2_C1 LSon1_A4

LSon2_A1 LSon1_C4

LSon1_C3 LSon1_B2 LSon1)_B3

Figure 3. MDS plot of just polychaetes at each station including singleton species with similarity rings in (%) to show how similar each site are from each other.

Within (figure 3) where crustacean species have been removed when comparing to (figure 2) it shows a similar pattern of variation between sites at 24% similarity and also between stations at 40% similarity. However it has also increased similarity between some stations for example station LSon2_A1 is now more similar to its surrounding stations, yet it has also decreased similarity between other LSon stations. This indicates that there is a strong crustacean presence at LSon which is causing variation between the sites and stations.

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Adundance
Transf orm: Log(X+1) Resemblance: S17 Bray Curtis similarity

Kb1a

2D Stress: 0.08

Site
TND(N) TND(S) TND KRa(N) KRa(S) KRa LSon2 LSon1 Kb1a Kb1b Kb2a Kb2b

Kb2a Kb2b

TND(N)

TND(S) TND KRa KRa(S)

KRa(N)

Kb1b

Similarity
LSon2 LSon1

37 41 60 68

Figure 4. MDS plots of total abundances at each site for just polychaete species with singleton species with similarity rings in (%) to show how similar each site are from each other.

Looking at (figure 4) which looks at the polychaete abundances at each site as a whole it shows that there is a large variation between the Northern, Central and Southern sites particularly Krabi. It also shows that the sites are more similar if within the same area for example TND (N), TND and TND(S) are reasonable similar with 60% similarity this is also true for KRa sites. Whereas the other stations LSon and Kb show this trend to a lesser extent. This shows that there is a geographical difference between the Northern, Central and Southern sites that is causing some variation which will be looked at within the discussion.

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Adundance

Group average
Transform: Log(X+1) Resemblance: S17 Bray Curtis similarity

Kb1b Kb1a Kb2a Kb2b KRa(N)

Samples

KRa(S) KRa TND(N) TND(S) TND LSon2 LSon1 100 80 60 Similarity 40 20

Figure 5. CLUSTER plot of site total abundances of polychaetes with singleton species with a 50% slice.

The similarity between each site shown in (figure 5) also shows the differences between the Northern, Central and Southern sites as shown in (figure 4). The TND and KRa Central sites are more closely related together, the Southern Krabi sites are more closely related to each other and the Northern LSon sites. From the above figures it shows that there are clear site specific variables causing variation.

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Abundance
Transform: Log(X+1) Resemblance: S17 Bray Curtis similarity 2D Stress: 0.22

Site
TND(N) TND(S) KRa(N) KRa(S) LSon1 LSon2 Krabi 1a Krabi 1b Krabi 2a Krabi 2b

Similarity
20 50

Figure 6. MDS plot of polychaete abundances at each station without singleton species with similarity rings in (%) to show how similar each site are from each other.

By removing species that only appeared once at a site or station the species that form the majority of a sites population can be compared and as a result can be more closely compared. from (figure 6) looking at polychaete species by removing the singletons it has decreased the similarity between sites as the Northern, Central and Southern sites have decreased similarity between each other. Furthermore comparing between stations at each site their similarity has also decreased. This shows that the addition of singleton species causes less variation between sites and stations, but also that the majority of the population present at each site is more varied between each other which by removing the singleton results have revealed this result.

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100% 90% 80% 70% 60% 50% 40% 30% 20% 10% 0% TND(N) TND(S) TND KRa(N) KRa(S) carnivore KRa LSon2 LSon1 Kb1a Kb1b Kb2a Kb2b surface deposit feeder deposit feeder filter feeder carrion feeder

Figure 7. The percentage of each feeding type found at each site shown comparatively against other stations (Fauchald & Jumars, 1979).

Analysing the feeding type (figure 7) shows that there is some variation with feeding type at each site as the Central cites TND and KRa have a higher percentage of deposit feeders than the other stations. However they vary from each other as TND has a larger percentage of carnivores than KRa. Yet KRa has a higher percentage of surface deposit feeders than TND. When looking at the LSon Northern sites has the largest percentage of carnivores present and has low percentages of other feeding type species. It also has a small percentage of filter feeders present as well. And lastly the Krabi Southern sites have a consistent carnivore percentage but have a high percentage of surface deposit feeders and deposit feeders. From these variations in feeding type between each site this could explain some of the variation between the sites. However as they are still very similar as shown by (figure 8) it cannot account for all of the variation and there are other factors causing the variation between the sites.

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Abundance Transform: Log(X+1) Resemblance: S17 Bray Curtis similarity

Kb1b

2D Stress: 0.11

Kb1a TND(N) TND(S) Kb2a LSon1 KRa(N) KRa(S) TND LSon2 Kb2b KRa

TND(N) TND(S) TND KRa(N) KRa(S) KRa LSon2 LSon1 Kb1a Kb1b Kb2a Kb2b

Similarity
85 90 95

Figure 8. MDS plot to show how similar the variation in feeding types between stations is using similarity rings in (%) to show how similar each site are from each other (Fauchald & Jumars, 1979).

Within (figure 8) it shows the similarity of each site with regards to each feeding type found at each site for polychaete species which shows that the sites are very similar with similarity between all sites above 80% as shown by (figure 9). It also shows there is still a larger difference between the Southern Krabi sites and the other Northern LSon and Central TND and KRa sites. This could account for some of the large species variations between Kb and the other three sites TND, KRa and LSon.

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Abundance

Group average
Transform: Log(X+1) Resemblance: S17 Bray Curtis similarity

Kb1a Kb1b Kb2b TND(N) TND(S)

Samples

TND KRa(S) Kb2a LSon2 LSon1 KRa(N) KRa 100 95 90 Similarity 85 80

Figure 9. CLUSTER plot showing the similarity of different feeding types at each station (Fauchald & Jumars, 1979).

From (figure 9) it further reinforces the high similarity shown by (figure 8) and that there is a site variation with regards to feeding type which could account for some of the variation of the study area of Thailand. It also shows that the variation between the Northern, Central, Southern site is the greatest. Also as the similarity of the feeding types is very high it means that the feeding type does not explain all of the variation between the sites and is therefore only a contributing factor.

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Station TND (N) TND (S) KRa (N) KRa (S) LSon1 LSon2 Kb1a Kb1b Kb2a Kb2b

Habitat Vegetated Vegetated Non-Vegetated Non-Vegetated Non-Vegetated Vegetated Vegetated Non-Vegetated Vegetated Non-Vegetated

Sand (%) 71 62 60 64 55 41 60 93 60 56

Table 1. Habitat state and sand coverage at each station (Barrio Frojan et al. 2006).

Within (Table 1) it shows what type of habitat is present at each site and the sand coverage of the seabed. From which you can see that the habitat state is variable as Vegetative state is regardless of sand coverage for example KRa (N) is not vegetated but Kb2a is vegetated where both have a sand coverage of 60%. However as the habitat state is variable between sites this could explain some variation in species found. The habitat state variation is most likely due to some other variable acting upon the system. Also when comparing the vegetative state against feeding type. Sites that have vegetation have a larger percentage of the more dominant feeding type and seem to account for a larger percentage of the feeding types present at a station. To summarise the results there is a large difference between the sites particularly between Kb and the other three sites TND, KRa and LSon. Also by removing the crustaceans it decreases the variation as and the sites become more similar. But removing the singletons increases variation between the sites. The stations within each site are also more similar to each other than compared to other station from other sites. There is also a variation of feeding type between the sites but are still very similar. And lastly the vegetative state varies from site to site with different coverages of sand. Furthermore the most common species of polychaetes to be found are from the families: Nereididae, Spionidae, Capitellidae and Paraonidae (Barrio Frojan et al. 2006).

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Discussion Analysing the results there are many different potential causes to the large variation between each site seen within the results which could be attributed to physical, chemical or biological factors. Furthermore there may be some anthropogenic influences that could cause these variations. In addition interactions between all these may produce further variations. Firstly possible variations that may be quantified by physical processes could be the presence of solitons caused when the waves hit the Nicobar Islands causing internal waves which allow the potential for high productivity. This causes turbulence and shear which has a potential of mixing and exposure to strong wave action which is documented on nearby beaches of Phuket Island (Dexter, 1996). However due being on the other side of Phuket island Krabi is sheltered from this and therefore is not exposed to these unlike LSon, TND and KRa. This therefore allows for some potential variation between these stations. Furthermore LSon is more exposed as it is not as sheltered as TND or KRa which are more sheltered by the island Ko Phra Thong (Nielsen et al. 2004). Furthermore Krabi is within a mangrove area which reduces water currents making it a sheltered area which is also similar for TND which has allowed the growth of sea grass and vegetation which may quantify for variation between the sites (Chansang and Poovachiranon 1994 & Cochard et al. 2008). Due to the sheltered nature of Krabi compared to the other stations which are more exposed via the internal waves this has implications towards larval dispersal as the larvae will not disperse as well compared to the other stations and therefore species at Krabi may be different compared to the other stations as they become more isolated (Beu and Kitamura, 1998). This could explain why the other sites are more similar compared to Krabi. Chemical processes could also quantify some of the variation between the sites as Thailand has a large amount of pig, poultry farm, fish ponds and paddy fields which causes a high amount of nutrients to run-off into the estuaries of Thailand which pollute the surrounding areas (Buranapratheprat et al. 2002). Where also industrialisation and use of pesticides has caused an increase in mercury contamination within the Andaman sea (Cheevaporn and Menasveta, 2003). Through different levels of nutrient inputs from different estuaries can cause variation between the sites through different levels of productivity. The different levels of nutrients available to the sites are also different if vegetation is present as the bioavailable carbon is much higher where vegetation is present compared to sites where no vegetation is present. As a result of sites with vegetation present more nutrients are available and therefore
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have a higher productivity allowing a larger carrying capacity for that site which will allow more species to exist and as a result have a higher diversity. This higher diversity will cause variation between the sites and could be a possible reason for the large difference seen between the sites (Duarte et al. 2005). Furthermore because Krabi is more sheltered it could cause eutrophication through high residence times and poor tidal flushing (Cheevaporn and Menasveta, 2003). In addition to that there is also a large amount of sewage discharge from Phuket which would fuel eutrophication (Reopanichkul et al. 2010). In addition due to the other sites being more exposed to wave action and in a tsunami vulnerable area they are prone to backwash where sediment is transported via tsunamis and causes large amounts of Suspended particulate matter to become available (Sugawara et al. 2009). In addition potential biological influences that could cause the large variations observed could be due to different predators present at different site which would cause different food webs to emerge and as a result cause different species to occur at different sites causing variation between the sites. This would arise through a variation of habitat caused by the above physical and chemical factors which cause different habitats to occur. For example the sheltered nature of Krabi surrounded by mangroves and some of the stations vegetated by sea grass provide a more complex habitat allowing prey items to hide more efficiently as a result predators adapted to hunting in vegetation would arise here. This would also be similar at TND but because it is more exposed a different predator adapted to hunting in more exposed conditions would arise here. Then compared to the sites like LSon and KRa which are exposed and low in vegetation predators adapted to catching prey items that dont have the option of hiding in vegetation would be found here as the habitat complexity would be lower. Through different levels of habitat complexity i.e. vegetated with sea grasses or nonvegetated will cause different predators which causes variations between the sites (Beukers & Jones, 1997). In addition the variation in habitat makes different niche environments available where different species will arise suited to their surrounding environment for example the presence of vegetation at the sites which is known to be sea grass (Barrio Frojan et al. 2006). The presence of sea grass provides a three dimensional habitat for macrofauna as they can feed amongst the sea grass and not just on the sediment as a result it provides a larger habitable environment for species to live in which will increase the diversity of the site containing sea grass. On the other hand the sites without sea grass will be restricted to feeding on the sediment and therefore be less diverse as the habitable environment is less without sea grass
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(Ansari et al. 1991). Which as shown within the results there are sites with sea grass and without sea grass present. Through the presence of different habitats this therefore causes variation between the sites. Furthermore from the seabed coverage of sand all sites had sand present which would indicate that the sea grass is most likely patchy which could explain the variation between the stations of sites that contained sea grass (Wiens, 1997). Furthermore patchiness also creates two habitats within the same area which would increase diversity further and allow new species that can utilise both the sea grass habitat and the exposed seabed environment which would cause greater variation between the sites depending on the patchiness of sea grass (Wiens, 1997). This could account for the differences between the Northern, Central and Southern areas. Where the Northern site is non-vegetated and exposed. The Central sites are both vegetated and non-vegetated and semi-exposed and the Southern Krabi site is both non-vegetated and vegetated but sheltered. With different habitats present at each location different species will occur and cause variation between the sites and could be the reason why the sites are so different from each other. One potential cause of variation between the sites could be due to the presence or absence of mangroves at a station as it is documented that mangroves are along the coast line (Barrio Frojan et al. 2006) which provide a different habitat for species to thrive in. however near Ranong the mangroves have been commercially exploited which is near the site LSon which is an exposed area. The removal of the mangroves from this area may have caused variation of LSon from the other sites by removing a habitat for prey items to hide amongst. This could explain why LSon has a large proportion of carnivores present compared to the other sites (Macintosh et al. 2002). Another biological influence that could be a contributing factor towards the the large variation between the sites is the feeding type of the species found at the sites. Due to the different habitats different food sources are available and therefore different feeding methods are used in order to obtain that food which causes variation as shown in the results. These different feeding mechanisms arise as a result of the habitats being different which as a result mean different species are present (Fauchald and Jumars, 1979 & Gutierrez and Iribarne, 2004). Looking at the results section other potential reasons for the large variability between stations could be due to the presence of crustaceans as well as polychaetes. As within the singleton section where crustaceans have been added the sites become more dissimilar from each other
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but when removed become more similar to each other. This shows that basing the similarity of a site based on just polychaetes is not enough to quantify the full extent of variability at a site as when adding other classes of species the variation increases between the sites as interactions between different species becomes more noticeable as species that are usually associated with another species will occur near them. After taking into account all the species that appear at each site there will be even more variation as it cascades through different trophic levels (Southwood, 1977). When the singleton species were removed it showed that the sites became more dissimilar indicating that the singletons reduced variability. This is most likely due to the singletons being present at more than one site and therefore by removing them it removes a common factor between sites and therefore decreasing similarity. Also it was documented from previous studies using the same dataset that only one species occurred at all stations and that 33% of the species found only occurred at one site therefore by removing singletons it would mean that there would not be the same species present at every station and unlikely that 1 species would appear at multiple stations as a result removing singletons causes more variability between the sites (Barrio Frojan et al. 2006). The possibility why the removal of a singleton species may decrease similarity is because if at one site a species is abundant but at another site it is a singleton, the presence of that singleton makes it more similar to the site where the same species is abundant. By removing it removes the link between the sites and makes them more dissimilar. On the other hand it would improve similarity if rare species were removed that only occurred at one site as it would remove a difference between sites. However as similarity decreased it would imply that singletons found and removed were most likely present in more abundance at other sites rather than being rare. This has most likely occurred due to larval dispersal where a few larvae have left the Krabi site and become singletons at other sites or vice versa where larvae have been carried into Krabi this would explain how you could have an abundant species at one site and then a singleton species at another site (Beu & Kitamura, 1998). Another source of potential variability is the genetics of species at each site. Where Krabi is more isolated there is the potential for limitation in the gene pool as conditions are more stable due to being sheltered. This would impact the site by reducing the fitness of individuals and whilst having a large species diversity there may be a low genetic diversity as a result of low larval dispersion. As a result at the other stations there may be smaller species diversity due to more exposed conditions but a greater genetic diversity and therefore a higher
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individual fitness due to a greater larval dispersal potential (Bell and Okamura, 2005 & Bulleri and Chapman, 2010). However this has not been tested and would be ideal for future study in order to test this as a hypothesis. Another potential topic for future study would be to look at the larval dispersal from each site as this data would reinforce or disprove the gene pool limitation hypothesis. Furthermore it would also further investigate the presence of singletons and their implications on variation between the sites. To conclude the main variation between the sites can be attributed to different habitats at each site which allow different species to occur as they occupy different niches created by the habitat causing the variation. However the variation in habitats is caused by variation of various contributing factors such as physical, chemical and biological interactions and anthropogenic inputs which create these different habitats and allow the wide variation of macrofaunal species to occur. References Ababio, T.K.G., Furstenberg, J.P., Baird, D., Vanreusel, A., 1999, Nematodes as indicators of pollution: a case study from the Swartkops River system, South Africa, Hydrobiologia, 397, pp155-169. Ansari, Z.A., Rivonker, C.U., Ramani, P.,Parulekar, A.H., 1991, Seagrass habitat complexity and macroinvertebrate abundance in Lakshadweep coral-reef lagoons, Arabian Sea, Coral Reefs, 10, pp127-131. Barrio Frojan, C.R.S., Hawkins, L.E., Aryuthaka, C., Nimsantijaroen, S., Kendall, M.A., Paterson, G.L.J., 2005, Patterns of Polychaete Communities in Tropical Sedimentary Habitats: a Case Study in South Western Thailand, The Raffles Bulletin of Zoology, 53, 1, pp.1-11. Barrio Frojan, C.R.S., Kendall, M.A., Paterson, G.L.J., Hawkins, L.E., Nimsantijaroen, S., Aryuthaka, C., 2006, Patterns of polychaete diversity in selected tropical intertidal habitats, Scientific Advances in Polychaete Research, 70,3, pp239-248. Bell, J.J., Okamura, B., 2005, Low genetic diversity in a marine nature reserve: re-evaluating diversity criteria in reserve design, Proceedings of The Royal Society, 272, pp1067-1074.

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