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Spring 2017 The iF sh Community of Misali Island: Recommendations for Conservation Management and Implications for Mixed-Use Conservation Areas Stuart Jones SIT Study Abroad

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Recommended Citation Jones, Stuart, "The iF sh Community of Misali Island: Recommendations for Conservation Management and Implications for Mixed- Use Conservation Areas" (2017). Independent Study Project (ISP) Collection. 2617. https://digitalcollections.sit.edu/isp_collection/2617

This Unpublished Paper is brought to you for free and open access by the SIT Study Abroad at SIT Digital Collections. It has been accepted for inclusion in Independent Study Project (ISP) Collection by an authorized administrator of SIT Digital Collections. For more information, please contact [email protected]. The Fish Community of Misali Island: Recommendations for Conservation Management and Implications for Mixed-Use Conservation Areas

Stuart Jones

Advisor: Dr. Narriman Jiddawi

Academic Director: Dr. Richard Walz

SIT Spring 2017

Zanzibar: Coastal Ecology and Natural Resource Management

Table of Contents:

Acknowledgements……………………………………………………………………3

Abstract…………………………………………………………………………………4

Introduction……………………………………………………………………………..5

Site Selection…………………………………………………………………………..9

Methods………………………………………………………………………………...14

Data……………………………………………………………………………………..17

Discussion……………………………………………………………………………...25

Recommendations…………………………………………………………………….29

Conclusion……………………………………………………………………………...35

Bibliography…………………………………………………………………………….37

Fish List…………………………………………………………………………………39

Acknowledgements:

This study would not have been possible without the logistical assistance, expert advice, and moral-boosting support from a number of people on the ground in Zanzibar. First I’d like to thank

Said, Richard, and the SIT office staff for their support throughout my time in Zanzibar, and Dr.

Richard, Dr. Jiddawi, and Dr. Richmond for their advice on picking, planning, and executing this project. I’d also like to thank the Misali park rangers, Mohammed, Juma, and Hamiss, for their welcome, their generosity, and their support during our time on Misali. We couldn’t have survived on that island for a week without them, and I everything I was able to accomplish to their help and kindness. Special thanks to Jeremy for bravely watching over me during transects from your black rubber throne. You radiated warmth, safety and security that I could feel all the way under the water. Shout out to Kendrick Lamar, Adu’s brother, mangoes, and the crunchy crew for making my time on Misali so enjoyable.

Abstract:

Misali, an uninhabitated island near Pemba Island off the Tanzanian Coast, is a historic biodiversity hotspot and was at one point home to 80% of the coral in East Africa. However, it has been nearly 10 years since any formal, published study has examined the fish or coral communities around the island, and new information is needed to inform local conservationists and fisheries managers. It was discovered that, despite a severe bleaching event in 1998 with (80% mortality) and long-term budgetary neglect, Misali’s fish community has remained fairly healthy and diverse, with over 200 species of fish observed within the limited scope of the survey area. An analysis of ordinance and beta diversity also strongly suggests that the island is home to a wide variety of habitats with distinct fish assemblages, and that the fully protected but unenforced non-extraction zone (NEZ) encompasses only one small part of the diversity of the greater Misali ecosystem. Futhermore, this zone is home to the highest abundances of fish, including several important commercial families. The NEZ’s current boundaries, therefore, are not only insufficient to protect the largest possible swath of diversity on the island, but also imposing an undue burden on local fisherman by depriving them of the most productive fishing ground. If these boundaries were more stringently enforced, these regulations may also endanger Misali’s fish stocks by concentrating fishing pressure on the other areas that may have been least able to support it. This study recommends that further research be done on the biodiversity and distinct habitats within Misali, and that the boundaries of the NEZ then be redrawn to encompass more of this diversity while opening up part of the current NEZ, and its productive fisheries, to local artisanal use.

Introduction:

Coral reefs are among the most diverse and productive ecosystems on Earth, and are known around the world for their rich and colorful life and their spectacular beauty. Less appreciated, though, are the benefits and ecosystem services that coral reefs provide to local communities almost everywhere that reefs and humans coincide. In the Zanzibar Archipelago, located in the Indian Ocean off the coast of Tanzania in East Africa, healthy coral reefs form the foundation of local industries, diets, cultures, and communities. Fish from coral reefs provide employment for thousands of local fisherman and a crucial protein source to most of the island’s inhabitants. The renowned beauty of coral reefs also attracts scores of foreign divers and tourists, whose dollars and euros flow into the local economy. These crucial sources of income, employment, and resources are crucial to the health and continued development of communities from those as large and cosmopolitan as Stone Town in Zanzibar and as small and local as Wesha in southern Pemba. Without these reefs, many communities, especially smaller and poorer fishing communities, could not survive. The loss of coral reefs, even in just one area, would therefore be not only an ecological disaster, but also an economic and human disaster as well.

It is therefore of great interest to local people and governments to protect and conserve healthy coral reefs, not only for their biodiversity, but also to protect and stimulate development of local communities. In Zanzibar, numerous Marine Parks and conservation areas have been implemented in the last twenty years as this importance to locals has been realized. However, as Zanzibar and Africa in general develops, more and more coral reefs have come under threat, and many reefs with poorly enforced or absent protections have been perhaps irreparably degraded. Global climate change has played a part, with coral bleaching a widespread and ever present threat, even to protected reefs. However, the costs of the onward march of development and explosive population growth have arguably had a greater toll in the region, particularly near major population centers. More development means more waste, more foreign visitors, more coastal development, more people, more mouths to feed, more men in need of work, and more lines in the water, and in many cases reefs have been unable to keep up. Many reefs have been polluted by solid and human wastes coming from towns and cities, and overfishing and destructive fishing practices have taken their toll as catches have increased to keep pace with demand. While responsible and well-planned tourism can offer an alternative source of income and incentivize conservation, the increase in foreign visitors can also be a double edged sword.

Tourism brings more money but it also brings more trash, more pollution, and more damage from careless fins and feet.

However, while Zanzibar and Dar es-Salaam have developed aggressively over the last twenty years, many areas of the Tanzanian coast remain rural, and even unprotected reefs remain fairly healthy. One such area is Pemba Island, north of Zanzibar. While Pemba’s population has grown substantially in the last 20 years, the island still remains fairly rural, and its population is just a fraction of that of Zanzibar Island, despite the two being roughly the same size. Pemba is also subject to much lower levels of tourism than Zanzibar, as it lacks an international airport and therefore requires an extra flight after arriving in Tanzania. While there are many noteworthy hotels and resorts on the island, these tend to follow the high-cost low- number model of tourism more than the hotels and resorts of Zanzibar. In Pemba, fewer, richer, and more conscientious tourists pay higher costs, which both decreases the environmental impact of tourism through reduced volume and allows hotels and businesses more room in their finances to adopt more sustainable practices. The island is renowned for its SCUBA diving, and its coral reefs, particularly those in the Pemba Island Conservation Area, remain fairly healthy.

One such reef is that around Misali Island. Officially protected in 1998, Misali was incorporated into the larger Pemba Channel Conservation Area in 2005 (Richmond and

Muhando 2001). Misali reef is a historic biodiversity hotspot, hosting over two-thirds of all coral genera in East Africa (Horrill et al 1994), and providing a home for marine mammals, vervet monkeys, flying foxes, and coconut crabs, as well as an important nesting site for Hawksbill turtles. Despite a catastrophic bleaching event in 1998, with coral mortality from 60-90%,

(Grimsditch et al 2009, Muhando 1999), the islands reefs have bounced back well. Reefs in the

Indian Ocean as a whole have recovered well from 1998 in terms of coral cover (Baker et al

2008), but not all reefs have done so as spectacularly as Misali. A recent study estimated the percent cover of live hard coral in the no-take zone at Misali at 82% (Grimsditch et al 2009), well into the high range of the 60-90% observed in surveys taken before 1998 (Horrill et al 1994,

Mohammed et al 2000). Anecdotal evidence from local scientists and conservationists also indicates that the island remained relatively unscathed by the recent global coral bleaching events in 2010 and 2015 (Jiddawi, Richmond, pers. comm.). The lack of a hotel or permanent settlement on Misali and its location over 16 kilometers from the nearest urban area has also helped to insulate it from the effects of pollution, motorized boat traffic, and destructive tourism, and reefs have remained in good overall health.

However, despite its advantageous location, Misali’s reefs remain somewhat imperiled moving forwards. While bleaching due to climate change remains an existential threat, the more pertinent local threat may be from overfishing. While the island is designated as a Conservation

Area, large parts of the reef are still open to fishing and are critically important to local communities (Richmond & Mohammed 2001). As a result, the island sees relatively high fishing pressure, even sometimes in areas that are meant to be off-limits. It is in this struggle against overexploitation that the island’s isolation and relative lack of tourists act as a double-edged sword. While the lack of tourists or a hotel protects the reef from pollution and mechanical damage from careless fins and feet, the lack of tourist interest and money flowing into Misali, particularly in the last five to ten years, means that the island is no longer considered a funding priority for the Zanzibari government. While there are rangers stationed on the island full-time, a lack of funding and proper equipment has been a persistent problem (Daniels et al 2004).

Rangers often lack the resources, including flashlights, footwear, working boats, and petrol, to adequately patrol their waters and enforce the law (Haji, Suleiman, Walz, pers. comm.). Furthermore, the lack of marker buoys and support from mainland law enforcement means that fisherman caught in the non-extraction zone have been able to escape the law by pleading ignorance (Klein & St. Denis 2016, Suleiman pers. comm.). While the no-take zones are essentially not enforced, the rangers have largely been able to halt destructive fishing practices through social pressures and their relationship with the community, and have also halted illegal, industrial foreign trawlers from exploiting local fishing stocks (Suleiman, pers. comm.)

Additionally, while the island does see a large quantity of fisherman, much of the focus is on pelagic fisheries such as sardines and tuna, and fisherman targeting reef species almost all fish using duma traps, which are both low-volume and non-selective in the fish that they catch, meaning that the fishing pressure is spread out among a very wide variety of species (pers. obs.). As such, while the park has failed to specially protect the non-extraction zone it has succeeded in keeping fishing from becoming overtly destructive. In its current state, the fisherman employ practices and equipment similar to that which have been traditionally used, and due to Pemba’s long history of human habitation, these fisherman have become as much a part of the natural environment and the normal state of affairs as the sharks and the corals themselves (Walley 2004).

As Tanzania and Pemba continue to develop over the next generation, preserving healthy reefs like those around Misali will become both more difficult and more important.

Sustainable economic growth and development is built on a healthy environment, and the continued health of reefs like Misali’s should be an economic and social priority for the Zanzibari

Government. This study will aim to build a reasonably large, complete, and quality data set inventorying the fish community around Misali, in order to demonstrate the diversity still present at the island, and to look for ways that the island’s conservation plan might be improved. This study will also be the first in almost a decade to study the fish community around Misali, and should provide information to local fisheries managers and conservation officials about the state of the island’s fisheries, and the efficacy of current conservation practices. Site Selection:

Misali Island is a unique environment, and was uniquely suited to this study in a number of ways. First, the island’s fish community has gone completely unstudied for the past 10 years, and due to Pemba’s remoteness and the low level of tourist interest there, an official study of the island’s fish is unlikely to happen anytime soon. By examining Misali, therefore, this project could contribute crucial data and information to fisheries managers and conservationists that would not have been collected Fig. 1: Map of Misali inset into map of Pemba otherwise. As stated above, the continued health and productivity of Misali’s reefs is critical to local fisherman and local communities, so, by contributing information that could aid in their management, this project can give back to the local community. Besides the value that the study site adds, the particular topography of the island made studying Misali feasible within the limitations of this study in ways that other islands and reefs around Zanzibar would not have been. The narrowness of the reef flat, particularly on the northern end of the island, meant that

Misali’s reef slopes could be studied from the shore without the need for boat transportation.

Combined with the severe 4.5-plus meter spring tides around Zanzibar and the relative shallowness of the reefs, most all of the reef crests and upper and even lower slopes were accessible from land and could be observed using snorkel-and-dive methods without requiring the use of SCUBA. Furthermore, most of the coral growth and fish life is concentrated on the very narrow, steep reef slopes along the various tidal channels and the Pemba Channel itself.

This concentration and the overall small size of Misali allowed a one-person study lasting just a month to achieve a relatively complete inventory of the island’s reefs and get a decent idea of what is going on with Misali’s fish community. Taken together, the lack of current and future research, easy access, and small concentration of reefs allowed for a relatively brief study to make a meaningful contribution to current knowledge with limited need for expensive equipment and technological support. Misali, therefore, was the perfect site for this study.

Within the context of the broader Misali area, the individual locations at which data were taken were chosen with great care. However, all sites were subject to the contraints of safety.

While many of Misali’s reefs are easily accessible from shore, some areas have incredibly large intertidal areas or were very long swims from shore, and would require long transport times.

Furthermore, the south side of Misali has no trails, and is not accessible over land from the

Fig. 2: Map of study sites and transect locations around Misali. General study sites are indicated in Red exact transect locations in Yellow ranger station. A surveyor, therefore, who was stranded by a rapidly rising tide or a sudden storm would have to swim all the way back to the ranger camp to reach safety, a distance of well over two kilometers. These safety and access considerations concentrated the study into the more accessible reefs on the northern side.

Beyond safety concerns, transect sites had to meet a number of constraints, and each site was chosen for a very specific reason. First, because of the goals of the study, taking good data inside the non-extraction area was an absolute must in order to gain the information required to make meaningful conclusions about conservation and the state of Misali. Within the

NEZ, sites had to be concentrated in the northern end near West Island. The NEZ is the deepest and most exposed area surveyed in this study, and reaching the reef slope required a long swim through often rough, unsheltered waters, often carrying heavy and unwieldy gear.

West Island can be reached over dry land at low tide, and while the island is still a fair distance from the slope, it provided an invaluable base from which transects and gear could be launched to survey the NEZ.

Outside of the NEZ there was a bit more freedom with site selection. Data needed to be taken at sites that corresponded to sites examined in previous studies, in order to allow for the best possible comparisons. The reefs at Mbuyuni Beach have been fairly well studied by recent

ISP’s done by SIT students, and combined with the site’s proximity to the ranger station and easy accessibility from shore, this area was an easy choice for survey. However, while the

Mbuyuni site promised good comparisons to past data, it would not be as useful as a present comparison to the NEZ. Mbuyuni beach was too different in terms of exposure, depth, width of shelf, and general environment to allow it to be compared one-to-one with the NEZ, so a third location was needed. This was to be another seaward site, as similar to the NEZ as possible, but outside the protection of the park, and thus a suitable control for the effects of the no-take zone’s designation. Ideally, the seaward reef of the fully submerged reef north of the channel would have been a perfect comparison; however, this site would have been completely inaccessible without boat transportation. As a result, I chose to survey the seaward reef along the northern tidal channel, because it was very similar oceanographically to the non-extraction zone, particularly in the southern parts that are less protected by the reefs on the opposite side of the channel.

Finally, a site had to be taken on the lagoon side of Misali. Lagoon reefs represent vastly different habitats from seaward reefs, and the East side therefore represented a potentially distinct environment from all the other areas. In order to gain a more complete idea the current state of Misali’s fish community and avoid biasing the data toward species more common on seaward reefs, a site had to be taken on the eastern side of Misali. However, the challenges associated with this area were many. While the lagoon side was generally calmer and shallower than the seaward side, it was also an enormous area, and had a huge intertidal zone between the reef and the island. A swiftly rising tide could therefore strand surveyors in water too deep to stand more than a kilometer away from shore. With the storms and altered currents common during the rainy season on Misali, this could be very unsafe. Furthermore, most of the southern part of the lagoon reefs were rendered inaccessible, simply because the lack of trails on the island made the southern beaches inaccessible by land. The location of transects on the lagoon reefs therefore had to be concentrated on the areas close to shore and those most easily accessible at low tide. This meant that the transects were primarily concentrated in the northern part of the lagoon flat, with the southern reef almost completely unstudied.

As a result of limitations due to safety and accessibility, the sites and transect locations were mostly constrained to the northern half of the island, and excluded the submerged reef to the north of the lagoon channel. However, the survey area did include seaward, channel, and lagoon reefs, and covered the full span between the Pemba Channel on the oceanic side and the eastern extreme closest to landward Pemba. As a result, despite the aforeward limitations, the survey area is likely fairly representative of the greater Misali fish community, and priority was given to developing a more thorough data set at sites that were sampled rather than attempting to survey the whole island.

Methods:

The methodology of this study was designed to match that of previous studies, to ensure the most valid possible comparison to previous results (Horrill 1992, Horrill et al 1994, Mohammed and Muhando 2000, Frontier-Tanzania 2004). The biggest consequence of this was the decision to use belt-transects run parallel to shore across the reef flat and slope rather than line- transects conducted perpendicular to shore. While this decision was made to agree with previous studies, it proved also to be the best way to inventory fish communities that were almost always strongly concentrated along the narrow reef slopes. While line transects would have been more representative of the total biomass and abundance per m2 around Misali, it would have significantly reduced the amount of reef slope that was surveyed, from 700 m2 per site to only 150 m2 per site. Because the slope and the deeper parts of the reef flat contained most of the coral, and almost all of the fish and diversity, for the purposes of this study, it was more important to use belt-transects to quickly gain an idea of what fish and diversity were present, rather than focus on representative shore-to-slope biomass or abundance. For this reason, belt-transects proved to be the most prudent surveying method, even for reasons beyond the ones for which it was originally chosen.

Once the belt transect method was chosen, every attempt was made to match the standard methodology as closely as possible. However, due to the limitations of gear and technology available to this study, some small alterations had to be made. Each transect was

35m by 5m, cut down from standard 50m by 5m, in order to make transects more manageable to set up from the water and examined thoroughly within the time constraints created by the tides. Because these surveys were conducted on snorkel and reached without the aid of a boat, transects had to be done during low tide, and if transects ran too long the rising tide could make the return to shore more difficult and even dangerous. Therefore, priority was given to examining a smaller transect thoroughly rather than trying to record a larger area and having to rush. Transects were also marked by floats set out 35m apart at least an hour prior to conducting the survey, rather than laying out an underwater transect line immediately prior to surveying. This was done because many wary species of reef fish will flee or hide from human snorkelers, especially in areas where even light spear-fishing occurs. Most studies overcome this by using SCUBA and waiting, because fish will often return or re-emerge when SCUBA divers remain stationary on the bottom for long enough. Because this study could not use

SCUBA, this obstacle had to be overcome in different ways. Setting out the transects beforehand allowed surveyors to begin to inventory the fish community immediately upon arriving at the site, and since sites were accessed from the shore rather than from noisy motorboats, this afforded more than enough time to record secretive and wary fish before they had a chance to flee. Transects were inventoried in standard 5m by 5m segments, but because of the aforementioned time limits afforded by the tides, each 5m by transect was surveyed for three intervals of five minutes, rather than the standard five intervals of five minutes.

Six 35m by 5m transects were conducted at each site in total, four of which were along the reef slope, and two of which were on the lower reef flat. The only exception was the eastern study site, where most of the far eastern part of the reef had no definite steep reef slope. In this area, two ‘slope’ transects were conducted along the line of richest coral growth, and two ‘flat’ transects were conducted on the shallower side of this line.

At each site, data was recorded on a tri-fold slate featuring the names of 118 species of fish, based loosely on a list of 78 species given to volunteers in a previous study (Daniels et al

2004). Though this list was altered and added to over time as it was discovered which fish were present and abundant, all fish were recorded, regardless of whether their name was on the slate at the start of the transect. Known species not on the slate were simply recorded in the margins.

When unknown species were encountered, notes were taken on their body shape, coloration, distinctive markings, suspected family, and behavior. These species were then identified as soon as possible after the transect was completed, using the book “ Fishes” (Lieske and Myers 1996). Abundance data was recorded for all species present, and for larger and commercially important fish, such as parrotfish, goatfish, and large , the size of each individual was estimated to the nearest 5 cm and recorded. For smaller fish, such as surgeonfish, where most adults usually tend to be about the same size, and commercially unimportant fish, such as angelfish, size data was not recorded.

In addition to the 6 transects, one survey dive was conducted at each site, still using snorkel, to search for species that, because of their preferred depth, rarity, or limited range within the reef, were not recorded in any of the transects. Additionally, any unique species that could be definitively identified seen outside of the transects before and after the surveys were recorded as having been observed, but not included in transect data. These species were added to the overall species list, but not included in transect data or species richness or diversity comparisons between sites, because the sampling effort of these surveys and extraneous observations could not be adequately quantified or controlled for.

Data:

The first question to answer about this or any ecological data set is whether sampling was adequate. There are several ways to answer this by graphing and analyzing the statistical patterns within the data, and comparing them to see if they match the patterns found in all thoroughly sampled ecosystems. The first way is the Preston distribution. Preston assumes that the abundances of all species within an ecosystem follows a bell curve, with many species of intermediate abundance, and equal numbers of very rare and very common species. When inventorying species, most all very common species will be found, but only with thorough sampling will all of the rarest species be recorded. Only studies that have conducted thorough sampling and found all the species within an ecosystem, therefore, will have an equal number of very rare and very common species. Preston therefore groups all species found in a study into groups based on their abundance, and plots these groups as a histogram. If an ecosystem has been sufficiently sampled, then the Preston distribution should show an even bell curve, with even tails of both rare and dominant species. However, if sampling is not sufficient, then the left hand side of the bell curve, that representing the rarest species, will appear truncated, as sampling was insufficient to discover all of the rare species present in an ecosystem.

The Preston distribution for this study suggests that sampling was unlikely to have been sufficient to record every species present in the study area.

Fig. 3: Preston Distribution plots the number of species of high and low abundances. If study is well- sampled, graph should show a normal distribution Another method for determining whether sampling was sufficient is the Species

Accumulation Curve (SAC). The SAC assumes that, in any census, species will be discovered very quickly at first and more slowly as the study goes on until the number of species observed finally reaches the true number of species in a given community. Therefore, if the number of observed species is plotted against time, or samples collected, the graph will rise very quickly at first before slowing down and eventually plateauing once all species have been found. The graph, therefore, represents a curve, which flattens out as the observed species approaches the true species limit.

Fig. 4: Species Accumulation Curves plot the rate at which new species are discovered in a study. As Richness approaches the true Species Richness of the study area, the curve should flatten out and reach a plateau

This graph shows the study’s actual observed species richness together with two statistical estimates that attempt to show the true number of species in the community, including the species that are present but were not observed in the study. Using these estimates allows the curve to flatten out more quickly, so that it can reach the true species richness even with a lower sample size. While the observed species richness has not quite reached it’s plateau, the Chao index, which attempts to estimate the number of unseen species, has settled at around 200 species. Therefore, while this study was not able to physically observe every species in the community, the sample size of the study was sufficient to estimate the true species richness from the number of rare species observed in the available data. The total number of species in the survey area is estimated at just over 200. Species richness data were also compiled for each site, to determine whether there were substantial differences between different geographic areas. Somewhat surprisingly, all estimators showed roughly even levels of biodiversity across all sites, with the southeast flat Fig. 5 shows three statistical estimates for the true number of species in the being perhaps a fraction lower in study site terms of species richness

Fig. 6 shows the number of species observed (Species) and three estimates of the true number of species present (Chao, Jack.1, Bootstrap) for each site. The four indices show the species richness across sites to be relatively even. Fig. 7 Biodiversity was also even across the four sites

While each fish community had roughly equal diversity, this does not mean that all communities were identical. In fact, data on the species composition, the particular group of species present at each site, showed that the fish communities at each site were distinct. A technique known as ordinance is used to represent these differences visually in species composition..

Each yellow dot in the two graphs represents a transect, and the distance between two transect dots represents the difference in species composition between the two sites. The transects have been further grouped into their respective sites, with colored circles and

Fig 8. Each yellow point represents a transect, with the distance between two transects showing the difference in fish species present at each site. The four sites were relatively distinct, especially the NEZ, and Southeast Flat polygons used to approximate the overall composition of the fish communities at each site, and show how similar distinct each site is from the others. These two graphs show the fish communities at each site to be very distinct, particularly the southeast flat (SE) and the non- extraction zone (NEZ), which showed little to no overlap with the other sites.

These differences were not random but varied along with the distance of each site from the Pemba Channel, and the difference between seaward and lagoons reefs. Here, the Bray index for beta diversity, or the difference in species composition between two transects, is plotted against the difference in longitude between those same two sites, a proxy for the distance from the Pemba Channel.

Fig. 9: One factor influencing species composition of each site was the average longitude of the transects, a proxy for the distance from the Pemba Channel. This association was shown to be significant, with (p=.109, r2=.393). Distance from the

Pemba Channel, therefore, is almost certainly a factor in determining the species composition of a given reef, with clear differences between seaward reefs, those facing mainland Africa and the deep, open waters of the Pemba Channel, and lagoon reefs, those facing Pemba itself and the shallow, relatively calm waters of the bay.

However, the low R squared values (.39, .18) show that although seaward-lagoon position was certainly a factor affecting beta diversity, it explains only a small fraction of the total variance. There are therefore other, possibly stronger, variables affecting species composition.

Because this study sampled only one cross section between seaward and lagoon, it is likely that there are many more microhabitats and unique fish communities at Misali not covered by the scope of this study.

The second difference between the sites is the abundance of fish. Unlike diversity, which showed that the NEZ was not more important than the other sites, abundance was observed to be higher inside the zone than at the other sites. This was true both for fish in general and for several commercial families (Acanthuridae, Scaridae, Carangidae, Lutjanidae, etc.)

Fig. 11: While diversity varied evenly across sites, overall abundance and abundance of several commercially important families were higher in the NEZ These differences were again correlated with longitude, the proxy for each transect’s position on the seaward-lagoon spectrum. Since reef flat transects had vastly different abundances than slope transects, these were excluded for the purposes of this comparison.

Fig. 12: Fish abundance correlated strongly with longitude, with the highest abundances observed on seaward reefs Comparing both by site (p=.0047) and by transect (p=.057) yielded significant or near significant correlations, but the most important outcome is told by the Rsq values. An adjusted

Rsq value indicates how much of the variation in the y variable, in this case abundance, is explained by the x variable, in this case longitude. Longitude explained only 18 percent of the variation between sites, as there are many other stronger variables, such as coral cover, depth, etc., influencing fish abundance at that level, resulting in a weaker correlation (p=.057).

However, longitude explained almost all (Rsq=.986) of the variated between sites. Distance from the Pemba Channel, therefore, is more or less the only significant variable influencing the average fish abundance across geographic areas. This is reflected by the strength of the correlation (p=.004692).

Discussion:

While statistics show that the sampling intensity of this study fell just short of a complete inventory, it seems to have been adequate to allow us to estimate the overall species richness and make inferences about the Misali ecosystem. Data showed that all sites had approximately equal biodiversity; however, there are still important differences between the sampled communities. Ordinance and beta diversity analyses showed that the species compositions of each site varied significantly, and that the species present at each site varied according to how far the sample was taken from the Pemba Channel. Furthermore, while the NEZ was not more diverse, there were higher abundances of fish, particularly several important commercial families.

While the estimation of true species richness from this study, at around 210 species, is considerably lower than the ~300 observed in previous studies, there are a number of ways to explain this. First, it is important to note that this underestimation cannot be explained simply because of the lower sample size of this study. The chao, jackknife, and bootstrap estimators have already taken unobserved species into account, and the sample size, as indicated by the

SAC and Preston distributions, was sufficiently large and close to complete that these estimators should be reasonably accurate. What these estimators do not account for are species that went unobserved because they simply live outside the study area. While the 210 number likely represents all species, seen and unseen, within the four sites sampled, these four sites still represent a very small slice of the overall Misali ecosystem. While substantial spring tides, clear waters, and strong lungs allowed this study to inventory sites of depths up to 8-10m, there are still large areas of both the seaward and lagoon reefs that are simply too deep to survey using snorkel under any conditions. Depth is a strong factor influencing species composition on many coral reefs, and it is likely there could be dozens of unobserved species living below the depths examined by this study. Of the roughly 400 species in the species inventory created by the FTER report, (Daniels 2004), several dozen are exclusively or primarily found below the depths covered by this study. Furthermore, limitations on access and technologies mean that this study was forced to leave essentially the entire southern half of

Misali’s reefs unexamined, as well as the submerged reef to the North. While this study covered a good cross section of the seaward, channel, and lagoon reefs present on Misali, as the linear regression of longitude and species composition showed, there are other strong factors at play besides simply longitude, and there are likely more distinct habitats in areas not covered by this study. Another solution is that previous studies may have over-estimated the species richness of Misali by mis-identifying and listing species that were not actually present. Sixteen of the species listed in FTER are not found in East Africa, and many are not even known to exist in the

Indian Ocean.

However, even once these differences are accounted for, there still seems to be a sizeable gap between the species observed in this study and the species observed in Misali previously. This may simply mean that many species have disappeared from the island.

Specifically, large Angelfish (>10cm), Triggerfish (>20cm), and Groupers (>40cm) were either notably absent or represented by only one or two species. Interviews with park rangers, who conduct and accompany all of the recreational SCUBA diving that happens around Misali, support the loss of these species, as rangers stated that they had not observed any of the species of Angelfish or Triggerfish that were conspicuously absent from this study, and that the few large species of grouper that were still present were very rare. Overall, many of the largest representatives of each family had disappeared, including Scaridae (Bicolor, Bumphead),

Lutjanidae (Green Jobfish, Red/Twinspot, Humpback), and the families mentioned above. This may indicate that Misali’s ecosystem is simply no longer productive enough to support such large fish, or that overfishing has eliminated these species. Overfishing has been known to lead to local species extinctions and deplete several important families in particular, including

Scaridae, Pomacanthidae, and Balistidae (McClanahan 1998, Koslow et al 1988). However, the

1998 coral bleaching, and long term effects on the composition of the coral community may also be to blame. Acropora, or branching corals, is one of the coral genera most vulnerable to climate change (Loya et al 2001), and is known to be crucial habitat for juvenile fish (Floros and

Schleyer), especially in areas without extensive mangrove or seagrass areas such as Misali. A previous ISP conducted at the island confirmed that juvenile fish were correlated with the percent cover of branching corals (Katz 2006). Twenty years after the 1998 bleaching event, nearly all of the corals in shallow protected areas and flats around Misali are Porites, or boulder corals (pers. Observ.), while sketches from Horrill 1992 show that before there was a substantial amount of Acropora. Many large prominent species, such as Pomacanthidae (Angelfish), have long juvenile stages that rely on shallow, protected, and coral-rich and complex environments to grow and mature (Lieske and Myers 1996). Twenty years later, what we may be observing is a lost generation of adults, as all the fish that were mature in 1998 have died out, and have not been replaced by new recruits due to these changes in the shallow water habitat. Coral bleaching has been shown to create these kinds of lost generation effects, a delayed population crash without any initial decline in abundance (Graham et al 2007). Data from previous studies show that this seems to be exactly what has happened at Misali (Mohammed & Mohammed

2002).

The first statistical finding, that diversity was not higher in the non-extraction area than it was elsewhere around Misali, is surprising in that it contradicts data available from previous surveys (Daniels et al 2004). However, this disparity is due to faults and limitations in the methods of those previous studies. The 2004 Frontier-Tanzania report, for example, measured biodiversity by having volunteers record data on the presence and abundance of 78 pre- selected species. However, narrowing an ecosystem of 200+ species down to only 78 species inevitably created substantial bias in the results. Whole families, including Scaridae (parrotfish) and Pomacentridae (damselfish) were excluded, likely because they were deemed too difficult for volunteers to identify. Most of the fish and the species recorded in the Southeast flat, for example, were damselfish and small species of not included in the FTER report’s fish list. The selection of species to be recorded, therefore, created a bias that substantially underestimated the biodiversity of the lagoon and flats reefs outside of the non-extraction zone.

Furthermore, if the goal of creating a non-extractive area is to protect and increase biodiversity relative to areas with no protection, then either the current levels of enforcement or the current boundaries of the non-extractive area are insufficient to accomplish this goal, or both. Enforcement, certainly, has been an issue. The rangers have done a fantastic job using their relationships with fishermen and the community to end destructive fishing practices and ensure that fishing is local, artisanal, and that all fishermen must have a license in order to fish.

However, without adequate funding, petrol, and working boats, there is only so much they can do to enforce boundaries such as the non-extractive zone. Currently, there is no enforcement of the non-extractive area (Klein & St. Denis 2016), and fishing pressure is at least as high inside the NEZ as it is outside. Fisherman were seen inside the NEZ conducting their business every time a transect was done there, and active duma traps were frequently observed underwater and even a few times within transects as they were being conducted (pers. obs.)

However, these observations were not limited to the NEZ. Fisherman were seen on almost every visit to Fig. 13: Fishermen observed using duma traps during a transect in the NEZ all sites, and duma traps were observed in or near to transects at all four sites (pers. obs.).

Fishing, therefore, does not seem to be a significant difference between the sites, and is unlikely to have been a significant driver of differences in diversity, community structure, and fish abundance between different geographic areas.

The second result, that the species composition varied drastically as sites moved further inshore, should come as a surprise to no one. It has long been known that many species tend to prefer either seaward or lagoon reefs and that factors such as exposure to waves, sediment type, depth, and steepness of slope, all of which varied significantly across sites, significantly influence the type of habitat present (Lieske and Myers 1996). However, if these facts were known, they seem not to have been taken into account when Misali’s conservation area and non-extraction zone were created. The current NEZ protects only one type of habitat, pure seaward reef, across this spectrum, and the ordinance analysis shows that it’s fish community is very distinct from the other sites examined in this study. If the goal of creating this kind of no- take area is to protect the maximum possible biodiversity, the current boundaries of the NEZ are neither ideal nor sufficient.

Finally, the result indicating that the NEZ had higher levels of overall and commercial fish abundance may seem to simply confirm that halting fishing leads to increased levels of fish.

However, the NEZ on Misali is currently not enforced, as mentioned above. It is therefore unlikely that there are significant differences in fishing pressure between sites, and certainly not enough to create this kind of difference. Furthermore, fish abundance was shown to significantly correlate (p=.0047) with the distance of a site from the Pemba Channel, suggesting that factors other than fishing pressure may be at play. It is the conclusion of this study that the NEZ is simply the site with the highest naturally-occuring fish abundances, and that this has been the case since before the creation of the park (Horrill et al 1994). The current NEZ, therefore, was drawn around the area that was naturally the most productive fishing ground, forcing fisherman, if it had been enforced, to fish areas that are less productive and supply less income to them and their families. Indeed, if the NEZ had been rigidly enforced as planned, it might also have proved harmful to the overall health of Misali’s ecosystem and put its fish stocks in danger of collapse, by concentrating fishing pressure on the areas and populations least likely to be able to support it.

Recommendations:

The Misali ecosystem is home to a huge variety of fish and distinct habitats. While each of these habitats seems to contribute evenly to the overall biodiversity of Misali, the seaward reefs, such as those in the current non-extraction area, have the highest fish abundances and the greatest number of commercially important species. As such, enforcing the current NEZ would protect only one habitat and one small part of Misali’s diversity while also imposing an undue burden on local fisherman and potentially endangering fish stocks by concentrating fishing pressure on areas less able to support them. If the goals of a non-extractive zone and a mixed-use conservation area in general are to preserve the greatest swathe of biodiversity while imposing the least burden on local fishermen and their ability to make a healthy, sustainable living, then the current boundaries of the NEZ are both inadequate and overbearing. It is therefore the recommendation of this study that Misali’s non-extraction zone should be redrawn.

By changing the boundaries of Misali’s NEZ, local fisheries managers have a fantastic opportunity to protect more biodiversity while simultaneously opening up more productive fish stocks to local fisherman. One way this could be done is by redrawing the NEZ longitudinally, so that it runs from the outermost reefs on the Pemba channel on one side, across the island, and all the way into the lagoon on the opposite side. Data from this study show that the species present at a site were determined largely by where that site was positioned along this imaginary line between the open channel and the protected lagoon. Redrawing the NEZ in this way would therefore allow it to protect a smaller slice of all habitats present, rather than simply all of the seaward reef habitat inside the current NEZ. Such a strategy would allow the NEZ to protect a much larger cross section of Misali’s biodiversity, while also opening up more of the productive seaward reef to local fisherman. As for the exact location, one option for the new NEZ would be to designate the area between West Island on the seaward side, the lagoon channel to the north separating Misali from the submerged reef, and East Island on the lagoon side as protected. Fig. 14: A map of Misali showing the current NEZ outlined in red, and the proposed new NEZ outlined in white. The new NEZ would run along the northern end of the island from West Island to East Island.

This new designation would protect the full range of habitats observed in this study, while also officially opening up the large, productive area of the current NEZ south of West

Island to local fishermen. From a tourism perspective, the new NEZ would also continue to protect the diversity and abundance of fish at the best dive sites around Misali, which are in the area of the current NEZ north of West Island, and newly protect the areas most accessible to casual tourists. All visitors to Misali are dropped off on Mbuyuni beach, on the north of the island near the ranger station, and most of them do not leave this area (pers. obs., Suleiman pers. comm.). While many visitors are interested in and engage in snorkelling, many are unaware or unwilling to trek to Turtle Beach and cross the wide intertidal zone and reef flat to reach the richest coral areas of the current NEZ. The new NEZ boundaries would add a new protected area that is right in front of Mbuyuni Beach, close to where tourists are dropped off and picked up, and is shallow, protected, and easily accessible without a boat at both high and low tides.

Additionally, this designation would help address several practical issues endemic to the current set-up. First of all, by using natural markers, the two islands and the channel, this design would obviate the need to set out marker bouys to clearly show where the NEZ begins and ends. This has been a recurrent problem with the current set-up, as without marker buoys, which are often ripped out by the sea or stolen, fisherman have been able to successfully claim ignorance in order to escape charges for fishing in the NEZ. The second issue this would address is the issue of monitoring and accessibility for the rangers. Turtle Beach, near the current NEZ, is around a 30 minute walk from the ranger station, requires shoes and flashlights at night, and is quite far from the actual reef fishing grounds within the NEZ, which are near the dropoff. Most of the new NEZ, by comparison would be in view directly from the current ranger station, and along areas where the dropoff and the fishing grounds are very close to shore and to Mbuyuni

Beach, which is essentially the ranger’s front yard.

However, given the limited scope of this study and limited data on fisherman’s use of various parts of the island, much more research needs to be done before any actions are taken to redraw the boundaries of the NEZ. There is very little existing data on the fish communities in most of the south side of Misali and on the submerged reef to the north, and it may be that these are home to their own microhabitats distinct from those inside this study area. It may be more practical to protect the entire northern submerged reef, which is less used by local fisherman, but data are insufficient to say how much of Misali’s biodiversity is contained there.

Furthermore, the proposed new NEZ boundaries may also raise unintended practical obstacles for the fishermen, and further research is needed to ensure that this new designation will not impose an undue burden on their work. Most fishermen camp at People of Pemba beach on the southeast side of the island and, at least during the rainy season, travel around the northern side, through the proposed new NEZ, to access the rich pelagic and reef fisheries along the

Pemba Channel on the west side of the island. If this is because the route around the southern side of the island is unsafe or impassible during certain times of the year, the new NEZ would require them to make an even longer journey from their camp to their fishing grounds. Rangers would also have to take care to allow fisherman to travel through the area but not to fish in it. Fisherman also currently make heavy use of the areas along the channel, which can be fished with a line from shore and are easily accessible even by rowed boats. Since many fisherman rely on sails and paddles, rather than motors, to reach their fishing grounds, it may be that, for lower tech operators, the opening of new grounds on the opposite side of the island, in the current NEZ, is insufficient or impractical to compensate them for the loss of the more accessible reefs on the north side. While the current location of the NEZ seems clearly to be less than ideal, much more research is needed on the understudied parts of Misali’s reefs and the needs of the fisherman on the island before any decision can be made on where exactly would be best for the new zone.

Conclusions:

Misali Island, an uninhabited coral island off the coast of Pemba in East Africa, boasts a stunning and remarkably diverse ecosystem, supporting both local fishermen and tourist operators. However, despite Misali’s importance to the local community and economy, over the past ten years it has been largely neglected as a priority for scientific research and government conservation funding. As this study shows, despite this neglect, Misali has remained healthy productive, and diverse, even with the current levels of funding and protection. However, as

Pemba’s population continues to increase and the island inevitably develops to the level of

Zanzibar or even the mainland coast, the current measures and levels of enforcement will no longer be enough to protect Misali’s ecosystem and the people and villages who depend on it.

More action must be taken to smartly, efficiently, and permanently protect Misali’s fish and reefs, while allowing fishermen to use the island sustainably as they have for generations. The current non-extraction area is insufficient to protect diversity and unfair to local fisherman in that it blocks all the best fishing grounds. It should be moved in order to protect a wider swathe of biodiversity and to open up more of the productive seaward reefs to local fishermen. The island is also badly in need of greater funding from the Zanzibari government, so that the rangers can patrol the park and enforce any NEZ boundaries.

In terms of research too, Misali offers a unique environment for research and must receive more attention soon. Misali experienced a severe bleaching event in 1998, and uniquely has had a chance to recover, relatively undisturbed, for almost 20 years since then. It therefore offers a unique opportunity to learn about coral bleaching and how coral and fish communities respond to and recover from severe bleaching events like this one, an opportunity that could be lost almost at any time, as bleaching events are becoming more common, and another severe event at Misali would alter the environment and restart the clock of recovery. Furthermore, it has been suggested that the strong tidal currents around Misali, particularly in the channel to the north along the proposed non-extraction area, allow coral polyps from these reefs to travel great distances before settling, and that the island may therefore play a role in reseeding reefs miles away. The island therefore offers an opportunity to learn about coral dispersal and how communities reseed and regrow after disturbance. Misali, therefore, should be the focus of renewed efforts at conservation, research, and government funding, so that a healthy Misali can continue to benefit scientists, tourists, local fishermen, Pemban diets and businesses, and any government agency seeking to look out for the health, well-being, and economic outlook of its citizens.

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Appendix:

Fish List:

Acanthuridae Powder Blue Acanthuridae Convict Acanthuridae Blackstreak Acanthuridae Pale-lipped Acanthuridae Ringtail Acanthuridae Goldring Acanthuridae Orange-Socket Acanthuridae Striped Bristletooth Acanthuridae Dusky Acanthuridae Lieutenant Acanthuridae Elongate Acanthuridae Blue Tang Acanthuridae Brushtail Tang Acanthuridae Desjardin's Tang Acanthuridae Spotted Unicornfish Orangespine Acanthuridae Unicornfish Black Tongue Acanthuridae Unicornfish Acanthuridae Humpback Unicornfish Acanthuridae Horseface Unicornfish Apogonidae 5-lined Cardinalfish Apogonidae Lined Cardinalfish Apogonidae 8-Lined Cardinalfish Aulostomidae Trumpet Balistidae Orange-Striped Balistidae Halfmoon Balistidae Titan Balistidae Scythe Balistidae Arabian Belonidae Houndishh Mozambique Bleniidae Fangblenny Caesonidae Twinstripe Fusilier Caesonidae Yellowback Fusilier Carangidae Bluefin Carangidae Bigeye Carangidae Bar Jack Carangidae Brassy Carangidae Rainbow Runner Chaetodontidae Bennetts Chaetodontidae Meyers Chaetodontidae Saddleback Chaetodontidae Madagascar Chaetodontidae Zanzibar Chaetodontidae Threadfin Chaetodontidae Long-nosed Chaetodontidae Big Long-nosed Chaetodontidae Vagabond Chaetodontidae Purple Chaetodontidae Kleins Chaetodontidae Raccoon Chaetodontidae Spotted Chaetodontidae Somali Chaetodontidae Black-backed Chaetodontidae Chevroned Chaetodontidae Longfin Banner Chaetodontidae Schooling Banner Chaetodontidae Yellowhead Chaetodontidae Black Pyramid Cirrhitidae Freckled Hawkfish Cirrhitidae Arc-eye Hawkfish Dasyatidae Bluespotted Ray Diadontidae Porcupinefish Fistuariidae Cornetfish Gobiidae Twostripe Goby Gobiidae Leopard Goby Haemulidae Blackspotted Sweetlips Haemulidae Oriental Sweetlips Haemulidae Slatey Sweetlips Haemulidae Giant Sweetlips Holocentridae Bronze Soldierfish Holocentridae Crown Soldierfish Holocentridae Pearly Soldierfish Holocentridae Violet Soldierfish Holocentridae Red Soldierfish Holocentridae Tailspot Squirrelfish Holocentridae Clearfin Squirrelfish Kuhliidae Barred Flagtail Kyphosidae Highfin Rudderfish Labridae Barred Thicklips Labridae Blackedged Thicklips Labridae Diana's Hogfish Labridae Tripletail Labridae Red-Banded Labridae Floral Labridae Ring Labridae Slingjaw Labridae Queen Labridae Spottail Coris Labridae Cigar Labridae Twotone Labridae Goldbar Labridae Sixbar Labridae Fivestripe Labridae Goldstripe Labridae Tubelip Labridae Nebulous Labridae Vermiculate Labridae Yellowtail Labridae Dragon/Rockmover Labridae Crescent Labridae Bird Labridae Cleaner Labridae Bicolor Cleaner Lethrinidae Blackspot Emperor Lethrinidae Spangled Emperor Lethrinidae Orange-stripe Emperor Lethrinidae Bigeye Emperor Lutjanidae Bluelined Snapper Lutjanidae Blackspot Snapper Lutjanidae Scribbled Snapper Lutjanidae Twospot/Red Snap Monocanthidae Broom Filefish Mullidae Dash & Dot Mullidae Sidespot Mullidae Double bar Mullidae Yellowfin Mullidae Yellowstripe Mullidae Long-barbel Muraenidae Zebra Moray Nemipteridae Arabian Spinecheek Ostraciidae Yellow Boxfish Ostraciidae Spotted Trunkfish Pempheridae Vanikoro Sweeper Pempheridae Copper Sweeper Plotosidae Striped Catfish Pomacanthidae Regal Pomacanthidae Many Spined Pomacanthidae African Pygmy Pomacentridae Indo-Pacific Seargent Pomacentridae Scissortail Seargent Pomacentridae False-eye Seargent Pomacentridae White Belly Damsel Pomacentridae Ternate Damsel Pomacentridae Webers Damsel Pomacentridae Dusky Gregory Pomacentridae Black Damsel Pomacentridae Jewel Damsel Pomacentridae Johnston's Damsel Pomacentridae Dick's Damsel Pomacentridae Onespot Damsel Pomacentridae Sulfur Damsel Pomacentridae Doublebar Chromis Pomacentridae Blue-Green Chromis Pomacentridae Black-Axil Chromis Pomacentridae Two-tone Chromis Pomacentridae Footballer Dascyllus Pomacentridae Humbug Dascyllus Pomacentridae Indian Dascyllus Pomacentridae Caerulean Damsel Pomacentridae Clark's Anemonefishh Pomacentridae Allard's Anemonefish Pomacentridae Skunk Anemonefish Scaridae Bullethead Scaridae Russell's Scaridae Red-lipped Scaridae Swarthy Scaridae I-O Longnose Scaridae I-O Steephead Scaridae Tricolor Scaridae Dusky-Capped Scaridae Dark Crescent Scaridae Blue-Barred Scaridae Tail-barred Scaridae Palenose Scaridae Bridled Scaridae Greenlip Scaridae Greenthroat Scaridae Seagrass Scorpaenidae Lionfish Serranidae Redmouth Grouper Serranidae Peacock Grouper Serranidae Coral hind Grouper Serranidae Sixspot Grouper Serranidae Goldie/Lyretail Anthias Serranidae Yellowback Anthias Serranidae Bicolor Anthias Syngnathidae Pipefish Synodontidae Variegated Lizardfish Synodontidae Blackbloth Lizardfish Tetraodontidae Blackspotted Pufferfish Whitespotted Tetraodontidae Pufferfish Tetraodontidae Guineafowl Pufferfish Tetraodontidae Black-Saddled Toby Tetraodontidae Crown Toby Tetraodontidae Bennett's Toby Zanclidae Moorish Idol

Additional Species Not Observed in Transects:

Mbuyuni Beach:

African Coris Clearfin Lionfish Scorpionfish spp Barred Seargent

Non-Extraction Zone

Quakerfish Bicolor Toby Lyretail Hogfish Bandcheek Wrasse Thompson's Surgeonfish Yellowtop Fusilier Arrowhead Soapfish

Channel Reef

Black Snapper Somali Butterflyfish Picassofish/humuhumu Black-spot Seargent Goggle-eye Masked Bannerfish Yellowsaddle Goatfish Rosy Goatfish Striped Blanquillo

Southeast Flat

Halfmoon Triggerfish Yellowsaddle Goatfish Snowflake Moray Peppered Moray Longfin Spadefish Juv. Bumphead Parrotfish