BELIZE BEACH CONSERVATION,MANATEE CONSERVATION, MARINE

CONSERVATION AND DIVINGPROGRAMME

Caye Caulker, Belize

BZM Phase 184 Science Report

Dagny-Elise Anastassiou, Sophie Pipe, Bridie Matthews and James

McKay

September 2018 - December 2018

Staff Members

Dagny-Elise Anastassiou (DEA) Project Manager (PM) and Principal Investigator (PI)Sophie Pipe (SP) Research Officer (RO)Bridie Matthews (BM) Assistant Research Officer (ARO)James McKay (JM) Assistant Research Officer (ARO)

Acknowledgements

We would like to thank the Belize Fisheries Department (BFD) for the continued assistancethroughout the project. We would also like to thank the Frontier Research Assistants that tookpart in our BZM 184 projects.

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Contents

1 Introduction 31.1 Project Background and Location . . . . . . . . . . . . . . . . . . . . . . . . . . 31.2 Caye Caulker Marine Reserve . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51.3 Aims of the Project . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61.4 Training . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

1.4.1 Science training and volunteer briefings . . . . . . . . . . . . . . . . . . . 71.4.2 Dive Training . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

2 Research Projects 82.1 SMP Coral Reef Diving Surveys . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

2.1.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82.1.2 Methodology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112.1.3 Coral Abbreviations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132.1.4 Data Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

2.2 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142.2.1 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

2.3 Conch Survey . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222.3.1 Ecology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222.3.2 Habitat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222.3.3 Life Cycle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222.3.4 Predators and threats . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222.3.5 Aims & Objectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232.3.6 Methodology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232.3.7 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 242.3.8 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

2.4 Caribbean Spiny Lobster Surveys . . . . . . . . . . . . . . . . . . . . . . . . . . 282.4.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282.4.2 Methodology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 302.4.3 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

2.5 Lionfish Study . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 312.5.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 312.5.2 Lionfish Survey Methodology . . . . . . . . . . . . . . . . . . . . . . . . 322.5.3 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 342.5.4 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

2.6 Caribbean Spiny Lobster Surveys . . . . . . . . . . . . . . . . . . . . . . . . . . 352.6.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 352.6.2 Methodology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 362.6.3 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 372.6.4 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38

2.7 Seagrass and Manatee Monitoring . . . . . . . . . . . . . . . . . . . . . . . . . . 412.7.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 412.7.2 Methodology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45

2.8 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 482.8.1 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48

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Figure 1: Caye Caulker Marine Reserve (Google Maps, 2017).

1 Introduction

1.1 Project Background and Location

Belize, formerly known as British Honduras, is a Central American country on the Easterncoast. Belize is bordered by Guatemala from the west and Mexico from the north with theCaribbean Sea to the east. The small mainland area is 290 km long and 110 km wide. TheMesoamerican Reef is the largest barrier reef in the Northern Hemisphere and the secondlargest barrier reef in the world (Seijo,2007). Large areas of the barrier reef are protected bythe Belize Barrier Reef Reserve System (BBRRS) and therefore it has been designated a worldheritage site by UNESCO for the past 20 years. Included in the BBRRS are seven marinereserves, 450 Cayes and three of the four atolls present: Turneffe Atoll, Glovers Reef Atolland Lighthouse Reef Atoll (home to the Great Blue Hole dive site made famous by JacquesCousteau) (Gibson et al., 1998). The Belize Barrier Reef (BBR) is the largest single sectionof the Mesoamerican Barrier Reef System (MBRS) and extends 998 km from the tip of theYucatan Peninsula to Honduras; making it the second largest barrier reef in the world. TheFrontier Belize Marine project (BZM) was established in April 2014 within the Caye CaulkerForest and Marine Reserve (figure 1). Caye Caulker (CC) is a small limestone island locatedapproximately 20 miles north-northeast of Belize City at 17o4433N88o130W . The FrontierBelize camp is located on the North island of Caye Caulker and hosts Marine Conservation,Diving and Beach Conservation volunteers.

The project aims to conduct long term monitoring of key habitats and species through theassistance of international volunteers and in collaboration with the BFD. CC spans approxi-mately 8 km from north to south and approximately 1.5 km east to west at its widest part. Theisland is separated by a narrow man-made waterway known as the Split creating a north-southdivide of the island (CZMAI, 2016). The majority of the islands infrastructure is located onthe smaller South Island which holds a population of approximately 1700 residents and manytransient visitors (CZMAI, 2016). Many tourists visit the island due to the abundance of exoticflora and fauna, which provide excellent snorkeling, diving and nature trail opportunities. Theisland is particularly popular for backpackers due to its easy-going reputation, a result of the

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strong Creole presence and community of Rastafarians. Tropical marine ecosystems worldwideare under threat from anthropogenic activities such as overfishing, unsustainable tourism prac-tices, irresponsible coastal development, climate change and pollution (Acosta, 2006). This hasled to degradation of coral reefs, mangrove forest and seagrass meadows and thus a decline inbiodiversity and ecosystem health (CZMAI, 2016). Caulker Cay is no exception as it providesa habitat for several endangered species such as the critically endangered staghorn (Acroporacervicornis) and elkhorn (Acropora palmata) corals as well as the hawksbill (Eretmochelys im-bricata), green (Chelonia mydas), and loggerhead turtles (Caretta caretta) (IUCN, 2015). Otherthreatened species found in the waters of CC include The West Indian Manatee (Trichechusmanatus) and the common bottlenose dolphin (Tursiops truncates) (Fisheries, 2010; IUCN,2015).

Global fisheries are dependent on near shore tropical coastal waters and reefs as more than25% of marine life is found on coral reefs (Acosta, 2006). The coastal waters around CCprovide rich fishery grounds for two of the most important commercial fisheries in Belize: theCaribbean spiny lobster (Panulirus argus) and the queen conch (Strombus gigas). Currentlythere is a strict management regime in place to regulate the fishing seasons for each specieswhich includes catch size and gear restrictions (CZMAI, 2016). Fishing practices include hand-line fishing, speargun, Hawaiian sling, lobster hook and conch fishing. These regulations aimto maintain healthy and sustainable lobster and queen conch populations within the area usingsize as an estimation of sexual maturity (Huitric, 2005). This data can be used to informfishery management and can be utilised within the marine park management assessments, toanalyse the effectiveness and success of the marine park. Furthermore, the data can be usedon a national scale when compiling national and international data. The coastal waters aroundBelize are highly productive and local scale conservation plays an important role in protectingthese at risk, delicate ecosystems.

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1.2 Caye Caulker Marine Reserve

Figure 2: Map of the zonation of the Caye Caulker Marine Reserve (BFD, 2008).

A large portion of the waters around CC are under marine spatial management (CZMAI,2016). The Caye Caulker Marine Reserve (CCMR) is 11 km long, extending to the BelizeBarrier Reef from the Northern tip of CC (CZMAI, 2016). The CCMR was established in theearly 1990s but did not become completely recognised as a marine reserve until 1998 at whichpoint it was included alongside the Caye Caulker Forest Reserve (CCFR) as part of a singlemanagement unit (CZMAI, 2016). CCMR and CCFR encompass an area of 40km2 and 0.5km2

respectively (CZMAI, 2016). This unit was set up with the aim of ensuring protection of thelittoral forest, reef lagoon, and reef crest and fore- reef areas (CZMAI, 2016).

CCMR encompasses five ecologically-related but distinct habitats: mangrove forest, littoralforests, lagoon marshlands, seagrass beds and coral reefs (BFD, 2008). The marine reserveis managed via three regulated zones; the General Use Zone (GUZ), the Conservation Zone(CZ) and the Preservation Zone (PZ) (BFD, 2008). Table 1 below outlines the description anduses of each zone. Tourists are charged a park fee of 10 Belize dollars which helps subsidizemanagement of the area, while locals may use the reserve for free. There are no charges tomarine users outside of the reserve and these areas are not often monitored by the BFD. Despitethe fishery management practices that are in place, overfishing and illegal fishing are still majorissues in CC and across Belize (Fujita et al, 2018). As a result, many fishermen have reportedthat their catch per unit effort has decreased noticeably in recent years (CZMAI, 2016).

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Table 1: Description of each zone within the Caye Caulker Marine Reserve, Belize.

Zone name Size of zone Usage

General use zone (GUZ)Licensed commercial Fishing

25km2 Sport FishingRecreational Use

Conservation Zone (CZ)No-take zone

8.2km2 Recreational Use Allowed

Preservation Zone (PZ)No-Catch Zone

5.8km2 No recreational ActivitiesNo boat-traffic

The CCMR is critical to Belizes economy as a source of tourism revenue (on which manylivelihoods are entirely dependent) as well as a provider of vital ecosystem services, such asprotection from storm damage and wave erosion (Belize Fisheries Department, 2008). Thereis little research into the sustainability and effectiveness of the reserves current managementplan; however, this information is essential for the stakeholders involved (Polunin, 1993). Man-agement of the Belize Barrier Reef was originally envisioned through the creation of marineprotected areas (MPAs), but the influence from land-based activities was not accounted forwithin these programs. As a result, the focus was shifted towards the previously describedintegrated, multi-sectoral approach currently used in Belize marine reserves (Cho, 2005).

The research conducted by Frontier within the CCMR lies within the MBRS which ispart of the Mesoamerican Barrier Reef Systems Project, a regional SMP. This program has astandardised manual of methods (MBRS Project Description, 2004) for monitoring changes inreef ecosystem health which Frontier has incorporated into its own methods for effective datacollection by volunteer citizen scientists. Surveys carried out as part of BZM include MBRSSMP surveys (fish, benthic, coral colony characterization, and invertebrates) as well as conch,lobster, mangrove, seagrass and beach clean surveys. It is important to note that some surveysare seasonal and/or volunteer dependent.

1.3 Aims of the Project

The aim of BZM is to conduct long term scientific monitoring of key habitats and specieswithin the forest and marine reserve with the assistance of international volunteers and toreport information on the health of the reef to our partner organisations. The current projectobjectives are:

• To establish new fixed survey sites in addition to the sites suggested and already used bythe BFD.

• Continue to collect data on the health of the coral reef via the MBRS SMP methodologythrough benthic point intercept transect surveys, coral colony characterisation surveys,and adult/juvenile reef fish surveys.

• Continue to collect data on the abundance, sex ratios, maturity and size-frequency distri-butions of the commercially important Caribbean spiny lobster throughout the year andsupplement the biannual data collection conducted by the BFD.

• Reinstate and establish sites for data collection on the abundance, sex ratios and size-frequency distributions of the commercially important queen conch alongside the BFD.

• Continue and improve seagrass surveys on species composition, percentage coverage,abundance and health across the four established survey sites.

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• Reinstate the manatee monitoring project during manatee observation season.

• Continue to build upon new connections with the local community, including local NGOOceana and the local private school Ocean Academy.

1.4 Training

1.4.1 Science training and volunteer briefings

To achieve the above aims, all volunteers and staff members receive a combination of briefings,science presentations and lectures and practical field training (table 2) before conducting anymarine surveys. For all tests a 95% pass mark is required and in case of any failures, thoseindividuals will have to re-sit a different version of the test. All marine conservation and divingvolunteers are PADI scuba trained to at least Advanced Open Water level.

Table 2: Complete list of science lectures, field training lectures, briefings and staff memberresponsible for training. All training was conducted by DEA, SP, BM and JM.

Lecture/Presentation/Test LecturerHealth and safety and medical presentations and tests DEADangers of the reef presentation DEAIntroductory science presentation for Frontier Belize Project DEAIntroduction to coral reefs presentation DEABenthic identification and survey methodology presentation DEACoral health presentation DEACoral flash, revision slides DEA/SPBenthic test DEA/SPIn water, practical benthic test DEA/SPFish identification presentation DEA/SP/BM/JMFish flash, revision slides DEA/SP/BM/JMFish families test DEA/SP/BM/JMFish ID test DEA/SP/BM/JMJuveniles and recruits fish ID test DEA/SP/BM/JMIn water, practical fish test DEA/SP/BM/JMPractice mock survey DEA/SP/BM/JMQueen conch presentation DEA/SP/BM/JMCaribbean spiny lobster presentation DB/SP/BM/JMSeagrass presentation DEA/SP/BM/JM

1.4.2 Dive Training

All diving volunteers receive training up to PADI Advanced Open Water. PADI dive trainingis currently outsourced to a local dive company on the South Island. This will continue untilFrontier is able to offer this training in-house.

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2 Research Projects

2.1 SMP Coral Reef Diving Surveys

2.1.1 Introduction

Coral reefs are highly valuable and productive marine ecosystems. They provide essentialhabitats and feeding grounds for a diverse array of marine fauna such as fin-fish, elasmobranchs,turtles, sea-birds and numerous invertebrates (Conservation International, 2008). Coral reefsalso provide an array of crucial ecosystem services such as fish production and buffers for currentand wave energy. Reefs also generate revenue through tourism and medicinal discoveries andare especially vital to human survival in developing countries (Costanza et al., 1997; Bryantet al., 1998; Hughes et al., 2010). The most recent global economic value placed on coralreefs is $9.9 Trillion US dollars (Conservation International, 2008). Despite this, coral reefsremain one of the worlds most heavily impacted marine ecosystems. There are many seriousanthropogenic threats to coral reefs, which include; overfishing, habitat destruction, increasesin sea surface temperature (SST), coral disease, invasive species, and poor land use-practiceswhich lead to coastal eutrophication and heavy sediment loading (Harvell et al., 2007; Hugheset al., 2010). These direct and indirect pressures result in nutrient pollution and sedimentationwhich enhances algal growth and reduces zooxanthellae photosynthesis efficiency (Pearse &Muscatine, 2013). Stresses on the mutualistic relationship between coral and the zooxanthellaecan result in coral bleaching (Pearse & Muscatine, 2013). The presence of stressors, such as anincrease in water temperature, causes coral to expel the zooxanthellae (Pearse & Muscatine,2013). The zooxanthellae are what provides corals colour, thus when the algae exits the coral,it will turn white, this is given the term coral bleaching. Coral is not dead when it becomesbleached; however, it increases corals vulnerability as it has lost its predominant food source(Pearse & Muscatine, 2013). As a consequence, it can lead to coral reef decline (Pearse &Muscatine, 2013).

The Belize Barrier Reef has experienced frequent and severe ecological changes due to coraldisease, coral bleaching, extreme weather events and human disturbance. This has resulted in adecline from 25-30% live coral cover in the mid-1990s to approximately 11% live coral cover in2006, and a subsequent increase in macroalgae (Wilkinson, 2004). The region was devastatedby hurricanes between 2000-2002, which destroyed many coral formations with recorded lossesof up to 75% in some parts of Belize (Almada-Villela et al., 2002). In 1998, the reefs werepreviously damaged by Hurricane Mitch, a category 5 storm, as well as an extreme bleachingevent which exacerbated the destruction caused by Mitch (Wilkinson, 2004). The bleachingevent was caused by unprecedented elevations in SST due primarily to a severe El Nio SouthernOscillation (ENSO) and was likely enhanced by global warming, resulting in a 48% reductionin live coral coverage (Goreau et al., 2001). All habitats along the BBR experienced bleachingbecause of these thermal anomalies (Mumby & Harborne, 1999). By 1999, some fore-reefhabitats demonstrated signs of recovery, while coral populations amongst the sheltered lagoonsof the back reef displayed reduced coral cover, low coral recruitment and little indication ofrecovery (Aronson et al., 2002). Aronson et al (2002) determined that during this prolongedperiod of elevated sea surface temperatures, anomalies peaked at 4OC above the local hotspotthreshold, significantly greater than the previous years sea surface temperature anomaly spikeof 1OC.

Currently coastal and marine activities in the CC area are relatively minor; however, this ischanging with the rapidly growing tourism industry (Almada-Villela et al., 2002; CZMAI, 2016).An increase in tourist numbers has caused an increase in waste, plastic pollution, boat traffic,coastal development, and human interactions with the marine wildlife. These disturbances havepromoted algal growth on the reefs which corresponds with a decline in coral cover (McClanahan

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et al., 2001; Schutte et al., 2010). The decline in coral cover and reef health will have negativeimplications for fish assemblages, for example, a correlation has been found between abundanceand diversity of reef fish and coral health (Jones et al., 2004). Moreover, a decline in coral covercan impact fish recruitment and post recruitment pressures such as competition and predation(Jones et al., 2004). Belize has the highest fish species richness in the MBRS region, with over320 recognised species, but intensifying fishing efforts are threatening fish diversity (Acosta,2006). This is largely due to changes to spawning aggregations; spawning location dependson the recognisable characteristics of Cayes and reefs, characteristics which may be lost withoverfishing and other human activities (FAO, 2002). A documented example of this is theNassau grouper (Epinephalus striatus) which during the full moons of December and Januaryform spawning aggregations annually in the same location (Aguilar-Perera, 2006). This allowsfishermen to become familiar with these cycles which can lead to overfishing of this species. TheNassau grouper is currently listed as endangered on the IUCN Red List of Threatened Speciesand despite its protected status in Belize it is seldom observed within the reefs of the CCMR(Almada-Villela et al. 2002; Hughes et al., 2007). It has been observed that the exclusion oflarge predatory and herbivorous fish, such as the Nassau grouper, led to a dramatic explosionof macro-algae which in turn suppressed the fecundity, recruitment and survival of coral andultimately reduced total coral cover within the reef systems (Almada-Villela et al., 2002).Consequently, the coordination and control of fish stocks is one of the primary components inpreventing phase shifts and managing reef resilience (Hughes et al., 2007).

Disturbances to the invertebrate community have also profoundly influenced reef health onthe BBR, for example; diseases, potentially caused by water-borne pathogens, have dramaticallyreduced populations of the herbivorous sea urchin Diadema antillarum (black long spine) inthe Caribbean, which resulted in the increased abundance of macro-algae and reduced coralcover (Harborne et al., 2009). Many other invertebrate species play important roles in reducingmacro-algae cover by grazing on the biofilms on substrate suitable for hard cover (Klumpp& Pulfrich, 1989). Furthermore, it is important to monitor the abundance and diversity ofinvertebrates which are vulnerable to overfishing, such as the commercially important queenconch (Strombus gigas) and Caribbean spiny lobster (Panulirus argus) (Theile, 2002; Perez &Garcia, 2012).

The MBRS SMP was designed to standardize data collection and management for ecosys-tem monitoring. It aims to monitor changes in ecosystem health within priority protectedareas, enabling quicker and more effective responses to changes in reef health (Gomez, 2004).Ultimately the utilization of this protocol coupled with the long-term monitoring work con-ducted by Frontier will provide much needed baseline data on the health of the BBR marineecosystem. In the past, Frontier conducted SMP surveys in four locations within the CCMR;however, due to weather conditions the fore-reef sites were rarely sampled. Currently, sincephase 173, (July 2017) ten locations have been surveyed to meet the first aim of the projectwhich is to establish new fixed survey spots. The back-reef survey sites are both shallow patch-reefs located approximately 3.2 km apart. Due to how shallow the survey sites are, coral islikely to experience relatively higher wave action as well as strong currents from the nearbychannels: South Channel and North Channel (Komyakova et al., 2013). In areas of high-watermovement, the slower growing massive corals such as Orbicella annularis thrive particularlywell as they can withstand moderate wave action, the larger they grow the more stable theybecome (Barnes & Hughes, 1999). In contrast branching colonies such as Acropora cervicor-nis and Acropora palmata grow much faster; however, as they grow, they become increasinglyunstable as the small area attached to the substratum has a heavier load to support. In areassuch as the back-reef survey sites, where wave action is higher, this structural instability is putunder greater pressure often resulting in branches snapping off (Barnes & Hughes, 1999). Inthe high wave action areas, the community diversity is expected to be low as few species are

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able to tolerate these extreme conditions. Additionally, low wave action areas also exhibit lowerspecies diversity, but this is due to competitive dominants excluding other species (Komyakovaet al., 2013).

The baseline coral reef studies here will be used for long term monitoring of the MPA.Through conducting comparative studies of the different management zones with various levelsof protection within the MPA, it enables evaluation of the productivity of the guidelines in placein each zone. For example, if a correlation between poor coral health and low fish abundanceis found within the GUZ over a long-term data set, this could support the proposal of rotatingthe zones within the MPA or extending the CZ or PZ to include a proportion of the currentGUZ to aid the recovery of coral species and sustainable fish populations. Moreover, if asignificant difference in the health of coral within the CZ and the rest of the MPA after longterm monitoring is not concluded, this could imply the regulations in place may require are-evaluation.

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2.1.2 Methodology

Survey Area In each management zone within the MPA, similar sites are chosen for compar-ison. Table 3 describes the locations of all the sites surveyed. In each zone there is a fore-reefsite, a channel site and a back-reef site. The CZ has two back-reef sites as this is the biggestzone.

Table 3: Sites for coral reef monitoring studied in Caye Caulker, Belize for the 183 Phase whichincludes the months of July to September 2018.

Site New or Existing Zone Reef Area SMP site Location1 Existing GUZ Back Elkhorn Point 16 Q 1968256 UTM 3944642 Existing GUZ Channel North Channel 16 Q 1967918 UTM 3944243 Existing GUZ Fore CCC Fore 16 Q1957698 UTM 3940794 Existing CZ Channel Island 16 Q 1966980 UTM 3948395 Existing CZ Back No-manns land 16 Q 1906367 UTM 3925796 Existing CZ Back Lobe City 16 Q 1906365 UTM 3920177 Existing CZ Fore Raggedy Ann 16 Q 1959791 UTM 3944198 Existing PZ Back Tap tap fish 16 Q 1966286 UTM 03938229 Existing PZ Channel The Swash 16 Q 1963507 UTM 39392710 Existing PZ Fore Corolitta 16 Q 0391885 UTM 1967918

Site 1: Elkhorn Point (EP) A shallow patch reef on the inside of the MBRS locatedjust north of the North Channel, with a maximum depth of approximately three metres andwithin the General Use Zone of CCMR. Current flow and wave energy are generally moderateto high on days with greater wind force.

Site 2: North Channel (NC) The North Channel has a wall of a maximum depth ofsix metres, the wall is surveyed and then the small patch of reef slightly further into the reef.This site is within the General Use Zone.

Site 3: Caye Chappel Channel Fore (CCC Fore) A relatively deep reef on theoutside of the MBRS, with a depth ranging between 14 metres to 25 metres. This site is onlyaccessible when the wind force is low and therefore the sea state is low.

Site 4: Island (I) A shallow patch reef on the inside of the MBRS which is situatedbetween the reef crest and the lagoon, with a maximum depth of approximately five metresand located in the Conservation Zone of CCMR. Due to moderately low current flow this siteis frequented regularly by tour operators and dive companies.

Site 5: No mannsland (NML) A barrier reef on the inside of the MBRS with amaximum depth of approximately eight metres in the Conservation Zone bordering the GeneralUse Zone of CCMR.

Site 6: Lobe City A patch reef on the inside of the MBRS with a maximum depth ofapproximately five metres in the Conservation Zone bordering the General Use Zone of CCMR.

Site 7: Raggedy Ann (RA) A barrier reef on the outside of the MBRS, with a depthranging between 15 metres to 30 metres. This site is only accessible when the wind force is lowand therefore the sea state is low.

Site 8: Tap tap fish (TTF) The North Channel has a wall of a maximum depth of sixmetres; the wall is surveyed followed by the small patch of reef slightly further into the reef.This site is within the General Use Zone.

Site 9: The Swash (S) A smaller channel on the MBRS with a maximum depth of twometres. This site is located in the Preservation Zone.

Site 10: Coralitta (PZC) A barrier reef on the outside of the MBRS within the Preser-vation Zone with a depth between 15 metres and 25 metres.

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Experiments and Survey Protocol

The protocol for SMP surveys is adopted from the MBRS SMP as the standard methodologyused to monitor coral reefs at established sites (Almada-Aleman et al., 2003). Survey methodswere adapted for the BFD, FAMRACC, and FRONTIER. For each survey a minimum of threedivers was required and each had a specific role (physical, fish, benthic) for which they hadundergone the required training. At each survey site five replicates were conducted, roughlyfive meters apart to prevent resampling.

Physical Surveyor

A physical surveyor collected coral community characterization data. Prior to each transectthe following information was recorded: date and site name and/or GPS coordinates. A 30m transect line was laid haphazardly within the general confines of the site in order to avoidany unconscious bias towards an area. The transect line was weighted at one end with a 2 lbdive weight so it remained in place. With the aid of an underwater compass the transect linewas laid parallel to the reef in a straight line to comply with the MBRS methodology. Coralmeasurements using a ruler began at the first colony located directly beneath the transectline which was at least 10 cm in diameter. For each colony surveyed the following data werecollected: the species of coral, diameter, and maximum height of the colony. Percentage of deadcoral, the presence of any diseases, and/or any bleached tissues were also estimated (table 4).These measurements were repeated for the rest of the transect, until at least 45 coral colonieshad been sampled at each site. Care was taken to ensure that no diver or equipment wasdamaging the studied coral samples.

Table 4: Definitions of the codes used for coral disease and bleaching.

Disease Codes Bleaching CodesBB= Black Band PB= Partially bleachedWB= White Band BL= Bleached, fully-bleachedRB= Red Band P= Pale, signs of colour loss/ colour changeYB= Yellow BandWP= White PlagueDS= Dark Spot

Fish Surveyor

The Atlantic and Gulf Rapid Reef Assessment (AGRRA) protocol for coral reef fishes has beenadopted by the MBRS SMP as the standard methodology for assessing overall fish communitycomposition. The first method involved belt transects which is designed to measure the densityand size of fish species. A fish surveyor swam alongside the physical surveyor whilst the transectline was reeled out, in order to minimize any changes in depth. While the full 30 m transectline was laid out the surveyor counted and recorded the fish species observed within a 2 m widevisual estimate (1 m either side of the line), counting while deploying minimizes disturbance tothe fish prior to their being counted. The size of each fish was estimated by assigning them tothe following size categories (< 5, 6 − 15, 16 − 30, 31 − 40, > 40cm). A two-minute pause wastaken once the surveyor reached the end of the transect line before the surveyor returned alongthe line looking for any additional recruits (only during summer months) (Almada-Villela etal., 2003). On the return journey the surveyor swam along the 30 m transect and recordedonly selected juvenile fish, fish species < 5cm and Diadema sp. Sea urchins. The latter wasrecorded as part of the fish survey whenever an invertebrate surveyor was not present. Eight

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transects were conducted at each site.The data will be used to examine the three zones within the CCMR, the Conservation Zone(CZ), General Use Zone (GUZ) and the Preservation Zone (PZ). The CZ and PZ are the no-take zones. The data will be used to analyse the densities of fish by family, the total densitieswithin each of the zones, and to compare habitat type of the fore-reef and back-reef. Thedensities of adult fish will be analysed separately from the density of recruitment. Recruitmentrefers to the juvenile population. It is the addition of this population to the adult populationwhich is studied. The identification of the areas of highest recruitment may identify the larvaldistribution sources (Almada-Aleman et al., 2003). Furthermore, this will give an understandingof the adult species distribution and abundance (Doherty & Fowler, 1994). This can then beanalysed across the different zones. To be able to determine which families of fish are moresusceptible to changes in the community through either fishing pressures or habitat changesand environmental changes the study of the recruitment is vital (Planes et al., 2009). Theresults are shown by the species known to be connected to reef health (Almada-Aleman et al.,2003).

Benthic Surveyor

A benthic surveyor swam the 30 m transect line to identify and count the benthos every 25 cm,providing 120 records of data. Substratum percentage coverage was calculated from the dataas (numberofrecords/120) ∗ 100. SMP target benthos includes coralline algae (code = COR),turf algae (TURF), target macro algae species, sponges (SPN), gorgonians (GG) and targetstony corals (see appendix for full species list). Any abiotic substrate, including sand (SN),bare rock (BR) and dead coral (DC), was also recorded. Five PIT (Point Intersect transects)were conducted at each site. A PIT is a 30 m transect along the benthic cover, recording thebenthic cover and coral every 25 cm.

2.1.3 Coral Abbreviations

All target coral species are abbreviated from their scientific names as per Atlantic and GulfRapid Reef Assessment (AGRRA), where the first letter of the genus is the first letter in theabbreviation and the last three letters of the abbreviation are from the first three letters of thespecies. For example: lettuce coral, Undaria agaricites UAGA. For a full list of target coralsand their abbreviations see appendix 1.

2.1.4 Data Analysis

The data analysis was done using Microsoft Excel and the statistical programme Rstudio version1.1.456 (R Core Team, 2018). Microsoft Excel was used for data input and Rstudio was usedto run statistical tests to determine significant relationships between fish densities, coral cover,and habitat type. Packages used in Rstudio include: dplyr, ggplot2, gridExtra, zoo, reshape,car, pscl, scales, extrafont, FSA, rcompanion, nortest, lattice, and WRS2 (R Core Team, 2018;Augie, 2017; Chang, 2014; Mair & Wilcox, 2018; Mangiafico, 2019; de Mendiburu, 2017; Dinno,2017; Fox & Weiberg, 2011; Gross & Ligges, 2015; Jackman, 2017; Ogle, 2018; Pinheiro et al.,2018; Sarkar, 2008; Wickham, 2007; Wickham, 2016; Wickham, 2018; Wickham et al., 2018;Zeileis & Grothendieck, 2005). A robust independent factorial one-way ANOVA was used to seeif there was a significant difference between fish species density in different management zones.A post-hoc tukey test was then used to determine which species were significantly differentfrom each other. A one-way independent ANOVA was used to determine whether there wasany significant difference between the recruitment densities in different phases throughout 2018.A post-hoc tukey test was then used to determine where the significance was. A Pearson

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correlation test was used to determine if there was a correlation between percentage algaecoverage and grazer densities in phases 182 and 184, other phases were not included in thisanalysis due to no surveys taking place during those time periods. An ANCOVA was used todetermine if there was a significant correlation between total fish density and coral cover indifferent management zones. Again, only phases 182 and 184 were included. All tests weredone on a 95% confidence interval.

2.2 Results

Coral Results

In total, seven full PIT surveys were conducted within the CCMR. In the past, up to 10 siteshave been surveyed, due to inclement weather conditions only seven sites were possible tosurvey. In the CZ four PIT surveys were conducted, providing n=2400 points of data, 600 foreach survey. Within the PZ, one PIT survey was conducted providing n=600 points of data.In the GUZ, two PIT surveys were conducted providing n=1200 points of data. All percentagedata is represented as the mean SD. This is displayed in table 5.

Table 5: Average % coverage ±SD of coral colonies across three different types of managementsite in Caye Caulker Marine Reserve (Conservation zone, General use zone and PreservationZone). Samples were taken in Belize during October 2018 to December 2018.

Zones Sites % Cover ± SD

CZCZ Fore-reef 16.5 6.78CZ Black-reef 18.4 7.43

GUZ GUZ Black-reef 11.8 5.25PZ PZ Black-reef 9.58 5.14

When analysing the habitat type of the back-reef the CZ displayed the highest percentagecover of coral colonies of 18.4 ± 7.43, followed by GUZ of 11.8 ± 5.25 and the lowest coveragefound in the PZ with 9.585.14. With only one fore-reef site surveyed this phase it was notpossible to see which management zone had the highest percentage cover. The CZ fore-reefshowed a coverage of 16.5 ± 6.78. This is displayed in figure 3.

Figure 3: Average % coverage SE of coral colonies across three different types of managementsite in Caye Caulker Marine Reserve (Conservation zone, General use zone and PreservationZone). Samples were taken in Belize during October to December 2018.

Within the CZ the most dominant hard coral showed to be OANN (Orbicella annularis),commonly known as lobed star coral making up 2.6% of coral cover. In the PZ and the GUZ,the most dominant hard coral was PAST (Porites astreiodes) at 1.92% on the PZ and 0.67%in the GUZ, most commonly known as Mustard Hill. The majority of the coverage across the

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Figure 4: :Average % coverage SE of algae across three different types of management site inCaye Caulker Marine Reserve (Conservation zone, General use zone and Preservation Zone).Samples were taken in Belize during October to December 2018.

three zones was GG (Gorgonian) and algae such as DT (dictyota), HA (Halimeda) and LOB(lobophora). The fore-reef CZ exhibited an algal coverage of 59 ± 69. On the back-reef habitattype the CZ demonstrated 15.2±26 coverage, the GUZ 18.3±38 coverage and lowest percentagecover was in the PZ 3.75 ± 11.9. This is displayed in figure 4 below.

Throughout the CCMR, the Acropora sp. made up 0.78 ± 4.47% of coral coverage in theall three management zones of the CZ, PZ and GUZ. The GUZ exhibited the highest coverageof 0.42 ± 0.10%, with the majority species found at one site, Elkhorn point. The PZ showedthe lowest coverage of 0.13 ± 0.06%, and the CZ showing results of 0.23 ± 3.49% of Acroporacover.

Soft corals are classed as Gorgonian, throughout the CCMR 411 total gorgonian colonieswere counted in the phase. The CZ had the highest percentage coverage of 14%, followed bythe GUZ with 6.8% and the lowest found in the PZ with a mere 1.8% coverage. The resultsare displayed in figure 5 below.

The second and last phase of each year include studies of bleaching across the CCMR.This phase looks at the bleaching evidence after the summer month bleaching event. Figure 6demonstrates the percentage of corals affected by bleaching across the CCMR comparing thetwo habitat types of the back-reef and fore-reef. A coral was noted if it showed signs of paling(P), partial bleaching (PB) or bleached (B). On the back-reef 16% of corals showed signs ofbleaching, this is almost a 12% increase from the phase before the summer.

Fish Results Across seven sites within the CCMR, a total of 2107 adult fish were counted.The GUZ fore-reef showed a total fish density of 0.35 ind/100m2 and the back-reef a fish densityof 0.66 ind/100m2. The CZ fore-reef showed a total density of 0.88 ind/100m2 and back-reef1.39 ind/100m2. The PZ back-reef gave a density of 0.46 ind/100m2. There is no value for thePZ fore-reef as the site was not surveyed this phase due to adverse weather conditions. The CZholds the highest average fish density of 2.39 ± 1.811, followed by the GUZ with an average of1.47/pm1.19, and then the PZ which has an average density of 1.34/pm1.69. The size structureacross the CCMR showed the most common sizes being 6-10cm and 11-20cm. The GUZ showeda decrease in fish density from the previous phase of 35%. Table 6 shows a breakdown of the

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Figure 5: Average % coverage of Gorgonian across the three different management zones in CayeCaulker Marine Reserve on the back reef (Conservation zone, General use zone and Preservationzone). Samples were taken in Belize during October to December 2018.

density of fish by family in each of the habitat types. Pomacentridae has the largest densityin the back-reef (45%) followed by Scaridae making up (19%) and Labridae making (9%). Onthe fore-reef Pomacentridae also has the largest density (31%), followed by Scaridae (23%)and Acunthuridae (11.6%). Overall across the entire CCMR the back-reef habitat boasts thehighest density ind/100m2 of 0.84 ± 0.49, compared to the fore-reef with a density ind/100m2of 0.41 ± 0.44.

Table 6: Density (ind/100m2) of each fish family in different habitat types

Habitat Type Back-reef Fore-reef

Lutjanidae 18.33 6.00Scaridae 116.33 25.00Balistidae 7.67 1.33Monacanthidae 0.00 0.00Haemulidae 58.00 5.00Epinephelidae 6.00 1.67Pomacanthidae 3.67 4.00Chaetodontidae 21.00 2.33Acanthuridae 39.00 12.67Pomacentridae 267.67 33.67Labridae 55.33 7.00Miscellaneous 19.00 10.00

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Figure 6: Average % coverage ±SE of coral colonies exhibiting bleaching across two types ofhabitat types three in Caye Caulker Marine Reserve (Back-reef and Fore-reef.) Samples weretaken in Belize during April to June and October to December 2018.

Statistical Analysis An ANOVA test was used to identify whether there was a significantdifference between the total fish densities in the three different management zones. Therewas no significant difference between the total fish densities in different management zones(F = 0.194, df = 2, P> 0.05; figure 7). A one way independent ANOVA was also used toassess whether there was a significant difference between fish densities and the two habitattypes (back-reef and fore-reef habitat). There was no significant difference between the totalfish densities between the back and fore reef (F = 0.0792, df = 1, P> 0.05). The third testran was a Kruskal Wallis test to identify whether there is a significant difference in densitybetween each family. There was a significant difference in density between each family (X2 =30.655, df = 10, P< 0.001; figure 8). A post hoc Tukey test was used to identify where thesignificance was. There was a significance between Pomacanthidae and Acanthuridae (t value= -3.588, P< 0.05), Pomacentridae and Epinephelidae (t value = 5.625, P< 0.01), Scaridaeand Epinephelidae (t value = 4.193, P< 0.01), Pomacanthidae and Labridae (t-value = -3.662, P< 0.05), Pomacentridae and Lutjanidae (t value = 4.600, P< 0.01), Pomacentridaeand Miscellaneous (t value = 3.464, P< 0.05), Pomacentridae and Pomacanthidae (t value =6.189, P< 0.01), Scaridae and Pomacanthidae (t value = 4.693, P< 0.01).

2.2.1 Discussion

Fish SMP and coral and benthic surveys Fish abundances have previously been found tobe greater in protected areas (Sedberry et al., 1996). The decline in the species seen within ourresults and a decrease in fish densities within this phase may reflect the impacts of overfishing.Particularly as the high season for tourism began within this phase, the demand for local fishmay have increased. The dominance of Scaridae, a protected species, also suggests overfishingof commercial species, reducing their competition. Pomacentridae is shown to be the mostabundant fish family, this coincides with phase 183 results. There was a greater abundanceof Scaridae observed in the conservation zone and the general use zone, this is likely to becaused by the increased fishing pressure removing commercial fish that are considered largerpredators, such as Lutjanidae and Hamulidae. Pomacentridae and Haemulidae are the mostdominant species on the back- reef habitat and Scaridae is dominant on the fore-reef habitat.

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Figure 7: Total fish densities per management zone in phase 184 (dark grey = CZ, mediumgrey = GUZ and light grey = PZ).

Moreover, a study, using the AGRRA methodology, concluded that Scaridae population sizecan be used as an indicator throughout the Caribbean for overfishing (Valles & Oxenford,2014). Pomacentridae is the most abundant on both the back-reef and fore-reef habitats asit is not a commercial fish and therefore has not got the additional pressures. Scaridae ismost likely to be the most dominant on fore-reef habitat due to the high demand for themas a commercial species, local fisherman fish on the back-reef and therefore the main pressureon the population is applied on the back-reef. A high population of Chaetodontidae on theback-reef is understood as they feed on coral polyps and sponges which are at a higher densityon the back-reef than fore-reef which was observed in the previous phases: BZM 172, 173 and174 (Motta, 1988). A very low density of Epinephelidae was found on both habitat types.With the Nassau Grouper recently being categorised as critically endangered by the IUCN, theminimal presence within CCMR should be of great concern. Not only is the species at highrisk of extinction but their low population numbers have the potential to alter fish communitystructure and cause a trophic cascade as Epinephelidae are high up within the food chain aspredators. The continuing decline in density will impact the rest of the whole food web, havingpotential catastrophic effects on the entire reef ecosystem (Pace et al., 1999). The overall totaladult fish count of the previous quarter and this quarter has decreased by 18.5%. However, itmust be taken into consideration that 8 out of the 10 survey sites were not completed due tocircumstances that could not be avoided. Though the preservation zone is the most protectedzone in the CCMR, the PZ showed the lowest density ind/100m(2). This result was also drawnin phase 183. This suggests it is not a new pressure causing this outcome. It is possible thatit is due to the habitats within the PZ, which provides less suitable habitat and live coral reefin comparison to the GUZ and CZ. Only one site within the PZ, the Swash, provides an ideal

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Figure 8: Total fish densities per fish families in phase 184.

habitat. This indicates that a change or extension in the zones is required in order to includea larger area of living reef. Moreover, the PZ is significantly smaller in size and therefore ifthe density per meter square was tested, perhaps there would be a difference. In addition, thepreservation zone is a smaller water body, as a result there is much higher competition betweenindividuals for food, territory and breeding partners.

Much like previous phases, the fish size community structure indicates that fish are notreaching sizes larger than 40 + cm very often. Due to low numbers of fish populations reachingthis size, extra stress is put on the smaller size fish, causing a possible shift in the communitystructure. The size structure is an indicator of the fishing pressure (Almada-Aleman et al.,2003). This data can be considered in the future when examining and adjusting the MPAmanagement scheme in the future as well as fishing regulations.

The contrast in the presence of bleaching from before and after the summer season isvast. This indicates that, though Caye Caulker does not experience large temperature changesthroughout the seasons, it is evident that it has an impact on the benthic environment. A long-term study suggested that coral is more susceptible to repeated bleaching from a long-termsteady sea temperature increase over rather than exposure to a season of dramatic temper-ate increase (Brown, 1997). Many studies have concluded that the most common cause forcoral bleaching is increased sea temperature, which supports the results in phases 182 and184 (Brown, 1997). Bleaching is episodic, previous mass bleaching events have been linkedto atmospheric phenomena, such as the El Nino-Southern Oscillation (ENSO). This supportsthe hypothesis that the bleaching increase is linked to the temperature increase of the summerseason in Caye Caulker. However, a larger data-set is required to make a full conclusion. Afurther study that could be done would be to record the sea temperature on monthly basis to

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Figure 9: Fish recruitment densities in 2018 phases.

identify the exact temperature threshold for coral health, then an exact temperature rise can beidentified that relates to when zooxanthallae is expelled from the mesenterial filaments of coralpolyps (Brown, 1997). Moreover, additional stressors that may potentially affect coral bleach-ing, which should be assessed is reduced salinity, solar radiation and water quality; includinginvasive bacteria (Brown, 1997).

The results displayed that the highest benthic cover in all three management zones wasalgae. This could potentially be related to the increased coral bleaching. Bleaching in thelong-term can lead to severe loss in coral cover (Baker et al., 2008). Furthermore, it can causea change in community structure (Baker et al., 2008). Bleaching leave coral colonies vulnerableto competitive overgrowth of algae. This has the potential to be catastrophic impact on coralhealth within CCMR (McCook et al., 2014). One of the biggest consequences of increase algaecover is bioerosion and abrasion. Acroprora species are shown to be susceptible to large growthreductions when exposed to competition with algae, this is of great concern as Acropora islisted as Threatened by the IUCN (Tanner, 1995; IUCN, 2015).

Future Research

It is critically important that the current AGRRA protocol and SMP surveys continue to beconducted on a quarterly basis to ensure long term monitoring which then can be used to re-assess the management zones in CCMR. Moreover, long term comparisons can be made alongthe MBRS. Future development in order to re-evaluate the sites selected for the studies couldbe conducted via drones. Accessibility to a drone would enable all sites, in all managementzones to be mapped out and compared. This can help us to understand whether the sites weare assessing are representative of the overall site. Moreover, we must question the efficiency of

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the size and location of each zone. For example, the preservation zone is the smallest the threezones but of the most importance. It provides the smallest ideal habitat for fish species, outof all the study sites. From the data collected this phase and in the past phases it has beenidentified that the conservation zone holds a higher density of fish. Perhaps this suggests thatthe preservation zones limited space has reached its threshold of carrying capacity and that apotential study testing the efficacy of a buffer or overflow zone between each zones could bebeneficial. Moreover, an evaluation of other MPA strategies could enable the improvement ofCCMR.

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2.3 Conch Survey

2.3.1 Ecology

The Queen Conch (Lobatus gigas) is the largest species of native gastropod in the Caribbean.They can grow to up to 35 cm in shell length (Appeldoorn, 1988). Conches are slow moving,relying on their protective shells as their only defense. Juvenile mortality is high from predationdue to their newly developing shell thickness (Stoner, 1997).

2.3.2 Habitat

Since Lobatus gigas are dependent on sea grass and algae cover as their habitat, they are limitedto certain depth ranges, 0.3-18 m and from 25-35 m (Welch, 2010). The Queen Conch is foundliving among sea grasses, usually they are found in association with turtle grass, Thalassiatestudinum, and manatee grass, Syringodium filiforome. They have also been found to inhabitsandy strata although juveniles tend to inhabit shallow sea grass meadows close to shore (Stoner,1988). The association with sea grass and algae is one of grazing and as a form of shelter frompredators as well. Queen Conch have been found to not only inhabit sea grass with densitiesfrom 10-50% and 50-90% cover sea grass, but the sea grass functions as a valuable nurseryhabitat for juveniles (Doerr & Hill, 2018).

2.3.3 Life Cycle

Queen Conch are gonochoristic, meaning an individual is either male or female, a female will beinternally fertilised by a male before producing long gelatinous strings of eggs. The spawningfrequency and number of egg masses has been shown to increase proportionally with increasingtemperature, meaning the egg production of Queen Conch increases in higher temperatures(Cala et al., 2013). In addition, a positive feedback relationship between spawning and game-togenesis in females has been shown (Cala et al., 2013).

2.3.4 Predators and threats

Fishing pressure is one of the main and most impactive threats to the population of QueenConch in the Caribbean. Studies have shown that over repeated surveys the densities of matureconch have declined and the populations have become younger over time (Kough et al., 2017).The reason for the high fishing pressure on Queen Conch is that it is an important localfood source and is economically important as an export product throughout the Caribbean(Randall, 1964). The conch fisheries in Belize have grown rapidly in the last few centuriesincreasing its exports from 437747.9 kgs in 2013 (Department, 2014). As with all exploitedfishery populations, total biomass decreased as the abundance of older reproductive adultsdeclined and the size distribution shifted towards immature conchs. The closed conch seasonwas implemented in 1978 alongside a catch size limit (Strasdine, 1998). Currently there is alegal minimum size, 18 cm, for conch catching (Stoner, 1997). However, conch reaches sexualmaturity at around 21 cm, which has led to an overall decrease of average conch size. This hashad a domino effect wherein conch egg production is in decline, due to the reduced numberof large adults that produce larger numbers of offspring (Jesus-Navarrete et al., 2003). Notonly has average shell length decreased but so has population density. Population density ofQueen Conch has decreased from 2.651 conch m(2) in 1997 to 0.006 conch m(2) in 1997 (Jesus-Navarrete et al., 2003). The fishing pressure on Queen Conch has therefore reduced the overallbiomass, age and size at sexual maturity. Conch as a commercial resource is highly valuedas a substitute to lobster income when the lobster season is closed (Joyce, 1997; Aiken et al.,1999). In Belize, conch fisheries are small-scale and characterized by skiffs which are owned or

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jointly owned by a senior operator with a middle-income level (Berkes et al., 2001). The use ofpoisons, dynamite, SCUBA, and hookah are illegal. Conch fishing grounds are thus limited towaters less than 30 m as this is the maximum depth for most skin divers. Skin diving in CayeCaulker is further limited by visibility which is better behind the barrier reef and on the atolls(Wetmore, 2014). Despite the regional management strategies in place, the Queen Conch is stilllisted as a commercially threatened species, and has been since 1992 by CITES (the Conventionon International Trade in Endangered Species of Wild Fauna and Flora), there has been nosubstantial recovery of the population of queen conch (Stoner & Ray, 1996; Stoner et al., 1996;Stoner, 1997; Jesus-Navarette et al., 2003). In 1998, the conch fishery accounted for 20% ofearnings from marine produce, approximately BZ $3 million (Belize Fishery Department, 2001).The conch fishery is dependent on national stocks, as opposed to pelagic stocks, and cannoteasily relocate when stocks decline. Even with the dependence on national stocks, Belizesconch fishery demonstrates a pathological resource use (Smith & Berkes, 1991; Hughes, 1994;McManus, 1988).

2.3.5 Aims & Objectives

This aims of this project are to assess the status of Queen Conch populations around CayeCaulker and provide data to the BFD to inform management decisions on the effectiveness ofthe conservation zones and fishery season management within the CCMR.

• Set up long-term survey monitoring of conch populations; including density, size andmaturity.

• Compare differences in the Queen Conch population between different conservation zoneswithin the CCMR.

• Evaluate the sustainability of local conch fishing practice, the effectiveness of multi-zonemanagement and the efficacy Queen Conch fishery season.

2.3.6 Methodology

The protocol for this methodology was adapted from the methodology used by the BFD. Toevaluate local Queen Conch fishing practices, which involve fishermen snorkeling and skin div-ing to collect conchs, snorkel surveys will be used. Size is a useful parameter for fisheriesmanagement because the Queen Conchs shell and lip size can be used to assess sexual maturity(Avila-Poveda & Baqueiro-Cardenas, 2006; Foley & Takahashi, 2017). As size of the conch isboth useful and easy to measure this is the main parameter used during Frontiers Queen Conchsurveys in Belize. Three zones have been selected for surveying annually during the week beforeconch season opens. Below is a description of each of the conservation survey zones:

Conch Site 1: (General Use Zone) No person is permitted to use long lines, spear guns ornets unless authorized by the Fisheries Department. Commercial fishing is allowed with a validfisher folk license, traps must be placed 100 m from corals. Sport fishing is allowed underlicense from the Fisheries Department, but spear fishing is not permitted. Catch and releasetours may only remove fish for subsistence purposes.

Conch Site 2: (Preservation Zone) No fishing, sport fishing, diving or any other water activ-ities are permitted within the preservation zone. Motor boats are not to be operated within thepreservation zone except for emergencies or with permission from the Fisheries Department.

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Conch Site 3: (Conservation Zone) No fishing or collection of marine items of any type. Onlynon-extractive recreational activities allowed in the conservation zone. No feeding of marinelife is allowed except from licensed tour guides. Water skiing and jet skiing are not allowed.

The BFD has allocated 21 sites for Frontier to survey around Caye Caulker. Each survey is100 m long, and 2 m wide, giving a total of 4200m(2) surveyed areas. The sites were randomlyselected, with some within the CCMR and some outside of the reserve on the west side of CayeCaulker.

For each survey a 100 m transect line is laid out by the boat. Once the transect line is laid,two surveyors search for Queen Conchs along a 4 m wide belt transect (2 m either side). Eachconch found inside the belt-transect was collected and carried to the boat for measurementand to avoid duplicate recording of individuals. Once every conch is collected the survey teammeasures the total length of each conch shell and the thickness of the lip, if present, usingcallipers. Once all the conchs are measured, they are placed carefully back on the seafloor atthe site where they were found.

Conchs are divided into three maturity status categories according to the measured lipthickness. Queen Conchs with lip thickness > 5mm were classified as adults, those with a< 5mm lip were classified as sub-adults, and those where the lip was absent were classified asjuveniles (Horsford et al., 2012).

2.3.7 Results

The conch survey data has so far been collected for 2017 and 2018 and both years they wererecorded in September for consistency. The surveys are completed in September because that isthe last month of the closed season for conch fishing. For 2017, there were a total of 20 transectswith 69 total conches found in the three sites surveyed, those sites being the: General use zone,Preservation Zone and Northwest Caye Caulker. In 2018 there were a total of 25 transectsrecorded across four sites, those being: General use zone, Preservation zone, Conservation zoneand South Caye Caulker. There were a total of 714 conches found across all the sites surveyedin 2018, which is a 934% increase from the year before (2017).

Figure 10 shows the mean abundance across all sites for both 2017 and 2018, and fromthose two years the most conches were found in the conservation zone in 2018. However, moredata is required before any correlations can be made. Unfortunately, due to the way MicrosoftExcel works there is some data on the graph that is hidden. In 2017, the mean abundance forNW CC was 0.25, for 2017 in the PZ the mean was 0.20 and in 2018 for S CC the mean was 0.Regarding the graph above, the W CC, CZ and S CC sites were not surveyed in 2017 neitherwas NW CC site in 2018.

Figure 11 shows the mean shell length (mm) of the Queen Conch, from the data displayed thePZ has the greatest mean shell length (mm) that is 242 mm, however, that is only one readingand so with future years may change that mean. The W CC site was surveyed; however, noconches were found. In addition, the CZ, W CC and S CC were not surveyed at all in 2017,so there is a lack of data. Lastly, the NW CC site was not surveyed in 2018, but will be in thefuture in order to compare with previous data.

A t test was used to compare the mean lip thickness from 2017 with 2018. There wasno significant difference between lip thickness in 2017 and 2018 (T = 0.0000195620252857935,P > 0.05; figure 12).

The 2017 survey has showed a total density from all sites of 0.8625 conch/100m(2), withat least 1 conch found in 70% of the sites. One site recorded 34 conchs, which accounted for46% of the total surveyed. Of the total, 26 conchs (35%) had a lip with 11% reaching adultsize. The 2018 survey showed a total density of 7.14 conch/100m(2), of six total sites five weresurveyed and there was at least one conch found in each site. Conch were found in 66.67% of

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Figure 10: Mean abundance of Queen Conch, Lobatus gigas, across six sites: West Caye Caulker(W CC), General use zone (GUZ), Preservation zone (PZ), Conservations Zone (CZ), SouthCaye Caulker (S CC), Northwest Caye Caulker (NW CC). The bars with diagonal lines aredisplaying data from 2018, the dotted bars are displaying data from 2017

Figure 11: Mean average shell length (mm) of Queen Conch, Lobatus gigas, across six sites:West Caye Caulker (W CC), General use zone (GUZ), Preservation zone (PZ), ConservationsZone (CZ), South Caye Caulker (S CC), Northwest Caye Caulker (NW CC). Plotted with errorbars displaying 1 standard deviation from the mean. Those without error bars only had onerecorded conch. The bars with diagonal lines are displaying data from 2018, the dotted barsare displaying data from 2017.

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Figure 12: Scatter plot showing shell length (mm) plotted against lip thickness (mm) for both2017 and 2018 across four sites: General use zone (GUZ), Preservation zone (PZ), Conservationzone (CZ) and South Caye Caulker (S CC). The data from 2017 is plotted as Xs and the datafrom 2018 is plotted as s. This plot only contains the data where both the shell length and thelip thickness were measured.

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Table 7: Summary table of results. Average density (conch/100m(2)) is calculated by firstcalculating the total area sampling and dividing that total area to calculate the number ofconch found in 100m(2).

Total conch Average Density Conch/100m(2) % Of Sites with Conch Highest Recorded Abundance Total Conch with Lip % Conch with Lip % Adult Conch with Lip2017 67 0.86 70 34 26 35 112018 714 7.14 66.67 187 139 19.5 25.17

all the sites. The highest recorded abundance was 187 conches found in the conservation zone,this accounted for 26.19% of the total surveyed. In total, 139 conches (19.5%) had a lip andout of that 19.5%, 25% reached adult size (table 7).

2.3.8 Discussion

This project is currently in development to build a long-term monitoring program on the statusof the Queen Conch in Caye Caulker, in order to compare the differences in the Queen Conchpopulation between different conservation zones within the CCMR. Additionally, this projectaims to evaluate the sustainability of local conch fishing practice, the effectiveness of multi-zonemanagement and the efficacy of Queen Conch fishery season.

Our preliminary observations are that the majority of conches are found in the no catchareas (CZ and PZ). The sites which were allocated around the island without protected statusyielded almost no sightings. Adult Queen Conch are classified as any individual less than 21cm in shell length. It is concerning that under 11% of conchs found were classified as adults,in 2017, as it may indicate that conchs are being harvested before reaching adulthood, andtherefore before being able to reproduce. Other studies have suggested a high percentage ofadults in a non-reproductive stage, along with 40% adult conch surveyed (Aldana Aranga etal., 2014). Currently there is no law stating that the conch must have a lip to be harvested.More data from this project may give an incentive to the BFD to implement a minimum lipsize for Queen Conch landings. In 2018, the number of conches classified as adults droppedto 2.94% of total conch found. This reduction in adult abundance since 2017 is concerningas it gives a preliminary insight into the possibility that the closed season is not working as apreventative method to protect the population of Queen Conch. Other management methodshave also been shown to actually increase illegal catches, including minimum size of 200 mmand permanent ban on coastal areas in Costa Sur (Jesus-Navarrete et al., 2003; Stoner, et al.,2018). The reduction of adult abundance will only escalate as the immature Queen Conchesreach sexual maturity and will then be harvested come open season. Already the concerns areproblematic because this means the amount of conch able to spawn is heavily reduced which willhave a heavy impact on the reproductive stock (Jesus-Navarrete et al., 2003). Along with thedecreasing population of Queen Conch, a reduction in commercial catch (Brownell & Stevely,1981).

The data we have collected so far has been collected in September, the last month before theopen season starts. Although this data is only preliminary and no patterns can be identifiedas of yet, this data will be improved year after year to identify a trend in the populationabundance, shell length and lip thickness/absence.

Future research Comparisons between the protected zones and the fishing grounds wouldbe an interesting project to research in the future. This may influence the decision to rearrangethe current protected zones to accommodate for the change in abundance of conch. Perhaps asocial science project could be undertaken, one which would involve discouraging the buying ofconch shells since this permanently removes the calcium carbonate which makes up the shell

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from the ecosystem. Currently the Conch study conducted only briefly assesses the habitatat each survey site. Conchs can be very preferential in their habitat selection. As the sitesare selected through random coordinate sampling, the preferential habitat selection by conchssuggest this might not be the most suitable method of selecting survey sites. For example,10 out of 18 sites may have no Conchs present but this may not be due to overfishing, thiscould be due to dense areas of seagrass and depths exceeding 7 meters which are not optimum/preferential habitat conditions for conch. Conch are known to avoid seagrass when they arereproducing, and juvenile Queen Conch select coral rubble as their habitat of choice (Glazer &Kidney, 2004; Stoner, 2003). In addition, it has been found that the Queen Conch are shown toweigh more when settling in sand than seagrass, however it is not known why (Ray & Stoner,1995). Moreover, the frequency of which conch surveys are conducted could increase to oncea month, much like with the lobster study, in order to provide a more detailed conclusion onconch populations and what and when the population is declining and/or recovering. Thismay also aid in identifying what the specific pressures are that cause the fluctuation in thepopulation size.

In addition, regular assessments of what is sold at fishermans cooperative on Caye Caulkercould also give a larger representation of population decline and the exact pressures from fishing.However, this would have to be conducted with the assistance of the Fisheries Department asdue to cultural difference, foreign entities question catch quantities is likely to give false results.

2.4 Caribbean Spiny Lobster Surveys

2.4.1 Introduction

Ecology The Caribbean Spiny Lobster (Panulirus argus) is one of the most prominent crus-taceans in the Caribbean Sea. This species is listed as data deficient on the IUCN red list,which means further study is required to ascertain whether conservation methods are working(IUCN Red List of Threatened Species, 2019). It is a decapod covered with a spiny exoskeletonwhich provides protection from some predators. Like most crustaceans, they must shed (ormolt) their exoskeleton at regular intervals as it does not expand with their own growth, thisprocess occurs approximately two to three times per year (Williams, 1984). During this processit is more vulnerable to predation whilst the new exoskeleton hardens. P. argus can be preyfor sharks and certain groupers who are able to digest the hard-shell-like exoskeleton (Oceana,2019). Maturity is generally estimated to occur around two years and full-grown adults canreach between 12 and 20 years of age (Chavez, 2001; Erhardt, 2005).

Commercial Species P. argus is the most important commercial marine species in theCaribbean due to its economic value, both as a source of direct income and employment for thelocal population, as well as foreign exchange for national governments (Acosta & Robertson,2003). Lobster fishing is the primary fishery for 24 Caribbean nations (Gnanalingam & ButlerIV, 2017). Furthermore, due to the high unit prices, the international trade of lobster providesimprovements to the livelihoods of fishery-dependent populations (Monnereau & Pollnac, 2012).

Habitat and nutrition These lobsters inhabit a wide range of habitats around Caye Caulker.The larval stage begins in seagrass meadows; they will then migrate to mangrove areas duringtheir juvenile stage; and eventually migrate again to rubble or reef areas after reaching adult-hood (Acosta, 1997). They detect food and prey using their front mounted antenna which areessentially olfactory sensors (Derby et al., 2001). The lobsters are important predators; andtheir primary diet consists of molluscs (gastropods, chitons and bivalves) and arthropods (Coxet al., 1997; Seudeal, 2015). They are also believed to show preference for certain foods based

28

on their own habitat and neural receptors. Ecologically, Caribbean spiny lobsters are not onlykeystone species they prey upon other organisms, but they also serve as prey for a wide rangeof marine animals, including sharks, rays, turtles and moray eels (Seudeal, 2013). Caribbeanspiny lobsters are found in a wide range of habitats including sand, seagrass, coral reefs andcoral rubble (Seudeal, 2013).

Status Due to lack of abundance data, their threatened status is currently assessed as datadeficient on the IUCN Red List, this is likely due to the cryptic nature of their behaviour andappearance (Butler et al., 2013). Unsustainable fishing of this species could lead to furtherpopulation declines and ultimately collapse of the industry, particularly in Belize, where adecline of 28% in catch per unit effort has been reported by fishermen, from 2.7 kg per fishingday to 1.94 kg per fishing day between the years 1999 and 2009 (Gongora, 2010; Butler et al.,2013; CZMAI, 2016).

Protection and monitoring P. argus are a social species and when there is an abundanceof food, they are known to form high population densities, similar to the American lobster(Homarus americanus) (Behringer & Butler, 2006). This suggests healthy, sustainable locallobster populations can be maintained through effective fishery management and protectedareas, like the conservation zone and preservation zone found within CCMR which can functionas a refuge for P. argus. In these protected areas lobsters are expected to be found in higherpopulations, have a larger mean size and thus are often more reproductively successful due toincreased fecundity (Acosta & Robertson, 2003). These protections should result in increasedregional larval supply and net movements of adult individuals from the reserve to adjacentfishing grounds. In addition to the protection granted by marine reserves, the Belize lobsterfishery is seasonal; the fishery is closed on February 14th until June 15th country-wide whichprotects the lobster populations throughout their reproductive season. Furthermore, P. arguslandings must adhere to a minimum size limit/carapace length of 7.6 cm and tail weight of 4ounces (113 g), which is applied throughout the year. There is also a ban on the use of SCUBAfor catching lobsters and other gear restrictions and license limitations. Despite this there is nocurrent catch quota for lobsters (Babcock, 2012), as such fishermen can catch as many lobstersas they are able to find of legal size in the general use zone. This can devastate the local lobsterpopulation in this zone, it has also been predicted that 2% of annual catches are undersizedand thus illegal (Gongora, 2004).

Given the importance of P. argus at both an economic and ecosystem level, it is essentialto assess the effectiveness of the conservation efforts mentioned above, to gain an insight intothe sustainability of the lobster fishery. Frontier aims to carry out long-term monitoring andassessment of the local population size structure and sex ratios of the P. argus. Surveys areconducted and compared between each zone of CCMR to provide data on the effectiveness ofspatial management on the local lobster population.

In addition to vital information on the sustainability of the lobster fishery, data on lobsterdistribution could potentially provide insight into the complexity and structure of the localcoral reef systems. This is because higher densities of P. argus have been associated with areasof higher ecosystem complexity, where the reef is intricate and creates crevices and hidden sitesfor lobsters (Rios-Lara et al., 2007). Higher lobster densities could therefore be used as anindicator of reef habitat complexity and general health.

This report aims to monitor the population of the P. argus throughout the year with theCaye Caulker Marine Reserve in order to monitor the populations of the commercial specieswhich hundreds of fishermen rely on. Further management legislations cannot be put in placeunless there is scientific evidence to support the claims.

29

2.4.2 Methodology

The methodology for this is taken from the LAMP (Long-term Atoll Monitoring Protocol),designed by a local NGO WCS (Wildlife Conservation Society) which is the nationwide tech-nique used to survey both lobster and queen conch (Acosta, 2006). In line with the aim ofthis study, surveys were carried out within the Caye Caulker Marine Reserve (CCMR). GPSlocation of the survey sites were recorded, to ensure the same sites were surveyed each time.The sites surveyed are; the Island, the Swash and the Graveyard. Surveys were carried outusing an active search approach, which consisted of a team of snorkelers actively searching forlobsters without the use of fixed transects. By splitting the site up, the surveyors make surethat the same lobster is not recorded twice. The rationale behind this method was to simulatelocal fishing practices and get an estimate of Catch Per Unit Effort (CPUE); the effort unitwas time (minutes).

Each individual snorkeled for 30 minutes looking for lobsters. Each individual lobster foundwas measured using a ruler to approximate its total carapace length. Sex was also recorded,which was carried out by determining if two extra claws were present on the back legs, or ifthere was an extra pair of swimming legs underneath the tail in females. Despite best effortsit was not always possible to visually determine the lobsters sex, as they are usually foundunderneath rock crevices with their abdomen not visible to the surveyor. Lobsters were nothandled in order to minimize disturbance. If lobsters escaped before measurements could bemade, then their carapace length was estimated by sight. During this quarter lobster surveyswere conducted monthly on the 14th-15th of each month to gain better insight into how thepopulation develops on a more regular basis. Each survey was conducted with between threeand six surveyors snorkeling the site and recording data and were always led by at least one staffmember. A thorough briefing and sufficient training was provided to ensure the data recordedwas as accurate as possible.

All data analysis was conducted using RStudio version 1.1.456 (R Core Team, 2018). Forthe statistical analysis of carapace length female and male Caribbean spiny lobster were testedseparately due to the difference in carapace lengths between the sexes. To determine whetherthere is a significant difference between the carapace lengths of female and male lobsters indifferent management zones throughout 2018 a Kruskal Wallis test was used. A one wayANOVA was used to determine if the carapace lengths of female and male lobsters in differentzones in just phase 184.

2.4.3 Results

A total of 135 individual lobsters were counted across all surveys with 84 being males and 51females. Overall the CZ and PZ contained the highest numbers of P. argus (CZ mean = 8.43,SD = 5.65; PZ mean = 8.38, SD = 3.66) with the GUZ has the lowest abundance throughoutthe year with only 2 individuals being found (mean = 1.1, SD = 1.36). The highest number oflobsters recorded during a single survey was in February in the CZ with 12 individuals. Themean and standard deviation of carapace length in male and female lobsters in each managementzone is compared in table 8.

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Table 8: The Mean and Standard deviation (SD) for carapace length of all lobsters and formale and female Caribbean spiny lobsters shown separately for each management zone.

CZ PZ GUZMean SD Mean SD Mean SD

All lobsters 92.48 60.89 140.1 73.83 112.34 57.07Females 92.97 61.71 104.03 65.52 141.11 60.09Males 92 61.01 116.40 78.45 82.57 33.99

2.5 Lionfish Study

2.5.1 Introduction

Lionfish (Pterois) are an invasive species of the Caribbean, which has been shown in previousstudies to cause deleterious impacts on the coral reef food web (Albins & Hixon, 2011). Thefocus of this study is to monitor the lionfish population throughout the management zones ofCCMR throughout the year. This study will also assess the efficiency of the current cullingmethods of lionfish and the recovery rate of the population after the Caye Caulker LionfishDerby annually in March.

FeedingLionfish (Pterois) are recognised as excellent generalist predators for two significant reasons.Firstly, lionfish move slowly, and they possess cryptic colouring making them appear as a loosetuft of seaweed, thus providing them with substantial camouflage (Hixon et al., 2016). Inaddition to this, while stalking their prey lionfish flare their large pectoral fins and encouragesmall and juvenile fish to herd towards them, after which they are rapidly consumed (Albins &Hixon, 2011). Furthermore, lionfish adopt an array of feeding strategies including ambushingprey, as well as ejecting jets of water at their prey, disorientating, and stunning them (Hixonet al., 2016). The means by which lionfish intake their prey allows for massive numbers to beconsumed at one time, it has been observed that they can consume 20 individual prey fish perhalf hour (Albins & Hixon, 2011), this is facilitated by the ability for their stomachs to expandby 30 times their original size (Morris Jr & Whitfield, 2009). Lionfish utilise a method ofsuction to intake prey, this process occurs through the extension of their buccal and opercularcavities along with a rapid forward swim, taking in prey within their path (Morris Jr & Akins,2009).

Furthermore, lionfish possess bilateral swim bladder muscles which give them the uniqueability to control their pitch, allowing them to fine-tune their position within the water. Thisallows them to orient themselves in upside down and lateral positions within the water whichfacilitates their ability to strike swiftly at prey (Morris Jr & Akins, 2009).

Lionfish are estimated to consume 2-3 times more native fish species than competing nativepredators within the Caribbean reefs (Rocha et al., 2015). The expansive number of speciesconsumed is estimated at over 40 different species (Albins & Hixon, 2011). As aforementionedthe diet of the lionfish is broad and comprises of both juvenile large fish species and small fishspecies alongside a large range of invertebrates. The native prey that are consumed by lionfishare of great concern as they consist of the commercially and recreationally important nativefish species, including but not limited to, groupers and snappers as well as the ecologicallyvital reef grazers including parrotfish and surgeonfish, both of which are protected under strictlaws within Belize (Hixon et al., 2016). These adaptations make lionfish highly deleteriousgeneralised invasive predators and a cause for concern within the Caribbean reefs.

Predators Within invaded regions lionfish lack natural predators allowing the populationsto grow uncontrollably. It has been suggested that their lack of predators may have arisendue to their cryptic appearance and movements causing any native predators to not recognise

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lionfish as potential prey (Albins & Hixon, 2011). Undetectable chemical cues further assistin the ability for lionfish to remain cryptic and avoid predation (Hackerott et al., 2017). Inaddition to this, lionfish are protected by long venomous spines which have the potential todeter predators from attack (Morris Jr & Whitfield, 2009). It has been observed that attackingsharks and groupers will almost immediately retreat when attacking lionfish (Albins & Hixon,2011). The lack of natural predators present within areas invaded by lionfish will allow for thepopulations to grow exponentially if management schemes are not introduced.

Reproduction and Spawning Lionfish have been observed to reach sexual maturitywithin their first year of life, with females maturing at around 180mm and males at 100mm(Morris Jr & Whitfield, 2009). Lionfish are gonochoristic pair spawners meaning that femalescan release two floating egg masses, one from each ovarian lobe at a single time (Butterfieldet al., 2015). Prior to spawn release lionfish will exhibit a courtship ritual which occurs be-fore dusk and may continue for many hours proceeding (Morris Jr & Whitfield, 2009). Postcourtship the female lionfish will ascend towards the surface to release the egg masses. Thiswill occur every four days throughout the year resulting in a total annual fecundity of approx-imately two million eggs (Morris Jr & Whitfield, 2009), leading to massive population growthand expansion rate.

The lionfish embryos develop at the oceans surface. Upon hatching the larvaes appearanceis described as having a large head, long triangular shaped snout, serrated head spines, largepelvic spines and pigmented pectoral fins and they are estimated to be a length of 1.5mm(Morris Jr & Whitfield, 2009). This larval stage duration is approximated at 26 days posthatching; however, this can be impacted by other factors such as temperature and nutrientlevels (Morris Jr & Whitfield, 2009).

During the 26-day post hatch interval the lionfish larvae can disperse long distances throughocean currents (Butterfield et al., 2015). The means of dispersal and their high reproductivefecundity allows lionfish populations to spread across massive regions, in the case of the currentinvasion this distance is as extensive as 7 million km2 (Dahl & Patterson III, 2014).

The Threat and Current Management An invasive species is defined by the successfulsurvival and reproduction of a non-native species into a new habitat (McCleery, 2011). Invasionwithin marine environments are exceedingly rare due to the complex systems and food websin place, however in the case of the lionfish their invasions have shown unprecedented levels ofdispersal and they have the ability to cause high levels of potential destruction in the invadedhabitats (Green et al., 2012). The means by which the lionfish invaded habitats is still underdebate however the most widely accepted hypothesis is that the lionfish were released or hadescaped from a marine aquarium, this is due to genetic marker analysis suggesting that theexpansive lionfish population occurred from a few individuals (Morris Jr & Whitfield, 2009).

As previously mentioned, lionfish adopt a generalist diet, lack effective predators, and theirextremely high reproductive fecundity means that their ecological impact will be unparalleled iftheir population numbers are not controlled and managed (Ferreira et al., 2015). Due to the highecological impact that this species has on coral reef habitats and ecosystems, several countriesglobally have introduced removal schemes by which fishermen and divers are encouraged tohunt the invasive lionfish as an alternate food source (Barbour et al., 2011). In Caye Caulker,Belize lionfish hunting is seen as a common recreational activity and Frontier, alongside a localbusiness, host an annual lionfish derby during which there is a competitive removal of lionfish.

2.5.2 Lionfish Survey Methodology

Lionfish surveys have been conducted since phase 173, these surveys were based off the roversurveys previously conducted. Lionfish surveys are an effective means to monitor the lionfish

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populations within the Caye Caulker reef, they are specific and introduce a new aspect to theproject as well as assist the local people to manage and control the invasive populations. Eachsurvey was a 45-minute rover survey, when on the back reef the divers could swim around thesites, looking closely in crevices and under rocks. A rover survey consists of actively look forthe focus species within the selected site for a given period, for the purpose of this study, theduration is 45 minutes. When on the fore-reef, the surveyors would swim along the wall of aspur and then swim back up on top of the spur to cover the entire area of the spur and grooveformation. Each lionfish and its size class, to the closest 10 cm, were recorded. This year thedata collection will be done every six weeks to track the population trends. A comparison ofthe fish counts will be made between each of the three zones and each habitat type.

From this phase lionfish population control dives have been introduced. The volunteersact as spotters and the staff will be equipped with Hawaiian slings and home-made holdingapparatus. Once a lionfish is spotted a staff member will collect the lionfish using the sling andstore them in containers called zookeepers. After the dive, the lionfish length is measured incentimetres, they are weighed, and the venomous spines are cut off by staff using scissors forhealth and safety precautions.

Data analysis was conducted using the statistical programme Rstudio version 1.1.456 (RCore Team, 2018). An ANOVA test was run to compare if there was a significant variance oflionfish counts across each of the phases (p=0.05). A post hoc Tukey test was then used toidentify where the significance was.

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2.5.3 Results

During phase 183 a total of eight lobsters were found in the GUZ, seven lobsters in the CZ, andfive lobsters in the PZ. There was a significant difference between the total number of lionfishfound in the different phases (f = 2.7, df = 4, p = 0.05). Phase 174 had significantly largerlionfish counts (Mean = 8, SD = 6.87) compared to phase 182 (Mean = 1, SD = 0.89, p =0.03).

2.5.4 Discussion

There appears to be a sudden drop in the population of lionfish during the 182 phase, comparedto the population count previously. This phase falls right after the annual lionfish derby held onthe Caye as well as Easter time. During the last lionfish derby hosted by the Barrier Reef SportsBar over 500 lionfish were caught. The FD does not permit any fishing within the CCMR atall for the derby, however all the lionfish caught during the derby come from around the island.Removing a great number of lionfish around the island may reduce the overall population oflionfish in the reserve, as the lionfish are caught before they can reach a mature age to migrateto the reef. These results may indicate the effects of anthropogenic efforts, such as the derby,on a fish population. The highest count was seen during 174, which is 6 months after the derby,this suggests there has been a sufficient amount of time after the derby to allow the populationto recover.

With only two years of population data, it is difficult to draw conclusions on lionfish pop-ulation within the CCMR. The lionfish season is all year round and fishermen are encouragedto kill lionfish whenever they can. The immediate benefit to the fishermen is that lionfish canbe sold to local restaurants on the island allowing for an income. If lionfish become a suc-cessful alternate fishery product the fishing pressure on the already overburdened native fishstocks could be lessened (Moore, 2012). The lionfish population in Belize was slow to increasewith the first sighting in 2008 and only 13 confirmed sightings by 2009. This gave the BelizeFisheries Department and ECOMAR a chance to prepare fishermen and start their lionfish:wanted dead or alive poster campaign. It is possible that this is the reason that Belize has, forthe moment, managed to mitigate the damage the lionfish could have caused if left unchecked(Moore, 2012). The PZ zone showed the highest number of lionfish potentially due to the banof all activity within the zone. Fishermen and tour guides have been authorised by the FisheriesDepartment to hunt lionfish in the CZ and GUZ which would explain the high abundance inthe PZ compared to the two other zones. Allowing controlled fishing unfortunately would givethe opportunity to illicit fishing activity so only controlled lionfish culling should be conductedby the Fisheries Department or the Frontier team, as it currently is.

Regular population surveys will allow for long term monitoring of the lionfish populations.This project meets a knowledge gap in the lionfish population around Caye Caulker, which canbe developed into understanding the site-specific lionfish feeding and behavioural ecology.

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2.6 Caribbean Spiny Lobster Surveys

2.6.1 Introduction

EcologyThe Caribbean Spiny Lobster (Panulirus argus) is one of the most prominent crustaceans

in the Caribbean Sea. This species is listed as data deficient on the IUCN red list, which meansfurther study is required to ascertain whether conservation methods are working (IUCN, 2000).It is a decapod covered with a spiny exoskeleton which provides protection from some predators.Like most crustaceans, they must shed (or moult) their exoskeleton at regular intervals as itdoes not expand with their own growth, this process occurs approximately two to three timesper year (Williams 1984). During this process it is more vulnerable to predation whilst thenew exoskeleton hardens. P. argus can be prey for sharks and certain groupers who are able todigest the hard-shell-like exoskeleton (Oceana, 2015). Maturity is generally estimated to occuraround 2 years and full-grown adults can reach between 12 and 20 years of age (Chavez, 2001;Erhardt, 2005).

Commercial SpeciesP. argus is the most important commercial marine species in the Caribbean due to its

economic value, both as a source of direct income and employment for the local population, aswell as foreign exchange for national governments (Acosta & Robertson, 2003). Lobster fishingis the primary fishery for 24 Caribbean nations (Gnanalingam & Butler IV, 2017). Furthermore,due to the high unit prices, the international trade of lobster provides improvements to thelivelihoods of fishery-dependent populations (Monnereau & Pollnac, 2012).

Habitat and NutritionThese lobsters inhabit a wide range of habitats around Caye Caulker. The larval stage

begins in seagrass meadows; they will then migrate to mangrove areas during their juvenilestage; and eventually migrate again to rubble or reef areas after reaching adulthood (Acosta,1999). They detect food and prey using their front mounted antenna which are essentiallyolfactory sensors (Derby CD et al., 2001). The lobsters are important predators; and theirprimary diet consists of molluscs (gastropods, chitons and bivalves) and arthropods (Cox etal., 1997; Seudeal, 2015). They are also believed to show preference for certain foods basedon their own habitat and neural receptors. Ecologically, Caribbean spiny lobsters are not onlykeystone species they prey upon other organisms, but they also serve as prey for a wide rangeof marine animals, including sharks, rays, turtles and moray eels (Seudeal, 2013). Caribbeanspiny lobsters are found in a wide range of habitats including sand, seagrass, coral reefs andcoral rubble.

StatusDue to lack of abundance data, their threatened status is currently assessed as data deficient

on the IUCN Red List, this is likely due to their cryptic nature of their behaviour and appear-ance (Butler et al., 2013). Unsustainable fishing of this species could lead to further populationdeclines and ultimately collapse of the industry, particularly in Belize, where a decline of 28%in catch per unit effort has been reported by fishermen, from 2.7kg per fishing day to 1.94 kgper fishing day between the years 1999 and 2009 (Gongora, 2010; Butler et al., 2013; CZMAI,2016).

Protection and MonitoringP. argus are a social species and when there is an abundance of food, they are known to

form high population densities, like the American lobster (Homarus americanus) (Behringer& Butler, 2006). This suggests healthy, sustainable local lobster populations can be main-tained through effective fishery management and protected areas, like the conservation zoneand preservation zone found within CCMR which can function as a refuge for P. argus. Inthese protected areas lobsters are expected to be found in higher populations, have a larger

35

mean size and thus are often more reproductively successful due to increased fecundity (Acosta& Robertson, 2003). These protections should result in increased regional larval supply andnet movements of adult individuals from the reserve to adjacent fishing grounds. In additionto the protection granted by marine reserves, the Belize lobster fishery is seasonal; the fisheryis closed on February 14th until June 15th country-wide which protects the lobster popula-tions throughout their reproductive season. Furthermore, P. argus landings must adhere to aminimum size limit/carapace length of 7.6 cm and tail weight of four ounces (113 g), which isapplied throughout the year. There is also a ban on the use of SCUBA for catching lobstersand other gear restrictions and license limitations. Despite this there is no current catch quotafor lobsters (Babcock, 2012), as such fishermen can catch as many lobsters as they are able tofind of legal size in the general use zone. This can devastate the local lobster population inthis zone, it has also been predicted that 2% of annual catches are undersized and thus illegal(Gongora, 2004).

Given the importance of P. argus at both an economic and ecosystem level, it is essentialto assess the effectiveness of the conservation efforts mentioned above, to gain an insight intothe sustainability of the lobster fishery. Frontier aims to carry out long-term monitoring andassessment of the local population size structure and sex ratios of the P. argus. Surveys areconducted and compared between each zone of CCMR to provide data on the effectiveness ofspatial management on the local lobster population.

In addition to vital information on the sustainability of the lobster fishery, data on lobsterdistribution could potentially provide insight into the complexity and structure of the localcoral reef systems. This is because higher densities of P. argus have been associated with areasof higher ecosystem complexity, where the reef is intricate and creates crevices and hidden sitesfor lobsters (Rios-Lara et al., 2007). Higher lobster densities could therefore be used as anindicator of reef habitat complexity and general health.

This report aims to monitor the population of the P. argus throughout the year with theCaye Caulker Marine Reserve in order to monitor the populations of the commercial specieswhich hundreds of fishermen rely on. Further management legislations cannot be put in placeunless there is scientific evidence to support the claims.

2.6.2 Methodology

The methodology for this is taken from the LAMP (Long-term Atoll Monitoring Protocol),designed by a local NGO, Wildlife Conservation Society (WCS), which is the nationwide tech-nique used to survey both lobster and queen conch (Acosta, 2006). In line with the aim ofthis study, surveys were carried out within the Caye Caulker Marine Reserve (CCMR). GPSlocation of the survey sites were recorded, to ensure the same sites were surveyed each time.The sites surveyed are; the Island, the Swash and the Graveyard. Surveys were carried outusing an active search approach, which consisted of a team of snorkelers actively searching forlobsters without the use of fixed transects. By splitting the site up, the surveyors make surethat the same lobster is not recorded twice. The rationale behind this method was to simulatelocal fishing practices and get an estimate of Catch Per Unit Effort (CPUE); the effort unitwas time (minutes).

Each individual snorkelled for 30 minutes looking for lobsters. Each individual lobster foundwas measured using a ruler to approximate its total carapace length. Sex was also recorded,which was carried out by determining if two extra claws were present on the back legs, or ifthere was an extra pair of swimming legs underneath the tail in females. Despite best effortsit was not always possible to visually determine the lobsters sex, as they are usually foundunderneath rock crevices with their abdomen not visible to the surveyor. Lobsters were nothandled in order to minimize disturbance. If lobsters escaped before measurements could be

36

made, then their carapace length was estimated by sight. During this quarter lobster surveyswere conducted monthly on the 14th-15th of each month to gain better insight into how thepopulation develops on a more regular basis. Each survey was conducted with between threeand six surveyors snorkelling the site and recording data and were always led by at least onestaff member. A thorough briefing and sufficient training was provided to ensure the datarecorded was as accurate as possible.

All data analysis was conducted using Rstudio version 1.1.456 (R Core Team, 2018). Todetermine whether there is a significant difference between the number of spiny lobsters foundbetween each zone and each month an Anova test was used followed by a post hoc Tukey testto determine where the significance was.

For the statistical analysis of carapace length female and male Caribbean spiny lobsterwere tested separately due to the difference in carapace lengths between the sexes. The non-parametric Kruskal-Wallis test was used to test whether there was significance between thecarapace lengths in females between months and the carapace lengths in males between months.Additionally, a Kruskal-Wallis test was also used to determine whether there was a significantdifference between the length of carapaces in females in different zones, and the length ofcarapaces in males in different zones.

2.6.3 Results

A total of 144 individual lobsters were counted across all surveys with 85 being males and 5females, and 9 lobsters which we were unable to sex. Overall the CZ and PZ contained thehighest numbers of P. argus (CZ mean = 8, SD = 5.37; PZ mean = 8.9, SD = 4.79) with theGUZ being the only zone to have counts of 4 and lower throughout the year (mean = 1.1, SD= 1.35). The highest number of lobsters recorded during a single survey was in February inthe PZ with 19 individuals.

The sex ratio of Caribbean spiny lobster each month for the CZ and the PZ is comparedin figure 9, in general more male lobsters were found than female however no obvious trendsappear between each month or between the CZ and PZ. To determine any seasonal trend insex ratio we will need to continue monitoring for at least a full year. The mean and standarddeviation of carapace length in male and female lobsters in each management zone is comparedin table 6.

Table 9: The Mean and Standard deviation (SD) for carapace length of all lobsters and for maleand female Caribbean spiny lobsters shown separately for each management zone.

CZ PZ GUZMean SD Mean SD Mean SD

All Lobsters 82.01 88.23 81.84 40.27 75.36 35.86Females 71.90 30.76 80.26 46.07 65 49.50Males 92.11 107.44 83.43 36.11 85.71 34.57

There was a significant difference between the number of spiny lobsters counted in each zone(F = 7.99, df = 2, P¡0.01). Further analysis revealed that the significant difference was foundbetween the GUZ and both the CZ (t = -3.25, P¡0.05) and PZ (t = 3.66, P¡0.01), howeverthere was no significant difference between the CZ and the PZ (t = 0.41, P¿0.05). Analysis ofthe number of Caribbean spiny lobsters counted each month revealed no significant differencebetween each month (F = 0.52, df = 7, P¿0.05).

There was no significant difference between the length of the carapace on females in differentmonths (X2 = 13.30, df = 7, P¿0.05; figure 10). There was no significant difference betweenthe length of carapaces on males between months (X2 = 13.19, df = 7, P¿0.05; figure 11).

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Figure 13: Sex ratios of Caribbean spiny lobsters throughout the study period in A) CZ and B)PZ (black = female, grey = male).

Additionally, there was no significant difference between the length of carapaces in females indifferent zones (X2 = 0.51, df = 2, P¿0.5). There was also no significant difference between thelength of carapaces in males in different zones (X2 = 0.69, df = 2, P¿0.5).

2.6.4 Discussion

The results showed a decrease of P. argus counts compared to previous surveys that wereconducted by Frontier in quarters 174 and 181. Overall the trend of lobster population decreasedover the last few months which was a surprising result, considering all surveys this year havebeen conducted during the closed season for lobster fishing. This may be because the spawningareas are outside our survey sites and they may migrate to different zones during this time.More study will need to be done to determine if this is a possible hypothesis and perhapsexpanding our sites to include greater variety of habitats to include more habitat complexityinto the surveys. Mangroves are used as a nursery for juvenile lobsters and this may resultin larger numbers being counted if we include them in our studies. Carapace length has alsodecreased overall except in the PZ, which has had the only increase in size and the GUZ whichhad a fairly steady average size recorded, although the least number were sighted in the GUZ.The ratio of males to females was almost 2:1 with nearly double the number of males sightedacross all sites. Females with a tar patch or eggs are illegal to catch, which makes this resultcurious. As expected, the least number was found in the GUZ which could indicate that thezonations are working with the CZ and PZ showing higher numbers which may be used asshelter during the open season. However, the overall decrease in size and number indicates aworrying trend which could have major impacts on local fisherman and marine environment.

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Figure 14: The carapace lengths (mm) of female Caribbean spiny lobsters in different monthsduring 2018.

Figure 15: The carapace lengths (mm) of male Caribbean spiny lobsters in different monthsduring 2018.

Due to their economic importance a decrease in stocks could lead to more illegal fishing and ahigher strain on other local resources. Taking more numbers out of the local area will also havean impact on marine life that relies on lobsters for prey. To collect more accurate and actionabledata, extra sites will have to be included for the future. Length frequency distribution graphscan also be created when more data is collected (Gongora, 2010).

The Fisheries Department of CC is currently understaffed compared to the size of the reefthey have to patrol, this leads to significant amounts of illegal lobster fishing of undersizedlobsters and even lobsters being removed from the conservation zone, as noted by the FisheriesDepartment. Community established MPAs have previously been found to be more successfulthan government MPAs, where local community fishermen (resource users/stakeholders) have

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greater man power (Buglassa et al., 2018). Creating No Take Zones (NTZs) in a marine reserveis often perceived negatively by the local community; therefore, it is vital that positive resultsare shown from it in the long term (Buglassa et al., 2018). The catch per unit effort hasdeclined by 28.15%, stock size by 25% and recruitment levels by 36.1% during the period of1999 to 2009 (Gongora, 2010). In recent years the region has reported declines in lobster catchescausing concern for the many livelihoods that could be affected (FAO, 2007). At the currentrate of lobster fishing, the Caribbean spiny lobster will likely become endangered within thisregion. A stronger level of management is required within the local fishing co-operative towork in tandem with the fisheries department in order to regenerate the increase of lobsterpopulations in the long term and to show positive results that benefit the local communityrather than just seeming to restrict fishing practices (McDonald et al., 2017). Currently moredata is needed to make an effective plan for mitigating the lobster fishery damage on the lobsterpopulation. If possible, the off season may need to be extended and/or increased policing ofillegal lobster fishing is needed as in a similar study on the Galapagos Islands (Buglassa etal., 2018). Additionally, recommendations currently include increasing the minimum lobstertail weight to 4.5 ounces instead of 4 ounces, a deep-water lobster stock assessment should becarried out and the traditional licensing system, which allows all fishermen to fish for lobsters,needs to be revised (Gongora, 2010).

One final factor that may have a large impact on these results is the ability of the recordersthemselves. The surveys are supposed to mimic that of a fisherman, hence are done throughsnorkelling. However, most of the volunteers who assist in these surveys have never seen alobster in the wild before and do not have the expertise to be able to spot them in the sameway a fisherman who has been doing this their whole lives can. Moreover, the fishermen areusually extremely talented free divers with the ability to stay underwater for a lot longer thanthe research assistants. A redesign in this survey needs to be contemplated to collect morereliable data. For example, working directly with the fishermen to see their catch and possiblysurvey their lobster shades may be more effective. More possible surveys and techniques thatshould be considered suggested by Gongora (2010) are surveys of deep-water lobster populationsand lobster traps and shade surveys. Night surveys may also need to be implemented whenlobsters are more active, but this would require additional resources like flashlights and a lackof experience in the surveyors may impact results (Buglassa et al., 2018).

The research conducted here gained valuable information on lobster stocks in Caye Caulker.A continued study is necessary to understand the ecological process and anthropological pres-sures effecting lobster populations. The information gained here and in future study can beused to inform local policy makers for the preservation of lobster communities and sustainablefishing practice.

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2.7 Seagrass and Manatee Monitoring

2.7.1 Introduction

Despite the importance of seagrass ecosystems compared to coral reefs and mangroves, thereis a lack of data to enable a better understanding of their importance. This has led to fewinter regional comparative studies due to the unfashionable nature of seagrass conservation(Duarte et al., 2008). Seagrass are primary producers and marine angiosperms that account for0.1-0.2% coverage of ocean habitat worldwide (Duarte, 2002). Acting as ecosystem engineers,seagrass profoundly influence the physical, chemical and biological environments within coastalwaters (Fourqurean et al., 2012) by altering water flow, enhancing nutrient cycling and foodweb dynamics, and stabilising sediments (Orth et al., 2006). Seagrass is also a vital food sourcefor mega-herbivores in the Caribbean, including the green, loggerhead and hawksbill sea turtles(Chelonia mydas, Caretta caretta and Eretmochelys imbricata respectively) and West IndianManatees (Trichechus manatus) all of which are of international conservation concern (O’Sheaet al., 2001; Hughes et al., 2009). Seagrass provides a critical habitat for juveniles of bothcommercially and recreationally important fishery species that depend upon the habitat fornursery and refuge areas (Heck et al., 2003).

Seagrass EcologySeagrass can be found across most continents where their predominant habitat is shallow,

coastal waters; however, there has been a noticeable loss of seagrass coverage worldwide formany decades equating to a scale of hundreds of square kilometres (Heck et al. 2003). This lossis mostly due to rapid environmental changes because of increased coastal development and agrowing global population (Lotze et al., 2006; Wilkinson & Salvat, 2012). The most widelyaccepted disturbances include sediment and nutrient runoff, physical damage, invasive species,disease, commercial fishing practices, aquaculture, overgrazing, algal blooms, and increasedsea surface temperatures (Orth et al., 2006). Such disturbances have resulted in the recentdecline of seagrass meadows in Belize, which consequently has increased the necessity for theprotection, as well as monitoring and management of seagrass meadows.

Coastal ecosystems can sequester huge amounts of carbon. These ecosystems include man-groves, coral reefs, salt marshes and seagrass, with seagrass having disproportionately largecarbon storage potential relative to their global area (Laffoley & Grimsditch, 2009). Theseecosystems cover ¡0.5% of the ocean floor and form part of the earths blue carbon sink, ac-counting for 10-16% of annual carbon storage in the ocean (Nellemann et al., 2009). Therehas been a recent surge in interest of carbon sequestration due to increasing global pressureto mitigate the effects of climate change brought on by carbon emissions. The recent focuson carbon trading and carbon pricing has resulted in considerable interest in quantifying thecapacity of the worlds ecosystems to trap and store carbon (Lavery et al., 2013).

Blue carbon is a new initiative that has a new strategic approach to make use of thelarge carbon capture and storage potential of coastal ecosystems. Mangroves, seagrass andsalt marshes are found on every continent in the world except for Antarctica and account forapproximately 49 mha. If this carbon could be quantified and sold on international carbontrading markets then this could help fund preservation and restoration projects, which wouldalso help capture more carbon and ease the effects of climate change (Crooks et al., 2010). Thereare three species of seagrass found in Caribbean waters; turtle grass (Thalassia testudinum),manatee grass (Syringodium filiforme) and shoal grass (Halodule wrightii), with turtle grassbeing the most abundant in the waters around CC.

Above the ground it may appear that seagrass are individual plants, but these plants haverhizomes that are the horizontally growing stems below the ground which connects each plantto each other, and where the roots sprout. The species Halophila spinulosa has fleshy rhizomes,suggesting high abundance of starch and carbohydrates (Anderson, 1994). The leaf Halodule

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univervis is fragile and often lost to drift (Anderson, 1998), and so the rhizomes have a highconcentration of digestible starch (Masini, 1982) and non-cellulose glucose (Masini et al., 2001).Thayer et al. stated in 1984 that fibre makes up a high percentage of the plant (Burn, 1985);this indicates a potential reason as to why seagrass and specifically the rhizomes are favouredby grazers.

The weekly seagrass surveys in CC allow us to monitor the health of the seagrass beds duringthe continued coastal development of CC (CZMAI, 2016) and their influence on species thatdepend on seagrass as forage. By comparing seagrass coverage and health between north CC(un-developed) and south CC (developed/developing), we aim to provide data on the adverseeffects of coastal development on seagrass communities. It is hypothesised that there will be adecrease in seagrass coverage in seagrass beds closest to coastal development. To understandhow seagrass health impacts other species, we can also understand anthropogenic effects onthese species. This study primarily focuses on the Antillean manatee (Trichechus manatusmanatus) and how its uses seagrass meadows as it is said to graze mainly on seagrass meadows.Several studies have already contributed to the known feeding ecology of this manatee such asAlvarez (2010); Bengtson (1981); Hartman (1979); Ledder (1986); Lefebvre et al., (2000); and(Reynolds, 1981). This study aims to document the Antillean manatee population and providehabitat evaluations of the seagrass meadows. Due to the high manatee population, studies inthis area can be used to aid the on-going conservation efforts in Belize (Alves-Stanley et al.,2010).

Manatee EcologyThe Antillean manatee (Trichechus manatus manatus) belongs to the order Sirenia which is

the only aquatic herbivorous order of mammals currently existing (Bertram & Bertram, 1973;Burn, 1985). These mammals form multispecies communities and partition seagrass resourcesfor feeding (Marsh et al., 2011). A combination of global cooling after the middle Miocene aswell as human predation in the North Pacific has reduced their diversity to only two livinggenera and four species (Marsh et al., 2011).

These two genera or families of the Sirenia are the Dugongidae (dugongs) and Trichechi-dae (manatee) (Aketa & Kawamura, 2001). These can further be divided into four speciesthrough skull examination: the dugong (Dugong dugong), the Amazonian manatee (Trichechusinunguis), the West African manatee (Trichechus senegulsensis) and the West Indian manatee(Trichechus manatus) (Hatt, 1934). The West Indian manatee is finally subdivided into an-other two subspecies, the Florida manatee (Trichechus manatus latirostris) and the Antilleanmanatee (Trichechus manatus manatus) (Marsh et al., 1986), both of which reveal a diversityof aquatic herbivore feeding patterns (Aketa & Kawamura, 2001). The latter subspecies is theone examined in this project.

The West Indian manatees can be found in both freshwater and saltwater habitats (Camp-bell & Irvine, 1977). The Antillean manatees habitat is along the coast of Mexico, CentralAmerica as well as South America, but can also be found in the Greater Antilles such as Cuba,Jamaica, Hispaniola and Puerto Rico (Alvarez-Aleman et al., 2007). Belize is believed to be oneof the biggest strongholds of the Antillean manatee (Morales-Vela et al. 2000). They are foundin shallow coastal areas, slow moving rivers, estuaries, saltwater bays, and canals, with preg-nant females seeking refuge in even shallower waters (Bengston, 1981). The manatees exhibitelusive and secretive behaviour, making them one of the hardest animals to study and observedirectly (Bertram & Bertram, 1973). Importantly, they can stay submerged under water forrelatively long periods of time, with a dive time between 16.3 to 24 minutes (Best, 1981).

The shallow waters in their habitat are essential for feeding, particularly as fewer shal-low waters do not require such deep dives and therefore they can save more energy and timespent foraging and grazing (Anderson, 1994). The seagrass communities in these shallow wa-ters are found to be the most ideal depth for the manatees dive time (Domning, 1982). The

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main species that make up the diet of the manatees are marine angiosperms (seagrass) (Divi-sion angiospermae) (Green & Short, 2003), leaves of mangrove (Rizophora mangle) and algae(Bengston, 1981). Previous studies have found that the more favoured seagrass is the Haloduleand Halophila (Marsh et al., 2012). The manatees have one of the slowest consumption andpassage rate of any mammalian herbivore (Lomolino & Ewel, 1984), allowing for a cellulosedigestion coefficient of 80% (Burn, 1985). Using their pectorals, the T.manatus excavates rhi-zomes from the ground, which it then feeds on (Packard, 1984; Anderson, 1994). Both thedugong and manatee share a similar cellular digestion (Murrey et al., 1977), which ultimatelyallows for comparison between these two subspecies.

Several studies indicate that manatees are opportunists at seagrass foraging, and specializein the readily available species (Anderson, 1994; Anderson and Birtles 1978; Boyle & Khan,1993; Domning, 1976). By foraging between six and eight hours per day (Bengston, 1981) theycan have a potential daily consumption of 42-65kg (Lomolino & Ewel, 1984). Most West Indianmanatee habitats have access to both salt water and freshwater and so they may choose to eatseagrass, despite this not being their preferred food. This is because it will be the most readilyavailable food source, as seagrass meadows are found in the majority of the manatees habitats(Domning, 1982). Importantly, the manatee has continuous dental replacement; new molarsgrow regularly at the back of the jaw, forcing old, worn teeth out, similar to elephant teeth,which means they may choose less fibrous seagrass, as low fibre content requires less chewing(Domning & Hayek, 1984). Thus, the seagrass species preferred are Thallassia sp., Halodulesp., Halophila sp. and Syringodium sp. (Deutsch et al., 2008).

While there are 60 species of seagrass (Green & Short, 2003), manatees feed on a limitedrange of species. Hartman (1979) suggested that Thalassia testudinum, Syringgodium filiforme,Halodule wrightii and Halophila engelmanni are the main dietary preferences of the West IndianManatee. T. testudium is the most dominant species in the Caribbean due to their regenerationspeed (Dawes & Lawrence, 1979). However recent studies conducted in Cuba show that the onlyspecies preferred by the T.manatus was the H.wrightii, T. testudinum, S. filiforme, rhizomesand mangrove although the Caulerpa paspaloides and Halophila enegelmanni were occasion-ally consumed (Alvarez-German, 2010). The seagrass species found in Belize, and specificallyaround Caye Caulker, are H.wrightii, T. testudinum, S. filiforme, with T. testudinum being themost abundant, offering manatees their preferred choice of forage (Mudan et al., 2017). Theseseagrass species are likely favoured by manatees due to their rhizomes which play an importantrole in manatee feeding ecology. Each species of seagrass provides different nutrients that willbenefit manatees in different ways.

It is well documented that manatees will migrate to new feeding areas between seasons inBelize, however, it is less known why they choose to migrate. Anderson and Birtles (1978)discovered dugongs moved on to new feeding grounds five or six days after grazing 30% ofan area. Anderson (1994) also investigated why dugongs were diving deeper for H.spinulosa,and suggested the nutrient content compensated for the high cost of dive. Anderson (1998)further discovered the seagrass species Halodule uninervis rhizomes were rich in carbohydratesand therefore were selected for by dugongs. This corroborates with the study of Masini et al,(2001), who further discovered that dugongs excluded the dominant seagrass and chose earlygrowth Halodule, which were low in fibre and high in nitrogen (Ledder 1986). This characterizedthe feeding habits of the West Indian manatee Trichechus manatus latirostris ; Halodule wrightiiwas the dominant species in the diet in the summer and winter but changed seasonally in springto Syringodium filiforme. The above research may indicate why the manatees choose to migrateto new feeding areas and also indicates that when studying manatee feeding areas, the sedimenttype may also play a key role in the site choice. For example, if the rhizomes are unobtainabledue to a harder sediment type it could suggest the manatees are unlikely to choose this area,allowing for the expectation that the manatees will choose areas with a soft sediment type,

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with mainly H.wrightii or S.filiforme present.Examining the manatee feeding patterns allows an understanding of the composition and

preference as well as the role within their ecosystems. Seagrass meadows are highly overlookedbut extremely important habitats for providing shelter for fish, shellfish and nursery areas forother animals. The manatees provide important nutrient cycling within seagrass meadows andcommunities because their grazing may affect the plant traits, such as chemical and nutrientcomposition. Here, the nutrient content and palatability of the grass determines the plantsresponse to grazing (Adler et al., 2005). While nitrogen (N) is usually replenished into theterrestrial ecosystems through defecation, this is not possible in marine ecosystems due to themovement of water preventing any nutrient to settle on the ground and diffuse back into theecosystem. The manatee and dugong play a vital role regarding nitrogen recycling (Aragoneset al., 1997); through the dugong-grazing patterns the bacterial nitrogen fixation was found tobe disturbed due to their serpentine feeding behaviour (Anderson & Britles, 1978). Serpen-tine feeding behaviour refers to the pattern of the trail taken by Sirenia whilst feeding; theirsnout leaves a serpentine shaped course. Serpentine feeding plays a role in cultivation grazingin which the detritus aerates the sediment providing substrate for the bacterial nitrogen fix-ation (Aragones et al., 1997). Here, Adler et al. (2005), argues that the plant traits are theindependent variable in grazing. However, further study would be needed to find the causefor variation in plant traits. Overall the manatees are important in the nitrogen cycle withinseagrass meadows.

All the field experiments mentioned previously indicate that the Sirenia feeding habits havethe ability to change the seagrass communities, through composition alteration and nutrientmodification. For example, herbivores have been found to increase primary production bylimiting nutrient recycling resulting in grazing optimization, occurring when the lost nutri-ent proportion is smaller than the lost nutrient proportion throughout the overall ecosystem(Mazancourt et al., 1998). Preen (1995) suggested grazing may alter the species compositionwithin a community and act as cultivation for species, thereby preventing certain species grow-ing. For example, when Zostera capricorni is grazed, it allows the faster growing Halophila totake its place. Aragones et al, (1977) further found that nutrient levels in regrowth were affectedby grazing. Manatees have also been linked to important aquatic weed control (Campbell &Powel, 1976). Aragones et al, (1997) found that dugong grazing on seagrass communities inAustralia increased the speed of seagrass species regrowth, such as Halophila ovalis and Halod-ule inivervis, by 25-35% in one year. The nutrient concentration such as nitrogen (N) increasedby 30% and reduced the fibre concentration too. These findings are supported by Aragones etal., 2006 and Hik et al., (1991).

Manatee ThreatsHumans present the main threat to manatees (Bertram & Bertram, 1973). Anthropogenic

influences contributed to the manatee being listed as protected under the Wildlife ProtectionAct 1981 (Morales-Vela et al., 2000). The International Union for Conservation of Nature(IUCN) has now assessed and concluded the Antillean manatee to be endangered due to lowpopulation counts (Deutsch et al., 2008). The mature population size is less than 2500 with adecline of 20% expected to occur over the next two generations due to minimal conservationefforts and the increased threats to their environment. The main threats to their manateepopulations include hunting, habitat degradation, and entanglement in fishing gear, pollution,natural disasters, incidental catch and watercraft collisions.

Examining the history of the Stellers sea cow (Hydrodamalis gigas) gives the perfect exampleof how important conservation of the manatee is. By protecting the individual species, theirhabitat will additionally be protected as a result of the umbrella effect (Roberge & Angelstam,2004). Stellers sea cow was the first aquatic mammal to go extinct due to anthropogenic threats.Their size was described to be larger than the Blue whale. They were classified as extinct in

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1756, supposedly only resisting a mere 27 years of human pressure (Marsh et al., 2012). Theshort existence of such mammals shows how much of a threat humans pose and should be takenas evidence that intervention is necessary for the survival of several marine mammals.

In 1976, the Florida manatee population was estimated at 600-1200 mature individuals(Campbell & Powell, 1976); with decline expected (Marmontel et al., 1997) if no conservationefforts were implemented. The US took action through protection acts and today it is expectedthat the Florida manatee (Erickson, 2016) will be down listed from endangered to threatenedon the IUCN red list as the population size is estimated to be more than 6000. There arealmost no natural predators for manatees due to their ecology and size (Bertram & Bertram,1973).

Knowledge gap and rationale: future conservation and importance of studyTagging a manatee allows for the observation of their movement patterns, but additional

examination is necessary to determine reasoning for specific movements. From the literaturepreviously cited it is evident that there are three factors affecting the movement of the Antilleanmanatee; the temperature of the water they live in, their source of food, and lastly their sourceof fresh drinking water. The water temperature varies slightly in Belize separating manateesightings in a high season and low season, with high season being May until September andlow season being October to April (Morales-Vela et al., 2000).

The understanding of the feeding ecology of the manatee is very important for their futureconservation. If manatees lack the proper nutrients it can result in health issues, similar to thehealth issues the sea cows experienced. This was due to seagrass communities declining whichforced the sea cow to change their feeding habitat and resulted in the loss of their molars andteeth (Anderson, 1994). Not only are the seagrass meadows important habitats for manatees,the manatees are important grazers of seagrass (Courtene-Jones et al., 2014). Therefore, ifmanatee numbers reduce there will be a negative impact on seagrass meadows (Aragones etal., 1997). Also, if the manatees feeding habitat is lost due to over development this may havea direct impact on tourism.

Furthermore, in Vela et als study (2000) they recommend that more manatee surveys needto be completed around the Cayes off Belize City. The South Island of Caye Caulker is highlydeveloped, thus a comparison of the seagrass meadows found around the South Island and theNorth Island will indicate the effects that development has had on the meadows, the resultsof the study may be used effectively by the Fisheries Department when managing the forestreserves and possible future development on the north island. The main forms of conserva-tion that will for the Antillean manatee include policy-based actions; educational outreachprogrammes; protected areas; species-based actions (reintroductions, stranding networks); andresearch actions (site-specific and country-wide surveys) (Deustch, Self-Sullivan & Mignucci,(2008). This project will directly contribute to the site-specific research actions by studyingthe following aims:

1. Observe the distribution of the Trichechus manatus (Antillean manatee) around CayeCaulker, and their habitat

2. Test for a correlation between the manatees distribution and habitat choice

3. Create a photo ID data base of the manatee population around Caye Caulker

2.7.2 Methodology

The methodology for this project has not changed greatly from previous phases. Boat surveyswere conducted by using sites that are frequented by tourist boat tours. Using local knowledgefrom fishermen and tour guides, the transects completed between the phases 172 and 181

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allowed for identification of 3-4 hotspots in which manatees are spotted throughout the year,these sites were also used for manatee feeding habitat evaluations.

To meet aim two, habitat evaluations were conducted from phase 172. Using the datacollected from the boat survey transects 2-3 high use sites were compared to 2-3 low usesites (table 7). The sites are deemed as high use sites if manatees are frequently seen there.The seagrass composition of each area has been determined using a random quadrat samplingtechnique; additionally, the Index Foliage Area (IFA) of each site has been calculated. Paststudies by Clergy (2003) cited by Alvarez (2010) use the IFA in order to give the seagrassspecies of T.testudinum and H.wrightii a comparable quality data set. The IFA allows us tocompare the seagrass quality in different sites to understand the manatees habitat choice. Thesetechniques have been used in previous studies successfully when studying manatees around SanPedro (Courtene-Jones et al., 2014).

Table 10: Description of survey sites on seagrass monitoring around Caye Caulker, Belize

Zone 1 2 3 4 5 6 7 8No. Sites 3 2 3 4 3 2 1 2Description Behind Split In front of Split In front of Base North Channel East of North Tip South of South Channel South Channel West of South TipLat. 17o44′32.8”N 17o45′03.0”N 17o47′42.8”N 17o47′29.1”N 17o43′59.2”N 17o43′21.8”N 17o43′21.8”N 17o43′25.1”NLong. 88o01′57.6”W 88o01′24.2”W 88o01′17.1”W 87o59′46.1”W 88o02′28.1”W 88o00′30.8”W 88o00′30.8”W 88o02′22.7”WBottom Type Deep Silt Sand Sand Sand Deep Silt Deep Silt Sand SiltDepth 2-3m 1m 1m 4m 3m 5m 5m 2mManatee Use High Low Low High High High High LowCurrent High High Low High Low High High High

Within each site, 5 zones were chosen to sample. Four zones have been completed suc-cessfully with the IFA of each of the seagrass species calculated using the IFA equation below.Further mapping and profiling of each of the sites cannot be completed until more zones aresurveyed. Each profile will include the information shown in table 11, alongside the followinginformation collected through using 50cm x 50cm quadrat, repeated 25 times:

Variables:% Cover of 3 main seagrass species H. wrightii, S.filiforme & T. testudinum)% Cover of coralNo. Coral% Cover of spongesNo. sponges% Cover of gorgonian% Cover of echinodermsNo. echinoderms (starfish & sea urchins) present% of algae presentSpecies of algae

Using a 25cm x 25cm quadrat the average density and average canopy height was collectedfor the three main seagrass species variables. Average density was calculated by counting thenumber of individual shoots in the quadrat. The IFA was also calculated for the three mainseagrass species using the following equation:

The IFA is calculated by collecting 30 samples of each seagrass, for each plant the variablerecorded will be the number of grass blades, height and width of each blade. The IFA calculationwill be attempted for Syringodium filiforme using the same equation and additionally the bladewill be flattened in order to obtain the correct dimensions. The highest IFA result indicates

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Figure 16: Equation for Index Foliage Area (IFA) calculations; D = average density; A =average number of leaves; N0HD = average area m2 (average area/10000).

the area with the highest density of plants with the highest average number of leaves per plant;therefore, the plants will have the highest surface area per m2. This means that areas withthe highest IFA are likely to have the highest nutritional value and most abundance of specificspecies. The underwater evaluation generated profiles of the sites, allowing comparison to eachother. The IFA will be calculated to give the areas a quantitative comparison factor of seagrassto understand why the manatees choose a certain area. The habitat evaluation was conductedduring the high season and also the low season allowing seasonal comparison additional to thedistribution comparison.

Once all the profiles were constructed the data was then tested for normality using theShapiro-Wilkes test. The normally distributed data was then further analysed using theOne-Way ANOVA, and the variables not normally distributed were analysed using the non-parametric Kruskal-Wallis test. These tests determined whether there were any significantdifferences in any of the variables between each of the sites. Any variable which showed a sig-nificant difference was then analysed using the Mann-Whitney U Test to determine the specificsites which were significantly different.

Additionally, to the seagrass surveys, a volunteer from the Tampa Bay Estuary Programoffered to map out the seagrass using satellite imagery, which may complement the surveyswhen final profiles are made.

Photo-identification of manateesLastly, to meet aim 3 a photo identification database will be made of the current population.

This technique has been extremely successful in past studies in Florida by Beck and Reid(1995), Langtimm et al. (2004) and O’Shea et al. (2001). By creating a database of themanatee population, it allows for resident manatees to be identified and migratory behaviourto be monitored. Distinctive scarring and unique features are recorded as manatee identificationmarkers. The size of each manatee was also estimated where possible.

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2.8 Results

The seagrass species that were shown to be most dominant in each of the sites matches resultsfound in previous phases: H. wrightii (Hw), T. testidium (Tt) and S. filoforme (Sf). At lowuse sites Tt has the highest average IFA of 0.575 ± 0.29, followed by Sf which had an averageIFA of 0.075 ± 0.08, and lastly Hw which had an average IFA of 0.0 ± 0.0. When comparingthe high use sites, similarly Tt has the highest IFA average of 2.199 ± 2.16 followed by Sf of0.088 ± 0.09, with the lowest being Hw of 0.0 ± 0.0.

Statistical AnalysisThere was a significant difference between IFA values of Sf in different sites (F=10.94, df=6,

P¡0.001). There was also significant difference between the canopy heights of Sf in different sites(F=7.545, df=9, P¡0.01). There was a significant difference between the density of Sf in differentsites (F = 5.636, df = 6, P¡0.05). There was a significant difference between the percentagecover of Sf in different sites (F=27.68, df=6, P¡0.001). There was no significant difference inany variables with the other seagrass species.

The sites were then grouped together to test for a significant difference between the highuse sites and the low use sites. There was no significant difference between the highly used sitesand the low used sites by manatee (F=1.141, p=0.3035, p¿0.05). Further analysis was run forthe abiotic variables collected. There was a significant difference between the ground types inthe high use sites compared to the low use sites (F= 21, P¡0.001).

A photo-ID data base has been collected since April 2017, and now includes up to 13manatees recorded, see appendix 2.

2.8.1 Discussion

The manatee database is the first to be created of the population around Caye Caulker and isbeing used to monitor the movement patterns of any manatees that were recorded in previousseasons. This will allow us to determine whether any of the documented manatees are returningeach season. Having only one year of data on seagrass IFAs did not provide enough data todraw strong conclusions between high and low use sites, and so a long-term data set is requiredfor a statistical difference to be present. The difference between manatee use of sites with siltand no silt indicate that around Caye Caulker the profile for their feeding habitat will havesilt; this will allow the manatees to easily access the roots. As mentioned above, manatee areopportunistic feeders and may not choose a feeding site based on the seagrass species but theareas where they will use less energy to access the roots of the seagrass. Therefore, continuousmonitoring of this behaviour and the differences in ground type are required. The assessmentof the seagrass health around the CCMR is of huge importance as this could provide supportfor additional protected areas to be put in place in areas in which manatees use as feedinggrounds, particularly where anthropogenic disturbance may be impacting seagrass health. Inorder for this to be possible in the future, seagrass monitoring must continue over the comingyears to produce a long-term data set.

At the high use sites at site 4, 5, 6 and 7, Tt and Sf were the dominant species. Fromall sites, these sites were the ones with the highest percentage cover of Sf. Furthermore, thesesites had the highest IFA values of Tt. This suggests that the two species of choice for themanatees are Tt and Sf. Furthermore, looking at the sightings and the manatee behaviour, itis evident that the feeding ecology might not be the main deciding factor in the distribution ofthe manatees around Caye Caulker. Observations of manatees suggests that they choose theNorth Channel and South Channel for breeding purposes as mating seems to be a very commonbehaviour seen here.

Moreover, the health of the seagrass can be directly linked to the health of the mangrovesalong the coastline. Therefore, it is suggested it is possible to analyse the health and distribu-

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tion of the mangroves to investigate a possible correlation between the seagrass health to themangroves.

Lastly, understanding the distribution and health of the mangroves around Caye Caulkermay assist in further development on the island. Mangroves are directly linked to fisheriesand lobster recruitment (Vaslet et al., 2012). This means that understanding the health anddistribution of the mangrove forest around Caye Caulker is vital for the fisheries and lobstersurveys. Lobster recruitment boxes may be placed along the Caye in areas of high and lowcurrent as well as close to and far from mangrove.

Future ResearchA study by Adler et al, (2005) suggests that abiotic variables such as pollution, turbidity

and chemical composition of the water can affect the ability of a seagrass ecosystem to tol-erate grazing or not. Further studies could consider these environmental variables to furtherunderstand how healthy the seagrass meadows in CC are.

Another prospective project is to conduct GIS mapping of the seagrass beds around CC.The seagrass habitat has never been mapped in this area and so a detailed survey to examinethe extent of seagrass beds would prove extremely beneficial. This mapping would provide keylogistical information for the management of this habitat and start the collection of baselinedata, which will enable monitoring of changes over time. If a suitable area is found, this couldbe used as an incentive to allow the designation of an extended area of protection. The proposedextension includes an area of reef which includes the critically endangered Acropora sp. and anarea which manatees are known to frequently inhabit. A suitable area of seabed, for example,within the general use area in the northern extent of the MPA could be proposed as an areaof algae farming for the local fishermen in return for the extension of the protected zone area,this would be an alternative livelihood for fishermen. The area allotted for algae farming willneed to be where the seabed is barren substrate and not near seagrass or coral, which the algaecould smother and kill through inhibiting photosynthesis. An extensive mapping survey wouldnot only allow vital mapping of crucial habitats but would also allow GIS spatial analysis topresent any potential sites for algae farming. Lastly seagrass mapping allows us to monitor thefeeding habitats of the Antillean manatee.

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Appendices

Fish Species List

Angelfish - PomacanthidaeFrench Angelfish Pomacanthus paruGrey Angelfish Pomacanthus arcuatusQueen Angelfish Holacanthus ciliarisRock Beauty Holacanthus tricolorButterflyfish - ChaetodontideaBanded Butterflyfish Chaetodon striatusFoureye Butterflyfish Chaetodon capistratusLongsnout Butterflyfish Prognathodes aculeatusReef Butterflyfish Chaetodon sedentariusSpotfin Butterflyfish Chaetodon ocellatusSurgeonfish - AcanthuridaeBlue Tang Acanthurus coeruleusDoctorfish Acanthurus chirurgusOcean Surgeonfish Acanthurus tractusJacks - CarangidaeBar Jack Caranx ruberPermit Trachinotus falcatusGrunts HaemulidaeBlack Margate Anisotremus surinamensisBluestriped grunt Haemulon sciurusCaesar Grunt Haemulon carbonariumCottonwick Haemulon melanurumFrench Grunt Haemulon flavolineatumPorkfish Anisotremus virginicusSailors Choice Haemulon parraSmallmouth Grunt Haemulon chrysargyreumSpanish Grunt Haemulon macrostomatumStriped Grunt Haemulon striatumTomtate Haemulon aurolineatumWhite Grunt Haemulon plumieriiWhite Margate Haemulon albumSnappers - LutjanidaeCubera Snapper Lutjanus cyanopterusDog Snapper Lutjanus jocuGray Snapper Lutjanus griseusLane Snapper Lutjanus synagrisMahogany Snapper Lutjanus mahogoniMutton Snapper Lutjanus analisRed Snapper Lutjanus campechanusSchoolmaster Lutjanus apodusYellowtail Snapper Ocyurus chrysurusGroupers - SerranidaeBlack Grouper Mycteroperca bonaciConey Epinephelus fulvusGoliath Grouper Epinephelus itajaraGraysby Cephalopholis cruentata

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Nassau Grouper Epinephelus striatusRed Grouper Epinephelus morioRed Hind Epinephelus guttatusRock Hind Epinephelus adscensionisTiger Grouper Mycteroperca tigrisYellowfin Grouper Mycteroperca venenosaYellowmouth Grouper Mycteroperca interstitialisParrotfish - ScaridaeBlue Parrotfish Scarus coeruleusGreenblotch Parrotfish Sparisoma atomariumMidnight Parrotfish Scarus coelestinusPrincess Parrotfish Scarus taeniopterusQueen Parrotfish Scarus vetulaRainbow Parrotfish Scarus guacamaiaRedband Parrotfish Sparisoma aurofrenatumRedtail Parrotfish Sparisome chrysopterumStoplight Parrotfish Sparisoma virideStriped Parrotfish Scarus iseriYellowtail Parrotfish Sparisoma rubripinneTrunkfish - OstraciidaeSpotted Trunkfish Lactophrys bicaudalisTriggerfish & Filefish - BalistidaeBlack Durgon Melicthys nigerOcean Triggerfish Canthidermis sufflamenOrangespotted Filefish Cantherhines pullusQueen Triggerfish Balistes vetulaScrawled Filefish Aluterus scriptusWhitespotted Filefish Cantherhines macrocerusMiscellaneousChub Kyphosus sectatrix/incisorGreat Barracuda Sphyraena barracudaHogfish Lachnolaimus maximusLionfish Pterois volitansSpanish Hogfish Bodianus rufusYellowtail Damselfish Microspathodon chrysurusJuveniles and Recruits Fish Species ListDamselfish (¡3.5cm)Bicolor Damselfish Stegastes partitusBlue Chromis Chromis cyaneaBrown Chromis Chromis multilineataCocoa Damselfish Stegastes variabilisDusky Damselfish Stegastes adustusLongfin Damselfish Stegastes diencauesThreespot Damselfish Stegastes planifronsSurgeonfish (¡5cm)Blue Tang Acanthurus coeruleusOcean Surgeonfish Acanthurus tractusButterflyfish (¡2.5cm)Banded Butterflyfish Chaetodon striatusFoureye Butterflyfish Chaetodon capistratus

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ParrotfishGreenblotch Parrotfish Sparisoma atomariumPrincess Parrotfish Scarus taeniopterusRedband Parrotfish Sparisoma aurofrenatumStoplight Parrotfish Sparisoma virideStriped Parrotfish Scarus iseriOthers (¡3.5cm)Bluehead Wrasse Thalassoma bifasciatumClown Wrasse Halichoeres maculipinnaFairy Basslet Gramma loretoRainbow Wrasse Halichoeres pictusSlippery Dick Halichoeres bivittatusSpanish Hogfish Bodianus rufusYellowhead Wrasses Halichoeres garnotiBenthos species listSponges SPNCoralsBranching Fire Coral Millepora cervicornis MALCBlade Fire Coral Millepora complanata MCOMGorgonian GGStaghorn Coral Acropora cervicornis ACERFused Staghorn Coral Acropora prolifera APROElkhorn Coral Acropora palmata APALClubtip Finger Coral Porites porites PPORPillar Coral Dendrogyra cylindrus DCYLBlushing Star Coral Stephanocoenia intersepta SINTLobed Star Coral Orbicella annularis OANNMountainous Star Coral Orbicella faveolata OFAVGreat Star Coral Montastraea cavernosa MCAVElliptical Star Coral Dichocoenia stokesi DSTOMassive Starlet Coral Siderastrea siderea SSIDLesser Starlet Coral Siderastrea radians SRADSymmetrical Brain Coral Pseudodiploria strigosa PSTR (was Diploria sp.)Knobby Brain Coral Pseudodiploria clivosa PCLI (was Diploria sp.)Grooved Brain Coral Diploria labyrinthiformis DLABMaze Coral Meandrina meandrites MMEARose Coral Mancina areolata MAREBoulder Brain Coral Colpophyllia natans CNATWhitestar Sheet Coral Agaricia lamarcki ALAMLettuce Coral Undaria agaricites UAGA (was Agaricia sp.)Low Relief Lettuce Coral Undaria humilis UHUM (was Agaricia sp.)Thin Relief Lettuce CoralUndaria tenufoila UTEN (was Agaricia sp.)Ridged Cactus Coral Mycetophyllia lamarckiana MLAMSinuous Cactus Coral Isophyllia sinuosa ISINSpiny Flower Coral Mussa angulosa MANGSmooth Flower Coral Eusmilia fastigiata EFASMarine algaeThalassia sp. THDictyota sp. DTLobophora sp. LOB

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Halimeda sp. HMBlue/Green Algae BGATurf Algae TURFMacro Algae MACCoralline Algae COROther SubstratesSand SNBare Rock BRAbbreviationsAtlantic and Gulf Rapid Reef Assessment AGRRABelize Barrier Reef BBRBelize Barrier Reef Reserve System BBRRSCaye Caulker CCCaye Caulker Forest Reserve CCFRCaye Caulker Marine Reserve CCMRConservation Zone CZConvention on International Trade in Endangered Species of Wild Fauna and Flora CITESForest and Marine Reserve Association of Caye Caulker FAMRACCGeneral Use Zone GUZMarine Protected Areas MPAMesoamerican barrier reef system MBRSNorth Back reef NBPreservation Zone PZSea Surface Temperature SSTSouth Back reef SBSynoptic Monitoring Program SMP

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Manatee Database

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