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3-29-2017 Distribution and Condition of Stony Corals in The Veracruz Reef System National Park: A Management Perspective Mauricio López Padierna Nova Southeastern University, [email protected]
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NSUWorks Citation Mauricio López Padierna. 2017. Distribution and Condition of Stony Corals in The Veracruz Reef System National Park: A Management Perspective. Master's thesis. Nova Southeastern University. Retrieved from NSUWorks, . (447) https://nsuworks.nova.edu/occ_stuetd/447.
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NOVA SOUTHEASTERN UNIVERSITY HALMOS COLLEGE OF NATURAL SCIENCE AND OCEANOGRAPHY
Distribution and Condition of Stony Corals in The Veracruz Reef System National Park: A Management Perspective
By Mauricio López Padierna
Submitted to the Faculty of Nova Southeastern University Halmos College of Natural Science and Oceanography in partial fulfillment of the requirements for the degree of Master of Science with specialty in:
Coastal Zone Management & Marine Biology
Nova Southeastern University
March 2017
I dedicate this work to my grandmas Abi and Abue who showed me what a life well lived looks like. Their loving kindness and persistence is an inspiration.
Acknowledgments
I am thankful to the myriad of people who were involved in this long project. First and foremost, my advisor Dr. Dave Gilliam for giving me the opportunity to work on a project I was passionate about in my own home country. His willingness to travel and keep an open mind, finding funding to continue the project for as long as it did. I also thank my committee members Dr. Richard Spieler and Dr. Brain Walker for helping get the project going and their continued work on several of our sampling trips. I also thank our partners at the Parque Nacional Sistema Arrecifal Veracruzano. Dr. Elvira Carvajal who invited us to survey the reefs in Veracruz, Dr. Tomas Camarena-Luhrs for his support during our visits, and of course the staff who accompanied us during surveys: Luis, Miguel, Marcos, Israel and Jacobo, and their supporting crew who helped us fuel the boats, get tanks and helped us navigate the reefs. This project could not have been done without the d many friends and lab mates who collected the data during our many trips: Melissa, Joanna, Vanessa, Danny, Allison, Paola, Stephanie, Ari, Kate, Amanda, Nicole, Chuck. And a special thanks to Liz for taking this project and making it even better with her inquisitiveness curiosity and support. I am also grateful for the great many people who crossed paths with me during my time at the Oceanographic Center and made my experience here a memorable one Zach, Matt, Nick (x3), Ashley, Andia, Jigu, Ana, Cody, Lystina, Katelyn, Naoko, Corinne to name a few, but there too many others to name, thanks to all. Last but not least I must thank my family for sticking with me while they wondered what I was doing thing whole time. Their love and support keeps me going every day.
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Table of Contents
Acknowledgments ...... ii Table of Contents ...... iii List of Figures ...... iv List of Tables ...... v Abstract ...... vi 1. INTRODUCTION ...... 1 1.1 Coral Reefs ...... 1 1.2 Marine Protected Areas ...... 2 1.3 Coral Reefs in Mexico ...... 4 1.4 The Veracruz Reef System ...... 5 1.5 Veracruz Reef System National Park ...... 10 1.7 Justification ...... 15 1.8 Objectives ...... 15 2. METHODS ...... 16 2.1 Field Methods ...... 16 2.2 Descriptive statistics ...... 17 2.3 Statistical analysis ...... 18 2.4 Temperature ...... 18 3. RESULTS ...... 20 3.1 Sites Surveyed ...... 20 3.2 Distribution ...... 24 3.2.1 Benthic Cover ...... 24 3.2.2 Species Richness ...... 28 3.3 Stony Coral Demographics ...... 30 3.4 Distance Analysis ...... 38 3.5 Temperature ...... 41 4. DISCUSSION ...... 43 4.1 Distribution ...... 43 4.2 Stony Coral Demographics ...... 47 4.3. Distance Analysis ...... 49 5. SUMMARY OF FINDINGS ...... 51 5.1 Distribution ...... 51 5.2 Stony Coral Demographics ...... 51 5.3 Management ...... 52 Literature Cited ...... 53
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List of Figures
Figure 1. Map of the reefs and reef areas in the Gulf of Mexico ...... 5 Figure 2. Satellite image of the reefs of the Veracruz Reef System ...... 7 Figure 3. Map of the Port of Veracruz from 1807 ...... 9 Figure 4. Map of the VRS showing the 1992 (yellow) and 2012 (red) park boundaries. 14 Figure 5. Location of survey sites in the Veracruz Reef System...... 23 Figure 6. Percent cover for each of the seven functional groups...... 25 Figure 7. Percent cover for each functional group at each reef ...... 27 Figure 8. Species richness for each of the reefs surveyed ...... 30 Figure 9. Relative abundance of coral species ...... 34 Figure 10. Stony Coral Condition in the VRS ...... 35 Figure 11. Stony Coral Condition by Species ...... 37 Figure 12. Linear regression analysis ...... 40 Figure 13. Mean daily temperatures from April 2008 to May 2009 ...... 42
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List of Tables
Table 1. Location, depth and status of temperature loggers ...... 19 Table 2. Summary of sites surveyed at each of the reefs in the Veracruz group ...... 21 Table 3. Summary of sites surveyed at each of the reefs in the Antón Lizardo group ..... 22 Table 4. Scleractinian and hydrozoan corals recorded in the Veracruz reef group ...... 29 Table 5. Scleractinian and hydrozoan corals recorded in the Antón Lizardo reef group. 30 Table 6. Stony coral species richness, abundance, Shannon’s H’ diversity and Pielou’s J’ evenness for each reef of the VRS...... 34
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Abstract
The Veracruz Reef System (VRS) is located in the southwestern Gulf of Mexico. It is comprised of 28 coral reefs in various stages of development and conservation. They are protected under the Parque Nacional Sistema Arrecifal Veracruzano National Park created in 1992. There are many threats to the reefs of the VRS, including the Port and city of Veracruz, which hosts half a million inhabitants and Mexico’s oldest active port. The inhabitants of Veracruz have used reef resources for thousands of years, as evidenced in archaeological sites on Sacrificios island, and constructions throughout the city, most notably in the San Juan de Ulúa Fort which was built entirely of coral skeletons. Despite the usage and protection given under the National Park, there is relatively little known about the health and condition of the stony corals in the System. There has only been one large scale study of 21 reefs conducted in the VRS in the late 1980’s. Since then, the National Park was created and 28 reefs are now recognized. This study performed point- intercept transects on 24 of these reefs including five reefs added to the official list in 2012. Point-intercept transects were surveyed at 63 sites between 2007 and 2014. Percent cover was calculated for seven functional groups. Additionally, demographic data of a subset of individual stony coral colonies were assessed on each transect. The functional group with the greatest cover in the VRS was crustose coralline algae (mean ± S.E.: 28.9% ± 1.97), stony corals had the second highest cover (21.5% ± 1.24). The Jamapa river divides the VRS into two groups the Veracruz group to the North and the Anton Lizardo group to the south of the river mouth. The Veracruz group had lower crustose coralline algae cover (28.1% ± 2.71) and coral cover (17.8% ± 1.55) than the Anton Lizardo group (29.6% ± 2.87 CCA and 25.3% ± 1.86 coral cover). The highest average coral cover on a reef was recorded at Ahogado Chico (45.5% ± 5.58), and the highest cover recorded on a single transect was 70% at Santiaguillo reef. The lowest coral cover was recorded at the fringing reefs on the north of the VRS, Punta Gorda and Punta Brava which had less than 1% coral cover. Coral colonies averaged 69.1 cm ± 3.10 in length at the VRS, 56.8 cm ± 2.98 in the Veracruz group and 81.7 cm ± 5.11 in the Antón Lizardo group. Old partial mortality was 25% ± 1.05 overall and similar between groups, recent partial mortality was 1.2% ± 0.21 and 1% at both groups. Disease prevalence was 3.9% for the VRS, 2.9% ± 0.88 in the Veracruz group and 4.9% ± 1.11 in the Antón Lizardo group. Overall, these reefs are faring slightly better than other reefs in the Caribbean having higher coral cover and larger colonies. However, the great variability in the health and condition of these reefs demands added attention and clear management goals to ensure their persistence in the face of ever growing threats. It is important to decrease the sources of stress, such as construction and poor waste water management in the area, better regulate fishing and approach a watershed wide management plan which takes into account upstream effects from the rivers that discharge into the Veracruz Reef System.
Keywords: coral reefs, stony coral, distribution, Veracruz, Mexico, Veracruz Reef System, Sistema Arrecifal Veracruzano, Gulf of Mexico
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1. INTRODUCTION
1.1 Coral Reefs
Coral reefs are one of the most diverse ecosystems in the world (Eakin et al. 2008). They sustain over 75% of the oceans’ species richness and provide a wide array of resources and services to humans such as food, coastal protection, genetic resources, and pharmaceuticals (Wilkinson 2008). Currently, the exploitation of these resources along with the effects of global climate change have put these ecosystems in peril and their importance cannot be overstated, many of the world’s fisheries, large sectors of the tourist industry and coastal infrastructure depend on healthy coral reefs, whether it be for their natural beauty, the fish and other food sources or the protected bays that reefs create (Asafu- Adjaye & Tapsuwan 2008).
For thousands of years humans have exploited natural resources, and coral reefs have been no exception. Wilkinson (2008) estimated that 500 million people partially or wholly depend on coral reefs for their daily food and resources. Coral reefs provide food, construction materials, medicinal products, sand for beaches, protection from storm surges and natural beauty that many people enjoy. As with so many other natural resources, overexploitation is reaching a point where the survival of these ecosystems is seriously threatened. Furthermore, local stressors are being compounded by the effects of global climate change caused by increased levels of carbon dioxide in the atmosphere. The increase in this greenhouse gas is causing a rise in atmosphere and ocean temperatures pushing corals closer to or above their 32ºC tolerance level. Carbon dioxide is also affecting
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the acidity of the ocean which can alter the environmental conditions that allow calcification, the process by which corals and other organisms deposit their calcium carbonate skeletons and protective structures (Bell, 2006). Other effects of climate change include changes in storm severity and frequency and rising sea levels, both of which can have detrimental impacts on coral reefs.
The biological diversity and value of these ecosystems cannot be understated. Greater biological diversity is contained within marine systems than their terrestrial counterparts. Nearly all of the 32 phyla of animals occur within coral reefs, while only about half are represented anywhere on land; no phyla is entirely terrestrial (Tunnell 2007). Coral reefs are particularly diverse harboring almost a quarter of all the marine diversity despite covering only 1% of the world’s oceans. Their distribution is limited by their strict environmental requirements: oligotrophic, warm waters between 26-28 °C, and salinities between 33-36 psu, with minimum turbidity and sedimentation (Salas-Perez & Granados- Barba 2008). This constrains coral reefs to the tropics, and often to the eastern coasts of continents compared to the western coasts which generally suffer from cold water upwelling that inhibit the development of coral reefs.
Coral reefs can cover tremendous areas, for example, the Great Barrier Reef in Australia extends 350,000 km2 (Chin et al. 2008) and can be seen from space. Which is amazing considering that coral reefs are built by colonial cnidarians that are less then 1 cm in diameter. As an example, the corallites of Montastraea cavernosa, which are some of the largest, have a diameter between 5.5–7.5mm (Veron 2000). Most coral species grow on the order of millimeters per year, meaning that to reach a colony diameter of 1 meter it may take 100 years. Therefore, it will take a coral community hundreds of years to create a coral reef; one in which hundreds of plant, animal and algae species reach a size and density where they can alter the environmental conditions surrounding them.
1.2 Marine Protected Areas
In an attempt to mitigate or reduce human impacts to these fragile ecosystems, governments and communities around the world have started protecting parts of the oceans.
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Marine Protected Areas (MPAs) are an administrative tool used to protect coastal and oceanic environments. The International Union for the Conservation of Nature (IUCN) defines these as “any area of intertidal or subtidal terrain, together with its overlying water and associated flora, fauna, historical and cultural features, which has been reserved by law or other effective means to protect part or all of the enclosed environment” (Kelleher & Phillips 1999). MPAs have many different objectives and are defined by protection priorities ranging from total exclusion areas, no-take zones, to a mere declaration that the area is being managed-, but with little enforcement or guidance on what activities can be carried out in the protected area.
The success of MPAs has been variable and depends on a myriad of factors including funding, enforcement, community involvement, institutional support, and the people’s general understanding and desire to protect marine resources. Recently, focus has been on creating MPAs which limit the removal of marine species, especially fishes because of their overexploitation and the subsequent impact on the health of the coral reef ecosystem. Thus, establishing fishing regulations can also protect or assist the recovery of coral reef benthic communities (Mumby & Harborne 2010).
Establishing successful MPAs remains a challenge because of lack of funding, coordination with stakeholders and enforcement of regulations. A study by Edgar, et. al (2008) found that the most effective MPA’s have five common traits: 1) the removal of organisms is prohibited (no take zones), 2) well enforced, 3) old (>10 years), 4) large (>100 km2), and 5) isolated by deep water or sand. Currently, there is a push towards protecting more of our environment. In 2010, the Convention on Biological Diversity set a target that “By 2020 at least 17% of terrestrial and inland water areas, and 10% of coastal and marine areas, especially areas of particular importance for biodiversity and ecosystem services, are conserved through effectively and equitably managed, ecologically representative and well connected systems of protected areas and other effective area-based conservation measures, and integrated into the wider landscapes and seascapes” (United Nations Environment Programme 2010). This target makes the protection of marine ecosystems a priority, and the use of MPAs as the preferred tool to achieve it. Meeting the requirements
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set forth by Edgar et. al (2014) and the target of protecting 10% of marine areas is a great challenge that governments around the world must meet to preserve some of the most threatened marine ecosystems (Wilkinson 2004).
1.3 Coral Reefs in Mexico
Due to its location, much of Mexico’s coasts have the necessary conditions for corals to thrive. Coral communities are found on both the Pacific and Atlantic coasts. However, strictly speaking, coral reefs are found exclusively along the eastern coast, in the Caribbean zoogeographic region. The most important reefs, in terms of size and diversity are those in the Gulf of Mexico off the coast of the state of Veracruz and off the Yucatan Peninsula in the state of Quintana Roo (Gutiérrez et al. 1993).
The general climate of the Gulf of Mexico is subtropical to tropical with mean annual air temperatures ranging between 26°C and 28°C and annual rainfall ranging from 1,100 - 2,000 mm. The dominant winds for this region blow from the northeast and east, however during the summer months periodic changes in weather patterns can shift the wind to the southeast. Additionally, polar air invasions known as “Nortes” lasting between 2-6 days are common between October and March, with average wind speeds of 12-45 km/h with gusts up to 110-120 km/h. The region suffers an average of nine hurricanes a year between August and October which provide most of the rain for the period (Ferre-D’Amare 1985).
The Gulf of Mexico is an area of substantial terrigenous sedimentation, with large amounts of freshwater discharges from rivers and coastal lagoons as well as urban runoff. These environmental conditions are not ideal for the development of coral reef communities; however, several reefs have been described the Gulf of Mexico’s shallow continental shelf (Salas-Perez & Granados-Barba 2008). Carricart-Ganivet and Horta-Puga (1993) describe three reef areas in the Gulf of Mexico: 1) North Veracruz, three reefs off of the Tamiahua lagoon (Blanquilla, Medio and Lobos) and three reefs located at the mouth of the Tuxpan river (Tangüijo, Enmedio and Tuxpan), 2) South Veracruz which is
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represented by the Veracruz Reef System (VRS) and its 28 reefs that are divided into two groups by the Jamapa River, and 3) The Campeche Bank zone which includes Alacranes reef, Cayo Arenas, Triangulo West, Triángulo East, Triangulo South and Cayo Arcas (Figure 1).
Figure 1. Map of the reefs and reef areas in the Gulf of Mexico. The black line shows the continental shelf. Modified from Sanvicente-Añoreve (2014).
1.4 The Veracruz Reef System
The VRS is bounded by the La Antigua River to the north and the Papaloapan River to the south and the Jamapa River divides the reefs in two groups (Figure 2). The runoff from these rivers cause the surrounding waters to be turbid especially during the rainy season (Ferre-D’Amare 1985). There are 28 reefs in two groups, the northern group consists of 13 smaller reefs close to the coastal city of Veracruz: five fringing reefs Punta Brava, Punta Gorda, Hornos, Bajo Paducah and Ingeniero; three reefs with islands Blanquilla, Isla Verde and Isla Sacrificios; and 5 platform reefs Gallega, Galleguilla, Anegada de Adentro, Pájaros, Mersey. The southern group is made up of 15 larger reefs
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offshore the town of Antón Lizardo (Tunnell 2007). There are two fringing reefs Giote and Punta Coyol; four islands Santiaguillo, Cabezo, Isla de Enmedio and Chopas; three sunken bank reefs La Palma, Sargazo and Periférico, and six platform reefs Anegada de Afuera, Topatillo, Polo, Blanca, Anegadilla, Rizo (Tunnell 2007).
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The reefs of the VRS developed on the continental shelf after the last glacial period between 9,000 to 10,000 years ago. The reefs have well developed frameworks extending to depths of about 30 m on the windward sides which face to the north and east and historically had higher coral cover than the leeward side. The west facing lee sides extend to about 21 m depth and have lower coral cover but a higher diversity as some more delicate species can only be found here (Horta-Puga & Carricart-Ganivet 1993). Stony coral richness, 36 species, on these reefs is similar to that of other remote Atlantic reefs like the Flower Garden Banks and Bermuda, which is lower than that found in the Mexican Caribbean (47 coral species) (Beltrán-Torres & Carricart-Ganivet 1999).
The VRS reefs and their resources have been known and exploited for centuries. Fish, echinoderms and coral remains have been found at pre-Hispanic burial sites. The reefs also created a natural harbor that allowed for the Spanish to land in Mexico and found the city of Veracruz, the first city in Mexico during the Spanish conquest in 1519. The reefs provided protection from bad weather and were also mined for construction material as can be seen in many of the buildings in downtown Veracruz and perhaps most famously at the San Juan de Ulúa Fort and the Santiago Bulwark (Carricart-Ganivet 1998). One of the first detailed maps of Veracruz was produced by Alexander von Humboldt in 1811 (Figure 3). The map shows the reefs in front of the city of Veracruz before the first major construction of the Port in the early 1900’s. The construction of the Port of Veracruz closed off the area behind Gallega reef, buried Lavandera reef and altered water flow at Hornos reef.
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Figure 3. Map of the Port of Veracruz from 1807 and published in (Humboldt 1811). Notice the presence of the reefs of La Caleta, Lavandera and Hornos (red circles) which were used to expand the port in the early 1900s.
Angelo Heilprin (1890) is credited with the first descriptions of the coral reefs of Veracruz. Over the next 60 years very little was published until K. O. Emery visited Veracruz during the Twentieth International Geological Congress in Mexico in 1956 (Emery 1963). He found that the fore reefs had 90% coral cover, the most abundant species were Acropora, Porites, and brain corals. By the 1970’s the reefs became a popular research subject for both Mexican biologists from the Universidad Nacional Autónoma de México and students from Texas A & M University, unfortunately little of this work was published
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in easily accessible journals, more recently the Universidad Veracruzana has started to dedicate lots of resources to studying the Veracruz Reef System.
1.5 Veracruz Reef System National Park
The VRS’s long history has brought several attempts to protect the reefs and their inhabitants at different times. The first conservation action came in 1975 when Blanquilla reef was declared a flora and fauna protection area. In 1982 Sacrificios island and the surrounding reef were closed to public access and fishing was prohibited, due to the presence of historical artifacts from prehispanic times. As the value of the natural resources was further recognized, protection was expanded to the remaining reefs with the creation of the Parque Marino Nacional Sistema Arrecifal Veracruzano (Veracruz Reefs System National Marine Park) in 1992. At the time, 23 reefs were listed and regulations were put into place that restricted activities to only those related to the preservation of aquatic ecosystems, research, recreation and environmental education, and approved uses of natural resources. Among the approved uses was fishing using: hand lines, gill nets, trolling, traps and diving, but capture and collection of coral, coralline algae and mollusks was prohibited. Additionally, the decree stipulated that a management plan must be published within 180 days of the decrees publication, and must include: 1) a catalogue of the flora and fauna found in the Park, 2) the requirements for permits for the extraction or use of natural resources, 3) the activities which are permitted in the Park, 4) restrictions on the construction, occupancy and functioning of marine facilities (Diario Oficial de la Federación 1992).
In 1994, a management plan had not been published and under pressure from the local fishers (Jiménez-Badillo 2008b), the sixth article of the decree was reformed to reopen the octopus, conch and clam fisheries. The list of permitted fishing gear was removed and instead stated that the seasons, gear, areas and limits should be specified in the Management Plan (DOF, 1994). As time passed Mexico’s environmental legislation was updated, an in 2000 the National Park designations were simplified, eliminating the distinction between marine and terrestrial parks unifying them under National Parks (Diario Oficial de la Federación, 2000). Meanwhile, efforts from the Park managers to
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protect the reefs continued registering the in two international treaties: The Ramsar Convention on important wetlands in 2004 (Ramsar Convention Secretariat 2013) and as a Biosphere Reserve in the Man and the Biosphere program in 2006 (UNESCO). Then in 2008 an agreement was published allowing the National Commission for Natural Protected Areas (CONANP) to carry out protection, restoration, conservation, education and sustainable non-extractive uses of resources on the 48,333 m2 of emerged land within the Park, at the Enmedio, Verde, Sacrificios and Salmedina islands (Diario Oficial de la Federación, 2008). The most recent and dramatic change was made to the Park with a new decree in 2012. This new decree increased the Park’s from 52,238-91-50 ha. to 65,516-47- 08.05 ha. and added two nucleus zones, Blanca and Santiaguillo named after the reefs they protect. Although the overall area of the Park was enlarged, an area known as Bahía de Vergara, to the north of the current port of Veracruz, was removed along with a portion of Punta Gorda reef. It was determined that this area no longer deserves protection because the ecosystem within it was too degraded and was not representative of the rest of the reefs in the Park, and resources could be better used elsewhere (Figure 4). Additionally, the list of reefs was updated ton include 5 reefs which were not mentioned in the previous decree: Punta Brava, Mersey, La Palma, Sargazo and Periférico, bringing the total number of reefs in the VRS to 28. Article 2 and 3 of the decree states that it is the Department of the Environment’s responsibility to administer and manage the resources in the Park by producing a Management Plan which includes input from governmental and non- governmental organizations and must be published within a year of the publication of the decree. The list of permitted activities was updated for each zone; activities allowed in the nucleus zones were: 1) preservation of marine and terrestrial ecosystems, 2) environmental monitoring, 3) scientific research, 4) ecosystem restoration and species reintroduction, 5) buoy and signal installation, 6) maintenance of existing infrastructure, 7) others permitted under the current environmental legislation. Prohibited activities in the nucleus zones: 1) touching or manipulating corals and other organisms, 2) dumping of waste of any kind, 3) cleaning vessels, emptying bilges or discharging ballast water, 4) stirring the bottom or cause sediment to become suspended, 5) interrupt or alter marine currents, 6) mining or prospecting for oil or minerals, 7) hunting on Santiaguillo island, 8) extractive and non- extractive use of resources, 9) fishing, 10) introduction of non-native species or genetically
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modified organisms, 11) construction of infrastructure, 12) tourism, 13) harass, capture, remove or take coral species or other wildlife, 14) use of explosives, 15) anchoring boats on coralline structures, 16) others under the current environmental legislation. Activities allowed in the buffer zone: 1) scientific research and collections, 2) environmental monitoring, 3) environmental education, 4) sustainable tourism, 5) fishing and aquaculture, 6) non-extractive use of wildlife, 7) ecosystem restoration and species reintroduction, 8) eradication of species that may become pests, 9) construction of support facilities, 10) signal buoy installation, 11) maintenance of existing facilities, 12) large and small vessel navigation, 13) others prohibited under the current environmental legislation. Article nine further defines the conditions under which the previous activities can be done. Highlights include: sport fishing is only allowed away from the reefs and only catch and release, the use of permanent traps or techniques that damage the bottom for fishing are prohibited, fishing will be done in a sustainable manner and is allowed as long as the proper permits are obtained, dredging shall be done with strict containment guidelines and the dredge materials shall be disposed of on land, and small craft refueling shall be done in designated areas. Prohibited activities in the buffer zone: 1) touching or manipulating corals and other organisms, 2) dumping of waste of any kind, 3) construction of spaces for solid waste on the islands, 4) hunting on the islands, 5) fishing from large vessels, 6) using chemicals as aids in fishing, 7) spearfishing using SCUBA, 8) use of permanent traps or gear that alters the bottom for fishing, 9) removal, transplantation, refilling, trimming or any activity that alters the original ecosystems, 10) any private activity which involves the construction of infrastructure, 11) mining or prospecting for oil or minerals, 12) use of any sound source that could alter wildlife behavior, 13) changing land use on the islands, 14) anchoring on coral structures, 15) lighting of fires on the islands, 16) repair or maintenance of vessels or motors, 17) cleaning of vessels, 18) emptying bilges or dumping ballast water, except in an emergency for large vessels, 19) stirring the bottom or causing sediment to become suspended, 20) use of explosives, 21) others under current environmental legislation. An area of influence was defined for the Park which includes the watershed beyond the Park’s boundaries that have an influence on the health of the VRS and suggests collaboration with the entities that are part of the watershed to protect the VRS (Diario Oficial de la Federación 2012).
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A side effect of the of the modification of the park boundaries is that a large port expansion project which has been in development for the better part of a decade (APIVER,
2005) can now proceed in the area that was excluded in Bahía de Vergara. In 2012 construction of the land portion of the project had already begun with the flattening of an area a for dry docks and the paving of access roads. By 2017 the construction of the breakwaters for the new port were well under way.
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nucleus zones zones nucleus arkboundaries. Thetwo e 1992e (yellow)and 2012 (red) P in green. in shown . Map of the VRS showing th showing VRS the of . Map 4 Figure Blancaand Santiaguilloare
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1.7 Justification
Although the VRS have been subject of sporadic research through the years, most studies have been on a smaller scale, focusing on a few reefs at a time. There has only been one previous large-scale study, where Lara (1992) surveyed 21 VRS reefs before they were declared a National Park. Therefore, some of the requirements set forth in the decrees have not yet been fulfilled. For example, in 2006, there wasn’t a complete list of flora and fauna in the Park, and a management plan had not been produced. In light of this a research group from the Nova Southeastern University Halmos College of Natural Sciences and Oceanography (NSU) were invited to a meeting with the Park managers and a partnership was created where NSU researchers would update the VRS coral and fish species catalog, and aid in characterizing all the reefs in the Park, to gain a better understanding of the health of the reefs.
1.8 Objectives
The overall objective of this study was to characterize the stony coral community of the VRS. This was done in two ways: 1) Assessing the benthic community of the reefs by estimating percent cover of 7 major functional groups: stony corals, substrate, turf algae, macro algae, crustose coralline algae, gorgonians and sponges and 2) Assessing the stony coral community by collecting demographic data on species diversity, relative species abundance, and stony coral colony size and condition. Benthic cover of the 7 functional groups and the size and condition of stony coral colonies were compared between the two reef groups in order to inform managers if these groups should be managed differently. In addition, distance from three potential sources of pollution (the Port of Veracruz, the Jamapa River and Antón Lizardo) was tested for effects on coral cover, colony size and disease prevalence. This will help determine if there is a gradient of stress or deterioration on the reefs in relation to these potential sources of pollution.
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2. METHODS
2.1 Field Methods
From April 2007 to July 2014, a total 84 surveys at 63 distinct sites carried out during annual research trips to the VRS. Surveys were completed each year during one to two weeks in the summer months. Because benthic habitat maps were not available, sites were chosen using the Park staff’s historical knowledge, depth range and location on the reef, targeting areas where corals were likely to be found on different parts of the reef, focusing on the outer windward and leeward zones. The crest and lagoons were not surveyed. As priorities shifted during the project so did the site selection. In 2010, sites were surveyed on the northern end of the Park at Punta Brava and Punta Gorda because discussions were starting about removing them from the Park. In 2011, there was also a shift towards including four submerged bank reefs (Mersey, La Palma, Periférico and Sargazo), so we surveyed sites on those reefs which were added to the 2012 decree along with Punta Brava.
Thirty meter transects were surveyed at two depth intervals 3–10 m and 15–20 m where habitat permitted. A total of 10 transects, 5 at each depth interval were targeted however, the actual layout of transects depended on the number of surveyors and the topography of the site. Transects were separated by at least 2 m, ran parallel to the contour of the reef and maintained a constant depth. Some sites were surveyed on more than one occasion if they were of particular interest or required further inspection.
16 Methods
The point-intercept method was used to determine the percent benthic cover of turf algae, crustose coralline algae, macro algae, substrate (sand, hard bottom and dead coral), sponge, gorgonian and stony coral species along each transect. Turf algae was defined as algae shorter than 2 cm in height and with no identifiable structure, macroalgae were recorded as fleshy algae taller than 2 cm. Substrate included sand and areas of hard bottom that were not colonized by living organisms. Gorgonians were recorded when any part of the organism was directly below the tape, encrusting or branching (i.e. canopy cover). The functional group directly under the tape measure was recorded every 25 cm for a total of 120 points per transect. Percent cover was calculated by dividing the total number of points for each functional group by 120 and multiplying by 100%.
Stony coral demographic data were collected on the first 10 colonies encountered along each transect. Only colonies with live tissue under the tape were included. Demographic data included species name, length, width and height, percent old partial mortality, percent recent partial mortality, presence of disease, and bleaching. Partial mortality is the percent of the colony which was dead, recent mortality was defined as areas where the coral skeleton was stark white with minimal overgrowth and the corallite structure was still clearly discernible. Old mortality was defined as any area of the colony with no coral tissue present and significant overgrowth. Disease prevalence was calculated by dividing the number of colonies with signs of disease by the total number of colonies surveyed.
2.2 Descriptive statistics
The point-intercept data were used to calculate percent cover for each functional group and stony coral species richness. Colony data were used to calculate mean colony length, width and height, mean percent recent and old mortality, and disease prevalence. Relative abundance of stony coral species was calculated by summing the number of colonies of each species and dividing by the total number of colonies recorded for each grouping level. Data were grouped at three levels: by the whole VRS, by reef group (Veracruz (Vz) and Antón Lizardo (AL)) and by reef, pooling transects from all depths.
17 Methods
Colony data were used to calculate the Shannon-Weiner diversity index and Pielou’s evenness. Only the first survey for each reef was used to reduce possible bias from the additional sampling effort.
2.3 Statistical analysis
Normality tests indicated that parametric tests were appropriate for the data. A t- test was used to test for differences in mean coral colony length and mean partial mortality between the two reef groups and an analysis of variance (ANOVA) was used to determine differences in colony length and mean partial mortality between reefs. Tukey’s Honestly Significant Difference (HSD) test was performed post hoc to identify which reefs drove the differences. Linear regressions were used to determine if there was a correlation between distance from three points of interest (Port of Veracruz, Jamapa River mouth and Antón Lizardo) and mean coral cover, mean colony maximum diameter and disease prevalence.
2.4 Temperature
Ten HOBO Tidbit® temperature loggers were deployed in April/May 2008. The loggers were placed on 7 reefs at depths ranging from 3 m to 19 m to obtain temperature profiles for the VRS over a one year period. Six were placed at reefs in Antón Lizardo and four in Veracruz reefs. Nine were recovered in May 2009 and data was obtained from seven, four from Antón Lizardo and three from Veracruz. Daily average temperatures were calculated by grouping the data by reef group and two depth intervals: shallow (3 -11 m) and deep (>10 m) (Table 1).
18 Methods
Table 1. Location, depth and status of temperature loggers. Loggers were deployed in 2008.
19 Discussion
3. RESULTS
3.1 Sites Surveyed
Between April 2007 and July 2014, 670 transects across 63 sites were surveyed on 26 reefs (Table 2 & Table 3 and Figure 5). The sites were distributed on 24 of the 28 reefs mentioned in the PNSAV’s 2012 decree (Diario Oficial de la Federación 2012), additionally two were surveyed sites on the lee of Anegada de Adentro in the Veracruz group (Vz) (Ahogado Chico and Ahogado Grande). These two reefs were analyzed separately because they did not appear to be physically connected to the Anegada de Adentro reef and their coral cover also appeared distinct (). Nine sites were surveyed on the five reefs that were added in the 2012 decree (1 on Punta Brava, 1 on Mersey, 2 on la Palma, 2 on Sargazo and 2 on Periférico). These surveys were, to the best of my knowledge, the first on these reefs. Four reefs were not surveyed, two submerged bank reefs Bajo Paducah and Giote and two fringing reefs Punta Coyol and Ingeniero. They are mentioned in the decrees but do not appear on maps, and we were unable to determine it even with Park guidance. Giote was not surveyed because the Park already had a monitoring station and suggested other sites were higher priority. In total, my study added data for seven reefs that had not been surveyed previously Punta Brava, Mersey, Ahogado Chico and Ahogado Grande in the Veracruz group and La Palma, Periférico and Sargazo in the Antón Lizardo group. Sampling effort was similar between the Veracruz and Antón Lizardo reef groups, as can be seen from the number of sites (31 and 33), times surveyed (15 days and 16 days), number of reefs (13 at each) and number of transects surveyed (337 at Vz and 333 at AL) (Table 2 & Table 3).
20 Discussion
Table 2. Summary of sites surveyed at each of the reefs in the Veracruz group. Some sites were visited more than once and therefore more transects were surveyed. Average depth was calculated from all transects at that site. Distance from each of the sites to 3 possible sources of pollution are listed.
21 Discussion
Table 3. Summary of sites surveyed at each of the reefs in the Antón Lizardo group. Some sites were visited more than once and therefore more transects were surveyed. Average depth was calculated from all transects at that site. Distance from each of the sites to 3 possible sources of pollution are listed.
22 Discussion
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3 y v B 4 e ! ( . Location of survey sites in the Veracruz Reef System. The red line marks the VRS National Park’s 2012 2012 Park’s National VRS the marks line red The System. Reef Veracruz the in sites survey of . Location 4 a v k ! ( r r r 4 5 u a B
S P a t 5 ! ( n 5 ! ( u µ P Figure boundary.
23 Discussion
3.2 Distribution
3.2.1 Benthic Cover
Overall, crustose coralline algae (CCA) had the greatest benthic cover, followed by stony corals, turf algae, substrate, macroalgae, sponges and gorgonians (Figure 6). Crustose coralline algae had the greatest cover in both reef groups (28% in Vz and 29.5% in AL). The three algal functional groups (coralline, turf and macro algae) contributed more than half of the benthic cover (55% Vz, 59% AL) (Figure 6). In Antón Lizardo, stony corals had the second greatest cover (25%) while in Veracruz they had the third greatest cover (18%). Veracruz had more sponges (7%) and gorgonians (4%) than Antón Lizardo (2.5% and 3% respectively). Antón Lizardo had over twice the macroalgae cover (12%) of Veracruz (5%). Bare substrate had higher cover in the Veracruz than in Antón Lizardo (Figure 6).
24 Discussion
Figure 6. Percent cover for each of the seven functional groups. The top panel shows data for the whole VRS, the middle panel for Veracruz and the bottom panel shows Antón Lizardo.
Coralline algae were recorded at every reef, and cover ranged from 2.5% (Punta Gorda) to 38% (Pájaros) in Vz. At AL, the range was from 17% (Topatillo) to 53% (Sargazo) (Figure 7). Stony coral cover ranged from <1% (Punta Brava and Punta Gorda) to 46% (Ahogado Chico). Three reefs in AL had at least 30% stony coral cover (Enmedio, Santiaguillo and Rizo). Turf algae was present at all reefs and had a percent cover ranging from 14% (Blanquilla) to 47% (Ahogado Grande). Reefs with over 30% turf algae cover were Ahogado Chico (34%), Sacrificios (34%), Palma (30%) and Periférico (30%). Punta Gorda and Blanquilla in Vz had the highest cover of bare substrate with over 20%. Meanwhile Afuera, Cabezo, Periférico and Rizo had over 14% in AL. Macroalgae cover was generally low at most reefs, and wasn’t recorded at Hornos. The two reefs with the greatest macroalgae cover were Punta Gorda (47%) and Topatillo (45%). Anegadilla
25 Discussion
(30%), Anegada de Afuera (19%) and Palma (18%) also had above average macroalgae cover. Sponges were recorded at all reefs but were relatively uncommon. They had less than 7% cover throughout Antón Lizardo, but had over 20% cover at Hornos and Mersey in the Vz group. Gorgonians were rare in our surveys and were absent at five reefs. Blanca reef was an exception with 21% cover. Sacrificios and Blanquilla in Vz also had above average gorgonian cover of 5% and 9% respectively (Figure 7).
26 Discussion
. Percent cover for each functional group at each reef. Veracruz reefs on the left and Antón Antón and left the on reefs Veracruz reef. each at group functional each for cover Percent . 7 Figure Lizardo on the right. Colors differentiate each functional group.
27 Discussion
3.2.2 Species Richness
Thirty-three stony coral taxa were identified during this project, 29 scleractinian corals and two hydrozoans (Millepora alcicornis and Stylaster roseus). Eighteen colonies were identified only to genus, four as Oculina and 14 as Mycetophyllia. Thirty-one coral taxa were recorded in Veracruz and 28 in Antón Lizardo. Two species which were not recorded in the Veracruz group Mussa angulosa and Mycetophyllia aliciae. Dichocoenia stokesii, Isophyllia sinuosa, Oculina and Stylaster roseus were absent in Antón Lizardo. The reef with the most species was Pájaros (25) while Punta Brava and Punta Gorda only had one species (Figure 8). Siderastrea siderea was the most widely distributed species, present on all reefs, and Montastraea cavernosa was present at all but two reefs. Colpophyllia natans, Orbicella faveolata and Porites astreoides were present at all reefs except at the three fringing reefs (Punta Brava, Punta Gorda and Hornos). Dichocoenia stokesii, Isophyllia sinuosa, Mycetophyllia aliciae, Oculina varicosa and Stylaster roseus were only recorded once, Mussa angulosa was recorded twice (Table 4 & Table 5).
28 Discussion drozoan corals recorded in the Veracruz reef group. Presence is indicated by the number 1. number bythe indicated is Presence group. reef inVeracruz the recorded corals drozoan . Scleractinian and hy and Scleractinian . 4 Table
29 Discussion
. Scleractinian and hydrozoan corals recorded in the Antón Lizardo reef group. Presence is indicated by the number 1. number the by indicated is Presence group. reef Lizardo Antón the in recorded corals hydrozoan and Scleractinian . 5 Table
30 Discussion
Figure 8. Species richness for each of the reefs surveyed. Veracruz group reefs are in blue and Antón Lizardo reefs in orange. The mean for each group is depicted by a gray line.
3.3 Stony Coral Demographics
The health of 6,225 colonies belonging to twenty-three stony coral taxa was assessed in the VRS. Four species represented 70% of the colonies surveyed C. natans, O. faveolata, M. cavernosa and S. siderea (Figure 9). Ten species had less than 1% relative abundance and could be considered as rare, A. palmata, Oculina spp., S. radians, A. fragilis, Mycetophyllia spp., M. alcicornis, A. humilis, P. porites and S. cubensis. However, observations outside of the transects proved that A. palmata is still quite abundant in the shallow lagoons and crest areas. Mean ± SD maximum colony diameter was 71.3 cm ± 72.8. Mean old partial mortality was 25.5% ± 26.02 and mean recent partial morality was 1.2% ± 5.4. Five species had over 30% old partial mortality M. decactis (44% ± 28), S. intersepta (39% ± 29), O. annularis (37% ± 26), S. siderea (33% ± 29) and O. faveolata (33% ± 24). Three species had no partial mortality M. alcicornis, A. humilis and S. cubensis. Three species had over 2% recent partial mortality Oculina spp. (8% ± 21), P. porites (5% ± 7) and O. annularis (2.5% ±8). The prevalence of disease was 4.2%. The species with the highest prevalence of disease was Siderastrea spp. which had a high
31 Discussion incidence of dark spot disease. A. cervicornis (5.4%), A. palmata (7.5%) and O. faveolata (6.3%) had above average prevalence of disease. Nine species did not have any signs of disease (Figure 11).
The Shannon-Weiner index for the Veracruz group reefs was 2.13 and Pielou’s J’ was 0.84, H’ for AL was 2.25 and J’ was 0.85 (Table 6). Colpophyllia natans and M. cavernosa were the most abundant species in the Vz group, comprising 50% of the colonies surveyed, when S. siderea and O. faveolata are included, the total contribution of these four species was 75%. Similarly, for AL reefs, C. natans and O. faveolata made up 41% of the coral community and adding M. cavernosa, S. siderea and P. astreoides the total contribution increases to 75%. Oculina spp., S. siderea and S. intersepta were more abundant in the Vz group than at AL group (Figure 9). Colonies in Antón Lizardo had significantly larger maximum diameters (mean ± SD: 82.6 cm ± 86.8) than in Veracruz (58.9 ± 50.4) (t = 13.314, d.f. = 5321.5, p <0.05) (Figure 9). Four species had mean maximum diameters greater than 1 m in the AL group O. faveolota (148.2 cm ± 104.3), O. annularis (145.5 cm ± 173.3), O. franksi (123.1 cm ± 78.3) and A. palmata (108.5 cm ± 100.4). In Vz, only O. faveolata (106.7 cm ± 87.3) had mean maximum diameter over 1m, the next largest colonies were of M. cavernosa (75.4 cm ± 54.3) (). Old partial mortality in Veracruz was (25.6% ± 26.1) and (25.4% ± 26.0) at Antón Lizardo, which was statistically similar between reef groups (t = -0.39111, d.f. = 6159.9, p = 0.6957). The species with the highest old partial mortality in Vz were M. decactis (46.1% ± 28) and S. intersepta (41.7% ± 28.5), and in AL it was M. decactis (42.3% ±28.6) again, and O. annularis (37.3% ± 24.1). Seven species in the Veracruz group lacked any signs of recent mortality, and nine species in Antón Lizardo. In the Antón Lizardo group only O. faveolata had more than 2% recent partial mortality (Figure 11). Prevalence of disease was 3% at Vz and 5% at AL. Siderastrea spp. had the highest prevalence of disease in both groups (10.9% at Vz and 22.6% at AL), and O. annularis had the second highest prevalence in Vz (4.5%), while A. palmata had the second highest in AL (8.7%).
The highest H’ value per reef was obtained for Periférico (2.24), the lowest values were found at Punta Brava, Punta Gorda and Hornos which all had values less than 1.
32 Discussion
Evenness was similar at all reefs (around 0.7) which indicates the species are relatively equally represented with some dominating. The highest evenness was found on Sargazo, the lowest on Hornos where 4 species were identified but the site was dominated by M. cavernosa (Table 6). The reef with the largest mean diameter colonies in the Vz group was Ahogado Grande (93.3 cm ± 60.3). The smallest colonies were found on the fringing reefs Punta Brava and Punta Gorda (11.9 cm ± 7.3 and 8.7 cm ± 5.2). The reef with the largest colonies overall was La Palma (113 cm ± 92.3). Three reefs had colonies with mean maximum diameter larger than 1 m: Anegadilla, Santiaguillo and Topatillo, all of them in the AL group (Figure 10). The highest mean percentage of old partial mortality was recorded at Ahogado Grande (47.9% ± 30.2) in Veracruz group. The greatest old partial mortality in Antón Lizardo group was recorded at Topatillo (39.4% ± 29.5). Punta Gorda stood out as the reef with the highest recent partial mortality (11% ± 25.3). Rizo was the reef in the Antón Lizardo group with the highest recent partial mortality (2.2% ± 8.1). Topatillo had the highest prevalence of disease (18%), the most prevalent disease at this reef was dark spot disease on S. siderea. Santiaguillo (7.3%) and Anegadilla (8.4%), which are the closest reefs to Topatillo also had higher prevalence of disease. In the Veracruz group disease was 3% on average, Galleguilla (4.5%), Hornos (4.9%) and Sacrificios (5.4%) had above average prevalence of disease.
33 Discussion
Table 6. Stony coral species richness, abundance, Shannon’s H’ diversity and Pielou’s J’ evenness for each reef of the VRS.
34 Discussion
izardo L ón nt z (middle,blue) and A eracru . Relative abundance of coral species for the VRS (top, red), V red), (top, VRS the for species coral of abundance Relative . 9 Figure
35 Discussion
onright. the
reefs
centpartial mortality and prevalence of Veracruz reefs and Antón Lizardo Antón and reefs Veracruz
the middle shows middle the . Mean and SD colony maximum diameter, percent old partial mortality, percent re percent mortality, partial old percent diameter, maximum colony SD and . Mean 10 Figure group reef per values has panel left The disease.
36 Discussion
or by species overall overall species by or recent,and prevalence ofdisease f old and and old
. . Mean ± SD length (cm), percent partial mortality partial percent (cm), length SD ± Mean . 11 Figure forthe VRS
37 Discussion
3.4 Distance Analysis
The linear regression models are an attempt to link some of the traits observed on the reefs with their distance away from three potential land based sources of pollution: the two main human settlements (Veracruz and Antón Lizardo) and the Jamapa river which divides the reefs into two groups. The two most northern fringing reefs, Punta Brava and Punta Gorda were excluded from the analysis because I considered that they are in a state that is not comparable to the rest of the reefs. Topatillo was excluded from the disease analysis because there seems to have been an outbreak of disease at this reef having a prevalence completely out of proportion with the rest of the reefs, and Ahogado Chico was removed from the coral cover analysis since it is an outlier with its very high coral cover and will be discussed separately.
The linear regression model for distance from the Port gave significant results for positive relation with all three variables. There was a weak relationship with coral cover (R2 = 0.23, p = 0.02), and incidence of disease (R2 = 0.27, p = 0.01), and a stronger correlation to the length of colonies (R2 = 0.45, p < 0.001) (Figure 12).
Something to notice in the middle column of Figure 12 is that all the points are shifted towards the right. The closest reef is 8 km away from the river mouth. This gives a rough indication of the area that is affected by the river discharge which is at least 16 km wide. We can also see that the reefs are mostly found between 8 and 20 km away from the river, except Anegada de Afuera, Topatillo, Santiaguillo and Anegadilla which are over 25 km away.
Distance from the Jamapa river mouth did not influence coral cover (R2 = 0.07, p = 0.23), but the model did indicate a strong positive relationship with colony size (R2 = 0.43, p <0.001) and disease (R2 = 0.38, p <0.05). The four reefs which are over 25 km had the highest incidence of disease (although Topatillo was excluded from the model because it incidence of 18% was extreme).
38 Discussion
Distance from Antón Lizardo did not have any statistically significant relationship with any of the variables measured. There is a weak negative correlation with coral cover (R2 = 0.15, p = 0.05). The R2 values for the other two variables was less than 2%.
39 Discussion
tón Lizardo. Reefs tón Lizardo. coral cover(top to bottom). Eachvariable is plotted
o in orange in o . Linear regression analysis for disease prevalence, colony length and stony stony and length colony prevalence, disease for analysis regression . Linear 12 Figure againstthe distance fromthree land based sources of pollution fromleft to right: Port of Veracruz,the Jamapa River and An are colorcoded by group Veracruzin blue and Antón Lizard
40 Discussion
3.5 Temperature
Fourteen months of temperature data (April 2008 - June 2009) were obtained from 7 temperature loggers deployed throughout the VRS (Table 1). The annual mean temperature in the Veracruz group was 25.5 ºC. The highest daily mean temperature was 29 ºC recorded in September 2008 for both deep and shallow. The minimum mean temperature of 22 ºC was recorded in June 2008 at the deep sites and January 2009 in the shallow sites (Figure 13). The mean annual temperature at Antón Lizardo was 26.2 ºC with a maximum mean daily temperature of 30 ºC in September 2008 for both shallow and deep sites. The minimum mean temperature of 23 ºC was recorded in February 2009 at both depths (Figure 13).
Although temperature patterns follow seasonal trends there are periods when the water column is stratified and there were differences of up to 2 ºC between the deep and shallow measurements. This is most clearly seen during the late spring and early summer months when the deeper waters are cooler, and has been previously documented in this region (Salas-Perez & Granados-Barba 2008; Salas-Monreal et al. 2009). The cooling of the shallow waters during February 2009, when the shallow waters were 0.7 ºC cooler than the deeper water, can be explained by the winter “Nortes” which can occur frequently in this region. During the summer and fall of 2008, there were periods of up to 20 days between peak temperatures in which water temperatures dropped to more than 6 ºC. This could be evidence of upwelling or cold-water intrusions, while not an uncommon phenomenon the frequency of these periods could be affecting coral health and survival, especially when they are occurring during a period when corals should be spawning.
41 Discussion
red. ón Lizardo in ón Lizardo Veracruz temepratures are shoen in green and Ant
. Mean daily temperatures from April 2008 to May 2009. Shallow temperatures were recorded at approximately 10 m and deep deep and m 10 approximately at recorded were temperatures Shallow 2009. May to 2008 April from temperatures daily . Mean 13 Figure 18 m. 12 and between
42 Discussion
4. DISCUSSION
4.1 Distribution
In 2007, Horta-Puga compiled many of the historical stony coral species lists for the VRS (Villalobos-Figueroa 1971; Rannefeld 1972; Kühlmann 1975; Tunnell 1988; Lara et al. 1992; Beltrán-Torres & Carricart-Ganivet 1999; Horta-Puga 2003; Jones et al. 2008), resulting in a list of 36 stony coral species (Horta-Puga et al. 2007) . Four genera, Solenastrea, Dendrogyra, Isophyllia and Isophyllastrea, and five species, D. labyrinthiformis, Eusmilia fastigiata, Madracis auretenra ex mirabilis, Meandrina meandrites and Mycetophyllia aliciae, were absent from these list, but are commonly recorded in Western Atlantic reef studies. However, during our surveys we recorded M. auretenra at Isla Verde, Pájaros, Blanca and La Palma, Mycetophyllia aliciae at Rizo and Isophyllia sinuosa at Anegada de Adentro, adding these three species to the VRS species list. Ten species on the compiled species list were not recorded during our surveys, Favia fragum, Manicina areolata, Oculina valenciennesi, Mycetophyllia danaana, M. ferox, Scolymia lacera, Porites branneri, P. colonensis, P. furcata and P. divaricata. Some of these species can be easily confused or overlooked for example F. fragum can be confused with Dichocoenia stokesii, and M. areolata can be confused with small colonies of Colpophyllia natans. Species of the Oculina genus are difficult to differentiate in the field and although we identified two species (O. diffusa and O. varicosa) further study could aid in clarifying if there are additional species of this genus present. Further inspection of the Mycetophyllia genus is needed as well to confirm the presence of M. aliciae. We did not record S. lacera because there is debate over whether it is truly a distinct species (Veron 2000). Similarly, we did not differentiate between Porites divaricata, P. furcata and P.
43 Discussion porites as some authors consider these to be different forms of a single species (Veron 2000). Both I. sinuosa and M. auretenra have been mentioned in previous studies (Jones et al. 2008; Villalobos-Figueroa 1971), leaving M. aliciae as a first record for the VRS. An additional species was recorded on three sites was the hydrozoan, Stylaster roseus. This species is not often included in species list because it is a hydrozoan, however it was initially described by Horta-Puga in 1990. A species that was not included in the transect data, but that was observed and photographed during the surveys was Acropora prolifera this coincides with early descriptions of the area by (Heilprin 1890) and (Kühlmann 1975), who recorded this species during their surveys.
Stony corals and crustose coralline algae covered 50% of the benthos in the VRS. This is could be an indicator of good health at the reefs. The presence of coralline algae at both reef groups could be cause for hope for a sustainable stony coral population in the VRS because coralline algae has been found to attract coral larvae and promote settlement (Negri et al. 2001; Vermeij & Sandin 2008; Ritson-Williams et al. 2009). The high coralline algae cover, in conjunction with the low macroalgal cover appear to favor the settlement and survival of stony corals since macroalgae are corals’ main competitors for space on the reef (Hughes 1994; Connell et al. 1997; Jompa & McCook 2003; Puglisi et al. 2014). In order for recovery to happen, there needs to be space for corals to settle and grow on, but there also needs to be a source of new corals which can colonize that space. Spawning has been observed for O. faveolata (Beaver et al. 2004), and connectivity studies show that the reefs in the southwestern Gulf could disperse larvae between them and to a much lesser degree with the Campeche Bank reefs (Sanvicente-Añorve et al. 2014; Johnston & Akins 2016; Jordán-Dahlgren 2002). However, recruitment is relatively low according to Horta- Puga (2009), 2.6 individuals/m2 compared to an average of 4.4 individuals/m2 for the Western Tropical Atlantic (Kramer 2003). During our surveys, I also noticed that coralline algae is most abundant in shallow areas, which at many sites were quite barren except for standing dead colonies of A. palmata, and other eroded corals. Rannefeld (1972) states that there were reports of a chemical spill in the 1960s that killed many of the corals in the shallow waters, possibly poisoning these areas. Additionally, the continuous input of fresh
44 Discussion water and terrigenous sediments from the Jamapa river (up to 2 kg/m2/day) (Pérez-España, 2008) could be preventing any form of biota from taking a foothold on the available space.
Stony coral cover is 7% higher in Antón Lizardo than in Veracruz, this difference is countered by bare substrate and turf algae in Veracruz. This could be due to more shallow transects being surveyed on the Veracruz reefs. The topography of the reefs in Veracruz tends to be shallower and at some sites habitat was not available to complete deep transects. Additionally, three fringing reefs, with very low coral cover, were surveyed in the Veracruz group while none were surveyed in Antón Lizardo. There are two fringing reefs mentioned for Antón Lizardo in the decrees but we were unable to survey them. The fringing reefs in the Veracruz reef are heavily impacted by sedimentation, runoff and the improper disposal of waste waters (Horta-Puga 2007).
Gorgonians were quite rare in our surveys, this could be due to the location of our survey sites, as gorgonians were mainly found in the shallow leeward areas of the reefs, which he didn’t sample as intensively as the windward sides. The two reefs with high abundance of gorgonians Blanca (Anton Lizardo) and Sacrificios (Veracruz), had survey sites on the leeward side. The gorgonian fauna seems to be less diverse than in the Caribbean (Jordán-Dahlgren 2002) possibly due to low connectivity (Sanvicente-Añorve et al. 2014; Johnston & Akins 2016).
Sponges are another competitor of space on the reef and in the case of boring sponges can degrade reef structure (López-Victoria et al. 2006). They can also be an indicator of phase shifts in coral community structure (Lapointe et al. 2007; McMurray et al. 2010) from a coral dominated community to one where either macroalgae or sponges become the dominant groups (Aronson et al. 2008). The low incidence of sponges on most reefs did not support this idea at the VRS, although it is important to note that the fringing reefs, Hornos and Punta Brava, which are under severe stress due to human uses, had some of the highest sponge cover, therefore, these shifts might be being expressed at individual reefs.
45 Discussion
Bare substrate should be an uncommon site on a healthy reef, this is because high diversity and competition for space are the prime indicators of a thriving ecosystem (Connell 1978; Box & Mumby 2007). The average amount of bare substrate was lower in the Antón Lizardo reefs pointing to more of the space being utilized by other functional groups. This was not only coral but macroalgae in some cases. In Veracruz, higher amounts of bare substrate were recorded, especially at Punta Gorda. The amount of substrate available may indicate that there are other environmental issues which are inhibiting settlement of organisms on the reef, whether it be lack of reproduction and settlement, or low light conditions caused by high sediment loads in the water, or chronic stresses that need further study (Cacciapaglia & Van Woesik 2016).
Mean macroalgal cover on Antón Lizardo reefs was higher than on Veracruz reefs, this difference was mainly driven by Topatillo reef where macroalgae cover was the highest for the whole reef system. High macroalgae cover at Topatillo and its neighbors, Anegadilla and Anegada de Afuera was surprising because I would have expected these reefs to be healthier i.e. lower macroalgae cover and higher stony coral cover, due to their distance from land. At about 20 km away from the coast, these reefs are furthest from any land based sources of pollution. It is possible that as sedimentation from the Jamapa river decreases in these reefs algae can outcompete the corals thanks to clearer waters. Macroalgae are generally considered a negative sign on coral reefs as they deter coral larvae settlement and compete for space with juveniles and adult corals (Jompa & McCook 2003; Hughes et al. 2002; Pandolfi et al. 2005; Box & Mumby 2007).
The range of stony coral cover for the Park was large and varied greatly between reefs. Four reefs, Ahogado Chico, Chopas, Enmedio and Santiaguillo had coral cover above 30%. These values are amongst the highest recorded for the Caribbean in recent times (Kramer 2003). The average for the whole system is 22% which is more in line with values from throughout the Caribbean. Furthermore, forty-four transects (or 6% of all transects) had over 50% coral cover, 10 in the Veracruz group and 34 in the Antón Lizardo group. This indicates that some sites are in a state similar to what was reported historically for the VRS and the Caribbean (Kühlmann 1975; Heilprin 1890; Jackson et al. 2014).
46 Discussion
Acropora palmata and A. cervicornis were observed on 20 reefs (Larson et al. 2014), the presence of these species throughout the VRS are of particular importance because they have suffered heavy declines at many Caribbean reefs in the 1970s and 80s (Adey 1978; Davis 1982). These species, and their hybrid are widely distributed in the VRS, growing to sizes that would permit reproduction (Williams, 2008) which indicate that this is a relatively healthy population which may be self-sustainable. Both Acropora species are listed as Critically endangered in the IUCN Red List, as well as subject to special protection under the Mexican law NOM-059-SEMARNAT-2010 which lists endangered species. The VRS being part of these species distribution range makes it an area of high conservation priority.
4.2 Stony Coral Demographics
The VRS is dominated by three species, C. natans, O. faveolata and M. cavernosa. Interestingly, M. cavernosa and O. faveolata switch dominance between reef groups. The dominance of M. cavernosa as well as the increased abundance of Oculina spp., S. siderea and S. intersepta on the reefs of the Veracruz group could indicate that this area is more degraded as these species tend to be found in areas of high environmental stress (Lasker 1980; Loya 1976; Edmunds et al. 2014). Another possible cause for this shift is the general topography of the reefs. There are more shallow fringing reefs in the Veracruz group which tend to be favored by M. cavernosa, compared to the deeper Antón Lizardo reefs.
The coral demographic data seem counterintuitive, the reefs of Antón Lizardo had indicators of better reef health than the Veracruz reefs as they had higher coral cover and larger colonies, however, they also had higher percentage of old partial mortality, and a higher prevalence of disease which point towards a community which is not in optimal conditions. These results could also simply reflect natural processes. Where there are larger colonies, there is more tissue to be lost and disease can spread more readily as there are more colonies to spread to.
47 Discussion
The average coral colony length for the VRS (71.3 cm ± 72.8) was larger than the 57 cm average calculated for the initial AGRRA surveys in the late 90s and early 2000s (Kramer 2003). The difference is likely greater, because the AGGRA protocol only surveyed colonies over 25 cm in size skewing the results towards larger colonies, while our study included colonies over 10 cm. The size of these colonies is important because their presence can be sign of “stable old growth” (Hughes 1984) and a lower extinction rate (Johnson et al. 1995). Large colonies are also usually indicative of at least the potential for reproduction (Szmant 1986) which means that if they are reproducing, of which there are indications (Beaver et al. 2004), this population could maintain itself.
Partial mortality is prevalent throughout the VRS, the vast majority of the colonies surveyed had partial mortality. This is not uncommon in corals, and the larger the colonies the more likely it is that they get partial mortality (Hughes 1984; Meesters et al. 1997). This may explain why we saw higher levels of partial mortality in Antón Lizardo versus Veracruz, since the colonies tend to be larger in the former group. There are some exceptions to this, particularly M. decactis and S. intersepta, Oculina spp. and Siderastrea spp. The growth form of M. decactis makes measuring the size of the colonies and estimating partial mortality difficult which could lead to an over estimation of partial mortality when the colony boundaries can’t be reliably defined. The other species had smaller colony sizes but higher partial mortality. This could be due to a few reasons, there is evidence that small colonies tend to be more vulnerable to partial mortality due to their high circumference to surface area ratio (Meesters et al. 1997). Stephanocoenia intersepta, Oculina spp. Siderastrea spp. also tend to live in with high sedimentation rates resulting in high rates of partial mortality, either on the tops of the colonies or around the edges closest to the substrate where they may be partially buried during periods of high sedimentation. The branching morphology of Oculina spp. also creates a trap for debris and colonies were often seen being smothered by fishing line and other trash (pers. obs.).
Recent mortality was not observed frequently in our surveys which, could be a positive indicator. However, the transition from recent mortality where the coral skeleton is completely denuded of live tissue and has not been colonized by turf algae or other
48 Discussion organisms, can be fairly short, so unless there is an active cause of mortality present at the time of the surveys, it is unlikely that it would be captured by sampling once a year. Still, recent mortality was observed more frequently on the Veracruz reefs which may be due to unfavorable conditions, such as high turbidity, runoff and pollution.
Disease was present on 14 species of stony coral in the VRS, and was most prevalent on Siderastrea spp., A. palmata, O. faveolata and O. annularis. Siderastrea spp. Siderastrea spp. were mainly afflicted by dark spot disease which has been previously reported as the most common disease in the VRS (Carricart-Ganivet et al. 2011; Horta- Puga & Tello-Musi 2009). Acroporids were affected by white band, rapid tissue loss and white pox, this is concerning because these species have already suffered decline in their population densities, and although the population in the VRS may not be as affected (Larson et al. 2014), the above average prevalence of disease could be cause for concern. The other genus which was affected was Orbicella with black band and yellow band diseases affecting mainly larger colonies, this is of potential concern because they are the main framework builders of these reefs (Alvarez-Filip et al. 2011). On the other hand large colonies may be able to withstand these infections better than smaller colonies as they do with partial mortality (Meesters et al. 1997).
4.3. Distance Analysis
The linear regression graphs show that distance away from the Port of Veracruz and the Jamapa river were positively correlated with the size of the corals and the incidence of disease. This is a counterintuitive result, since one would expect to have larger, healthier colonies as one moved away from the sources of stress. One of those turned out to be true, colonies did get larger as we moved away from these two points, but disease also increased. This could be a point of concern since, considering that Topatillo had the highest prevalence of disease (18%) and is located between the other three reefs with the highest prevalence of disease: Santiaguillo, Anegadilla and Cabezo, it is possible we found the source of an outbreak. When we surveyed Topatillo for the first time in 2010 there was already a high prevalence of disease then we witnessed a decline in A. cervicornis and A.
49 Discussion palmata cover at Santiaguillo and Rizo between 2011 and 2012 (Larson et al. 2014), which could have been caused by Tropical Storm Harvey and Hurricane Nate. It is possible that the disease may have spread from Topatillo to these other reefs with the increased water movement brought on by these storms.
Stony coral cover, colony length and disease prevalence all increased as distance from the Port of Veracruz increased. This would indicate that the urban area around the Port is a source of stress for the corals. Many human activities have been observed in the VRS: use of corals for construction material , coastal runoff and improper waste water management, overfishing, extraction for souvenirs (Jiménez-Badillo et al. 2006; Jiménez- Badillo 2008a; Horta-Puga 2007; Zamudio-Zamudio et al. 2003; Carricart-Ganivet 1998; Vargas-Hernández et al. 1993; Tunnell 1992) which are affecting the health of the reefs. These anthropogenic impacts could lead to this gradient of increased coral cover and larger colonies further away from the sources of stress, of which the Jamapa river seems to have the largest impact on. Therefore, as is mentioned in the 2012 decree, a much broader approach to the Park management must be adopted (Diario Oficial de la Federación 2012), where the whole area of influence is addressed and managed in a way that helps protect the natural resources not only within the Park but the areas surrounding it.
50 Summary of Findings
5. SUMMARY OF FINDINGS
5.1 Distribution
1. M. aliciae was added to the list of species found in the VRS 2. Confirmed the presence of M. auretenra and Isophyllia sinuosa in the VRS. 3.all 5 reefs added under the 2012 Decree plus an additional two undescribed sites were surveyed, of which Ahogado Chico deserves special attention as it had the highest coral cover in our study. 4. Values for coral cover are within published ranges for the VRS, but found areas which maintain high coral cover. 5. The reefs of Antón Lizardo have higher coral cover, but they also have higher macroalgae cover which could inhibit recruitment of new corals to those reefs. 6. The high cover of crustose coralline algae shows potential for recovery.
5.2 Stony Coral Demographics
1. Mean coral colony size for the VRS is among the larger sizes for the western tropical Atlantic (WTO). 2. Old partial mortality at the VRS similar to the reefs of the WTO. 3. Recent partial mortality is low and was found to be below the WTO average. 4. Disease prevalence in the VRS is average for values found for the WTO. 5. The abundance of large Orbicella colonies could be an indicator of stable old growth. 6. The presence of large coral colonies also implies the existence of reproductive potential.
51 Summary of Findings
5.3 Management
1. There is still uncertainty over how many reefs are in the Park. Five reefs were added 20 years after it’s initial creation and we propose two more. Better mapping and clarification on what is required for reefs to be listed are necessary. 2. Immediate threats to the reefs: urban sprawl, waste water management, influx of terrigenous sediments from a large watershed, overfishing. 2. Identify “hope spots”: areas of high coral cover, large colonies, high diversity. E.g. Santiaguillo, Blanquilla, Ahogado Chico. 3. Must publish a management plan that prevents the further modification of the Park boundaries and that clearly states what activities are permitted within the Park. 4. Coordinate with other agencies within the watershed to develop a catchment wide plan for managing the use of resources and reduce pollution, mainly through rivers and streams. 5. Develop a strategy for offering local fishermen and population at large opportunities to use the resources in a sustainable manner. 6. Deploy outreach programs to spread awareness of about the natural goods and services the reefs provide.
52
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