Goals

On completing this module, students are expected to be able to:

• describe the physical, chemical and biological controls on the livelihood of coral reefs, calcareous plankton, dinoflagellates and jellyfish;
• explain how human activity is causing ocean acidification, coral bleaching, red tides and blooms of jellyfish
• interpret evidence for the decline in reef health and its relationship with changing sea surface temperature

Learning Outcomes

After completing this module, students should be able to understand the following topics:

• projected change in pH in 2100 under SRES A2
• significance of pH, omega
• dissolution and precipitation reactions
• diversity of reef environments
• biology of corals, polyps, hard and soft corals, reproduction, role of zooxanthellae
• changes in calcification in the Great Barrier Reef
• causes and effects of bleaching
• coralline algae and their significance
• impact of acidification on plankton, pteropods, foraminifera, and coccolithophores
• causes of HABs
• life cycle of dinoflagellates and diatoms
• general impacts of toxins on fish and humans
• Pfiesteria controversy
• origin of CyanoHABs
• origin of giant jellyfish blooms
• impact of giant jellyfish blooms on coastal ecosystems

Below is an overview of your assignments for this module. The list is intended to prepare you for the module and help you to plan your time.

Background

Increasing levels of CO₂ in the atmosphere are slowly causing the surface of the ocean to become more acidic. This is because the ocean absorbs some of the CO₂, forming a weak carbonic acid. At present, the ocean absorbs about a third of fossil fuel emissions, but this amount is likely to increase to 90% in the future. Over the last century, the average pH of the ocean has decreased, and there are hints that the current levels are beginning to impact organisms that make their shells out of the minerals aragonite and calcite (both composed of CaCO₃). Coral reefs that are made of the mineral aragonite are highly vulnerable to ocean acidification. A recent study has found, for example, that the area of coral covering the Great Barrier Reef in Australia has been cut in half since 1985. However, coccolithophores and foraminifera, organisms that serve a vital role at the base of the marine food chain that are composed of calcite, are also susceptible. Moreover, the future appears to be even more bleak; some CO₂ projections suggest an additional pH decrease of 0.3 by 2100, a 150% increase in the ocean’s acidity compared to preindustrial times. Here we review the chemical changes in seawater that result from increasing CO₂, and then we discuss the impact on reefs and planktonic organisms in the ocean. Finally, we discuss the evidence for acidification in ancient oceans and its impact on life in the past.

Changes in atmospheric CO2 and consequences on the oceans.

Credit: Used by permission from Monterey Bay Aquarium Research Institute (MBARI [3])

The following video provides a thorough overview of the potential impact of acidification on the oceans.

Video: Acid Test: The Global Challenge of Ocean Acidification (21:34)

Chemistry 101

The ocean contains a massive reservoir of dissolved CO₂, hundreds of times more than in the atmosphere, and, actually, by contrast, the amount derived from fossil fuel burning is relatively modest. Since the beginning of the industrial revolution, about 340 to 420 petagrams carbon (a petagram or Pg is 10¹⁵ grams) in the form of CO₂ has been emitted to the atmosphere, with about a third of that amount absorbed by the ocean, approximately 118 Pg. Seawater today may already contain more CO₂ than at any time in many millions of years.

As we discussed in Module 5 on the Carbon Cycle, the absorption of CO₂ in the ocean forms weak carbonic acid (H₂CO₃). Some of this acid dissociates in seawater releasing H⁺ ions, which make the water more acidic, as well as HCO₃- (bicarbonate ions) and CO₃^2- (carbonate ions). This reaction is as follows:

CO₂(aq) + H₂O = H₂CO₃ = H⁺ + HCO₃- = 2 H⁺ + CO₃^2-

Behavior of carbonate species in the ocean with the addition of CO2 from anthropogenic sources.

Credit: National Academies

The CO₂, H₂CO₃, HCO_3- and CO₃^2- represent the dissolved inorganic carbon reservoir of the ocean; approximately 90% of this is HCO₃- and 9% is CO₃^2-.

Going back to your elementary chemistry course, you might remember that a pH of greater than 7 is regarded as alkaline whereas a pH of less than 7 is acidic. Surface ocean waters have a pH of between about 7.9 and 8.3, which means that they are, by definition, alkaline. Anthropogenic CO₂ is thought to have decreased the mean pH of the ocean by 0.1 unit since 1800. This may not sound like that much, but more ominous is the projection that if CO₂ levels continue to rise unabated (i.e., projections based on SRES A2 “business as usual”, pH levels will drop a further 0.3 by 2100. As we will see below, in parts of the ocean, these levels would be extremely damaging to organisms that build their skeletons out of CaCO₃, which is very sensitive to CO₂ addition.

CaCO₃ is the dominant material used by invertebrate organisms to build their skeletons. There are two different minerals made of CaCO₃, known as polymorphs: calcite and aragonite. These minerals have the same composition but different crystal lattice structure and thus their properties and behavior in seawater differ, including their ability to dissolve. To understand how CaCO₃ dissolves and precipitates, we need to introduce a term Ω that represents the saturation state of the water. Where waters are highly saturated with respect to CaCO₃ and Ω is high, calcite and aragonite are less likely to dissolve than where these waters are less saturated or even undersaturated and Ω is low. Likewise, calcite and aragonite are more likely to precipitate under higher Ω values. The dissolution and precipitation reactions are as follows:

Dissolution reaction: CaCO₃ (solid) = Ca²⁺ + CO₃^2-

Precipitation reaction: Ca²⁺ + CO₃^2- = CaCO₃ (solid)

The solubility of calcite or aragonite in seawater represents its ability to dissolve and is a function of K'sp, the solubility product where:

[Ca²⁺] x [CO₃^2-] = K'sp and Ω = [Ca²⁺][ CO₃^2-] / K'sp.

Both Ω and K'sp differ between calcite and aragonite, and K'sp is also affected by the temperature and pressure of seawater in which the CaCO₃ mineral is growing.

An increase in CO₂ from the atmosphere presents a double whammy for skeletons formed from CaCO₃, both aragonite and calcite. The H⁺ ions and carbonate ions (CO₃^2-) that derive from the dissociation of carbonic acid combine to form bicarbonate ions (HCO_3-). This rapid reduction in available carbonate ions decreases Ω and limits calcification by organisms with aragonite- and calcite-based skeletons. However, here we need to dispel two myths. The first myth is that the precipitation of CaCO₃ is directly controlled by pH. In fact, precipitation is affected principally by the decrease in CO₃², which is coincident with the addition of H⁺ ions, and reduction in pH. The second myth is that precipitation of CaCO₃ can occur in any water that is oversaturated with respect to the particular CaCO₃ mineral. In fact, both corals and coccolithophores have been shown to have difficulty calcifying in environments when waters were actually oversaturated. Different organisms can calcify at very different Ω values, but for most the decrease in saturation that results from decreasing CO₃^2- content is a direct threat to calcification.

The following video explains the threat of ocean acidification to the calcareous plankton.

Video: The Other Carbon Dioxide Problem (3:57)

The saturation of CaCO₃ in the oceans is also a function of temperature and pressure. A delicate balance exists between the production of CaCO₃ via the formation of skeletons in the shallow part of the ocean and the dissolution of this aragonite and calcite in the colder and deeper realms of the ocean where waters are less saturated. In most parts of the ocean, undersaturation occurs far below the surface. However, recent increase in dissolved CO₂ is leading to a shoaling of the saturation horizon of CaCO₃, and, in the future, this will impact especially the organisms that live at depth or in colder waters as well as those that make their shells of the mineral aragonite, which is more soluble in seawater than calcite.

Introduction

Coral reefs have existed for hundreds of millions of years and provided a habitat for some of the richest diversity on the Earth’s surface. They are the marine version of tropical rainforests. Reefs harbor a slice of the marine food chain all the way from tiny autotrophic protistans (autotrophs fix carbon through photosynthesis) to large, predatory fish. Hundreds of millions of humans live near reefs and receive important resources from them. Reefs host productive fisheries; they also provide protection to low-lying coastal areas from storms and are vital for a number of key habitats, including mangrove forests.

Examples of Reef Corals

[4] [5] [5] [5]

Examples of Deep Water Corals

[6] [7]

Click on the images above to see a full-size image and complete sources information.

The organisms that have constructed reefs, largely corals, have evolved over time, and with that change so have the locations of reefs and the dynamics of the reef community changed. Over their long history, reefs have had several intervals of crisis; in particular, they almost ceased to exist at the Permian-Triassic boundary, where over 90% of marine species became extinct, and during the Cretaceous about 100 million years ago when giant clams took over these structures for several tens of millions of years. Both of these ancient times were potentially characterized by ocean acidification. However, reefs have been remarkably resilient over geologic time and generally have been able to adapt to environmental change. For example, as we will see in Module 10, they are able to grow fast enough to keep up with very rapid rates of sea level rise.

Impact of Ocean Acidification on coral reef ecosystems

Credit: Great Barrier Reef Foundation, Australia [8]

With this background, recent human activity has placed reefs in as precarious a position as at almost any time in their history. The last fifty years have witnessed an extremely dramatic decline in the health of many of the major reefs around the world, including reefs of the Caribbean, the Bahamas, and the Florida Keys as well as those in the Indian and Pacific Oceans, including the massive Great Barrier Reef of Australia. The outlook for these rich and complex ecosystems is about as bleak as any ecosystem on Earth. As it turns out, ocean acidification is one of several environmental threats to reefs, with warming, pollution, overfishing and physical destruction all exerting major threats to reefs in the future. As we will see, acidification is perhaps the greatest of all of these threats.

Before we start, let's consider how reefs grow. The main organism that constructs modern reefs, the coral, includes a number of species belonging to the Cnidaria, a phylum of organisms that uses stinging cells to capture their prey. Modern corals are colonial structures of millions of individual polyps that grow primarily in shallow and clear tropical and subtropical waters, restricted to these areas by light levels and temperatures as well as by nutrients.

Red and yellow soft coral from Fiji.

Credit: Flickr, Derek Keats [9] / CC-BY 2.0 [10] (Creative Commons)

Both types of coral reproduce both sexually and asexually. Asexual reproduction involves simple cell division, or budding, and takes place within the colony, whereas sexual reproduction involves the release of gametes into seawater. This is an amazing process that for many species happens once a year timed by the lunar cycle. The fertilized egg forms a larval planula that settles before forming a new colony.

Video: Coral Growth (00:58)

Large reef structures including fringing and barrier reefs, as well as atolls, represent the growth of these colonies over many thousands or millions of years.

Evolution of an Atoll

Examples of modern corals living in symbiosis with dinoflagellate algae called Zooxanthellae

[11] [12] [13]

Click on the images above to see a full-size image and complete sources information.

Branching coralline algae from California

Credit: Steve Lonhart/SIMoN [14]

The algae live within the coral polyp and receive CO₂ from the polyp that it requires for photosynthesis. Through photosynthesis, the Zooxanthellae convert the CO₂ to O₂ and provide this vital gas to the coral as well as crucial nutrients. Reefs are made up of much more than corals and their algal symbionts. Other organisms, including coralline algae (algae that secrete high magnesium calcite and conduct photosynthesis on their own), are active framework builders in modern reefs. Today, sponges, sea anemones, sea urchins, a diverse array of fish, and many other organisms live within reefs, some playing a vital role in building reefs and keeping them healthy and others taking advantage of their decline.

Examples of Sea Urchins, Sponges, Sea Anemone and Coral Reef

[15] [16] [17] [18]

Click on the images above to see a full-size image and complete sources information.

Although, as we will see, ocean acidification is only just beginning to affect reef growth, there is already another very serious affliction in many reefs around the world---coral bleaching. One of the key changes in corals over the last few decades has been an increase in the frequency and severity of bleaching events. Bleaching occurs when the dinoflagellate temporarily or permanently leaves the polyp, resulting in a loss of color.

[4] [5] [5] [5]

Click on the images above to see a full-size image and complete sources information.

Coral in Florida invaded by blooms of red and brown algae.

Credit: Flickr [19], eutrophication&hypoxia [20] / CC-BY-NC-SA 2.0 [21] (Creative Commons)

Bleaching today is common when temperatures increase very slightly (by about 1^oC) in summer months and thus is often an annual summer event. Bleaching is accompanied by slower growth and increased coral mortality. The response to bleaching differs considerably between different species. Some species are able to recover normal growth rates quite rapidly, whereas others are more vulnerable and unable to recover. Bleaching also changes the ecology of a reef, promoting the growth of algae that blanket corals, making the recovery of corals more difficult.

Impact of bleaching on coral livelihood

Credit: NOAA National Ocean Service [22]

One of the most significant bleaching events took place in 2016 and 2017 in the Great Barrier Reef. This catastrophic event caused by warmer than average water caused up to 50% coral mortality in some areas. The extent of bleaching shocked ecologists and is clearly a window into the future.

Map of Great Barrier Reef bleaching in 2016 and 2017

Credit: Australian Research Council Center for Excellence for Coral Reef Studies. Australian Government, Great Barrier Reef Marine Park Authority [23] 

[24] [25]

Click on the images above to see a full-size image and complete sources information.

Deep Water Corals

Not all corals live in shallow water. A very diverse group of corals are able to thrive in the deep realms of the oceans. These corals lack Zooxanthellae and are able to calcify at cooler temperatures and lower levels of saturation than exist in the shallower waters. The number of known deep-water species has increased dramatically with new methods of exploring the deep oceans, including ROVs (remotely operated vehicles) and AUVs (autonomous underwater vehicles).

Examples of Deep Water Corals

[6] [7]

Click on the images above to see a full-size image and complete sources information.

These organisms exist in isolated patches at depths down to 2000 meters. In these conditions, the corals grow considerably slower than shallow water reefs. They feed on zooplankton and in some cases use chemicals coming out of the sea floor for a source of nutrition. Like their shallow-water counterparts, deep coral reefs provide a habitat for a diverse array of creatures. However, at the same time, it is clear that deep corals are in a particularly precarious situation, as we will see later on.

Corals and Calcification

The growth rate of corals, technically known as the rate of calcification, corresponds more closely to aragonite Ω rather than any other variable. Cores of corals from the Great Barrier Reef indicate a 14% decrease in calcification rates from 1990 to 2005. Corals generally maintain a high Ω at the site of calcification, but with decreasing CO₃^2-, precipitation of aragonite requires more energy, hence the decrease in calcification rate. Moreover, lab experiments show that the larval stages of corals grow more slowly under elevated CO₂. Overall, the net calcification rates of reefs decrease to zero when the Ω of aragonite is <1, and thus we can predict that worldwide reefs will switch from actively growing via calcification to shrinking via dissolution when CO₂ doubles to 560 ppm (the year 2050 in emission scenario A1B). The same logic leads to predictions of decreases in calcification rates of 10-50% by 2050. Experiments show that modern coral species have very different abilities to grow in water with lower saturation, with some species able to continue growing at lower pH and CO₃^2- levels.

Many of the predictions about calcification and saturation are based on experiments in the laboratory. Experiments are one of the best ways to forecast into the future, but they have significant limitations. In particular, it is impossible to replicate natural growing conditions in the lab, and further, experiments are conducted at intervals that are significantly shorter than the changes that are occurring in nature. In fact, saturation experiments conducted over longer time periods demonstrate less significant decreases in calcification than those that are conducted at short intervals. Moreover, experiments show that certain species of coral and coralline algae are less sensitive to decreasing levels of saturation and thus may be more likely to survive in the future.

Field studies and experiments often show very different results. Strangely, field results indicate much higher susceptibility of growth rates to decreasing levels of saturation than do laboratory experiments. These studies suggest that there is much left to learn about calcification and how it will change in the future. At the same time, it is clear that there is significant variability between species; some species having a much greater likelihood of survival in the future. Moreover, even within individual species, it is apparent that some reefs likely have the ability to survive better than others. Factors such as community composition, growth rate, nutrient levels and local variations in seawater chemistry and sediment composition will also play a vital role in determining which reefs will survive.

Since the coralline algae are more tolerant of colder waters, they have a very widespread distribution in the oceans, extending from the tropics to polar regions. The algae are also able to use low light levels for photosynthesis and therefore live considerably deeper than most corals. Their distribution suggests that they have great potential to adapt to variable environments. As it turns out, however, they may be in more precarious position than the corals.

The recent decrease in CO₃^2- has also begun to lower calcification rates of the coralline algae. These species are composed of high Mg calcite, which is the most soluble form of CaCO₃ (more so than low Mg calcite and aragonite), so they are particularly prone to ocean acidification. Experimental work confirms that calcification of the coralline algae is particularly sensitive to CO₂ levels with growth rates slowing significantly, and actually, dissolution beginning at moderately high CO₂ contents.

Perhaps the coralline algae will be the “canary in the coal mine” for dissolution of all framework structures under rising CO₂. For the corals, it is very apparent that the threat varies considerably from reef to reef. Deep-water corals are in particular jeopardy because calcification rates are lower in colder waters, because saturation levels at these depths are lower, and because, in the future, saturation stands to decrease more rapidly in deep than in shallow waters. However, the fate of reefs in the tropics is also likely to vary significantly.

Future Predictions

Making predictions about the exact impact of ocean acidification in the future of coral reefs of all types is inherently difficult. Acidification will occur in parallel with other deleterious effects such as bleaching and sea level rise. Species that are most susceptible to the effects of bleaching may turn out to also be more susceptible to extinction. For example, by impacting zooxanthellae, bleaching impacts calcification and thus may exacerbate the impact of acidification. However, these same corals also have the ability to recover from bleaching events more quickly, possess shorter generation times, and thus may have the ability to evolve more rapidly to tolerate bleaching in the future. At the same time, nutrient levels in reef environments have and will continue to decrease as stratification of the upper ocean increases, and this will put considerable strain on future reefs. Temperatures, nutrient levels, local rates of sea level rise, and human activities are factors that will likely decide which reefs are the most vulnerable, possibly with acidification pushing the most threatened systems over the edge into extinction.

Probably the largest uncertainty with respect to the future of reefs is whether corals will be able to adapt to lower saturation levels, increased temperatures, and lower nutrient levels, and if so, how rapidly. As we have seen earlier, corals have shown great resilience in surviving numerous environmental threats in the past. So, why do reefs recovered after times of environmental stress in the past, at times when temperatures were even warmer than they are today, and CO₂ levels higher, appear to be in such dangerous territory today? The answer to this question is that in the past, temperature and CO₂ perturbations occurred slowly enough that ultimate increases in weathering (remember feedbacks we discussed in Module 3) essentially decreased the rate of CO₂ addition and buffered the ocean with CO₃^2- before extinction occurred. What concerns scientists is that warming today and CO₂ addition are rapid enough that the weathering feedback will lag the decrease in saturation by several thousand years.

So, we are left with a lot of questions: will the rates of saturation decrease and temperature rise be too rapid for modern species to adapt? Will algae take over the niche of shallow water corals and dominate the low pH oceans of the future? Or will a few species of coral and possibly coralline algae develop the ability to calcify rapidly enough to survive the current threats and take over the niche of species that do not? Will Zooxanthellae themselves be able to adapt and assist corals in calcifying? Finally, when will feedbacks, largely through weathering, come into play and make conditions more favorable for calcifying organisms? For some of these questions, it’s a matter of wait and see. However, ongoing research should shed light on others. For example, the genetics of coral populations are currently being explored to understand the ability of corals to adapt to environmental change.

At this stage, however, any outcome is possible for corals, ranging from complete extinction by late in the 21st century to adaptation to a new set of environmental parameters. Ultimately, like many elements of the ecosystem, the fate of reefs may rest on how well we manage CO₂ emissions in the future.

The goal of this lab is to:

• observe and compare the health of reef by exploring different parts of the ocean.

Introduction

First, you will be watching videos from the Catlin Seaview Survey as well as photos to learn how to determine the health of reefs. Then you will be looking at before and after photos of reef bleaching events. Finally, you will be looking at the risk to reefs in the future.

Please Note: There is practice only for the video part of the lab. There are not enough pre and post photographs of bleaching events to use some for the practice, but the questions are going to be fairly straightforward, so practice is not necessary.

Please watch the videos below. To move around, insert your cursor in the videos to manipulate the camera and stop on particular items of interest or to change direction. The best way to move the camera is with your keyboard arrows (up, down and side to side).

The goal of this part of the lab is for you to show you can identify different types of coral as well as overall health of the reef at different locations. Make sure you have read the material on reefs in the module before attempting to complete the lab. Below are the different types of coral for you to identify. In addition, we show pictures of algae that colonize reefs as well as reef damage from storms. Healthy corals show a variety of colors from the different algal symbionts. Unhealthy corals show fewer colors, more algal colonization, more breakage and often are bleached white. Remember, algae are some of the key markers of an unhealthy reef.

Photographs of different types of coral

[26] [27] [28] [29]

Click on the images above to see full-size version and complete source information(from left to right: Staghorn, Brain, Table Bottom and Turf Algae).

Photographs of algal colonization

The following images show some of the range of morphologies and colors of colonizing algae

[30] [31] [32] [33]

Click on the images above to see full-size version and complete source information.

Photographs of reef damage from storms or people

[34] [35]

Click on the images above to see full-size version and complete source information.

Files to Download

Reefsall.kmz [36]

ReefsAtRiskRevisited2030.kmz [37]

ReefsAtRiskRevisitedPresent.kmz [38]

ReefsAtRiskRevisited2050.kmz [39]

Prep and Instructions

Load the ReefsAll. [36]kmz [36] file. The locations of reefs of interest are shown with flags. The videos are shown with diver markers, the still photographs are shown with wave markers. Before submitting your lab, let’s begin with the practice. Make sure you do this part of the lab to get comfortable with the tasks you will be asked to do and to receive feedback about your answers. Watch the videos at the following locations and answer the questions below. Make sure you maneuver up and (especially) down as well as side-to-side.

Practice Questions

Part A

In this section, you will be looking at the health of reefs using their color, the presence and abundance of algal overgrowth and the and the presence of bleaching. Go to the Agincourt Reef off Australia. Make sure you look at more than the first video as there is a lot of variety. Look around the reef and answer the following questions.

1. Is there living coral? (look for different colored coral) (Yes/No)
2. Is there brain coral? (Yes/No)
3. Which types of coral do you see?
   A. Table bottom
   B. Table bottom and staghorn
   C. Staghorn and brain
   D. Table bottom, staghorn, and brain
4. Is there any bleached coral? (Yes/No)
5. Is there any algal overgrowth? (Yes/No)
6. How would you describe the health of the reef?
   A. Very healthy
   B. Moderately healthy
   C. Not healthy

Part B

In this part of the lab, you will compare photographs from before and after major events that have impacted reefs. Answer the questions about the changes in the abundance of different type of coral, algal overgrowth or percent bleaching.

Go to Lizard Island. Please look at the two pictures and answer the following questions.

7. Which picture has the healthiest reef? (upper/lower)
8. What is most diagnostic of the health of the upper photograph?
   A. The healthy reef has very tall coral
   B. The healthy reef has a lot of different colors
   C. The healthy reef has bleached corals
9. What is most diagnostic of the health of the lower photograph?
   A. The unhealthy reef is bleached
   B. The unhealthy reef is growing quickly
   C. The unhealthy reef is covered by algae

Part C

In this section, you will be looking at the projected health of the reef in the future. Load the three Reef risk kmz files, the first is for today (ReefsAtRiskRevisitedPresent.kmz [38]), the second for 2030 (ReefsAtRiskRevisited2030.kmz [37]), and the third for 2050 (ReefsAtRiskRevisited2050.kmz [39]). When you look at these, make sure you only have one set of projections and keep the others switched off. Look at the Indian Ocean and answer the following questions.

10. Where is the threat highest at the current day? Select all that apply.
    A. Indonesia
    B. East Africa
    C. Maldives
11. Given the distribution of the threats in 2030 compared to high population areas around the margins of the ocean and the lower population areas in the ocean islands, which is most likely the largest threat to reefs? Select all that apply.
    A. Bleaching
    B. Ocean acidification
    C. Pollution and overfishing
12. Why is the threat is so high everywhere in 2050? Select all that apply.
    A. Ocean acidification
    B. Bleaching
    C. Pollution and overfishing

Introduction

Corals and coralline algae are not the only organisms highly susceptible to ocean acidification. Coccolithophores, foraminifera, pteropods, three very different groups of plankton (a term that refers to organisms that float passively in the upper ocean) are also threatened by increasing atmospheric CO₂ levels. Pteropods, often called sea butterflies, are tiny snails made of aragonite that thrive in shallow waters and play a particularly important role in polar ecosystems.

Pteropods

Pteropod on the right and a very small octopus on the left

Credit: Flickr/NOAA Photo Library

Change in pteropod shells subjected to seawater with pH and saturation projected levels for 2100

Credit: PMEL Carbon Program [40]

Click on the images above to see a full-size image and complete sources information.

Because of the susceptibility of aragonite to decreasing saturation levels combined with the effect of temperature (calcification requires more energy in cold water) on saturation, pteropods may cease to exist in polar latitudes by the middle of the 21st century. In some polar areas, these organisms account for over 60% of the zooplankton biomass, thus their extinction or migration to warmer regions could have major repercussions to organisms up the food chain.

Foraminifera and Coccolithophores

The coccolithophores and foraminifera are constructed of low-Mg calcite and therefore are more stable than pteropods under conditions of increasing CO₂.

As we have learned earlier, foraminifera are groups of zooplankton with habitats both near the surface of the ocean (planktonic foraminifera) and on the ocean bottom (benthic foraminifera).

Foraminifera

[41] [42] [43]

Click on the images above to see a full-size image and complete sources information.

The planktonics are sensitive to changes in CO₂, have shown subtle changes in shell mass during the Pleistocene in response to CO₂ fluctuations, and have already begun to get lighter in response to recent CO₂ increase. This recent thinning of shells is likely a result of the effect of decreasing CO₃^2- on calcification by foraminifera. Interestingly, many planktonic species also harbor dinoflagellate symbionts that play a role in calcification. Changes in calcification appear to be a threat to the planktonic foraminifera, but we do not yet know how serious this threat is. However, we do know that the foraminifera are also threatened by the potential loss of their dominant source of food, the coccolithophores, as a direct result of CO₂ addition.

Living coccolithophorids of Emiliania huxleyi

Credit: Jeremy Young, the Resilient Earth [44]

Coccolithophores

Coccolithophores are ubiquitous in the oceans, essentially serving as the dominant species of phytoplankton in vast regions of the open ocean that are characterized by lower nutrient levels. For this reason, considerable attention has been devoted to the potential effects of increasing CO₂ on the coccolithophores.

Coccolithophores are haptophyte or golden brown algae (similar to diatoms) that produce tiny calcite scales known as coccoliths during certain phases in their life cycle.

The plates are several microns in diameter and can remain attached to the cell covering the soft organelles with a protective shield or break off after they are fully grown. Coccolithophores reproduce asexually, and, given the right conditions have the potential to multiply rapidly. When conditions are suitable, coccolithophores can form blooms of millions of cells per liter of seawater. These organisms are consumed by foraminifera and copepods, and transported to the bottom of the ocean as marine snow.

Video: Marine Snow (00:33)

Because of their prolific production and global distribution, coccolithophores are a vital part of the carbon and carbonate cycle of the oceans. Modern coccolithophorids are dominated by the species Emiliania huxleyi, a species with very small (1-2 micron) and delicate coccoliths. Due to their size and ecology, coccolithophores are inherently more difficult to study in the natural environment than are corals. However, samples can be collected using filters and cores of sediments can be studied to determine the effects of past changes of CO₂ on coccolithophore morphology. Coccolithophores can also be cultured in the laboratory where CO₂, CO₃^2- and pH levels can be altered to observe the effect of these variables on calcification. Although the results of both field and lab studies are by no means simple, it appears that in the case of E. huxleyi, increasing CO₂ and decreasing CO₃^2- has the effect of causing thinner coccoliths with smaller masses. In addition, laboratory studies show that such conditions also lead to malformations in E. huxleyi coccoliths, which are a potential sign of difficulty in calcification. Such trends of decreasing mass are by no means universal, however. Some lab experiments and field collections in particular environments show that species other than E. huxleyi actually grow thicker in low CO₃^2- conditions and even E. huxleyi recently has increased in mass in parts of the ocean. More study is required to determine the precise outcome of the coccolithophores in a high CO₂ world. However, the current signs generally point to significant reduction in the rate of calcification, which could lead to significant changes both in marine ecosystems and in the carbon cycle.

The following video, Climate Change and Ocean Acidification, by San Francisco State University clearly captures this problem and explains the ramifications on the ocean and the atmosphere.

Video: Climate Change and Ocean Acidification (3:47)

Coccolithophores have a spectacular 220 million year fossil record. This record allows paleontologists to observe the effects of past climate change, including increasing CO₂, on the livelihood of this group of plankton. Ancient global warming events, including those at 120 million years before present and 55 million years before present, have been gleaned for evidence of ocean acidification in the morphology of coccolithophores. The event at 120 million years shows evidence for a decrease in coccolith size, but this change is not apparent in the 55 million year event. As the species existing during these ancient events was entirely different from those at the present time, it could be that ancient species responded in a different fashion from those living today. Alternatively, the changes in ancient plankton assemblages may be in response to environmental variables other than CO₃^2-, for example, temperature and nutrients. The coccolithophores, like the corals, have been able to survive intervals of great ecological upheaval in the past. However, given that the rates of modern environmental change are so rapid compared to ancient events, we cannot assume that these same groups will have the ability to adapt to the changes that are to come. Moreover, while the coccolithophores appear to be less endangered by increasing CO₂ than the corals, largely as a result of their mineralogy, the impact of the decline of these vital algae would likely be even more devastating to the oceans.

Red tides are common events in warm and polluted coastal oceans. They form when dinoflagellate algae explode to huge population levels. Because the dinoflagellates have red plastids, the waters literally turn red. Dinoflagellates take advantage of harsh environmental conditions that kill off other organisms. As you will find out in the pages to follow, these tides can be major public health hazards.

Red tide off the coast of California

Credit: Woods Hole Oceanographic Institution/NOAA [45]

I remember that the grouper was delicious. Blackened, spicy, with an ear of corn and some slaw. On a white paper plate. I polished it off with a Heineken. It was a warm early January day, a seafood festival in Florida City. We went canoeing after lunch. I remember waking up the next morning with an intense thirst and extreme nausea. When I tried to get out of bed, I sprawled on the floor, my left side was completely paralyzed! I rolled to the sink, crawled up, and poured myself a glass of water, I gulped it down, and to my horror, it felt boiling hot! The neurological symptoms soon went away, but nausea and fatigue lasted weeks, I lost at least ten pounds. The doctors took that long to diagnose the problem---I had Ciguatera poisoning. That delicious grouper had ingested a lot of toxic dinoflagellates and transmitted the toxins to me. My first experience with harmful algae.

Red tides represent one of the most serious threats to coastal ecosystems today. Most red tides result from the input of an excessive amount of nutrients from fertilizers, sewage, and soils of nearby land areas to bays, estuaries, and shallow seas. These nutrients cause explosive growth of microscopic species of algae, a number of which carry toxins that are harmful or even lethal to other organisms. Red tides can be caused by major storms such as hurricanes, which cause excess runoff from the land and resuspension of the seed stages of the algae.

Red tides occur when dinoflagellates, and rarely diatoms, grow in massive quantities in surface waters. The photosynthetic organelles of these organisms, known as plastids or chloroplasts, are red, or golden brown in the case of diatoms, and the profusion of cells in surface waters imparts a red or brown color. Some of the culprit dinoflagellate and diatom species produce by-products that are highly toxic to many other organisms living in the coastal zone, all the way from fish to turtles to large mammals such as dolphins, manatees, and whales, as well as to humans. In some cases, shellfish or small fish such as sardines that consume the plankton are not harmed by the toxin but concentrate it for organisms that feed on them, a process known as bioaccumulation. Ingestion of the toxins can result in developmental, immunological, neurological, and reproductive damage of the host organism. For this reason, red tides are also known as harmful algal blooms (HABs). In fact, we will use the term HAB here because these events are not associated with ocean tides, because many HABs are not associated with a red color, and because blooms of dinoflagellates are often not harmful.

Examples of Dinoflagellates and Diatoms

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Harmful algal blooms are a global phenomenon and have increased in frequency in the last thirty years. Part of the increase may result from awareness of the phenomenon, but increasing pollution is also considered responsible. There are a number of reasons why climate change may further increase the occurrence of HABs. For examples, the increase in the frequency of large storms such as hurricanes will lead to greater runoff and input of nutrients from land. In addition, changes in temperature, wind patterns, upwelling, and stratification will alter the distribution of species. Because the degradation of a large amount of cellular material produced in HABs consumes oxygen, HABs can result in hypoxic or anoxic conditions.

Dinoflagellates are a group of microscopic single-celled organisms or protists that are dominantly autotrophic (i.e., primary producers). Interestingly, many species are also mixotrophic, having the ability to ingest their prey as a source of energy. Some species are entirely heterotrophic, lacking chloroplasts or plastids, and have been termed carnivorous. In fact, there have been reports in the scientific literature that some species have the ability to consume fish after having paralyzed them with neurotoxins. These claims are controversial but have given dinoflagellates a near-mythical reputation among the oceanic plankton.

Simplified life cycle of a dinoflagellate.

Credit: Woods Hole Oceanographic Institute/NOAA [48]

Diatoms and dinoflagellates are most common in the coastal oceans but also have the ability to live in freshwater environments and in intermediate salinity environments where fresh and marine waters mix in estuaries. They are the most prolific group of primary producers in the ocean. Dinoflagellates have a highly complex life cycle that consists of an alternation between a motile stage and a resting or cyst stage. In short, dinoflagellates enter the resting stage via sexual reproduction when conditions in the surface ocean are not suitable for them to thrive. They can remain dormant for weeks, months or years before they “excyst,” when surface conditions improve, reproduce vegetatively, and populate the surface ocean. Excystment and repopulation are triggered by changes in temperature, light, or oxygen levels or even resuspension of cysts by storms. Dinoflagellates generally thrive when nutrient levels are elevated, and, under conditions of extremely high nutrient levels, cell division can be so rapid that extremely high cell counts (millions of cells per milliliter of seawater) are reached, resulting in red tides. The cyst stage acts as a very effective mechanism for seeding blooms.

Dinoflagellates

The following video summarizes the life cycle of the dinoflagellates.

Video: Life Cycle of the Dinoflagellates (1:27)

Dinoflagellates have a broad range of different ecologies. As we saw earlier in the module, they can be endosymbionts of corals, facilitating calcification in the host colony. They have this same role in foraminifera and radiolarian, a group of siliceous zooplankton.

Diatoms

Diatoms are autotrophic protists that produce a delicate, microscopic test of opaline silica.

Pseudo-nitzschia, a toxin-producing diatom.

Credit: Raphael Kudela, USCS Ocean Data Center [49]

They are non-motile, and, for most of their life cycle, they reproduce asexually. Many nearshore diatom species also have a resting stage akin to the dinoflagellates, allowing them to exit the surface zone when conditions are unfavorable for their growth. This may occur in winter when surface waters are cold or at times when nutrients are depleted. The resting spore stage actually may resemble the vegetative stage of dinoflagellates. Like the dinoflagellates, diatoms are able to reproduce extremely rapidly by simple cell division, and this allows them to rapidly dominate the surface ocean when nutrients are readily available.

Video: Diatom Life Cycle (1:38)

Species that are harmful belong to the pennate diatoms that are long and thread-like and have the ability to attach to a host, although the relationship is not symbiotic. Both dinoflagellates and diatoms cysts can move around the oceans by currents, storms, dredging of the ocean bottom, and when cysts act as ballast on ships or even higher-level organisms. Toxins are not known in the cyst stage of either group.

Of some 60 or so species that cause red tides, only a handful is known to be toxic. Dominant dinoflagellate HAB genera include Alexandrium, Karenia, and Pfiesteria. The diatom genus most commonly associated with HABs is Pseudo-nitzschia. Each of these genera produces a different toxin and thus has a different role on organisms further up the food chain. Next, we discuss some of the HAB species in detail.

Scanning electron microscope view of cysts of the toxic alga Alexandrium tamarense

Credit: Woods Hole Oceanographic Institute/NOAA [50]

Alexandrium spp. is the dominant taxon in coastal regions of New England and eastern Canada but it is also found from California to Alaska.

It is a heterotropic dinoflagellate that produces a saxitoxin, one of the most powerful known types of neurotoxins. These toxins destroy the function of nerve cells and can thereby cause paralysis. Saxitoxins are most effectively concentrated by shellfish such as clams, quahogs, mussels, scallops and oysters that filter large volumes of sea water to acquire their nutrition. Although the saxitoxin does not harm these shellfish, even in small quantities, the toxin can be extremely dangerous for humans, resulting in a serious illness known as paralytic shellfish poisoning (PSP). The saxitoxin attacks the human nervous system within 30 minutes of ingestion with symptoms that may include numbness, tingling, weakness, partial paralysis, incoherent speech, and nausea. In severe cases, the toxin can lead to respiratory failure and death within a few hours. Alexandrium spp. toxins have also been harmful to whales, sea otters and birds.

The following videos describe the causes and impacts of red tides as well as possible antidotes for shellfish poisoning.

Video: Red Tide - Death Comes to Gasparilla Island Florida (5:59) This video is not narrated.

Video: Prized Science Episode 4: Taming the Red Tides (4:28)

Karenia is the dominant taxon causing red tides in Florida and Texas, but rarely species of Karenia have also been found up the east coast in North Carolina.

Examples of Karenia

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It produces a toxin known as a brevetoxin (named after a species of Karenia, K. brevis). Like saxitoxins, brevetoxins damage nerve cells, leading to disruption of normal neurological processes and causing neurotoxic shellfish poisoning (NSP). In humans, gastrointestinal symptoms and a variety of neurological ailments result, but there are no known fatalities. However, in fish, the brevetoxins attack the central nervous system and cause respiratory failure. Karenia dinoflagellates are responsible for massive fish and bird kills in the Gulf of Mexico. The brevetoxins are colorless, odorless, and heat and acid stable, thus they survive food preparation.

Gambierdiscus toxicus a species that can produce ciguatera toxins

Credit: Smithsonian Institute [54]

The genus Gambierdiscus lives in tropical waters, usually in reefs, and produces a toxin known as Ciguatoxin that causes gastrointestinal problems followed by mild neurological symptoms. This syndrome is known as Ciguatera fish poisoning. Because the toxin is fat soluble, it gets concentrated up the food chain by bioaccumulation from seaweed to smaller fish than to larger fish. The larger fish, which are the most dangerous to eat because they have the highest toxin concentrations, include commercially available seafood such as grouper, snapper, and barracuda. Ciguatera is responsible for more human illnesses—estimated between 10,000 to 50,000 cases annually—than any other HAB toxin.

Scanning electron microscope image of Pseudo-nitzschia australis

Credit: Virginia Institute of Marine Science [55]

Pseudo-nitzschia and the species Nitzschia navis-varingica are common diatom genera in Californian red tides. These taxa produce domoic acid, which is concentrated by filter-feeding shellfish. This neurotoxin can also bioaccumulate in fish such as anchovies that feed directly on the diatoms. Domoic acid causes a variety of gastrointestinal ailments, memory loss and brain damage in humans and is hence referred to as amnesic shellfish poisoning. Rarely, the neurotoxin can be fatal. It can also affect marine mammals, causing seizures.

Scanning electron microscope image of Pfiesteria piscicida.

Credit: Virginia Institute of Marine Science [55]

The species Pfiesteria piscicida and P. shumwayae have been the most common dinoflagellate species in red tides in estuaries and bays along the east coast of the US from Delaware to Florida. The species occur in environments where freshwater and saltwater mix and have not been reported from freshwater environments or the open ocean. Pfiesteria blooms are restricted to summer months. Species have been associated with massive kills of menhaden and other estuarine fish in the Chesapeake Bay and the Tar-Pamlico and Neuse River Estuaries in North Carolina. The fish in contact with Pfiesteria rapidly develop bleeding lesions and have skin actively flake off them, and it has been proposed that the presence of live fish stimulates the production of toxin in the dinoflagellate. Ultimately the open lesions may destroy gill function and lead to death. Nevertheless, the connection between fish mortality and Pfiesteria is still doubted by some scientists. In fact, the effects of Pfiesteria on fish and human health has been one of the largest and nastiest controversies in marine science over the last 25 years and it does not appear that the conflict is anywhere near over.

The following video explains some of the research on Pfiesteria.

Video: Pfiesteria Update: An Enduring Debate (4:57)

At least part of the debate has been fueled by the popular press, who have focused attention on the organism after studies suggested that it was carnivorous. These studies indicated that Pfiesteria species ingested the skin of fish after it flaked off. In fact, lab studies have shown that when fish were left in tanks with Pfiesteria, the fish died within hours. For this to occur, however, the dinoflagellate and fish must be in direct contact. Without contact, the same studies show that fish suffered no ill effects. Some scientists alternatively point to water molds or fungi including the species Aphanomyces invadans as the pathogen cause of the ulcerative lesions, skin loss, and damage to gills. A. invadans and other fungi are universally present in fish with ulcers and skin loss. Also weakening the case for Pfiesteria, this genus is still known to exist in North Carolina estuaries, but fish kills have become less frequent recently. Moreover, where lesions on the fish menhaden were observed, nearby fish including catfish, perch, and carp were unaffected. These disparities have cast some doubt on whether Pfiesteria is harmful to fish at all. In fact, great differences exist among public health professionals, and warnings from different state agencies are in conflict.

Examples of Pfiesteria

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Click on the images above to see a full-size image and complete source information.

The health impact of Pfiesteria on humans is also uncertain, as is the method of transmission of the potential toxin. Scientists working with Pfiesteria in the laboratory have suffered from long-term neurological symptoms, such as memory loss, fatigue, and dermatological problems, and fisherman in contact with Pfiesteria-related fish kills have also suffered from similar ailments. However, other groups of fishermen who have come in close contact with lesion-covered fish have not reported adverse effects. There are reports that the hypothetical Pfiesteria toxin is transmitted via aerosols.

Until recently, the missing link in the Pfiesteria conundrum is that the toxin produced by this organism has been elusive. However, in 2007 scientists in a government lab claimed the first positive identification of a toxin associated with Pfiesteria. Even with this identification, questions remain; for one, the toxin is unstable in the natural environment, and second, it is not been proven to have adverse health effects. Thus, the controversy about Pfiesteria is far from over. In all reality, a number of factors may result in the fish kills; in particular, the fish in estuaries may have already been under great stress from other biological agents (bacteria, viruses, fungi, parasites), exposure to chemicals (pollutants, toxins), suboptimal water quality, and rapid water temperature change, that have the potential to cause lesions to form.

Far less controversial than the relationship of Pfiesteria with fish kills are studies that directly relate fish kills to low oxygen levels caused by algal blooms. In a number of estuaries along the eastern US, warmer waters in summer combined with increased production by algae, as a result of increased runoff and eutrophication, lead to severely decreased oxygen levels and major fish kills without the involvement of a toxin.

Finally, HABs are not always produced by dinoflagellates and diatoms. Cyanobacteria, or blue-green algae, another group of single-celled organism, but one that is prokaryotic rather than eukaryotic, is also known to produce extremely potent toxins that can cause illness in fish, birds, and mammals including humans. Because of their potential to be harmful, this group is known as CyanoHABs.

Samples of CyanoHABs

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Click on the images above to see a full-size image and complete sources information.

Although the full scale of health effects of CyanoHABs on humans is not yet determined, the toxins may have gastrointestinal, respiratory, allergic and neurological responses, and potentially lead to liver damage. In addition, prolific growth of cyanobacteria can block sunlight, which can harm other organisms, and use up oxygen which can lead to hypoxia and anoxia. As in the case of the HAB dinoflagellates, the growth of CyanoHABs may be stimulated by nitrogen loading from agricultural industrial runoff, as well as sewage disposal.

As we have seen, there is great variety in the biology and ecology of HAB species. However, they share one major thing in common: all of them have the ability to wreak havoc on coastal fisheries. Since the growth of most if not all of the species directly responds to nitrogen loading, limiting the harm on fisheries will require significant changes in agricultural practices combined with modifications of drainage in coastal regions. Such changes will be difficult, if not impossible, to accomplish in the near term. Thus, the best strategies to deal with HABs focus on the integration of highly detailed algal sampling programs, ecological forecasts, and resource management. For example, if HABs can be predicted, then warnings can be issued and areas placed off limits to fisheries. Such strategies are being employed in areas where HABs are common, including the Gulf of Maine, the Gulf of Mexico and the Pacific Northwest.

HAB forecasts integrate ecological models, based on the physics, chemistry, and biology of nearshore and offshore regions, with satellite data and in situ measurements of cell counts and toxin levels. For example, models can be used to predict the development and movement of a HAB in the region of interest. Models can be used to identify HAB triggers (i.e., nutrients or temperature), likely areas of cyst seedbeds, likely bloom toxicity based on cell density, and progression of the toxin through the food chain, as well as the ultimate decline of the HAB.

A new threat to fisheries around the world has developed over the last decade---a surge in the number of jellyfish in coastal waters. The most dramatic of these outbreaks is in Japanese waters where the giant Nomura’s jellyfish has increased significantly, wreaking havoc with fisheries in the Sea of Japan.

Jellyfish populations are normally held in check by fish, mostly because these two groups compete for the same food sources. However, overfishing in many parts of the ocean has led to increasing jellyfish populations. Jellyfish may also be aided by warming ocean temperatures, which favors their development, and by the destruction of habitats of other natural predators such as turtles.

The massive Nomura’s jellyfish is a great threat to Chinese, Japanese and Korean fisheries. These creatures can grow to two meters diameter (the size of large refrigerators) with a weight of 200 kg.

Examples of Giant Jellyfish

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Click on the images above to see a full-size image and complete sources information.

Because of their size, they consume massive amounts of zooplankton, depleting this vital part of the food chain for other organisms. The key threat of the Nomura’s derives from the fact that this jellyfish reproduces extremely rapidly. A mature jellyfish has the ability to produce billions of eggs at a time, and they can do this when they are attacked. Once fertilized, these eggs develop into a resting polyp stage that also has the ability to multiply rapidly, effectively carpeting areas of the seafloor. When conditions are suitable, the polyp reproduces asexually, developing into the medusa stage, which grows into the mature jellyfish.

Video: Jellyfish Life Cycle (1:41)

Once ideal conditions develop either by increasing nutrients or warming of the surface ocean, Nomura’s jellyfish populations literally explode and render fishing virtually impossible because nets become filled with jellyfish. These jellyfish can also continue to reduce the number of fish in the oceans by feeding on their eggs. Moreover, there is evidence that jellyfish can tolerate conditions, like hypoxia, that fish cannot.

The following video summarizes the impact of giant jellyfish on Japanese fisheries.

Video: Monster Jellyfish (2:35)

Blooms of other jellyfish species are being reported in many other parts of the ocean. In the Gulf of Mexico, for example, the last thirty years populations of two species of jellyfish, the sea nettle, and the moon jellyfish, have exploded especially in dead zones as these are one of the few organisms that can tolerate hypoxia. Jellyfish in the Gulf now swarm over hundreds and perhaps even thousands of square miles each summer.

A swarm of sea nettles (Chrysaora quinquecirrha) in the Gulf of Mexico.

Credit: CC-BY-NC-SA, Lyn Gateley

Here also, invasive species of jellyfish including the Australian jellyfish, have been reported. Several other factors besides hypoxia have caused the increase in Gulf of Mexico jellyfish. As in the Sea of Japan, overfishing has reduced one of the main jellyfish competitors. In addition, drilling platforms have provided habitats in which jellyfish polyps can multiply. As in the Sea of Japan, jellyfish in the Gulf of Mexico are impacting the fishing industry.

Jellyfish swarms, too, have plagued other regions; they include northern Australia where the highly venomous box jellyfish has expanded its range, the Black Sea, and the Bering Sea off Alaska. Worldwide, jellyfish are one of the few organisms that can thrive in dead zones. With the spread of such dead zones in the oceans as a consequence of marine pollution and climate change, we could be entering the age of the jellyfish.

End of Module Recap:

In this module, you should have learned the following concepts:

• factors that cause the pH of the oceans to change and how it impacts saturation of the minerals calcite and aragonite;
• the biology and ecology of reef environments and how they are likely to change in the future with acidification and rising temperature;
• the potential impact of acidification on calcareous plankton groups and implications for the marine food chain;
• the biology of harmful algae including dinoflagellates and diatoms;
• factors that cause blooms of harmful algae, why they are likely to increase in the future, how they can be forecasted, and how they impact human health;
• about Cyanobacteria and the factors that cause them to bloom;
• about large jellyfish and factors that have led to major increases in their populations.

Assignments

You should have read the contents of this module carefully, completed the lab and submitted in Canvas, and taken the Module Quiz. If you have not done so already, please do so before moving on to the next module. Incomplete assignments will negatively impact your final grade.

Labs

• Lab 7: Reef Ecology

This week we focus on the impact of ocean acidification on fisheries and marine life in general.

A. US:

• Acidification and other Ocean Changes [62]

B. Global

• Ocean Change [63]

Instructions

Step 1: Read the following.

• Chapter 13: Ocean Acidification and Other Ocean Changes [62]

• Chapter 6: Ocean Systems [63]

Step 2: Consider how the element of climate change you chose to read about impacts a place of interest to you, then write a small vignette (200-400 words). You should focus on why that place is vulnerable and what is being done about it. Your vignette should include answers to the following questions:

• What is the threat?
• Why is that place/regional vulnerable?
• What are the forecasts in terms of environmental change?
• What are the forecasted impacts on communities?
• What are the solutions? (If applicable)?

Step 3: Using Screencast-o-matic [64], record your voice reading your vignette.

Step 4. Place your video clip at the relevant place in Google Earth by attaching it to a Pin and save as a kmz file (see how to video on the Capstone Introduction [65] page).

Grading Rubric

Download Grading Rubric [66]